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TRANSACTIONS

OF THE

Pee yA SOCINT Y

EDINBURGH.

VOL. XXVI.

EDINBURGH:

PUBLISHED BY ROBERT GRANT & SON, 54 PRINCES STREET.
AND WILLIAMS & NORGATE, 14 HENRIETTA STREET, COVENT GARDEN, LONDON.

MDCCCLXXII.

ROYAL SOCIETY OF EDINBURGH.

THE KEITH, BRISBANE, AND NEILL PRIZES.

The above Prizes will be awarded by the Council in the following manner :—

I. KEITH PRIZE.

The Ketru Prize, consisting of a Gold Medal and from £40 to £50 in
Money, will be awarded in the Session 1873-74, for the “best communication
on a scientific subject, communicated, in the first instance, to the Royal Society
during the Sessions 1871-72 and 1872-73.” Preference will be given to a
paper containing a discovery.

Il. MAKDOUGALL BRISBANE PRIZE.

This Prize is to be awarded biennially by the Council of the Royal Society
of Edinburgh to such person, for such purposes, for such objects, and in such
manner as shall appear to them the most conducive to the promotion of the
interests of science; with the proviso that the Council shall not be compelled
to award the Prize unless there shall be some individual engaged in scientific
pursuit, or some paper written on a scientific subject, or some discovery in
science made during the biennial period, of sufficient merit or importance in
the opinion of the Council to be entitled to the Prize.

1. The Prize, consisting of a Gold Medal and a sum of Money, will be
awarded at the commencement of the Session 1874~75, for an Essay or Paper
having reference to any branch of scientific inquiry, whether Material or
Mental.

2. Competing Essays to be addressed to the Secretary of the Society, and
transmitted not later than 1st June 1874.
VOL. XXVI. PART IV. b

3. The Competition is. open to all men of science.

4, The Essays may be either anonymous or otherwise. In the former case,
they must be distinguished by mottoes, with corresponding sealed billets super-
scribed with the same motto, and containing the name of the Author.

5. The Council impose no restriction as to the length of the Essays, which
may be, at the discretion of the Council, read at the Ordinary Meetings of the '
Society. They wish also to leave the property and free disposal of the manu-
scripts to the Authors ; a copy, however, being deposited in the Archives .
the Society, unless the Paper shall be published in the Transactions.

6. In awarding the Prize, the Council will also take into consideration any
scientific papers presented to the Society during the Sessions 1872-73 and
1873-74, whether they may have been given in with a view to the Prize or not.

Ill. NEILL PRIZE.

The Council of the Royal Society of Edinburgh having received the bequest
of the late Dr Patrick Nett of the sum of £500, for the purpose of “the
interest thereof being applied in furnishing a Medal or other reward every
second or third year to any distinguished Scottish Naturalist, according as such
Medal or reward shall be voted by the Council of the said Society,” hereby
intimate,

1. The NEILL Prizg, consisting of a Gold Medal and a sum of Money, will
be awarded during the Session 1874-75.

2. The Prize will be given for a Paper of distinguished merit, on a subject
of Natural History, by a Scottish Naturalist, which shall have been presented
to the Society during the three years preceding the 1st May 1874,—or failing
presentation of a paper sufficiently meritorious, it will be awarded for a work
or publication by some distinguished Scottish Naturalist, on some branch of
Natural History, bearing date within five years of the time of award.

AWARDS OF THE KEITH, MAKDOUGALL BRISBANE, AND NEILL PRIZES,
FROM 1827 TO. 1872.

REIT PRIZE.

1sr Brennran Psriop, 1827-29.—Dr Brewster, for his papers “on his Discovery of Two New lnmis-
cible Fluids in the Cavities of certain Minerals,” published in
the Transactions of the Society.

2p Bieyniat Periop, 1829-31—Dr Brewster, for his paper “on a New Analysis of Solar
Light,” published in the Transactions of the Society.

3p Brenniat Psriop, 1831-33.—THomas Grauam, Esq., for his paper “on the Law of the Diffusion
ef Gases,” published in the Transactions of the Society.

47H Brennian Periop, 1833-35.—Professor Forsus, for his paper “on the Refraction and Polarization
of Heat,” published in the Transactions of the Society.

5a Brenniat Perron, 1835-37.—Joun Scorr Russet, Esq., for his Researches “on Hydrodynamics,”
published in the Transactions of the Society.

6TH BrenniaL Periop, 1837-39.—Mr Joun Suaw, for his Experiments ‘‘on the Development and
Growth of the Salmon,” published in the Transactions of the
Society.

77H Brenniat Periop, 1839-41.— Not awarded.

87H Brenyiau Periop, 1841—43.—Professor Forses, for his Papers “on Glaciers,” published in the
Proceedings of the Society.

97H Brenntat Prriop, 1843—-45.—Noi awarded.

107TH Bienniat Periop, 1845—47.—General Sir THomas Brispane, Bart., for the Makerstoun Observa-
tions on Magnetic Phenomena, made at his expense, and
published in the Transactions of the Society,

11tse Brmyniau Prrtop, 1847—49.—Not awarded.

127rH Brenntau Periop, 1849-51.—Professor Ketuanp, for his papers “on General Differentiation,
including his more recent communication on a process of the
Differential Calculus, and its application to the solution of
certain Differential Equations,” published in the Transactions
of the Society.

137TH Brenniau Periop, 1851-53.—W. J. Macquorn Ranking, Esq., for his series of papers ‘on the
Mechanical Action of Heat,” published in the Transactions of
the Society.

147H Brewntat Perron, 1853-55.—Dr Tuomas Anpurson, for his papers “‘on the Crystalline Con-
stituents of Opium, and on the Products of the Destructive

Distillation of Animal Substances,” published in the Trans-
actions of the Society.

vill THE KEITH, MAKDOUGALL BRISBANE, AND NEILL PRIZES.

15TH Branniat Periop, 1855-57.—Professor Boots, for his Memoir “on the Application of the Theory
of Probabilities to Question of the Combination of Testimonies
and Judgments,” published in the Transactions of the Society.

167TH Brenniat Pertop, 1857-59.—Not awarded.

177H Brenna Perron, 1859-61.—Joun Atuan Broun, Esq., F.R.S., Director of the Trevandrum
Observatory, for his papers ‘‘on the Horizontal Force of the
Earth’s Magnetism, on the Correction of the Bifilar Magnet-
ometer, and on Terrestrial Magnetism generally,” published in
the Transactions of the Society.

18H Binwntat Psriop, 1861-63.—Professor Witt1am THomson, of the University of Glasgow, for his
Communication “on some Kinematical and Dynamical
Theorems,” published in the Transactions of the Society.

197H Brennrat Periop, 1863-65.—Principal Forprs, St Andrews, for his ‘‘ Experimental Inquiry into
the Laws of Conduction of Heat in Iron Bars,” published in
the Transactions of the Society.

20TH Bienniat Perrov, 1865-67.—Professor C. Prazzi Smyru, for his paper ‘“‘on Recent Measures at
the Great Pyramid,” published in the Transactions of the
Society.

21st BrenniaL Psriop, 1867-69.—Professor P. G. Tarr, for his paper ‘‘on the Rotation of a Rigid
Body about a Fixed Point,” published in the Transactions of
the Society.

22D Brennrat Periop, 1869-71.—Professor Cirerk Maxwewt, for his paper ‘on Figures, Frames,
and Diagrams of Forces,” published in the Transactions of the
Society.

II. MAKDOUGALL BRISBANE PRIZE.

lst Brenntau Pertop, 1859.—Sir Roperick Impry Murcuison, on account of his Contributions to the
Geology of Scotland. :

2p Brenniat Perrop, 1860-62.—Wituiam Setter, M.D., F.R.C.P.E., for his “ Memoir of the Life
and Writings of Dr Robert Whytt,” published in the Trans-
actions of the Society.

3p BrenniaL Periop, 1862-64.—Jonun Drnis Macponatp, Esq., R.N., F.R.S., Surgeon of H.M.S.
“ Tearus,” for his paper ‘‘on the Representative Relationships
of the Fixed and Free Tunicata, regarded as two Sub-classes
of equivalent value; with some General Remarks on their
Morphology,” published in the Transactions of the Society.

47H Biennial Periop, 1864-66.—Not awarded.

5TH Brenntau Periop, 1866-68.—Dr ALtexanpER Crum Brown, and Dr THomas RicHarpD FrRasmr, for
their conjoint paper ‘“‘on the Connection between Chemical
Constitution and Physiological Action,” published in the
Transactions of the Society.

6TH Brennrau Pertop, 1868—70.—Not awarded.

71H Bienntau Peston, 1870-72.—Gzorce James Attman, M.D., F.R.S., Emeritus Professor of Natural
History, for his paper ‘on the Homological Relations of
the Coelenterata,’ published in the Transactions, which forms
a leading chapter of his Monograph of Gymnoblastic or Tubu-
larian Hydroids—since published.

ibs

Ill. NEILL PRIZE.

lst Triennrat Periop, 1856-59.—Dr W. Lauper Linpsay, for his paper “on the Spermogones and
Pyenides of Filamentous, Fruticulose, and Foliaceous Lichens,”
published in the Transactions of the Society.

2n Trienniat Pertop, 1859-62.—Ropert Kaye Grevitte, LL.D., for his Contributions to Scottish
Natural History, more especially in the department of Cryp-
togamic Botany, including his recent papers on Diatomacez.

3D TRIENNIAL Prriop, 1862-65.—ANpREW CrompBir Ramsay, F.R.S., Professor of Geology in the
Government School of Mines, and Local Director of the
Geological Survey of Great Britain, for his various Works and
Memoirs published during the last five years, in which he has
applied the large experience acquired by him in the Direction

a of the arduous work of the Geological Survey of Great Britain

to the elucidation of important questions bearing on Geological
Science.

4ra Trienniau Periop, 1865-68.—Dr Wittiam Carmicuant M‘Intosu, for his paper “ on the Structure
of the British Nemerteans, and on some New British Annelids,”
published in the Transactions of the Society.

5TH TRIENNIAL Periop, 1868-71.— Professor TurnmR, for his papers ‘on the great Finner Whale ; and
on the Gravid Uterus, and the Arrangement of the Foetal
Membranes in the Cetacea,” published in the Transactions of
the Society.

is)

VOL. XXVI. PART IV.

a aa eels

ay a a as rit
wll rs ey ry es ' ee 43°

ae
\ a

LAWS

OF THE

ROYAL SOCIETY OF EDINBURGH,

AS REVISED JANUARY 1873.

LAWS.

[By the Charter of the Society (printed in the Transactions, Vol. VI. p. 5.), the Laws cannot
be altered, except at a Meeting held one month after that at which the Motion for
alteration shall have been proposed. |

if
THE ROYAL SOCIETY OF EDINBURGH shall consist of Ordinary and
Honorary Fellows.

Jid

Every Ordinary Fellow, within three months after his election, shall pay Two
Guineas as the fee of admission, and Three Guineas as his contribution for the
Session in which he has been elected ; and annually at the commencement of every
Session, Three Guineas into the hands of the Treasurer. This annual contribution
shall continue for ten years after his admission, and it shall be limited to Two
Guineas for fifteen years thereafter.*

IIT.

All Fellows who shall have paid Twenty-five years’ annual contribution shall
be exempted from farther payment.

IV.

The fees of admission of an Ordinary Non-Resident Fellow shall be £26, 5s.,
payable on his admission ; and in case of any Non-Resident Fellow coming to
reside at any time in Scotland, he shall, during each year of his residence, pay
the usual annual contribution of £3, 3s., payable by each Resident Fellow ; but
after payment of such annual contribution for eight years, he shall be exempt
from any farther payment. In the case of any Resident Fellow ceasing to reside

* At the Meeting of the Society, on the 5th January 1857, when the reduction of the Contribu-
tions from £3, 3s., to £2, 2s., from the 11th to the 25th year of membership, was adopted, it was
resolved that the existing Members shall share in this reduction, so far as regards their future annual
Contributions.

A modification of this rule, in certain cases, was agreed to 3d January 1831.

VOL. XXVI. PART IV. d

Title.

The fees of Ordi-
nary Fellows resid-
ing in Scotland.

Payment to cease
after 25 years.

Fees of Non-Resi-
dent Ordinary
Fellows.

Case of Fellows
becoming Non-Re-
sident.

Defaulters.

Privileges of
Ordinary Fellows.

Numbers Un-
limited.

Fellows entitled
to Transactions.

Mode of Recom-
mending Ordinary
Fellows.

Honorary Fellows,
British and
Foreign.

XIV

in Scotland, and wishing to continue a Fellow of the Society, it shall be in the
power of the Council to determine on what terms, in the circumstances of each
case, the privilege of remaining a Fellow of the Society shall be continued to
such Fellow while out of Scotland.

Members failing to pay their contributions for three successive years (due
application having been made to them by the Treasurer) shall be reported to
the Council, and, if they see fit, shall be declared from that period to be no
longer Fellows, and the legal means for recovering such arrears shall be
employed.

None but Ordinary Fellows shall bear any office in the Society, or vote in
the choice of Fellows or Office-Bearers, or interfere in the patrimonial interests
of the Society.

VIL.

The number of Ordinary Fellows shall be unlimited.

VEIL

The Ordinary Fellows, upon producing an order from the Treasurer, shall
be entitled to receive from the Publisher, gratis, the Parts of the Society’s
Transactions which shall be published subsequent to their admission.

1b.

Candidates for admission as Ordinary Fellows shall make an application in
writing, and shall produce along with it a certificate of recommendation to the
purport below,* signed by at least four Ordinary Fellows, two of whom shall
certify their recommendation from personal knowledge. This recommendation
shall be delivered to the Secretary, and by him laid before the Council, and
shall afterwards be printed in the circulars for three Ordinary Meetings of
the Society, previous to the day of election, and shall lie upon the table during
that time.

Le
Honorary Fellows shall not be subject to any contribution. This class shall
* “A,B, a gentleman well versed in Science (07 Polite Literatwre, as the case may be), being

“to our knowledge desirous of becoming a Fellow of the Royal Society of Edinburgh, we hereby
“ recommend him as deserving of that honour, and as likely to prove a useful and valuable Member.”

consist of persons eminently distinguished for science or literature. Its number
shall not exceed Fifty-six, of whom Twenty may be British subjects, and Thirty-
six may be subjects of foreign states.

Personages of Royal Blood may be elected Honorary Fellows, without regard
to the limitation of numbers specified in Law X.

XII.

Honorary Fellows may be proposed by the Council, or by a recommenda-
tion (in the form given below*) subscribed by three Ordinary Fellows ; and in
case the Council shall decline to bring this recommendation before the Society,
it shall be competent for the proposers to bring the same before a General
Meeting. The election shall be by ballot, after the proposal has been commu-
nicated viva voce from the Chair at one meeting, and printed in the circulars
for two ordinary meetings of the Society, previous to the day of election.

XIII.

The election of Ordinary Fellows shall only take place at the first Ordinary
Meeting of each month during the Session. The election shall be by ballot,
and shall be determined by a majority of at least two-thirds of the votes, pro-
vided Twenty-four Fellows be present and vote.

XIV.

Royal Personages. °

Recommendation
of Honorary Fel-
lows.

Mode of Election.

Election of Ordi-
nary Fellows.

The Ordinary Meetings shall be held on the first and third Mondays of Ordinary Meet-

every month from November to June inclusively. Regular Minutes shall be
kept of the proceedings, and the Secretaries shall do the duty alternately, or
according to such agreement as they may find it convenient to make.

XV. :
The Society shall from time to time publish its Transactions and Proceed-
ings. For this purpose the Council shall select and arrange the papers which

* We hereby recommend
for the distinction of being made an Honorary Fellow of this Society, declaring that each of us from
our own knowledge of his services to (Literature or Science, as the case may be) believe him to be
worthy of that honour.

(To be signed by three Ordinary. Fellows.)

To the President and Council of the Royal Society
of Edinburgh.

ings.

The Transactions.

How Published.

The Council.

Retiring Council-
lors.

Election of Office-
Eearers.

Special Meetings ;
how called.

Treasurer’s Duties.

Auditor.

XV1

they shall deem it expedient to publish in the 7vansactions of the Society, and
shall superintend the printing of the same.

xXVT.

The Transactions shall be published in parts or Fasciculi at the close of
each Session, and the expense shall be defrayed by the Society.

XVII.

There shall be elected annually, for conducting the publications and regu- .
lating the private business of the Society, a Council, consisting of a President ;
Six Vice-Presidents, two at least of whom shall be resident ; Twelve Council-
lors, a General Secretary, Two Secretaries to the Ordinary Meetings, a Trea-
surer, and a Curator of the Museum and Library.

Xx VIEL.

Four Councillors shall go out annually, to be taken according to the order
in which they stand on the list of the Council.

XIX.

An Extraordinary Meeting for the Election of Office-Bearers shall be held
on the fourth Monday of November annually.

»:©.€

Special Meetings of the.Society may be called by the Secretary, by direction
of the Council; or on a requisition signed by six or more Ordinary Fellows.
Notice of not less than two days must be given of such Meetings.

XXI.

The Treasurer shall receive and disburse the money belonging to the Society,
granting the necessary receipts, and collecting the money when due.

He shall keep regular accounts of all the cash received and expended, which
shall be made up and balanced annually ; and at the Extraordinary Meeting in
November, he shall present the accounts for the preceding year, duly audited.
At this Meeting, the Treasurer shall also lay before the Council a list of all
arrears due above two years, and the Council shall thereupon give such direc-
tions as they may deem necessary for recovery thereof.

XXII.

At the Extraordinary Meeting in November, a professional accountant shall
be chosen to audit the Treasurer’s accounts for that year, and to give the neces-
sary discharge of his intromissions.

XVil

XXIII.

The General Secretary shall keep Minutes of the Extraordinary Meetings of General Secretary's
the Society, and of the Meetings of the Council, in two distinct books. He ee
shall, under the direction of the Council, conduct the correspondence of the
Society, and superintend its publications. For these purposes he shall, when
necessary, employ a clerk, to be paid by the Society.

XXIV.

The Secretaries to the Ordinary Meetings shall keep a regular Minute-book, Secretaries to _
in which a full account of the procedings of these Meetings shall be entered ; pane cg
they shall specify all the Donations received, and furnish a list of them, and of
the Donors’ names, to the Curator of the Library and Museum ; they shall like-
wise furnish the Treasurer with notes of all admissions of Ordinary Fellows.

They shall assist the General Secretary in superintending the publications, and
in his absence shall take his duty.

XXV.

The Curator of the Museum and Library shall have the custody and charge Curator of Museum
of all the Books, Manuscripts, objects of Natural History, Scientific Produc- eae
tions, and other articles of a similar description belonging to the Society ; he
shall take an account of these when received, and keep a regular catalogue of
the whole, which shall lie in the Hall, for the inspection of the Fellows.

XXVI.

All Articles of the above description shall be open to the inspection of the Use of Museum
Fellows at the Hall of the Society, at such times and under such regulations, Sm
as the Council from time to time shall appoint.

XXVIT.

A Register shall be kept, in which the names of the Fellows shall be Register Book.
enrolled at their admission, with the date.

VOL. XXVI. PART IV. €

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DIRECTIONS TO THE BINDER FOR PLACING THE PLATES IN THIS VOLUME.

Evite Tlustrating Mr J. Clerk Maxwell’s ee on poe Figures, Frames, and
ante Diagrams of Forces, . 5 : To face page
Illustrating the Rev. Thomas Brown’s Paper on the Old River Terraces of the
IV. Earn and Teith, viewed in connection with certain Proofs of the Antiquity
of Man,
wy
VI. ( Illustrating Professor Turner's Paper, Account of the Great Finner Whale
AGU (Balenoptera Sibbaldii) stranded at Longniddry. Part I.—The Soft Parts,
VIL
IX. ) Illustrating Dr W. C. M‘Intosh’s Paper on some Points in the Structure of
x) Tubifex, : : 3 , : ‘ ; : 5
XI.
XII.
XIIL Tlustrating Dr James Bell Pettigrew’s Paper on the Physiology of Wings,
XIV being an Analysis of the Movements m which Flight is produced in the
; Tnsect, Bat, and Bird, ; , , ’ : ;
XV.
XVI.
XVII. ) Illustrating Professor Turner’s Paper on the Gravid Uterus, and on the Arrange-
XVIIL. ment of the Feetal Membranes in the Cetacea, ; 5 : ;
XIX.
XX. ( Illustrating Professor Alexander Dickson’s Paper on Some Abnormal Cones of
XXI. Pinus Pinaster,
XXII.
XXIII.

XXIV ) Illustrating Dr Thomas R. Fraser’s Paper on an Experimental Research on the
XXV. f Antagonism between the Actions of Physostigma and Atropia, 3

XXVI Illustrating Mr J. A. Broun’s Paper on the Lunar Diurnal Variation of Magnetic

XXVIL Declination at Trevandrum, near the Magnetic Equator, deduced from
: Observations made in the Observatory of His Highness the ies of

XXVIII.
Travancore, G.C.S.L., 4 : ee : :

XXIX Illustrating Professor Turner’s Paper on the Occurrence of Ziphius cavirostris in
XXX. the Shetland Seas, and a Comparison of its Skull with that of Sowerby’s
j Whale (Mesoplodon Sowerby), ‘ : : 5 :

XXXI. } Illustrating Professor Balfour's Paper, Remarks on the Ipecacuan Plant (Cephaélis
XXXII. Ipecacuanha, Rich.), as cultivated in the Royal Botanic Garden, Edinburgh,

149

253

321

467

505

529

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CONTENTS.

PART I. (1869-70.)

PAGE
I.—On Reciprocal Figures, Frames, and Diagrams of Forces.
By J. Crerk Maxwett, F.R.SS. L. & E. (Plates LI,
tf and brie: : 1
IL—On Scientific Method in the Interpretation of Popular Myths,
with special reference to Greck ee By Professor
BLACKIE, : : ; ; 4]
Il. Onthe Extension of Brouncker’s Method to the Comparison of
Several Magnitudes. By Epwarp Sane, Esq., . E 59

IV.—On Green’s and other Allied Theorems. By Professor Tarr, . 69

V.—On the Heat Developed in the Combination of Acids and Bases.
Second Memoir. By THomas Anprews, M.D., F.R.S.,
Hon. F-.R.S.E., Vice-President of Queen’s College,
Belfast, . ; ; : : , ; 85

VI—The Genetic Succession of Zooids in the Hydroida. By
Professor ALLMAN, ; : ; i : 97

VII.—Injfluence of the Vagus upon the Vascular System. By
Wittiam Rutuerrorp, M.D., F.R.S.E., Professor of
Physiology, King’s College, London, . ; OZ,

VIII.—On the Old River Terraces of the Earn and Teith, viewed in
connection with certain Proofs of the Antiquity of Man.
By the Rev. THomas Brown, F.R.S.E. (Plate IV.), . 149
VOL, XXVI. PART IV. yf

CONTENTS.

TX.—On Spectra formed by the Passage of Polarized Light through
Refracting Crystals. By Francis Deas, M.A., LL.B.,
E.R.S.E., c : hy stp ;

Addition to the above Paper. By J. CLERK MaxweE t, LL.D.,
E.R.SS., L. & E.,

X. On the Oxidation of Products of Picoline. By James Dewar,
F.R.S.E., Chemical Demonstrator in the University of
Edinburgh, and Lecturer on Chemistry at the Edinburgh
Veterinary College, : : é

XI. An Account of the Great Finner Whale (Balenoptera Sib-
baldii) stranded at Longniddry. Part I.—The Soft Parts.
By Witiram Turner, M.B., (Lond.), Professor of Anat-
omy in the University of Edinburgh. (Plates V.—VIII.),

PART IL. (1870-712)

XII. On Some Points in the Structure of Tubifex. By W. C.
M‘Intosy, M.D., F.B.S.E. (Plates IX. and X.),

XIII—On the Place and Power of Accent in Language. By Pro-

_ fessor BLACKIE,

XIV.—On the Average Quantity of Rain in Carlisle and the Neigh-

bourhood. By Tuomas Barngs, M.D., F.R.S.E.,

XV.—On the Physiology of Wings, being an Analysis of the Move-
ments by which Flight is produced in the Insect, Bat, and
Bird. By James Bett Petticrew, M.D., F.R.S., Path-
ologist to the Royal Infirmary of Edinburgh, and Curator
of the Museum of the Royal College of Surgeons of
Edinburgh. Communicated by Professor TURNER.
(Plates XI. to XVI),

PAGE

177

185

189

197

253

269

313

321

CONTENTS.

XVI—Additional Note on the Motion of a Heavy Body along the
Circumference of a Circle. By Epwarp Sane, Esq.,
E.RS.E., :

XVII.—On the Homological Relations of the Coelenterata. By Pro-
fessor ALLMAN, ; : ; : ,

XVIII. On the Gravid Uterus and on the Arrangement of the Foetal
Membranes in the Cetacea. By Professor TuRNER.
(Plates X VIL and XVIII),

XIX.—On some Abnormal Cones of Pinus Pinaster. By ALEXANDER
Dickson, M.D., Regius Professor of Botany in the
University of Glasgow. (Plates XIX. to XXII),

PART III. (1870-71.)

XX.— Account of the New Table of Logarithms to 200000. By
Epwarp Sane, Esq., F.R.S.E., : :

XXI—An Experimental Research on the Antagonism between the
Actions of Physostigma and Atropia. By Tuomas R.
Fraser, M.D., Lecturer on Materia Medica and
Therapeutics at Surgeon’s Hall, Edinburgh. (Plates
XXIII-XXvV.), : ;

PART IV. (1871- 2)

XXII.—On the Decomposition of Forces externally applied to an Elastic
Solid. By W. J. Macquorn Rankine, C.E., LL.D.,
FRSS. L&E, . : ; : :

XXIII—On the Geometrical Mean Distance of Two Figures on a Plane.
By Professor J. CLERK MAXweELt, F.R.S.,

XXxili

PAGE

449

459

467

505

715

729

XXIV CONTENTS.

XXIV.—On the Lunar Diurnal Variation of Magnetic Declination at
Trevandrum, near the Magnetic Equator, deduced Jrom
Observations made in the Observatory of His Highness the
Maharajah of Travancore, G.C.SL. By J. A. Broun,
FERS. (Plates XXVI-XXVIIL),
>
XXV.—-On the Occurrence of Ziphius cavirostris in the Shetland Seas,
and a Comparison of its Skull with that of Sowerby’s
Whale (Mesoploden Sowerbyi). By Professor TURNER,
(Plates X XIX., XXX.), ak

XXVI—Remarks on the Lpecacuan Plant (Cephaélis Ipecacuanha,
hich.), as cultivated in the Royal Botanic Garden, Edin-
burgh. By Joun Hurron Batrour, M.D., E.R.S., Sec.
R.S.E., F.L.S., Hon. Mem. Pharm. Soc., and Professor
of Medicine and Botany in the University of Edinburgh...
(Plates XX XI. and XXXII), |

PAGE

735

759

781

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Trans. Roy Soc.Edin? Vol. XX VI

Plate Il

Trans. Roy. Soc. Edin’ Vol. XXVI

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TRANSACTIONS.

I—On Reciprocal Figures, Frames, and Diagrams of Forces. By J. CLERK
MaxwELL, F.R.SS. L. & E. (Plates I. I. IIT.)

(Received 17th Dec. 1869 ; read 7th Feb. 1870.)

Two figures are reciprocal when the properties of the first relative to the
second are the same as those of the second relative to the first. Several kinds
of reciprocity are known to mathematicians, and the theories of Inverse Figures
and of Polar Reciprocals have been developed at great length, and have led to
remarkable results. I propose to investigate a different kind of geometrical
reciprocity, which is also capable of considerable development, and can be
applied to the solution of mechanical problems.

A Frame may be defined geometrically as a system of straight lines connect-
ing a number of points. In actual structures these lines are material pieces,
beams, rods, or wires, and may be straight or curved ; but the force by which
each piece resists any alteration of the distance between the points which it joins
acts in the straight line joining those points. Hence, in studying the equilibrium
of a frame, we may consider its different points as mutually acting on each
other with forces whose directions are those of the lines joining each pair of points.
When the forces acting between the two points tend to draw them together, or
to prevent them from separating, the action along the joining line is called a
Tension. When the forces tend to separate the points, or to keep them apart,
the action along the joining line is called a Pressure.

If we divide the piece joining the points by any imaginary section, the
resultant of the whole internal force acting between the parts thus divided will
be mechanically equivalent to the tension or pressure of the piece. Hence, in
order to exhibit the mechanical action of the frame in the most elementary
manner, we may draw it as a skeleton, in which the different points are joined
by straight lines, and we may indicate by numbers attached to these lines the
tensions or pressures in the corresponding pieces of the frame.

The diagram thus formed indicates the state of the frame in a way which is

VOL. XXVI. PART I. A

2, MR CLERK MAXWELL ON

geometrical as regards the position and direction of the forces, but arithmetical
as regards their magnitude.

But, by assuming that a line of a certain length shall represent a force of a
certain magnitude, we may represent every force completely by a line. This
is done in Elementary Statics, where we are told to draw a line from the point
of application of the force in the direction in which the force acts, and to cut off
as many units of length from the line as there are units of force in the force, and
finally to mark the end of the line with an arrow-head, to show that it is a force and
not a piece of the frame, and that it acts in that direction and not the opposite.

By proceeding in this way, we should get a system of arrow-headed forces
superposed on the skeleton of the frame, two equal and opposite arrows for
every piece of the frame.

To test the equilibrium of these forces at any point of concourse, we should
proceed by the construction of the parallelogram of forces, beginning with two
of the forces acting at the point, completing the parallelogram, and drawing the
diagonal, and combining this with the third force in the same way, till, when all
the forces had been combined, the resultant disappeared. We should thus have
to draw three new lines, one of which is an arrow, in taking in each force after
the first, leaving at last not only a great number of useless lines, but a number
of new arrows, not belonging to the system of forces, and only confusing to
any one wishing to verify the process.

To simplify this process, we are told to construct the Polygon of Forces, by
drawing in succession lines parallel and proportional to the different forces, each
line beginning at the extremity of the last. If the forces acting at the point
are in equilibrium, the polygon formed in this way will be a closed one.

Here we have for the first time a true Diagram of Forces, in which every
force is not only represented in magnitude and direction by a straight line, but
the equilibrium of the forces is manifest by inspection, for we have only to
examine whether the polygon is closed or not. To secure this advantage, how-
ever, we have given up the attempt to indicate the position of the force, for the
sides of the polygon do not pass through one point as the forces do. We must,
therefore, give up the plan of representing the frame and its forces in one
diagram, and draw one diagram of the frame and a separate diagram of the
forces. By this method we shall not only avoid confusion, but we shall greatly
simplify mechanical calculations, by reducing them to operations with the
parallel ruler, in which no useless lines are drawn, but every line represents an
actual force.

A Diagram of Forces is a figure, every line of which represents in magnitude
and direction the force acting along a piece of the frame.

To express the relation between the diagram of the frame and the diagram
of forces, the lines of the frame should each be indicated by a symbol, and the

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 3

corresponding lines of the diagram of forces should be indicated by the same
symbol, accented if necessary.

We have supposed the corresponding lines to be parallel, and it is necessary
that they should be parallel when the frame is not in one plane ; but if all the
pieces of the frame are parallel to one plane, we may turn one of the diagrams
round a right angle, and then every line will be perpendicular to the corres-
ponding line.

If any number of lines meet at the same point in the frame, the ee
ing lines in the diagram of forces form a closed polygon.

It is possible, in certain cases, to draw the diagram of forces so that if any
number of lines meet in a point in the diagram of forces, the corresponding lines
in the frame form a closed polygon.

In such cases, the two diagrams are said to be reciprocal in the sense in
which we use it in this paper. If either diagram be taken as representing
the frame, the lines of the other diagram will represent a system of forces
which, if applied along the corresponding pieces of the frame, will keep it in
equilibrium.

The properties of the “triangle” and “polygon” of forces have been long
known, and a “diagram” of forces has been used in the case of the “funicular
polygon,” but I am not aware of any more general statement of the method of
drawing diagrams of forces before Professor RANKINE applied it to frames, roofs,
&c., in his ‘‘ Applied Mechanics,” p. 137, &c. The “polyhedron of forces,” or
the proposition that forces acting on a point perpendicular and proportional to
the areas of the faces of a polyhedron are in equilibrium, has, I believe, been
enunciated independently at various times, but the application of this principle
to the construction of a diagram of forces in three dimensions was first made
by Professor RANKINE in the “ Philosophical Magazine,” Feb. 1864. In the
“Philosophical Magazine” for April 1864, I stated some of the properties of
reciprocal figures, and the conditions of their existence, and showed that any
plane rectilinear figure which is a perspective representation of a closed poly-
hedron with plane faces has a reciprocal figure. In Sept. 1867, I communi-
cated to the British Association a method of drawing the reciprocal figure,
founded on the theory of reciprocal polars.

I have since found that the construction of diagrams of forces in which each
force is represented by one line, had been independently discovered by Mr W.
P. Taytor, and had been used by him as a practical method of determining
the forces acting in frames for several years before I had taught it in King’s
College, or even studied it myself. I understand that he is preparing a state-
ment of the application of the method to various kinds of structures in detail,
so that it can be made use of by any one who is able to draw one line parallel
to another.

7 MR CLERK MAXWELL ON

Professor FLEEMING JENKIN, in a paper recently published by the Society,
has fully explained the application of the method to the most important cases
occurring in practice.

In the present, paper I propose, first, to consider plane diagrams of frames
and of forces in an elementary way, as a practical method of solving questions
about the stresses in actual frameworks, without the use of long calculations.

I shall then discuss the subject in a theoretical point of view, and give a
method of defining reciprocal diagrams analytically, which is applicable to
figures either of two or of three dimensions.

Lastly, I shall extend the method to the investigation of the state of stress
in a continuous body, and shall point out the nature of the function of stress
first discovered by the Astronomer Royal for stresses in two dimensions, extend-
ing the use of such functions to stresses in three dimensions.

On Reciprocal Plane Kectilinear Figures.

Definition —Two plane rectilinear figures are reciprocal when they consist
of an equal number of straight lines, so that corresponding lines in the two
figures are at right angles, and corresponding lines which meet in a point in
the one figure form a closed polygon in the other.

Note.—It is often convenient to turn one of the figures round in its own
plane 90°. Corresponding lines are then parallel to each other, and this is
sometimes more convenient in comparing the diagrams by the eye.

Since every polygon in the one figure has three or more sides, every point in
the other figure must have three or more lines meeting in it. Since every line
in the one figure has two, and only two, extremities, every line in the other figure
must be a side of two, and only two, polygons. If either of these figures be taken
to represent the pieces of a frame, the other will represent a system of forces
such that, these forces being applied as tensions or pressures along the correspond-
ing pieces of the frame, every point of the frame will be in equilibrium.

The simplest example is that of a triangular frame without weight, ABC,
jointed at the angles, and acted on by three forces, P, Q, R, applied at the
angles. The directions of these three forces must meet in a point, if the frame
is in equilibrium. We shall denote the lines of the figure by capital letters,
and those of the reciprocal figure by the corresponding small letters ; we shall
denote points by the lines which meet in them, and polygons by the lines which
bound them.

Here, then, are three lines, A, B, C, forming a triangle, and three other
lines, P, Q, R, drawn from the angles and meeting in a point. Of these forces
let that along P be given. Draw the first line p of the reciprocal diagram
parallel to P, and of a length representing, on any convenient scale, the force
along P. The forces along P, Q, R are in equilibrium, therefore, if from one

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 5

extremity of p we draw gq parallel to Q, and from the other extremity r parallel
to R, so as to form a triangle pg7, then g and 7 will represent on the same scale
the forces along Q and R.

pees

To determine whether these forces are tensions or pressures, make a point
travel along p in the direction in which the force in P acts on the point of con-
course of PQR, and let the point travel in the same direction round the
polygon pgr. Then, the direction in which the point travels along any side
of the polygon will be the direction in which the force acts along the corres-
ponding piece of the frame on the point of concourse. If it acts from the
point of concourse, the force is a tension; if towards it, it is a pressure.

The other extremity of P meets B and C, and the forces along these three
pieces are in equilibrium. Hence, if we draw a triangle, having p for one side
and lines parallel to B and C for the others, the sides of this triangle will
represent the three forces.

Such a triangle may be described on either side of p, the two together would
form a parallelogram of forces; but the theory of reciprocal figures indicates
that only one of these triangles forms part of the diagram of forces.

The rule for such cases is as follows :—Of the two extremities of p, one cor-
responds to the closed figure PRB, and the other to the closed figure PQC,
these being the polygons of which P is a side in the first figure.

We must, therefore, draw 6 parallel to B from the intersection of p and 7,
and not from the other extremity, and we must draw ¢ parallel to C from the
intersection of p and q.

We have now a second triangle, pbc, corresponding to the forces acting
at the point of concourse of P, B, C. To determine whether these forces are
tensions or pressures, we must make a point travel round pdc, so that its
course along p is in the opposite direction to its course round pqr, because the
piece P acts on the points PBC and PQR with equal and opposite forces.

If we now consider the equilibrium of the point of concourse of QC and A,
we shall find that we have determined two of these forces by the lines g and ¢,
and that the third force must be represented by the line a which completes the
triangle gca.

We have now constructed a complete diagram of forces, in which each force

VOL. XXVI. PART I. B

6 MR CLERK MAXWELL ON

is represented by a single line, and in which the equilibrium of the forces meet-
ing at any point is expressed visibly by the corresponding lines in the other
figure forming a closed polygon. P

There are in this figure six lines, having four points of concourse, and form-
ing four triangles. To determine the direction of the force along a given line
at any point of concourse, we must make a point travel round the corresponding
polygon in the other figure in a direction which is positive with respect to that
polygon. For this purpose it is desirable to name the polygons in a determi-
nate order of their sides, so arranged that, when we arrive at the same side in
naming the two polygons which it divides, we travel along it in opposite direc-
tions. For instance, if pgr be one of the polygons, the others are pbc, qca, rab.

Note.—It may be observed, that after drawing the lines p, g, 7, 6, ¢ with the
parallel ruler, the line a@ was drawn by joing the points of concourse of q, 7
and b, ¢; but, since it represents the force in A, a is parallel to A. Hence the
following geometrical theorem :—

If the lines PQR, drawn from the angles of the triangle ABC, meet in a point,
then if pgr be a triangle with its corresponding sides parallel to P, Q, R, and if
a, b, ¢ be drawn from its corresponding angles parallel to A, B, C, the lines
a, b, e will meet in a point.

A geometrical proof of this is easily obtained by finding the centres of the
four circles circumscribing the triangles ABC, AQR, BRP, CPQ, and joining
the four centres thus found by six lines.

These lines meet in the four centres, and are perpendicular to the six lines,
A, B, C; P, Q, R; but by turning them round 90° they become parallel to the
corresponding lines in the original figure.

The diagram formed in this way is definite in size and position, but any
figure similar to it is a reciprocal diagram to the original figure. I have
explained the construction of this, the simplest diagram of forces, more at
length, as I wish to show how, after the first line is drawn and its extremities
fixed on, every other line is drawn in a perfectly definite position by means of
the parallel ruler.

In any complete diagram of forces, those forces which act at a given point
in the frame form a closed polygon. Hence, there will be as many closed
polygons in the diagram as there are points in the frame. Also, since each
piece of the frame acts with equal and opposite forces on the two points which
form its extremities, the force in the diagram will be a side of two different
polygons. These polygons might be drawn in any positions relatively to each
other ; but, in the diagrams here considered, they are placed so that each force
is represented by one line, which forms the boundary between the two polygons
to which it belongs.

If we regard the polygons as surfaces, rather than as mere outlines, every

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 7

polygon will be bounded at every point of its outline by other polygons, so
that the whole assemblage of polygons will form a continuous surface, which
must either be an infinite surface or a closed surface.

The diagram cannot be infinite, because it is made up of a finite number of
finite lines representing finite forces. It must, therefore, be a closed surface
returning on itself, in such a way that every point in the plane of the diagram
either does not belong to the diagram at all, or belongs to an even number of
sheets of the diagram.

Any system of polygons, which are in contact with each other externally,
may be regarded as a sheet of the diagram. When two polygons are on the
same side of the line, which is common to them, that line forms part of the
common boundary of two sheets of the diagram. If we reckon those areas
positive, the boundary of which is traced in the direction of positive rotation
round the area, then all the polygons in each sheet will be of the same sign as
the sheet, but those sheets which have a common boundary will be of opposite
sign. At every point in the diagram there will be the same number of positive
as of negative sheets, and the whole area of the positive sheets will be auks to
that of the negative sheets.

The diagram, therefore, may be considered as a plane projection of a closed
polyhedron, the faces of the polyhedron being surfaces bounded by rectilinear
polygons, which may or may not, as far as we yet know, lie each in one plane.

Let us next consider the plane projection of a given closed polyhedron.
If any of the faces of this polyhedron are not plane, we may, by drawing
additional lines, substitute for that face a system of triangles, each of which is
necessarily in a plane. We may, therefore, consider the polyhedron as bounded
by plane faces. Every angular point of this polyhedron will be defined by its
projection on the plane and its height above it.

Let us now take a fixed point, which we shall call the origin, and draw from
it a perpendicular to the plane. We shall call this line the axis. If we then
draw from the origin a line perpendicular to one of the faces of the polyhedron,
it will cut the plane at a point which may be said to correspond to the projec-
tion of that face. From this point draw a line perpendicular to the plane, and -
take on this line a point whose distance from the plane is equal to that of the
intersection of the axis with the face of the polyhedron produced, but on the
_ other side of the plane. This point in space will correspond to the face of the
polyhedron. By repeating this process for every face of the polyhedron, we
shall find for every face a corresponding point with its projection on the plane.

To every edge of the polyhedron will correspond the line which joins the
points corresponding to the two faces which meet in that edge. Each of these
lines is perpendicular to the projection of the other; for the perpendiculars
from the origin to the two faces, lie in a plane perpendicular to the edge in

8 MR CLERK MAXWELL ON

which they meet, and the projection of the line corresponding to the edge is the
intersection of this plane with the plane of projection. Hence, the edge is
perpendicular to the projection of the corresponding line. The projection of
the edge is therefore perpendicular to the projection of the corresponding line,
and therefore to the corresponding line itself. In this way we may draw a
diagram on the plane of projection, every line of which is perpendicular to the
corresponding line in the original figure, and so that lines which meet in a point
in the one figure form a closed polygon in the other.

If, in a system of rectangular co-ordinates, we make z=0 the plane of pro-
jection, and x=0 y=0 z= —e the fixed point, then if the equation of a plane be

z= Az+By+C,
the co-ordinates of the corresponding point will be
hr ” = cB tek | ie
and we may write the equation
cz+0) = 2& + yn .

If we suppose &, n, ¢ given as the co-ordinates of a point, then this equation,
considering 2, y, z as variable, is the equation of a plane corresponding to the
point.

If we suppose 2, y, z the co-ordinates of a point, and &€, , ¢ as variable, the
equation will be that of a plane corresponding to that point.

Hence, if a plane passes through the point xyz, the point corresponding to
this plane lies in the plane corresponding to the point ayz.

These points and planes are reciprocally polar in the ordinary sense with
respect to the paraboloid of revolution

2ce2 = a? + 7?.

We have thus arrived at a construction for reciprocal diagrams by consider-
ing each as a plane projection of a plane-sided polyhedron, these polyhedra
being reciprocal to one another, in the geometrical sense, with respect to a cer-
tain paraboloid of revolution.

Each of the diagrams must fulfil the conditions of being a plane projection
of a plane-sided polyhedron, for if any of the sides of the polyhedron of which
it is the projection are not plane, there will be as many points corresponding to
that side as there are different planes passing through three points of the side,
and the other diagram will be indefinite.

Relation between the Number of Edges, Summits, and Faces of Polyhedra.

It is manifest that after a closed surface has been divided into separate faces
by lines drawn upon it, every new line drawn from a point in the system, either
introduces one new point into the system, or divides a face into two parts,

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 9

according as it is drawn to an isolated point, or to a point already connected
with the system. Hence the sum of points and faces is increased by one for
every new line. If the closed surface is acyclic, or simply connected,* like that
of a solid body without any passage through it, then, if from any point we draw
a closed curve on the surface, we divide the surface into two faces. We have
here one line, one point, and two faces. Hence, if e be the number of lines,
s the number of points, and / the number of faces, then in general

e—s—f=m

when m remains constant, however many lines be drawn. But in the case of a
simple closed surface

m=—2.

If the closed surface is doubly connected, like that of a solid body with a
hole through it, then if we draw one closed curve round the hole, and another
closed curve through the hole, and round one side of the body, we shall have
e=2,s=1,f=1,s0 that ~=0, If the surface is n-ly connected, like that
of a solid with »—1 holes through it, then we may draw n closed curves
round the 7 — 1 holes and the outside of the body, and m — 1 other closed curves
each through a hole and round the outside of the body.

We shall then have 4(2—1) segments of curves terminating in 2(” — 1)
points and dividing the surface into two faces, so that e= 4(n—1),
2 (7 — 1), and f= 2, and

e—s—f=2n—-4,

and this is the general relation between the edges, summits, and faces of a
polyhedron whose surface is -ly connected.
The plane reciprocal diagrams, considered as plane projections of such

* See Riemann, Crelle’s Journal, 1857, Lehrsdtze aus der analysis situs, for space of two dimen-
sions; also CayLEy on the Partitions of a Close, Phil. Mag. 1861; Hrtmuotrz, Crelle’s Journal, 1858,
Wirbelbewegung, for the application of the idea of multiple continuity to space of three dimensions ; J.
B. Listine, Gottingen Trans., 1861, Der Census Rdéumlicher Complexe, a complete treatise on the
subject of Cyclosis and Periphraxy.

On the importance of this subject see Gauss, Werke, v. 605, “ Von der Geometria Situs die LErBNITZ
ahnte und in die nur einem Paar Geometern (EuLpR und VaNDERMONDE) einen schwachen Blick zu thun
vergonnt war, wissen und haben wir nach anderthalbhundert Jahren noch nicht viel mehr wie nichts.”

Note added March 14, 1870.—Since this was written, I have seen Listrne’s Census. In his
notation, the surface of an n-ly connected body (a body with 7 — 1 holes through it) is (2n — 2)
cyclic. If 2n — 2 = K, expresses the degree of cyclosis, then Lisrine’s general equation is—

s—(e—K,)+(f/—K,+ =) -w—K,+ 2,-—wv) =0,
where s is the number of points, e the number of lines, K, the number of endless curves, / the number
of faces, K, the number of degrees of eyclosis of the faces, 2, the number of periphractic or closed
faces, v the number of regions of space, K, their number of degrees of cyclosis, 2, their number of
degrees of periphraxy or the number of regions which they completely surround, and w is to be put
= | or = 0, according as the system does or does not extend to infinity.
VOL. XXVI. PART I. C

10 MR CLERK MAXWELL ON

polyhedra, have the same relation between the numbers of their lines, points,
and polygons. It is manifest that since

ee =f,» and f, = 2,
where the suffixes refer to the first and second diagrams respectively
Nl, = Ny»

or the two diagrams are connected to the same degree.

On the Degrees of Freedom and Constraint of Frames.

To determine the positions of s points in space, with reference to a given
origin and given axes, 3s data are required; but since the position of the origin
and axes involve 6 data, the number of data required to determine the relative
position of s points is 3s — 6.

If, therefore, the lengths of 3s — 6 lines joing selectéd pairs of a system of
s points be given, and if'these lengths are all independent of each other, then
the distances between any other pair of points will be determinate, and the
system will be rigidly connected.

If, however, the lines are so chosen that those which join pairs of points of
a system of s’ of the points are more than 3s’ — 6 in number, the lengths of
these lines will not be independent of each other, and the lines of this partial
system will only give 3s’ — 6 independent data to determine the complete system.

In a system of s points joined by ¢ lines, there will in general be 3s — 6 — ¢
= p degrees of freedom, provided that in every partial system of s’ points joined
by é’ lines, and having in itself p’ degrees of freedom, p’ is not negative. If in
any such system p is negative, we may put g = — p, and call g the number of
degrees of constraint, and there will be g equations connecting the lengths of
the lines ; and if the system is a material one, the stress along each piece will
be a function of g dependent variables. Such a system may be said to have
q degrees of constraint. If p’ is negative in any partial system, then the
degrees of freedom of the complete system are p — p’, where p and p’ are got
from the number of points and lines in the complete and partial systems. If s
points are connected by ¢ lines, so as to form a polyhedron of / faces, enclosing
a space 2 times connected, and if each of the faces has m sides, then

My == 2a

We have also
e—s-f=dm-—4,
and
3s — 6 — 1) a Ole
whence

p=6—n) +(2-< e

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. un

Tf all the faces of the polyhedron are triangles, m = 3, and we have
p=6(1—n).

If m = 1, or in the case of a simply connected polyhedron with triangular
faces, p=o, that is to say, such a figure is a rigid system, which would be no
longer rigid if any one of its lines were wanting. In such a figure, if made of
material rods forming a closed web of triangles, the tensions and pressures in the
rods would be completely determined by the external forces applied to the figure,
and if there were no external force, there would be no stress in the rods.

In a closed surface of any kind, if we cover the surface* with a system of
curves which do not intersect each other, and if we draw another system inter-
secting these, and a third system passing diagonally through the intersections of
the other two, the whole surface will be covered with small curvilinear triangles,
and if we now substitute for the surface a system of rectilinear triangles having
the same angular points, we shall have a polyhedron with triangular faces
differing infinitely little from the surface, and such that the length of any line
on the surface differs infinitely little from that of the corresponding line on the
polyhedron. We may, therefore, in all questions about the transformation of
surfaces by bending, substitute for them such polyhedra with triangular faces. -

We thus find with respect to a simply connected closed inextensible surface
—1st, That it is of invariable form ;+ 2d, That the stresses in the surface depend
entirely on the external applied forces ;{ 3d, That if there is no external force,
there is no stress in the surface.

In the limiting case of the curved surface, however, a kind of deformation is
possible, which is not possible in the case of the polyhedron. Let us suppose
that in some way a dimple has been formed on a convexo-convex part of the
surface, so that the edge of the dimple is a plane closed curve, and the dimpled
part is the reflexion in this plane of the original form of the surface. Then the
length of any line drawn on the surface will remain unchanged.

Now let the dimple be gradually enlarged, so that its edge continually
changes its position. Every line on the surface will still remain of the same
length during the whole process, so that the process is possible in the case of
an inextensible surface. In this way such a surface may be gradually turned
outside in, and since the dimple may be formed from a mere point, a pressure
applied at a single point on the outside of an inextensible surface will not be
resisted, but will form a dimple which will increase till one part of the surface
comes in contact with another.

In the case of closed surfaces doubly connected, p = — 6, that is, such sur-

* On the Bending of Surfaces, by J. Crurk Maxwett, Cambridge Transactions, 1856.
t This has been shown by Professor Jutturr, Trans. R.I.A., vol. xxii. p. 377.

¢ On the Equilibrium of a Spherical Envelope, by J. C. Maxwert. Quarterly Journal of
Mathematics, 1867.

12 MR CLERK MAXWELL ON

faces are not only rigid, but are capable of internal stress, independent of
external forces, and the expression of this stress depends on six independent
variables.

In a polyhedron with triangular faces, if a number of the edges be taken
away so as to form a hole with e, sides, the number of degrees of freedom is

p=¢,—6n+3.

Hence, in order to make an n-ly connected polyhedron simply rigid without
stress, we may cut out the edges till we have formed a hole having 6 — 8 edges.
The system will then be free from stress, but if any more edges be removed, the
system will no longer be rigid.

Since in the limiting case of the inextensible surface, the smallest hole may
be regarded as having an infinite number of sides, the smallest hole made in a
closed inextensible surface connected to any degree will destroy its rigidity.
Its flexibility, however, may be confined within very narrow limits.

In the case of a plane frame of s points, we have 2s data required to deter-
mine the points with reference to a given origin and axes; but since 3 arbitrary
data are involved in the choice of origin and axis, the number of data required
to determine the relative position of s points in a plane is 2s — 3.

If we know the lengths of ¢ lines joining certain pairs of these points, then
in general the number of degrees of freedom of the frame will be

p= 2s—e-—3.

If, however, in any partial system of s’ points connected by e’ lines, the quantity
p =2s' — e —3 be negative, or in other words, if a part of the frame be self-
strained, this partial system will contribute only 2s’— 3 equations independent
of each other to the complete system, and the whole frame will have p— p’
degrees of freedom.

In a plane frame, consisting of a single sheet, every element of which is
triangular, and in which the pieces form three systems of continuous lines, as at
p. 11, if the frame contains ¢ pieces connecting s points, s’ of which are on the
circumference of the frame and s, in the interior, then

w— 5 = 6 toe
Hence

Cee oe) ee
anegative quantity, or such a frame is necessarily stiff; and if any of the points
are in the interior of the frame, the frame has as many degrees of constraint as
there are interior points—that is, the stresses im each piece will be functions of

s, variables, and s, pieces may be removed from the frame without rendering it
loose.

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 13

_If there are 7 holes in the frame, so that s’ points lie on the circumference of
the frame or on those of the holes, and s, points lie in the interior, the degree
of stiffness will be

—p=s, + 3n.
If a plane frame be a projection of a polyhedron of / faces, each of m sides, and
enclosing a space 7 times connected, then

mf = 2e
e—s—f=2a—-—4
28 —e¢=p+3,

whence
4
pes dns (1-S)e.

If all the faces are quadrilaterals m= 4 and p=5 — 4n, or a plane frame which
is the projection of a closed polyhedron with quadrilateral faces, has one degree
of freedom if the polyhedron is simply connected, as in the case of the projec-
tion of the solid bounded by six quadrilaterals, but if the polyhedron be doubly
connected, the frame formed by its plane projection will have three degrees of
stiffness. (See Diagram IT.)

Theorem.—lf every one of a system of points in a plane is in equilibrium
under the action of tensions and pressures acting along the lines joining the
points, then if we substitute for each point a small smooth ring through
which smooth thin rods of indefinite length corresponding to the lines are
compelled to pass, then, if to each rod be applied a couple in the plane, whose
moment is equal to the product of the length of the rod between the points
multiplied by the tension or pressure in the former case, and tends to turn the
rod in the positive or the negative direction, according as the force was a tension
or a pressure, then every one of the system of rings will be in equilibrium. For
each ring is acted on by a system of forces equal to the tensions and pressures
in the former case, each to each, the whole system being turned round a right
angle, and therefore the equilibrium of each point is undisturbed.

Theorem.—In any system of points in equilibrium in a plane under the
action of repulsions and attractions, the sum of the products of each attraction
multiplied by the distance of the points between which it acts, is equal to the
sum of the products of the repulsions multiplied each by the distance of the
points between which it acts.

For since each point is in equilibrium under the action of a system of attrac-
tions and repulsions in one plane, it will remain in equilibrium if the system
of forces is turned through a right angle in the positive direction. If this opera-
tion is performed on the systems of forces acting on all the points, then at the
extremities of each line joining two points we have two equal forces at right

VOL. XXVI. PART I. D

14 MR CLERK MAXWELL ON

angles to that line and acting in opposite directions, forming a couple whose
magnitude is the product of the force between the points and their distance, and
whose direction is positive if the force be repulsive, and negative if it be attractive.
Now since every point is in equilibrium these two systems of couples are in
equilibrium, or the sum of the positive couples is equal to that of the negative
couples, which proves the theorem.

In a plane frame, loaded with weights in any manner, and supported by
vertical thrusts, each weight must be regarded as attracted towards a horizontal
base line, and each support of the frame as repelled from that line. Hence the
following rule :—

Multiply each load by the height of the poimt at which it acts, and each
tension by the length of the piece on which it acts, and add all these products
together.

Then multiply the vertical pressures on the supports of the frame each by
the height at which it acts, and each pressure by the length of the piece on
which it acts, and add the products together. This sum will be equal to the
former sum.

If the thrusts which support the frame are not vertical, their horizontal
components must be treated as tensions or pressures borne by the foundations
of the structure, or by the earth itself.

The importance of this theorem to the engineer arises from the circum-
stance that the strength of a piece is in general proportional to its section, so
that if the strength of each piece is proportional to the stress which it has
to bear, its weight will be proportional to the product of the stress multiplied
by the length of the piece. Hence these sums of products give an estimate
of the total quantity of material which must be used in sustaining tension and
pressure respectively.

The following method of demonstrating this theorem does not require the
consideration of couples, and is applicable to frames in three dimensions.

Let the system of points be caused to contract, always remaining similar
to its original form, and with its pieces similarly situated, and let the same forces
continue to act upon it during this operation, so that every point is always in
equilibrium under the same system of forces, and therefore no work is done by
the system of forces as a whole.

Let the contraction proceed till the system is reduced to a point. Then the
work done by each tension is equal to the product of that tension by the distance
through which it has acted, namely, the original distance between the points.
Also the work spent in overcoming each pressure is the product of that pressure
by the original distance of the points between which it acts; and since no work
is gained or lost on the whole, the sum of the first set of products must be
equal to the sum of the second set. In this demonstration it is not necessary

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 15

to suppose the points all in one plane. This demonstration is mathematically
equivalent to the following algebraical proof :—

Let the co-ordinates of the » different points of the system be 2, y, 2,,
Ly Yo %oy Bp Yn Zp, &C., and let the force between any two points p, q, be P,,,
and their distance 7,,, and let it be reckoned positive when it is a pressure, and
negative when it is a tension, then the equation of equilibrium of any point p

with respect to forces parallel to z is
Coe EE Gee” op Bree (a, a) + die, = 0,
"p, "ps "pq
or generally, giving ¢ all values from 1 to n,
us efi) Ls
Si { x v1) | i

Multiply this equation by z, There are nm such equations, so that if each is
multiplied by its proper co-ordinate and the sum taken, we get

se id ane} ey
=P x14 Gp a) ae 0,
and adding the corresponding equations in y and Z, we get
=P > ¢ (Pn rt) = OF

which is the algebraic expression of the theorem.

General Theory of Diagrams of Stress in Three Dimensions.
First Method of Representing Stress in a Body.

Definition —A. diagram of stress is a figure having such a relation to a
body under the action of internal forces, that if a surface A, limited by a closed
curve, is drawn in the body, and if the corresponding limited surface a@ be drawn
in the diagram of stress, then the resultant of the actual internal forces on the
positive side of the surface A in the body is equal and parallel to the resultant
of a uniform normal pressure p acting on the positive side of the surface @ in
the diagram of stress.

Let 2, y, z be the co-ordinates of any point in the body, € », ¢ those of the
corresponding point in the diagram of stress, then €, y, ¢ are functions of a, y, z,
the nature of which we have to ascertain, so that the internal forces in the body
may be in equilibrium. For the present we suppose no external forces, such
as gravity, to act on the particles of the body. We shall consider such forces
afterwards.

Theorem 1.—If any closed surface is described in the body, and if the stress
on any element of that surface is equal and parallel to the pressure on the cor-

*%.

16 MR CLERK MAXWELL ON

responding element of surface in the diagram of stress, then the resultant stress
on the whole closed surface will vanish ; for the corresponding surface in the
diagram of stress is a closed surface, and the resultant of a uniform normal
pressure p on every element of a closed surface is zero by hydrostatics.

It does not, however, follow that the portion of the body within the closed sur-
face is in equilibrium, for the stress on its surface may have a resultant moment.

Theorem 2.—To ensure equilibrium of every part of the body, it is necessary
and sufficient that

where F is any function of z y and z.
Let us consider the elementary area in the body dy dz. The stress acting
on this area will be a force equal and parallel to the resultant of a pressure p
acting on the corresponding element of area in the diagram of stress. Resolving
this pressure in the directions of the co-ordinate axes, we find the three com-
ponents of stress on dy dz, which we may call p,,.dy dz, p,, dy dz, and p,, dy dz,
each equal to p multiplied by the area of the projection of the corresponding
element of the diagram of stress on the three co-ordinate planes. Now, the
projection on the plane yz, is
dn dé dn dt
dy dz dz dy
Hence we find for the component of stress in the direction of x
dn dé dn d

which we may write for brevity at present
ae Pex = pS (7,03 Y2)-
Similarly,
Pry = pJ(S,E3y>2) Pac = PI(E,9; Y52) -

In the same way, we may find the components of stress on the areas dz dx
and da dy— ;
Pye = pS (0,3; 2,2) Pw = p3(6,€3 #2) Pye = pS (E,05 2%)
Paz = pS (,53 @Y) Py = pI(E,E5 v,Y) Pi = pS (Es; @,Y)-
Now, consider the equilibrium of the parallelopiped dx dy dz, with respect to
the moment of the tangential stresses about its axes.

The moments of the forces tending to turn this elementary parallelopiped

about the axis of x are
dz Ax Dy, .dy — dx dy py. dz.

To ensure equilibrium as respects rotation about the axis of 2, we must have

Pyz = Py -

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. Ls

Similarly, for the moments about the axes of y and 2, we obtain the equa-

tions
Pex = Paz and Pry = Pryzx i

Now, let us assume for the present

d d dé
Bails oF = Bee, , 7a C,,
dn dn _ dn

ge Pec eae ae gee
ag ag ae
age Casa ede

Then the equation p,, = p., becomes

Ae ete aa
Cede ~divdz) ‘\du dy dy dx
or

(By ar C,) (B, — 03) — Ay (B, + Cy) = (B, + C,) (B, + C5) — A, (B, — C,)

0=A,C, # B,C, + BO, .

Similarly, from the two other equations of equilibrium we should find
0= A.C, + B,C, + B,C,
O= A, 0. ARC 4 BiG,

From these three equations it follows that

1 =0 C,=0 C7 — 08
Hence

dy_ at ae _ age
dz lay? @a de * - dy a

and édx + ndy + Cdz isa complete differential of some function, F, of x, y and z,
whence it follows that

F may be called the function of stress, because when it is known, the diagram
of stress may be formed, and the components of stress calculated. The form
of the function F is limited only by the conditions to be fulfilled at the bound-
ing surface of the body.

The six components of stress expressed in terms of F are

_ (VEGF (PF = (CECE _ =) _ (PF ae? il Oe
tee SE dy? dz =! » Pu -I\ EE da® dzda ), Pa =P da® dy? sit)

CE EP GEOR), _ (GF AL AEP), (CREE OF PY
Pea =P = dydz dy? ig) BIE Cas de® dady
VOL. XXVI. PART I. E

18 MR CLERK MAXWELL ON

dF ; : ‘ ,
lie = z,F becomes Arry’s function of stress in two dimensions, and we have

a?F d*F d?F
Paw =P Gye > Py oe a, in = ~ P acdy *

The system of stress in three dimensions deduced in this way from any
function, F, satisfies the equations of equilibrium of internal stress. It is not,
however, a general solution of these equations, as may be easily seen by taking
the case in which p,, and p,, are both zero at all pomts. In this case, since
there is no tangential action in planes parallel to ay, the stresses p,,., Pn, and py,
in each stratum must separately fulfil the conditions of equilibrium,

Gy Oy th en
de Pee t dy? ™ ae deb + dyh es
The complete solution of these equations is, as we have seen,

Pf d2f df

Pzz = dy? > Pay = ~ dx dy D Pa = Fe >

where / is any function of 2 and y, the form of which may be different for every
different value of z, so that we may regard fas a perfectly general function of
ay and z.

Again, if we consider a cylindrical portion of the body with its generating
lines parallel to z, we shall see that there is no tangential action parallel to z
between this cylinder and the rest of the body. Hence the longitudinal stress
in this cylinder must be constant throughout its length, and is independent of
the stress in any other part of the body.

Hence

Da = (ey),
where ¢ is a function of 2 and y only, but may be any such function. But

expressing the stresses in terms of F under the conditions p,,=0, p,,=0, we
find that if F is a perfectly general function of # and y

a’F d?¥
dat dz = 0 and dy dz => 0 )

whence it follows that “= and a are functions of z and y only, and that = is a

function of z only. Hence
F=G6G+4+2Z,
when G is a function of z and y only, and Z a function of z only, and the com-
ponents of stress are
PG AZ 2 Ordre fo Ce 0G \)
Pes =P Gye dg? Pw Pq qe? Pe = (Te dy? dudy

. | ae ez
En ee Por =~ P dedy de’

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 19

Here the function / which determines the stress in the strata parallel to zy is
d?Z

Now, this function is not sufficiently general, for instead of being any function of

x,y and 2, it is the product of a function of # and y multiplied by a function of z.
Besides this, though the value of 7,, is, as it ought to be, a function of 2 and

y only, it is not of the most general form, for it depends on G, the function which

determines the stresses p,,, Px,» and p,,, whereas the value of p., may be entirely

independent of the values of these stresses. In fact, the equations give

— Dey"
Pz =P Pele p ?
This method, therefore, of representing stress in a body of three dimensions
is a restricted solution of the equations of equilibrium.

On Reciprocal Diagrams in Three Dimensions.

Let us consider figures in two portions of space, which we shall call respec-
tively the first and the second diagrams. Let the co-ordinates of any point in
the first diagram be denoted by 2, y, z, and those of the corresponding point in
the second by & 7, ¢, measured in directions parallel to a, y, z respectively.
Let F be a quantity varying from point to point of the first figure in any con-
tinuous manner; that is to say, if A, B are two points, and F,, F, the values
of F at those points; then, if B approaches A without limit, the value of F,
approaches that of F, without limit. Let the co-ordinates (€, 7, ¢) of a point in
the second diagram be determined from 2, y, z, those of the corresponding point
in the first by the equations

d¥ d¥ d¥
E = ae > 7 = dy > G = GE ° . ° . (iL).
This is equivalent to the statement, that the vector (p) of any point in the
second diagram represents in direction and magnitude the rate of variation of F
at the corresponding point of the first diagram.
Next, let us determine another function, ¢, from the equation

wE+yn+20=F+ od . : j : : (2),

@, as thus determined, will be a function of x, y, and z, since €, y, ¢ are known
in terms of these quantities. But, for the same reason, ¢ is a function of €, », ¢
Differentiate @ with respect to €, considering 2, y and z functions of €, n, &

a! dx di dz - dF
emer ceah hae ar. ak

20 MR CLERK MAXWELL ON

Substituting the values of € 1, ¢ from (1)

a6 _ 4, Pde | dB dy | hide
dé. da dé dy dé dz dé dé
=>“2+ ce = =
meer oar
= #

Differentiating @ with respect to y and ¢, we get the three equations
d d. d 2
$ $ ie ; (3),

ye y= da z= at
or the vector (7) of any point in the first diagram represents in direction and
magnitude the rate of increase of ¢ at the corresponding point of the second
diagram.

Hence the first diagram may be determined from the second by the same
process that the second was determined from the first, and the two diagrams,
each with its own function, are reciprocal to each other.

The relation (2) between the functions expresses that the sum of the func-
tions for two corresponding points is equal to the product of the distances of
these points from the origin multiplied by the cosine of the angle between the
directions of these distances.

Both these functions must be of two dimensions in space. Let F’ be a
linear function of xyz, which has the same value and rate of variation as F
has at the point a, y, z
dF,
dz

‘ d¥ dF
i =F, + («—%) ie + Y—%) Gy + (¢—%) (4).

The value of F’ at the origin is found by putting 2, y and z=0

F=F,-—x&-—yn-a4~l=—-¢ . ; : : (5),
or the value of F’ at the origin is equal and opposite to the value of ¢ at the
point & », ¢

If the rate of variation of F is nowhere infinite, the co-ordinates € y ¢ of the
second diagram must be everywhere finite, and vice versa. Beyond the limits
of the second diagram the values of z, y, z,m terms of & », ¢, must be impossible,
and therefore the value of ¢ is also impossible. Within the limits of the second
diagram, the function ¢ has an even number of values at every point, corre-
sponding to an even number of points in the first diagram, which correspond
to a single point in the second.

To find these points in the first diagram, let , p be the vector of a given point
in the second diagram, and let surfaces be drawn in the first diagram for which
F is constant, and let points be found in each of these surfaces at which the
tangent plane is perpendicular to p, these points will form one or more curves,
which must be either closed or infinite, and the points on these curves corres-

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. Al

pond to the points in the second diagram which lie in the direction of the
vector p. If p be the perpendicular from a point in the first diagram on a
plane through the origin perpendicular to p, then all those points on these curves

at which os =p correspond to the given point in the second diagram. Now,
since this point is within the second diagram, there are values of p both greater

and less than the given one; and therefore ap neither an absolute maximum

nor an absolute minimum value. Hence there are in general an even number
of points on the curve or curves which correspond to the given point. Some of
these points may coincide, but at least two of them must be different, unless
the given point is at: the limit of the second diagram.

Let us now consider the two reciprocal diagrams with their functions, and
ascertain in what the geometrical nature of their reciprocity consists.

(1.) Let the first diagram be simply the point P,, (z,, y,, z,), at which F=F,,
then in the other diagram

$=HE+y,n+246-—F, . : : A : (6),

or a point in one diagram is reciprocal to a space in the other, in which the.
function ¢ is a linear function of the co-ordinates.

(2.) Let the first diagram contain a second point P,,, (2, y,, 2,) at which F=F,,
then we must combine equation (6) with

Pe ea gal Pao Ny Oe | Ke,
whence eliminating ¢,

(@—%)E + Yy— Y2)n + (%—%) 6 = F,—-F,.

If 7,, is the length of the line drawn from the first point P, to the second P,:
and if /,. m2 2. are its direction cosines, this equation becomes
Le& + Mo7 + Myo = eae
12
or the reciprocal of the two points P, and P, is a plane, perpendicular to the line
joining them, and such that the perpendicular from the origin on the plane
multiplied by the length of the line P,P, is equal to the excess of F, over F,.
(3.) Let there be a third point P, in the first diagram, whose co-ordinates are
X,Y, 2, and for which F = F,; then we must combine with equations (6) and (7)
p= a,& + ¥,n + %6— F, , : ° : (8).
The reciprocal of the three points P, P, P, is a straight line perpendicular to
the plane of the three points, and such that the perpendicular on this line from
the origin represents, in direction and magnitude, the rate of most rapid increase
of F in the plane P, P, P,, F being a linear function of the co-ordinates whose

values at the three points are those given.
VOL.. XXVI. PART I. F

22 MR CLERK MAXWELL ON

(4.) Let there be a fourth point P, for which F = F,.

The reciprocal of the four points is a single point, and the line drawn from
the origin to this point represents, in direction and magnitude, the rate of
greatest increase of F, supposing F such a linear function of zyz that its values
at the four points are those given. The value of ¢ at this point is that of F at
the origin.

Let us next suppose that the value of F is continuous, that is, that F does
not vary by a finite quantity when the co-ordinates vary by infinitesimal
quantities, but that the form of the function F is discontinuous, being a
different linear function of zyz in different parts of space, bounded by definite
surfaces.

The bounding surfaces of these parts of space must be composed of planes.
For let the linear functions of zyz in contiguous portions of space be

BL >= 4,24 By +942 —-— 9,
Fy = a,¢ + Boy + Yo% — dy,

then at the bounding surface, where F, = F,
(a,—a,)x + (B,—Bo)y + (i —-12)2 = di—-Gy- : . (9),

and this is the equation of a plane.

Hence the portion of space in which any particular form of the value of F
holds good must be a polyhedron or cell bounded by plane faces, and therefore
having straight edges meeting in a number of points or summits.

Every face is the boundary of two cells, every edge belongs to three or more
cells, and to two faces of each cell.

Every summit belongs to at least four cells, to at least three faces of each
cell, and to two edges of each face.

The whole space occupied by the diagram is divided into cells in two different
ways, so that every point in it belongs to two different cells, and has two values
of F and its derivatives.

The reciprocal diagram is made up of cells in the same way, and the
reciprocity of the two diagrams may be thus stated :—

1. Every summit in one diagram corresponds to a cell in the other.

The radius vector of the summit represents the rate of increase of the func-
tion within the cell, both in direction and magnitude.

The value of the function at the summit is equal and opposite to the value
which the function in the cell would have if it were continued under the same
algebraical form to the origin.

2. Every edge in the one diagram corresponds to a plane face in the other,
which is the face of contact of the two cells corresponding to the two extremities ~
of the edge.

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 23

The edge in the one diagram is perpendicular to the face in the other.

The distance of the plane from the origin represents the rate of increase of
the function along the edge.

3. Every face in the one diagram corresponds to an edge in which as many
cells meet as there are angles in the face, that is, at least three. Every face
must belong to two, and only two cells, because the edge to which it corresponds
has two, and only two extremities.

4. Every cell in the one diagram corresponds to a summit in the other.
Every face of the cell corresponds and is perpendicular to an edge having an
extremity in the summit. Since every cell must have four or more faces, every
summit must have four or more edges meeting there.

Every edge of the cell corresponds to a face having an angle in the summit.
Since every cell has at least six edges, every summit must be the point of
concourse of at least six faces, which are the boundaries of cells.

Every summit of the cell corresponds to a cell having a solid angle at the
summit. Since every cell has at least four summits, every summit must be the
meeting place of at least four cells.

Mechanical Reciprocity of the Diagrams.

If along each of the edges meeting in a summit forces are applied propor-
tional to the areas of the corresponding faces of the cell in the reciprocal
diagram, and in a direction which is always inward with respect to the cell,
then these forces will be in equilibrium at the summit.

This is the “ Polyhedron of Forces,” and may be proved by hydrostatics.

If the faces of the cell form a single closed surface which does not intersect
itself, it is easy to understand what is meant by the inside and outside of the
cell; but if the surface intersects itself, it is better to speak of the positive and
negative sides of the surface. A cell, or portion of a cell, bounded by a closed
surface, of which the positive side is inward, may be called a positive cell. If
the surface intersects itself, and encloses another portion of space with its
negative side inward, that portion of space forms a negative cell. If any portion
of space is surrounded by m sheets of the surface of the same cell with their
positive side inward, and by m sheets with their negative side inward, the space
enclosed in this way must be reckoned —m times.

In passing to a contiguous cell, we must suppose that its face in contact
with the first cell has its positive surface on the opposite side from that of the
first cell. In this way, by making the positive side of the surface continuous
throughout each cell, and by changing it when we pass to the next cell, we may
settle the positive and negative side of every face of every cell, the sign of
every face depending on which of the two cells it is considered for the moment
to belong to.

24 “MR CLERK MAXWELL ON

If we now suppose forces of tension or pressure applied along every edge of
the first diagram, so that the force on each extremity of the edge is in the
direction of the positive normal to the corresponding face of the cell corres-
ponding to that extremity, and proportional to the area of the face, then
these pressures and tensions along the edges will keep every point of the
diagram in equilibrium.

Another way of determining the nature of the force along any edge of the
first diagram, is as follows :—

Round any edge of the first diagram draw a closed curve, embracing it and
no other edge. However small the curve is, it will enter each of the cells which
meet in the edge. Hence the reciprocal of this closed curve will be a plane
polygon whose angles are the points reciprocal to these cells taken in order.
The area of this polygon represents, both in direction and magnitude, the whole
force acting through the closed curve, that is, in this case the stress along the
edge. If, therefore, in going round the angles of the polygon, we travel in the
same direction of rotation in space as in going round the closed curve, the stress
along the edge will be a pressure; but if the direction is opposite, the stress will
be a tension.

This method of expressing stresses in three dimensions comprehends all cases
in which RANKINE’s reciprocal figures are possible, and is applicable to certain
cases of continuous stress. That it is not applicable to all such cases is easily
seen by the example of p (18).

On Reciprocal Diagrams in Two Dimensions.

If we make F a function of # and y only, all the properties already deduced
for figures in three dimensions will be true in two; but we may form a more
distinct geometrical conception of the theory by substituting cz for F and c¢ for
¢. We have then for the equations of relation between the two diagrams

nog yf
~~ “da a ay
ae. nae oy pene CE
Fmues2 came

LE + yn = 8 + OC .

These equations are equivalent to the following definitions :—

Let z in the first diagram be given as a function of 2 and y, z will lie ona
surface of some kind. Let 2,, y, be particular values of 2 and y, and let z, be the
corresponding value of z. Draw a tangent plane to the surface at the point
Lor Yor Xo, and from the point € = 0,7 = 0, €=—c; in the second diagram draw
a normal to this tangent plane. It will cut the plane €=0 at the point € » cor-
responding to zy, and the value of ¢ is equal and opposite to the segment of the

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 25

axis of z cut off by the tangent plane. The two surfaces may be defined as recip-
rocally polar (in the ordinary sense) with respect to the paraboloid of revolution

Rel Yer Dee : : : , ; (lays

and the diagrams are the projections on the planes of xy and &y of points and
lines on these surfaces.

If one of the surfaces is a plane-faced polyhedron, the other will also be a
plane-faced polyhedron, every face in the one corresponding to a point in the
other, and every edge in the one corresponding to the line joining the points
corresponding to the faces bounded by the edge. In the projected diagrams
every line is perpendicular to the corresponding line, and lines which meet in a
point in one figure form a closed polygon in the other.

These are the conditions of reciprocity mentioned at p. 8, and it now
appears that if either of the diagrams is a projection of a plane-faced poly-
hedron, the other diagram can be drawn. If the first diagram cannot be a pro-
jection of a plane-faced polyhedron, let it be a projection of a polyhedron whose
faces are polygons not in one plane. These faces must be conceived to be filled
up by surfaces, which are either curved or made up of different plane portions. -
In the first case the polygon will correspond not to a point, but to a finite por-
tion of a surface ; in the second, it will correspond to several points, so that the
lines, which correspond to the edges of such a polygon, will terminate in several
points, and not in one, as is necessary for reciprocity.

Second Method of representing Stress in a Body.

Let a, 6 be any two consecutive points in the first diagram, distant s, and a, 8
the corresponding points in the second, distant o, then if the direction cosines
of the line a 0 are /, m, n and those of a B, X, p, v

Gh = sie WE ope
dz

+ sm +
dy
dn dn dn i
= sl —- ea —— : , ‘ 12).
of = lay + omy + sive (12)
_ 6 dt dt
oy = slr + a + sn
Hence
Y
2(r.+ mp +nv) =128 us +n m2 nie + mn( FE + ae) aloe +E )an n( e+ +7) (13).

If we put /\ + mp + mv = cos «, where ¢ is the angle between s and oa, and
if we take three sets of values of /mn, corresponding to three directions at right
angles to each other, we find

dé dn dg oe d?F . TAD in d?F
deo * dy + ie da dy? ~ dz
VOL. XXVI. PART I. G

O71
=) cos €, + —* cose, + —® cos es = (14).
1 8

26 MR CLERK MAXWELL ON

Hence this quantity depends only on the position of the point, and not on the
directions of s, s, s, or of # y z, let us call it A’F.

Now, let us take an element of area perpendicular to s, and let us suppose
that the stress on this element is compounded of a normal pressure = pA’F,

and a tension parallel to o and equal to p - é

By the rules for the composition of stress, we have for the components of the
force on this element, in terms of the six components of stress,

X = [pyr + Mpzy + Mor = ( [A?F — 2)
Y = pay + Mpy + Mpy: = p( mA?7F — 2) ; ‘ (15).

L = Woe, + Mpy + Me = of nd?F — 12)

Hence,
= Fh = a op _ UF oh fT ys (OE
Pre = r( A?*F ee ( A’F dx? ») ae CS 2 dz?
ar | a?F Le d?F d?F
= oo 9G eo) Pu = P + de? 2): iad & ce dy? (te
nt - d?F BA dF
Pu =~ P aedy a; y” Pie Poeda? Pah aady

By substituting these values in the equations of equilibrium

Apex , Way | Ure
tet dent dg =O? & - ,

it is manifest that they are fulfilled for any value of F.
The most general solution of these equations of equilibrium is contained in
the values

Be ae CO AAA _ @A oe d?B
ee Ae of dye Pai sa + mee Pu = dy? ae

1 GA ow d?B =e =
De i dydz Pea ~~ edz ia dxdy

By making A = B= C= ?7F we get a case which, though restricted in its
generality, has remarkable properties with respect to diagrams of stress.
We have seen that a distribution of stress according to the definition above
(16), is consistent with itself, and will keep a body in equilibrium. Since the
stresses are linear functions of F, any two systems of stress can be compounded
by adding their respective functions, a process not applicable to the first method
of representation by areas.

Let us ascertain what kind of stress is represented in this way in the case
of the system of cells already considered.

Since F in each cell is a linear function of 2, y, z, there can be no stress at any

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 2G

point within it. Let us take a and } two contiguous points in different cells,
then a and 6 will be the points at a finite distance to which these cells are

reciprocal, and A’F a , which becomes infinite when a vanishes.

If a and 6 are in the surface bounding the cells, a and B coincide. Hence
there is a stress in this surface, uniform in all directions in the plane of the
surface, and such that the stress across unit of length drawn on the surface is
proportional to the distance between the points which are reciprocal to the two
eells bounded by the surface, and this stress is a tension or a pressure according
as the two points are similarly or oppositely situated to the two cells.

The kind of equilibrium corresponding to this case is therefore that of a
system of liquid films, each having a tension like that of a soap bubble, depend-
ing on the nature of the fluid of which it is composed. If all the films are
composed of the same fluid, their tensions must be equal, and all the edges of
the reciprocal diagram must be equal.

On Airy’s Function of Stress.

Mr Arry, in a paper “On the Strains in the Interior of Beams,’* was, I-
believe, the first to point out that, in any body in equilibrium under the action
of internal stress in two dimensions, the three components of the stress in any
two rectangular directions are the three second derivatives, with respect to these
directions, of a certain function of the position of a point in the body.

This important simplification of the theory of the equilibrium of stress in
two dimensions does not depend on any theory of elasticity, or on the mode in
which stress arises in the body, but solely on the two conditions of equilibrium
of an element of a body acted on only by internal stress

d d ad d iz
dae an dy?” — 0) and dal” + dy?" =0 . : (19),
whence it follows that
d?F d2F hay
Prez = dy? Pry = — dady and Py = Aas . . (20),

where F is a function of w and y, the form of which is (as far as these equations
are concerned) perfectly arbitrary, and the value of which at any point is in-
dependent of the choice of axes of co-ordinates. Since the stresses depend on
the second derivatives of F, any linear function of w and y may be added to F
without affecting the value of the stresses deduced from F. Also, since the
stresses are linear functions of F, any two systems of stress may be mechanically
compounded by adding the corresponding values of F.

The importance of Airy’s function in the theory of stress becomes even more

* Phil. Trans. 1863.

28 MR CLERK MAXWELL ON

manifest when we deduce from it the diagram of stress, the co-ordinates of whose
points are
dF
—&= — and = . : . . . (21).

For if s be the length of any curve in the original figure, and o that of the cor-
responding curve in the diagram of stress, and if Xds, Yds are the components
of the whole stress acting on the element ds towards the right hand of the

curve s

dy aE dy , d& dy dé
a da Phi dy? ds “° — dy Faden: Mba

and (22).
os dz, | @Ede, dyda, ay
SAN a OS Tat Z,4= dn ds 8 = do

Hence the stress on the right hand side of the element ds of the original curve
is represented, both in direction and magnitude, by the corresponding element
do of the curve in the diagram of stress, and, by composition, the resultant
stress on any finite arc of the first curve s is represented in direction and
magnitude by the straight lme drawn from the beginning to the end of the
corresponding curve o.

If P,, P, are the principal stresses at any point, and if P, is inclined a to the

axis of z, then the component stresses are

Pr = P, cos? a + P, sin? a |
Dey = (P, — P,) sin a cos a : (23).
Dy = P, sin* a + P, cosa J
Hence
dF
lxda
t 9 = Pry — a Y
ers Pex ah Puy iy d?F
dat dy?
@F @F (24).
P,+P,= Pax 7 Poy pg ap

on pte CP ER_ EF
PP = Pex Poy — Pry” = dx? dy? dady

Consider the area bounded by a closed curve s, and let us determine the sur-
face integral of the sum of the principal stresses over the area within the curve.

The integral is
2 2
(P, + P,) dads oat) A Ho Me lt ROSE
2 Uy — ae.

By a well-known theorem, corresponding in two dimensions to that of GREEN in
three dimensions, the latter expression becomes, when once integrated,

dF d« dF dy
eae a6 ee a eee

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 29

or S(e& ait): ah rr ari CL.

These line integrals are to be taken round the closed curve s. If we take a
point in the curve s as origin in the original body, and the corresponding point
in o as origin in the diagram of stress, then € and y are the components of the
whole stress on the right hand of the curve from the origin to a given point.
If p denote the line joining the origin with the point &y, then p will represent in
direction and magnitude the whole stress on the arc o.

The line integral may now be interpreted as the work done on a point which
travels once round the closed curve s, and is everywhere acted on by a force
represented in direction and magnitude by p. We may express this quantity
in terms of the stress at every point of the curve, instead of the resultant stress
on the whole arc, as follows :—

For integrating (27) by parts it becomes,

d dy |
-f (2 e+ ot) ds=— f (Xe + Vy) ds ee oR

or if Rds is the actual stress on ds, and 7 is the radius vector of ds, and if R
makes with 7 an angle e, we obtain the result ,
Jf@Gi + 2, )dady=—f Rr eoseds. : ; (29).
This line integral, therefore, which depends only on the stress acting on the
closed curve s,is equal to the surface integral of the sum of the principal
stresses taken over the whole area within the curve.

If there is no stress on the curve s acting from without, then the surface
integral vanishes. ‘This is the extension to the case of continuous stress of the
theorem, given at p. 13, that the algebraic sum of all the tensions multiplied
each by the length of the piece in which it acts is zero for a system in equili-
brium. In the case of a frame, the stress in each piece is longitudinal, and the
whole pressure or tension of the piece is equal to the longitudinal stress multi-
plied by the section, so that the integral //(P, + P,)dxdy for each piece is its
tension multiplied by its length.

If the closed curve s is a small circle, the corresponding curve o will be an
ellipse, and the stress on any diameter of the circle will be represented in direc-
tion and magnitude by the corresponding diameter of the ellipse. Hence, the
principal axes of the ellipse represent in direction and magnitude the principal
stresses at the centre of the circle.

Let us next consider the surface integral of the product of the principal
stresses at every point taken over the area within the closed curve s.

aa Pata)
2 =
SN fl Piedad ah ah (‘a e — Tai) dady . : (30),
dé d dé d
“SI Gea — aya)

WOLD PXEXGVAL PARI 1. H

30 MR CLERK MAXWELL ON

or by transformation of variables

=f f ea

Hence the surface integral of the product of the principal stresses within
the curve is equal to the area of the corresponding curve o in the diagram of
stress, and therefore depends entirely on the external stress on the curve s.
This is seen from the construction of the curve o in the diagram of stress, since
each element do represents the stress on the corresponding element ds of the
original curve.

If p represents in direction and magnitude the resultant of the stress on the
curve s from the origin to a point which moves round the curve, then the area
traced out by p is equal to the surface-integral required. If Xds and Yds
are the components of the stress on the element ds, and 7 the whole length of
the closed curve s, then the surface integral is equal to either of the quantities.

1 s a
SLX fXis.0s, or —f Xf Vas. as.

In a frame the stress in each piece is entirely longitudinal, so that the pro-
duct of the principal stresses is zero, and therefore nothing is contributed to
the surface integral except at the points where the pieces meet or cross each
other. To find the value of the integral for any one of these points, draw a
closed curve surrounding it and no other point, and therefore cutting all the
pieces which meet in that point in order. The corresponding figure in the
diagram of stress will be a polygon, whose sides represent in magnitude and
direction the tensions in the several pieces taken in order. The area of this
polygon, therefore, represents the value of // P,P,dady for the point of concourse,
and is to be considered positive or negative, according as the tracing point
travels round it in the positive or the negative cyclical direction.

Hence the following theorem, which is applicable to all plane frames, whether
a diagram of forces can be drawn or not.

For each point of concourse or of intersection construct a polygon, by draw-
ing in succession lines parallel and proportional to the forces acting on the
point in the several pieces which meet in that point, taking the pieces in cyclical
order round the point. The area of this polygon is to be taken positive or
negative, according as it lies on the left or the right of the tracing point.

If, then, a closed curve be drawn surrounding the entire frame, and a poly-
gon be drawn by drawing in succession lines parallel and proportional to all the
external forces which act on the frame in the order in which their lines of
direction meet the closed curve, then the area of this polygon is equal to the
algebraic sum of the areas of the polygons corresponding to the various points
of the frame.

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. ol

In this theorem a polygon is to be drawn for every point, whether the lines
of the frame meet or intersect, whether they are really jointed together, or
whether two pieces simply cross each other without mechanical connection. In
the latter case the polygon is a parallelogram, whose sides are parallel and pro-
portional to the stresses in the two pieces, and it is positive or negative accord-
ing as these stresses are of the same or of opposite signs.

If three or more pieces intersect, it is manifestly the same whether they
intersect at one point or not, so that we have the following theorem :—

The area of a polygon of an even number of sides, whose opposite sides are
equal and parallel, is equal to the sum of the areas of all the different parallelo-
grams which can be formed with their sides parallel and equal to those of the

polygon.
This is easily shown by dividing the polygon into the different parallelograms.

On the Equilibrium of Stress in a Solid Body.

Let PQR be the longitudinal, and STU the tangential components of stress, |
as indicated in the following table of stresses and strains, taken from THomson
and Tart’s “ Natural Philosophy,” p. 511, § 669 :—

Components of the Planes, of which io
Relative Motion, or | _ Direction of
; ————] across which Force, | Relative Motion
Strain. Stress. is reckoned. or of Force.
e P Ye
we Q 200 y
g R wy Zz
1
a S { Y% y
eae Z
(! 4
b "ap 2y Zz
“LY ae
ty U aR a
yz y

Then the equations of equilibrium of an element of the body are, by § 697
of that work,

ee OL aa

da dy dz |

dU dQ. tacds ua

Gp ee Se
dT dS dk |

Tel Ay IE a ea ')

oe MR CLERK MAXWELL ON
If we assume three functions ABC, such that

__@a Manne orang
dydz dela dady
and put SAPD
‘tad dV dV

ae a ee lei

then a sufficiently general solution of the equations of equilibrium is given by
putting

an “aC

= ge tye
PC ee

= ahaa ©),
GA dB |

i dy? + J

I am not aware of any method of finding other relations between the com-
ponents of stress without making further assumptions. The most natural
assumption to make is that the stress arises from elasticity in the body. I
shall confine myself to the case of an isotropic body, such that it can be deprived
of all stress and strain by a removal of the applied forces. In this case, if
a By are the components of displacement, and 7 the co-efficient of rigidity, the
equations of tangential elasticity are, by equation (6) {§ 670 and 694 of THomson
and TAIT,

ec eee ee ea
Wa a cel ice eae . . . C (A),

‘with similar equations for bandc. A sufficiently general solution of these equa-
tions is given by putting

ae = (a- oe c) )
8 = 5, q(B-C-A) 42: a res
y= == (c- ee B)

The equations of longitudinal elasticity are of the form wie in § 693,

2 (84 be) eee 6),

where & is the co-efficient of cubical elasticity, with similar equations for Q and
R. Substituting for P, «, 8 and y in equation (6) their values from (3) and (5),

2 2 2 2 2 2 2 2 2 2 2
(es PC -v)= (i+! {\(C ee (2G PO dA dC dA ss

dz Ce? ea ae dy? dy? at 2” dA

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 55)

If we put

d7A 7B +70 d? d? a
at + aye oo yeaa he and 72 + cr ma 5

this equation becomes
€ + 3”) (47A+A°B+A°0 ) a ( k + 3 )2p— 2nV=2nA2A, soe cca)

We have also two other equations differing from this only in having B and
C instead of A on the right hand side. Hence equating the three expressions
on the right hand side we find
MA=A°B=A7C=D2, say, . . . .~ (8),
Ch np = Cr +2n)8D*= nV,
and
9k D?—2V _ p—3v

P+Q+R=5 Sy a ie Dey

(10).

These equations are useful when we wish to determine the stress rather than
the strain ina body. For instance, if the co-efficients of elasticity, 4 and m, are
increased in the same ratio to any extent, the displacements of the body are
proportionally diminished, but the stresses remain the same, and, though their —
distribution depends essentially on the elasticity of the various parts of the
body, the values of the internal forces do not contain the co-efficients of elasti-
city as factors.

There are two cases in which the functions may be treated as functions of
two variables.

The first is when there is no stress, or a constant pressure in the direction
of z, as in the case of a stratum originally of uniform thickness, in the direction
of z, the thickness being small compared with the other dimensions of the body,
and with the rate of variation of strain.

The second is when there is no strain, or a uniform longitudinal strain in the
direction of z, as in the case of a prismatic body whose length in the direction
of z is very great, the forces on the sides being functions of # and y only.

In both of these cases S = 0 and T = 0, so that we may write
HC a?C d?C

This method of expressing the stresses in two dimensions was first given by
the Astronomer Royal, in the “ Philosophical Transactions” for 1863. We shall
write F instead of C, and call it Atry’s Function of Stress in Two Dimensions.

Let us assume two functions, G and H, such that

2 2
= 7 and V = only : : : ; ; (12),
VOL. XXVI. PART I. I

34 MR CLERK MAXWELL ON

then by THomson and Tart, § 694, if a is the displacement in the direction of a

2n (o + 1) 7 =P-c(Q+R) . : : (13).
Case I.—If R = 0 this becomes

d? {i d?G \.

da
ane tae Gag dy da +(o-—1)H

Integrating with respect to 2 we find the following equation for a—

2n(o+1l)a=

204
aut? ais +Y : (14),

dy \ dy? ° da?

where Y is a function of y only. Similarly for the displacement 8 in the
direction of y,

Qn (o + 1)B = a\Ge ee

“ane att eat ae . “Gar

where X is a function of 2 only. Now the shearing stress U depends on the
shearing strain and the rigidity, or

Ven(S = oe Rees

Multiplying both sides of this equation by 2(o + 1) and substituting from (11),
(14), and (15),

“Ge dG dG d*G d7H 4 CH a =
= Ale tt a dardy? dy* — ~° da®dy? tae age = 7 ee
Hence
a = -
+ in) G+ G+ 4 = -o) (= pe) Ea

an equation which must be fulfilled by G when the body is originally without
strain.
Case II.—In the second case, in which there is no strain in the direction of
z, we have
dY

Substituting for R in (13), and dividing by o + 1,
an 52 = (1 —o)P—oQ

oe d?G H
=Gy lO -Gr og ton}... 0,
with a similar equation for 6. Proceeding as in the former case, we find

7 +o = wa fie ee )

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 30

This equation is identical with that of the first case, with the exception of the
coefficient of the part due to H, which depends on the density of the body, and
the value of o, the ratio of lateral expansion to longitudinal compression.

Hence, if the external forces are given in the two cases of no stress and no
strain in the direction of z, and if the density of the body or the intensity of the
force acting on its substance is in the ratio of o to (1 —o)’ in the two cases, the
internal forces will be the same in every part, and will be independent of the
actual values of the coefficients of elasticity, provided the strains are small.
The solutions of the cases treated by Mr Arry, as given in his paper, do not
exactly fulfil the conditions deduced from the theory of elasticity. In fact, the
consideration of elastic strain is not explicitly introduced into the investigation.
Nevertheless, his results are statically possible, and exceedingly near to the
truth in the cases of ordinary beams.

As an illustration of the theory of Arry’s Function, let us take the case of

uy
ii Dp 7°? cos 2p0 : ; ; ; : (22).

In this case we have for the co-ordinates of the point in the diagram corres-
ponding to (zy)
g = F = 1108 (2p—1)8 =p =" 18in (p18. . 23),

and for the components of stress

Par = = === p= Lm” eos Gp—2)0 =— = =— Pw ]
pe Ve en Od):
Bey = ging = 2p— Wr? sin (2p—2)8 |
If we make
= F rv cosp@ and H == 7? sin p@ ; : (25),
then Seen hoes
Ge ee

(26).
aG dG
(2p—1) 2 pan dy = Pry

Hence the curves for which G and H respectively are constant will be lines of
principal stress, and the stress at any point will be inversely as the square of

the distance between the consecutive curves G or H.

If we make
E=pcosd and yn=psing
then we must have Seen (27):
Die ease and = = (2p—1)0
If we put

1
for i then —
oe =1 P

$7=2 and (2p—1)(2qg—1) =.1,

36 MR CLERK MAXWELL ON

so that iff, g, 4 in the diagram of stress correspond to F, G, H in the original
figure, we have

f= yy prtcos 2¢ Me et cos gp h= Aa singp (28).

Case of a Uniform Horizontal Beam.

As an example of the application of the condition that the stresses must be
such as are consistent with an initial condition of no strain, let us take the case
of a uniform rectangular beam of indefinite length placed horizontally with a
load = / per unit of length placed on its upper surface, the weight of the beam
being & per unit of length. Let us suppose the beam to be supported by vertical
forces and couples in a vertical plane applied at the ends; but let us consider
only the middle portion of the beam, where the conditions applicable to the ends
have no sensible effect. Let the horizontal distance x be reckoned from the
vertical plane where there is no shearing force, and let the planes where there
is no moment of bending be at distances +, from the origin. Let y be
reckoned from the lower edge of the beam, and let 4 be the depth of the beam.

;
Then, if U =— iy is the shearing stress, the total vertical shearing force

through a vertical section at distance 2 is

oh vay = (Fe), “Gs

and this must be equal and opposite to the weight of the beam and load from 0
to x, which is evidently (h + £)z.

Hence
= =—(h+ keg) where $0)—-¢(0)=1. . . (29).

From this we find the vertical stress
an k
Q= Fat zy=—-hthowMr+zy-

The vertical stress is therefore a function of y only. It must vanish at the
lower side of the beam, where y = 0, and it must be — / on the upper side of
the beam, where y = 6. The shearing stress U must vanish at both sides of the
beam, or ¢’(y) = 0, when y = 0, and when y = 6.

The simplest form of ¢(y) which will satisfy these conditions is

$y) = “ (3by? — 2y*) .

Hence we find the following expression for the function of stress by integrating
(29) with respect to 2,

a Lk Gy eie= oer. 16 Wee. (30),

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 37

where @ is a constant introduced in integration, and depends on the manner in
which the beam is supported. From this we obtain the values of the vertical,
horizontal, and shearing stresses,

a eae h+k

_k 3
Se ne AU a D2) a ae ee
1 VE etch ; d2Y 3
LS Fhe Re ee a aie Ae oes (32),
a7 By eae 2

The values of Q and of U, the vertical and the shearing stresses, as given by
these equations, are perfectly definite in terms of / and &, the load and the
weight of the beam per unit of length. The value of P, the horizontal stress,
however, contains an arbitrary function Y, which we propose to find from the
condition that the beam was originally unstrained. We therefore determine
a and £, the horizontal and vertical displacement of any point (a, y), by the
method indicated by equations (13), (14), (15)

2n(o + 1)a= ne _ B

I, 2
| (a 27 — u*®)(b — 2y)— ou (Bby? —2y%)} ==. ny oe
an(ot1ye=— "Bf (iy 594) +30 -oy-v)} +5 Fro F +X (89),
where X’ is a function of 2 only, and Y’ of y only. Deducing from these dis-

placements the shearing strain, and comparing it with the value of the shearing
stress, U, we find the equation

h+k { , E é \ k TENE SGD ONG :
78 6a2x — 2x? + 12x(by — y?) Pie ane aye eter ag, 5 (GIO)
Hence
aby h = k
dy> = 1G, (by — y") . . 5 ° . rn (37),
Ae I ahh : k . ie i .

If the total longitudinal stress across any vertical section of the beam is zero,
the value of a must be the same when y = 0 and wheny = 0. From this con-
dition we find the value of P by equation (32)

{ 3 =) 4: Dy? — Ody — ub my). eee

The moment of bending at any vertical section of the beam is

b
wi Pydy=(h +k) G (a? — a?) + : i”) : : . (40).
0

VOL. XXVI. PART I. K

38 MR CLERK MAXWELL ON

This becomes zero when v = + a, where

Of ate SP cecal fin edaeeal
If we wish to compare this case with that of a beam of finite length supported
at both ends and loaded uniformly, we must make the moment of bending zero
at the supports, and the length of the beam between the supports must therefore
be 2a,. Substituting a, for a in the value of P, we find

h+k
a 78

(805 - Ba2 4+ Dy? — Qby + i) O—my o- 3 ~ e

If we suppose the beam to be cut off just beyond the supports, and supported
by an intense pressure over a small area, we introduce conditions into the
problem which are not fulfilled by this solution, and the investigation of which
requires the use of Fourrer’s series. In order that our result may be true, we
must suppose the beam to extend to a considerable distance beyond the sup-
ports on either side, and the vertical forces to be applied by means of frames
clamped to the ends of the beam, as in Diagram Va, so that the stresses arising
from the discontinuity at the extremities are insensible in the part of the beam
between the supports.

This expression differs from that given by Mr Arry only in the terms in the
longitudinal stress P depending on the function Y, which was introduced in
order to fulfil the condition that, when no force is applied, the beam is un-
strained. The effect of these terms is a maximum when y = ‘127884, and is
then equal to (2 + #)°314, or less than a third of the pressure of the beam and
its load on a flat horizontal surface when laid upon it so as to produce a uniform
vertical pressure h + &,

RECIPROCAL FIGURES, FRAMES, AND DIAGRAMS OF FORCES. 39

EXPLANATION OF THE DIAGRAMS (Puatzss I. II. III).

Diagrams I.a and I.d illustrate the necessity of the condition of the possibility of reciprocal
diagrams, that each line must be a side of two, and only two, polygons. Diagram I.a is a skeleton of
a frame such, that if the force along any one piece be given, the force along any other piece may be
determined. But the piece N forms a side of four triangles, NFH, NGI, NJL, and NKM, so that if
there could be a reciprocal diagram, the line corresponding to N would have four extremities, which is
impossible. In this case we can draw a diagram of forces in which the forces H, I, J, and K are each
represented by two parallel lines.

Diagrams IL.a and II.d illustrate the case of a frame consisting of thirty-two pieces, meeting four
and four in sixteen points, and forming sixteen quadrilaterals. Diagram Il.a may be considered as a
plane projection of a polyhedron of double continuity, which we may describe as a quadrilateral frame
consisting of four quadrilateral rods, of which the ends are bevelled so as to fit exactly. The pro-
jection of this frame, considered as a plane frame, has three degrees of stiffness, so that three of the
forces may be arbitrarily assumed.

In the reciprocal diagram II.d the lines are drawn by the method given at p. 7, so that each
line is perpendicular to the corresponding line in the other figure. To make the corresponding lines
parallel we have only to turn one of the figures round a right angle.

Diagrams III.a and III. illustrate the principle as applied to a bridge designed by Professor F.
Jenkin. ‘The loads Q, Q,, &c., are placed on the upper series of joints, and R, R,, &c., on the lower
series. The diagram III.6 gives the stresses due to both sets of loads, the vertical lines of loads being
different for the two series.

Diagrams IV.a and IV.d illustrate the application of Artrys Function to the construction of
diagrams of continuous stress.

IV.a@ represents a cylinder exposed to pressure in a vertical and horizontal direction, and to
tension in directions inclined 45° to these. The lines marked a, J, ¢, &c., are lines of pressure, and
those marked 9, p, q, are lines of tension. In this case the lines of pressure and tension are rectangular
hyperbolas, the pressure is always equal to the tension, and varies inversely as the square of the
distance between consecutive curves, or, what is the same thing, directly as the square of the distance
from the centre.

IV.b represents the reciprocal diagram corresponding to the upper quadrant of the former one.
The stress on any line in the first diagram is represented in magnitude and direction by the corres-
ponding line in the second diagram, the correspondence being ascertained by that of the corresponding
systems of lines a, 0, c, &c., and 0, p, g, &e.

We may also consider IV.0 as a sector of a cylinder of 270°, exposed to pressure along the lines

=o
a, 6, c, and to tension along 0, p, g, the magnitude of the stress being inthiscase r *. The upper

quadrant of IV.a is in this case the reciprocal figure. This figure illustrates the tendency of any _
strained body to be ruptured at a re-entering angle, for it is plain that at the angle the stress becomes
indefinitely great.

In diagram IV.a—

F= 79 cos 40 G = 51? cos 20 H = 5 7?sin 20.
In diagram IV.6—
4a 3 2 2
=i pcos <6 Oe cose h = 3p * sin 26

Diagrams V.a and V.b illustrate Arry’s theory of stress in beams.

V.a is the beam supported at C and D by means of bent pieces clamped to the ends of the beam
at A and B, at such a distance from C and D, that the part of the beam between C and D is free from
the local effects of the pressures of the clamps at A and B. The beam is divided into six strata by

40 MR CLERK MAXWELL ON RECIPROCAL FIGURES, FRAMES, ETC.

horizontal dotted lines, marked 1, 2, 3, 4, 5, 6, and into sixteen vertical slices by vertical lines marked
a, b,c, &e. -

The corresponding lines in the diagram V.d are marked with corresponding figures and letters.
The stress across any line joining any two points in V.a is represented in magnitude by the line in V.2,
joining corresponding points, and is perpendicular to it in direction.

These illustrations of the application of the graphic method to cases of continuous stress, are
intended rather to show the mathematical meaning of the method, than as practical aids to the engineer.
In calculating the stresses in frames, the graphic method is really useful, and is Jess liable to accidental
errors than the method of trigonometrical calculation. In cases of continuous stress, however, the
symbolical method of calculation is still the best, although, as I have endeavoured to show in this
paper, analytical methods may be explained, illustrated, and extended by considerations derived from
the graphic method.

( 41 )

Il.—On Scientific Method in the Interpretation of Popular Myths, with special
reference to Greek Mythology. By Professor BLACKIE.

(Read 17th January 1870.)

Of all the branches of interesting and curious learning, there is none
which has been so systematically neglected in this country as mythology—
a subject closely connected both with theology and philosophy, and on
which those grand intellectual pioneers and architects, the Germans, have
expended such a vast amount of profitable and unprofitable labour. The
consequence of this neglect has been, that of the few British books we have
on the subject, the most noticeable are not free from the dear seduction of
favourite ideas, which possess the minds of the writers as by a juggling witch-
craft, and prevent them from looking on a rich and various subject with that -
many-sided sympathy and catholic receptiveness which it requires. In fact,
some of our most recent writers on this subject have not advanced a single
step, in respect of scientific method, beyond Jacos Bryant, unquestionably the
most learned and original speculator on mythology of the last century; but
whose great work, nevertheless, can only be compared to a grand chase in the
dark, with a few bright flashes of discovery, and happy gleams of suggestion by
the way. For these reasons, and to make a necessary protest against some
ingenious aberrations of Max MuLiEer, GLApsTone, Inman, and Cox in the
method of mythological interpretation, I have undertaken to read the present
paper ; which, if it possess only the negative virtue of warning people to be
sober-minded and cautious when entering on a path of so slippery inquiry,
cannot be deemed impertinent at the present moment.

For the sake of distinctness and compactness, I will state what I have to
say in a series of articulate propositions.

I. By the mythology of a people, I understand the general body of their
traditions, handing down from the earliest times the favourite national ideas and
memories, in a narrative form, calculated to delight the imagination and stimu-
late the affections of love and reverence.

II. The dress of all mythology, as appealing to the imagination, is neces-
sarily poetical; the contents of it are generally four fold—(1.) Theological ;
(2.) Physical; (3.) Historical; and (4.) Philosophical and Moral.

VOL. XXVI. PART I. it

42 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

ITI. In the theological and moral myth, the idea is the principal thing, the
narrative only the medium ; in the historical and physical myth, the fact is the
principal thing ; what goes beyond the fact is mere scenic decoration or imagi-
native exaggeration.*

TV. A myth intended to convey an idea is distinguished from an allegory or
parable by the consciousness of purpose with which allegories and parables
strictly so called are put forth and received.

V. As it has been well said of popular proverbs, that they are the wisdom
of many and the wit of one, so theological and moral myths grew up in the
popular imagination, and were nursed there till in happy season they received a
definite shape from some one representative man, whose inspiration led him to
express in a striking form what all felt to be true and all were willing to
believe.

VI. The first framers of myths were, no doubt, perfectly aware of the real
significance of these myths; but they were aware as poets, not as analysts.
It is not, therefore, necessary to suppose that in framing these legends they
proceeded with the full consciousness which belongs to the framers of fables,
allegories, and parables. A myth is always a gradual, half-conscious, half-
unconscious growth ; a parable is the conscious creation of the moment.

VII. During a certain early stage of national life, which cannot be accurately
defined, but which always precedes the creation of a regular written literature,
the popular myth—like a tree or a plant—becomes subject to a process of growth
and expansion, in the course of which it not only receives a rich embellishment,
but may be so transformed by the vivid action of a fertile imagination, and by
the ingrafting of new elements, that its original intention may be altogether
obscured and forgotten. How far this first significance may in after times be
rightly apprehended, depends partly on the degree of its original obviousness
partly on the amount of kindred culture possessed by the persons to whom it is
addressed. |

VIII. As of essentially popular origin and growth the myth cannot, in the
proper sense, be said to have been the creation of any poet, however distin-
guished. Much less could a popular minstrel, like Homer, using a highly
polished language, and who manifestly had many predecessors, be said to have

* Sometimes, however, a historical person, like Faust, may be seized on by the people, merely as
a convenient vehicle for embodying a floating mass of mythological notions. In this case the person
is really a secondary consideration : a real person he remains, no doubt; but, for a legendary nucleus,
any other person would have done as well.

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 43

either created the characters or invented the legends about the Greek gods,
which form what the critics of the last century used to call the machinery of
his poems. In regard to theological myths, which are most deeply rooted in the
popular faith, such a poet as Homer could only turn to the best account the
materials already existing, with here and there a little embellishment or expan-
sion, where there was no danger of contradicting any article of the received
imaginative creed.

IX. The two most powerful forces which act on the popular mind, when
engaged in the process of forming myths, are the physical forces of external
nature, and the more hidden, though fundamentally more awful powers of the
human will, intellect, and passions. It is to be presumed, therefore, that all
popular myths will contain imaginative representations of both these powers ;
and, in their original shape, they are in fact nothing more than the assertion of
the existence of these two great classes of forces in a form which speaks to the
imagination—that is, in the form of personality ; and there will be a natural pre-
sumption against the adopting of any system of mythological interpretation
which ignores entirely either the one or the other of these elements. If this
proposition be correct, the objections of Max MULiEr (Chips, ii. 156) to the
Greek derivation of "Epwvs, from the old Arcadian épwrvew (PAUSAN. vill. 25, 6),
are unfounded.

X. The most fertile soil for purely theological myths is polytheism ; and the
most obvious as well as the largest field for a religion of multiform person-
alities, is external nature. In the interpretation of such myths, therefore, we
shall be justified in searching primarily for the great forces and phenomena of
the physical world, as underlying the imaginative narrative and imparting to it
its true significance ; and in proportion to the prominence of these phenomena,
and the potency of these forces, will the probability be that we shall find them
fully represented in any body of polytheistic theology.

XI. As the essence of polytheism thus consists in the habitual elevation
of what we call physical facts and forces into divine personalities, the line
betwixt a purely physical myth and a theological myth will naturally be
extremely difficult to draw. Zeus, for instance, as the Thunderer, represents
a physical fact as well as a theological doctrine ; nevertheless, it would be
wrong to assume that there is no such element in tradition as a strictly physical
myth. Certain striking facts of volcanic action or geological change, strange
and grotesque shapes of rocks and other natural objects, unusual conforma-
tions of landscape, not to mention the occasional discovery of gigantic fossil
bones, and even entire skeletons of animals no longer existing, might well

a4 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

form the basis of what is properly termed a physical or a geological, rather than a
theological myth; and, as Hartrune well remarks (Gr. Myth. i. p. 168), notable
recurrent events in nature, such as the heavy rains at the end of summer, are
peculiarly calculated to impress the popular imagination, and to produce
myths.

XII. But as to man there is, after all, nothing more interesting and more
important than man, it is in the highest degree unreasonable, in the interpreta-
tion of myths, to proceed on the assumption that all myth is idea, and that no
myth contains any historical element. It may be true, no doubt, that in the case
of some particular nation, all action of the popular imagination on human per-
sonalities has been excluded ; but such a one-sided action is not to be presumed ;
it must be proved ; and that in such a rich and various mythology as the Greek
all reference to human characters and human exploits should be systematically
excluded is in the highest degree improbable. In a country where the gods
descended so easily into humanity, it were strange if men had not occasionally
ascended into godhood.

XIII. In a theology so thoroughly anthropomorphic as the Greek, the
distinction between the divine and human element will sometimes be difficult
to trace; for the same feelings, situations, and actions will necessarily belong to
human gods and to godlike men. But this state of the case, in the interpretation
of any particular myth, is a ground for doubt, not for dogmatism. It includes
the possibility or the probability of one or two explanations, but the certainty
of neither.

XIV. The incredible exaggerations or embellishments with which the name
of any national hero may have been handed down im a popular myth afford no
presumption against the genuine historical character of its nucleus. On the
contrary, it is Just because extraordinary characters have existed, that extraor-
dinary and incredible, miraculous and even impossible stories are invented about
them. <A plain, sober, critical, matter-of-fact account of its early popular heroes
is not to be expected from any people. .

XV. The error of certain ancient rationalising interpreters of the Greek
myths did not consist im presuming historical fact as the nucleus of some
myths, but in the indiscriminate application of the historical interpretation to
all myths, and that often in a very prosaic and altogether tasteless way.

XVI. The error of certain modern idealismg interpreters of the Greek
mythology does fot consist in endeavouring to recover the ideas which

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 45

originally lay at the root of some myths, the full significance of which had been
lost so early as Homer, but in the partial and one-sided application of a few
favourite ideas to all physical facts, and in the broad denial of any historical
elements underlying any personality of early tradition.

XVII. Among the ancients, the extreme of the rationalising interpretation
of the Greek theological myths is what may be called the irreligious, godless,
and altogether prosaic system of EKUHEMERUS (B.C. 300), who wrote a book to
prove that all the Greek gods, not even excepting Jove, had been originally
dead men deified. The error of this system consisted, not in the assertion that
the elevation of extraordinary human characters to a divine rank with religious
honour after death, is an element traceable in the Hellenic, as in some other
popular theologies, but in the wholesale declaration that religious worship had
no other origin, and that this element, which is always secondary and derivative
in the popular creed, is primitive and exclusive.

XVIII. In order to ascertain how far the principle of EUHEMERUS may apply
to any particular case, the general religious tendencies and habits of the nation
or people must be considered in the first place, and then the whole circum-
stances and features of the mythical narrative must be accurately surveyed and
carefully weighed, and a separation of the canonised man from the deified nature
element with which he may have been mixed up, made accordingly.

XIX. EvHemerus, however, was altogether wrong in supposing that this
system of interpretation could be applied on any extensive scale to the mythical
theology of the Greeks; and the few French and English writers who, in the
flatness of the last century, gave a limited currency to this idea, have found no
followers in the present.

XX. An opposite theory to that of EuHEMERuS, much in fashion with the
Germans, is that, whereas he said the gods were elevated men, we ought rather
to say that many men, perhaps all the heroes of legendary story, are degraded
gods. That in the course of religious development, especially when mixed up
with great changes in the political relations of different races, such a degrada-
tion may have taken place is certain ; that it has taken place in certain special
cases will be a just conclusion from an analysis of the character and worship of
certain heroes, when a cumulative view of the myths connected with them
suggests the theory of a divine rather than a human significance ; but there is
no scientific warrant for the assertion which it is now the fashion to make
(Barinc Goup, Rel. bel. vol. i. p. 167), that the old heroic names of a country,
as King Artuur, for instance, are in the mass to be treated as degraded gods.

VOL. XXVI. PART I. M

46 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

XXI. The best authorities for the facts of a myth are not always the poets,
nor even the most ancient poets, as Homer, who in the exercise of their art
often took large liberties with sacred tradition; but the reliable witnesses are
rather such as PAusantAs, who record the old temple lore in its fixed local
forms. This distinction, often forgotten, has given rise to not a little con-
fusion, and created some needless difficulty in mythological interpretation ;
and HartunG (i. 184) has done important service to comparative mythology by
drawing attention emphatically to the difference between sacred LEGENDS as
believed by the people, and religious mytus freely handled by the poets.

XXII. In the interpretation of any popular myth, the first thing to be done
is to ascertain carefully what the thing to be interpreted actually is ; and this
can only be done by collecting all the facts relating to it, working them up
into a complete, and if possible consistent picture, and not till then attempting
an explanation. Now, as the facts relating to any single god, let us say in the
Greek Pantheon, are scattered over a wide space, and come from various sources,
to attempt the explanation of these facts without the previous labour of critical
and well-digested scholarship, may be an ingenious amusement, but never can
be a scientific procedure. All the facts must be collected, and all the criticisms
weighed, before a verdict can be pronounced.

XXIII. But the mere collection of facts will never help a prosaic or an
irreverent man to the interpretation of what is essentially poetic and devout.
A book supplies what must be read; but the eye that reads it can see only
what by natural faculty and training it is fitted to see. As the loving and rever-
ential contemplation of nature was the original source of the polytheistic myths,
so the key to them will often be recovered by a kindred mind acting under influ-
ences similar to those which impressed the original framers of the myth ; and if
this may be done with a considerable amount of success by a poetical mind,
acted on by nature in any country, much more will such success be achieved by
such a mind in the country where the myths were originally formed. But as
the aspects of nature are various, and the fancies of poetic minds no less so, it
will always be necessary to verify any modern notion of an ancient deity, thus
acquired, by confronting it accurately and continuously with the traditional
materials contained in books and works of art. Highly poetical minds, such as
SHELLEY, Keats, and Ruskin, when dealing with Greek mythology, without the
constant correction of accurate scholarship, are not seldom found using Greek
myths to represent modern ideas, rather than human ideas to interpret Greek
myths. And the example of the Germans proves, that in minds naturally fertile
and ingenious, no amount of erudition affords a safeguard against the besetting
sin of mythological interpreters, to find in all myths a select field and enclosed

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 47

hunting ground for the pleasant disport of an unfettered imagination. Dis-
coveries are easy to make in a region where plausibility so readily gains currency
for proof.

XXIV. An important aid in the interpretation of myths will often be
supplied by the etymology of the names of the mythological personages ; and
in this way new deities will sometimes be found to have arisen from the mere
epithets of old ones, as Jacos Bryant saw clearly nearly 100 years ago; nay,
even magnificent myths may at times be traced to no more sublime origin than
a false etymology which had taken possession of the popular ear. The signi-
ficance of divine names must, of course, be sought in the first place in the
language to which the mythology belongs; but in applying this test, with
the view of obtaining any scientific result, great care must be taken to avoid
treating doubtful etymologies in the same way that certain ones may be treated.
For where the etymology is uncertain, that is, does not shine out plainly from
the face of the word (as in the case of the Harpies in Hxstop), then the elements
of doubt are often so many, that it is wiser to abstain altogether from this
aid, than to attempt founding any serious conclusions upon it. For, in the
first place, we may not have the word in its original form; and, in the second
place, two or three etymologies may be equally probable. The best etymologies,
whatever Bryant, and InMAN, and Max MULLER may say to the contrary,
are only accessories of scientific mythological interpretation.

XXV. If the mythological names have no significance in the language
to which they belong, then reference may be made to cognate languages ;
and in the case of European tongues, with propriety to the Eastern sources
from which they are demonstrably derived. But here a double caution is
necessary ; for accidental resemblances may be found in all languages, and
extensive learning, coupled with a vivid imagination, may readily supply the
most plausible foreign derivations, which are merely fanciful.

XXVI. By referring to another, and it may be a more primitive and ancient
language, for the etymological key to a religious myth of any people, we are
treading on historical ground extrinsic to the people with whose myths we may
be dealing. For comparative philology, like archeology, recovers the earliest
history of a people before writing was known; and this raises the inquiry,
whether a mythology which bears a foreign nomenclature on its face may not
convey foreign ideas in its soul—that is, to take an example, whether the
Greek mythology, if the names of its personages are more readily explained in
Hebrew or Sanscrit than in Greek, may not in respect of its ideas and legends
be more properly interpreted from original Hebrew or Sanscrit, than from native

48 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

Greek sources? And may we not hope, in this way, in the Hebrew Scriptures,
or the Sanscrit Vedas perhaps, to put our fingers on the ancient germs of
those anthropomorphic myths which Homer and Heston present to us in adult
completeness and full panoply? and thus the highest end of scientific research
will be obtained, not only to dissect the flower, but to trace it to the seed, and
follow it through every stage of its rich and beautiful metamorphosis.

XXVII. The prospect this holds out of tracing famous European religious
myths to their far home in the East is extremely inviting.* It satisfies at once
scientific minds by the promise of going to the root of a matter which has
hitherto been treated superficially, and that not inconsiderable class of literary
men and scholars who have a keener eye for an ingenious novelty, than for
a stable truth. When we bear in mind also the significance of the homely
proverb, that “far birds have fair feathers,” and the well-known fact, that
every mother is apt to prefer her own bairn to others which may be more healthy
and beautiful, we shall see reason to proceed, not without hope indeed, but with
more than Scottish caution, in this Oriental adventure. There is a class of
persons in the world who have a strange pleasure in travelling a thousand leagues
to quarry out a truth, which they might have picked up from beneath their nose.
Against these seductions therefore, in the first place, while prosecuting this
foreign chase, we must be on our guard. We ought to know that we are hunting
on very deceitful ground ; that we are dealing with a class of phenomena, that,
like clouds and kaleidoscopic figures, are very apt to change their shape, not
only by their own nature, but specially also according to the position of the
observer ; and that the same nebulous conglomerate may at one moment
look very like a whale, at another moment very like Lord Brovenam, and at a
third moment very like Olympian Jupiter. And in the prospect of such a
possible ridiculous conclusion to the sublime adventure on which he is starting,
every inquirer into the remote origin of European myths ought to take with
him these cautions—

(1.) That there is no necessity and no scientific warrant for seeking a foreign
explanation of deities, which already sufficiently explain themselves by the
character which they bear, or the symbols which they exhibit in their own
country.

(2.) That the formative power by which myths were created, viz., the imagina-
tion, possesses a wonderful magic, in virtue of which the materials on which it
acts, especially with a quick and vivid people unfettered by formal creeds, are
subjected to a perpetual process of transmutation, which renders the recogni-
tion of the original identity of two diverging myths an extremely difficult and

* “The whole theology of Greece was derived from the East.’”—Bryant, vol. i. p. 184.

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 49

not seldom an altogether hopeless task. In this respect the recognition of the
original identity of different words in cognate languages by comparative
philology is a much more safe and scientific process than a similar recognition
of the identity of different persons of two Pantheons through the shifting masks
of comparative mythology.

(3.) That the principal relations under which the great objects of nature,
such as the sky, the sun, the sea, &c., may appear, when subjected to the process
of imaginative impersonation, are in many cases so obvious that two different
polytheistic peoples may easily hit upon them without any historical connection.
Even in the free exercise of poetical talent in the case of individual poets of
highly potentiated imagination, we constantly stumble on comparisons which
have been made independently by other poets at other times or in distant
countries, and which superficial critics are sometimes eager to fasten on as
plagiarisms ; much more, in the vulgar exercise of the imagination, by the mass
of the people on certain given natural objects may we expect frequent instances
of coincidence without connection. This consideration will restrain a prudent
investigator in this department from building any theory of foreign origin of
myths on a few points of natural similarity.

Taking these cautions along with us, we now observe, in reference to the
probable Eastern origin of certain Greek myths—

XXVIII. That the borrowing of one nation from another in the province of
mythological ideas, as in the case of philological materials, may take place in a
twofold fashion, either in the way of original descent from a common stock, far
back in the cradle of the race, or by importation through the medium of com-
merce or great religious revolutions and invasions. Of these two methods of
borrowing, it is impossible to say, a priori, which promises the greater amount
of gain to the adventurous inquirer ; for, while the advantage of greater close-
ness belonging to the original identity of stock may be in a great measure
neutralised by the distance of time and place, and the changes which they
induce, the disadvantage of a more loose connection which belongs to the foreign
importer may be amply compensated by the firm hold which the commerce, and
polity, and intelligence of a superior people may take of an inferior people.

X XIX. It must be borne in mind, also, that the recognition of a supposed
identity between the gods of any two polytheistic peoples may easily take place
without any real borrowing. For the desire of harmonising and classifying dis-
cordant phenomena, which belongs to the very nature of intellectual action, is
particularly displayed in the field of popular religion—to such an extent, indeed,
that it became a fixed habit of the Greek and Roman mind to identify the
deities of foreign countries with their own native deities by certain signs more

VOL. XXVI. PART I. N

50 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

or less superficial. The testimony of the Greeks, therefore, with regard to the
supposed identity of certain personages in their Pantheon with certain gods or
goddesses in the Egyptian or Phoenician, and their consequent foreign extrac-
tion, will require to be examined with the severest scrutiny.

XXX. In deriving any god from a foreign source, even though his foreign
origin should appear in some respects perfectly certain, we must not conclude
that all the phenomena which his person and character present are to be
explained from abroad. Nothing is more natural than that he should be
a compound god, one half native and one half foreign, or even a monstrous
conglomerate of many gods.

XXXI. Of all the foreign sources to which the Hellenic mythology has at
different times been referred by the learned, Egypt is at once the most reputable
and the least likely. For here we have neither original connection by identity
of stock, nor any such commercial or political action of the more ancient over |
the more modern people, as would lead to the importation of religious ideas.
The ancient Greeks had a great respect, and a sort of awful reverence for the
wisdom and the antiquity of the Egyptians ; but this respect and reverence was
more likely to lead them, as in fact it often did, to the recognition of super-
ficial resemblances (as in the case of Io and Isis), than to the trace of original
identity. Modern researches have added nothing to the probability of the
favourite notion of Bryant and BLACKWELL, that the principal persons and
legends of Hellenic mythology came directly from the land of Ham.

XXXII. For the Hebrew origin of some of the Greek theological ideas—the
darling notion of Church Fathers and Protestant theologians, and which has
been recently revived by a statesman of distinguished character, talent, and
erudition—there is even less to be said. For, in the first place, here we
are comparing a polytheistic system with a monotheistic, where antagonism
rather than similarity is to be looked for; the elements of original or super-
induced connection between the two peoples are altogether wanting; and the
original unity of the human family, which is the only link that binds the Greek
to the Jew, is so remote that it requires no inconsiderable amount of hardihood
to drag them into the arena of the present comparison. This hardihood, how-
ever, has never been wanting; and besides its own virtue, has always found
great favour with the religious public, which is pleased with nothing so much
as the idea that everything good, beautiful, or excellent in any way that
heathen religions may be allowed to possess must have come either from the
Hebrew Scriptures directly, or from some more ancient source of primeval
revelation. And no doubt there may be a certain truth in this view; but it is

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 51

a truth which affects the monotheistic element, that in the person of Zeus lies
the background of the Hellenic polytheism, rather than the polytheistic per-
sonages to whom it hasbeen applied. A consciousness of this, no doubt, led the
early mythological interpreters of this school to apply the principle of EUHEMERUS
largely to the Old Testament, in such a way only as to recognise the venerable
Hebrew patriarchs under various masks of old Pelasgic gods or demigods.

XXXIII. For the Pheenician influence on the formation of the early Greek
theology there is much more to be said. We can, indeed, scarcely imagine a
race of such distinguished merchants and navigators, commanding the Greek
seas in the early ages of European civilisation, without supposing some such
contagion and ingrafting of religious ideas, as the genius of polytheism was on
all occasions prone to invite. We shall, therefore, be disposed to receive
favourably any distinct proof, or even probable indication, of the derivation of
Greek gods from a Phoenician source ; but we must bear in mind at the same
time, that the Phoenicians were known to the Greeks as mere traders, with
temporary settlements on the coast of the Mediterranean, and that their
character, as exhibited in the Odyssey, was by no means possessed of such
attractions as might aid to allure the Greeks to the adoption of any of their
peculiar objects of worship.

XXXIV. The last source of Greek myths, for which a strong claim has
recently been put forth by a German of distinguished talent, taste, and
learning in this country, is Sanscrit. And here at last some people seem to
think, that with all certainty we have got at the true source of the many-winding
mythological Nile. But after looking into this matter with all possible care,
and with no prejudice whatever (for nothing would please me so much as to
catch the infant Mercury in the bosom of a cloud, floating over the shining
peak of the Hindoo Koosh, or to hook Proteus in one of his many forms at the
mouth of the Ganges), I must honestly confess, that hitherto the interpreters
of Hellenic myths from Sanscrit roots and Vedic similes have inspired me
rather with distrust than with confidence. The principal characters of the
Hellenic Pantheon tell their own story, to a poetical eye, more obviously and
effectively than with the help of a Sanscrit root ; and those few of them which
are more doubtful, such as Hermes and Athena, seem to be precisely those in
which the Sanscritizing mythologers have most egregiously failed. I consider,
therefore, that, while the Vedic mythology, preferably to any other polytheistic
system, presents an ample field from which some of the Hellenic legends may
be aptly illustrated, and a few, perhaps, correctly interpreted, the attempt to
explain the great and prominent phenomena of the Greek Pantheon, by an
ingenious application of a few favourite physical ideas variously impersonated

52 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

by the fancy of the Vedic poets, must be regarded in the meantime, at least, as
a failure.*

XXXV. Without, therefore, in the slightest degree wishing to throw
discouragement on the delightful and interesting study of comparative mytho-
logy,—a study that promises the most fruitful results in the domain of theology
and moral philosophy,—the procedure of exact science seems to demand that,
before venturing on extensive excursions into foreign regions, we should, in
the first place, carefully survey and exhaust our home domain—that is to say,
that the Greek traditions with respect to their gods, interpreted by themselves,
and the general principles of mythical interpretation laid down in the above
propositions, afford a surer basis for this branch of mythological science than
hints suggested by Oriental etymologies, or analogies from the Vedic hymns.
And in order to make this more clear, I will select a few examples of person
ages from the motley theatre of Hellenic legend, which may be best adapted
for testing the value of the different methods of interpretation.

XXXVI. As examples of how the elemental significance of the Hellenic
gods reveals itself to a sympathetic eye, from the mere presentation, epithets,
attitudes, and badges of the mythologic personages, we need do no more than
mention Zeus, Poseidon, and Apollo, in whom all the ancients, who exercised
reflection at all on the matter, recognised, with one voice and by an unerring
instinct, the great elemental powers of the sky, the sea, and the sun. And
these are precisely the powers which, from their prominence, might @ priori have
been predicated as certain to obtain a conspicuous place in an anthropomorphic
Pantheon of elemental origin. Of these three great gods also, be it noted, that
the first is the only one of which we can trace the etymology with any certainty ;
but neither does this one etymology, when recognised in the Sanscrit word
Diva, to shine, add anything to the already recognised idea of the Hellenic
Zeus, nor does the lack of an etymon in the other two cases render our percep-
tion of the character of the two gods less clear, or our knowledge of their
significance more certain. With regard to Poseidon, Mr GLApsTone’s recent
attempt to fix on him a Pheenician pedigree must be regarded as unsuccessful.
The people who at an early period sailed to Colchis and to Troy, did not
require to borrow a lord of the flood from the merchants of Tyre and Sidon.

XXXVII. In Hera who, to the people and the people’s poet, was simply.
the spouse of Zeus, a large class of ancient speculators, as is well known, were

* It may be proper to state, that the interpretation of certain personages in the Greek Pantheon
from sources of Sanscrit etymology, to which Max Mutxumr has given currency, is not at all con-
firmed by the judicious sobriety of our countryman Dr Muir. See his paper in our Transactions,
vol. xxi. p. 078.

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 53

inclined to recognise the lower region of the atmosphere, of which Zeus repre-
sented the ai?yp, or upperregion. But a little consideration has convinced most
modern interpreters that this idea was a mistake. When by the completion of
the anthropomorphic process, the original ovpavés had become “ Father Jove,”
it was most natural that his elemental counterpart 7, Mother Earth, should
become the matron Hera; and with this supposition, the well-known description
of the sacred marriage of Zeus and Hera (Il. xiv. 345), together with the
cow-symbolism belonging to the Booms, and her Argive priestess Io, notably
harmonise. It is no objection to this view, that Ceres or Demeter is also the
anthropomorphised earth; for “the many names of one shape” (modhav
dvouatav jopdy pia), characteristic of the oldest elemental theology, could easily,
and did often crystallise into two or more shapes of one power. We shall,
therefore, say with no rash confidence, that the Hellenic Hera means the earth;
and we readily allow the etymological conjectures connected with her name
to remain conjectures.

XXXVIII. On Athena, Max MUuuEr says, “ The Sanscrit root An, which
in Greek would regularly appear as Acu, might likewise then have assumed
the form of ArH; and the termination Enz, is Sanscrit Ana” (“Science of
Language,” vol. ii. p. 503) ; and again, “ How Athena being the Dawn, should
have become the goddess of wisdom, we can best learn from the Vedas. In
Sanscrit, Budh means to wake and to know” (Do. p. 504).

But this is manifestly following out a favourite idea upon theories of the
most flimsy texture. If any etymology is to be sought for the syllable AO, the
native root ai# which signifies to glow, corresponding as it does with the familiar
epithet of yhavxems, or “ flashing-eyed” (which I think WELcKER suggests), is
preferable to that suggested by the distinguished Sanscrit scholar. But here,
as in other slippery cases, the principles laid down in the preceding propositions
lead me to set etymology aside, and to look at the finished figure of the goddess,
with her badges, relations, and actions, as the natural and sure index to her
significance. Now if Zeus, according to the Greek conception, was the strong,
stormy, and thunderous element of the sky—as his epithets xedawedyjs, and
epiBpeuerns, and reprixépavvos, sufficiently declare—his flashing-eyed daughter,
who alone is privileged to wield his thunderbolt (A®scuyi. Eumen., 814), must be
some action or function of the sky. Let her, therefore, be the flashing lightning,
or the bright rifted azure sky between the dark rolling thunder clouds, or both
if you please, and you have at once an elemental theory which explains
adequately her anthropomorphic parentage and presentation. As to her moral
and mental significance, that follows necessarily from her Jovian fatherhood.
When the all-powerful was recognised as at the same time the all-wise, and the
great counsellor (uyriera Zevs), his daughter, as a matter of course, became the

VOL. XXVI. PART I. 0)

54 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

goddess of practical wisdom, that is, of the great arts of peace and labour (as
the vases largely show), and the patron and protector of all men of valour
like Achilles, and of sagacity like Ulysses.

XXXIX. The Hellenic Hermes is one of those mythological personages
who from an originally simple root, has grown up into such a rich display of
graceful ramification, that, when we approach him from his most familiar side
we are the least likely to interpret his true significance. But if we attend to
the earliest indication of his functions as found in Homer, and as displayed in
the familiar phallic symbol (Heron. ii. 51), we can have no difficulty in evolving,
by a series of graduated expansions, his final avatar as a god of eloquence, from
his original germ as a pastoral god of generation and increase (Hom. II. xiy.
491). As the god of shepherds and mountaineers, he was necessarily the
guide of all wanderers through the many winding glens, and across the many-
folded hills of the Arcadian Highlands. This early function accordingly appears
in Homer: he is the friendly guide of all persons who have lost their way or
who wander in the dark (Od. x. 277; Il. xxiv. 334). His connection with
music and with wrestling, the natural recreations of a pastoral people, of course
belong to this his earliest Hellenic character. Afterwards, when in the
necessary progress of society, the patriarchal shepherd of the hills resigned
his social position into the hands of the rich merchant of the great towns,
Hermes became the god of gain generally ; and, with gain, of all those arts of
adroitness and sharpness which belong to the career of a successful trader.
The kindly guide of night-wandering shepherds has now become the expert
negotiator, and the trusty messenger; he is the winged servant of the gods
above ; and among men his oaten pipe is exchanged for the charm of winged
words, which sway the counsels of the wise, and soothe the clamours of the
turbulent. With this natural and obvious interpretation of a purely Hellenic
deity, as given within the bounds of Greece itself, we shall raise only a brilliant
confusion, if we follow Max MULLER across the Hindoo Coosh, and ingeniously
attempt to find the germ of the Pelasgic shepherd god in the breeze of the
early dawn, which ushers in the march of the busy day. Such remote
conjectures may be both beautiful and ingenious, but they are a mere play of
fancy, and travel obviously far out of the way of a sober, a scientific, and
a stable interpretation.

XL. Dionysus was a god of comparatively recent introduction into Greece
(Heron. i. 49), confessedly of Asiatic origin, and in whom the union of fervid
wine with the phallic symbol and violent orgies, can leave no doubt as
to his true character. He is the male god of generation, according to the
Asiatic conception, as the Syrian goddess of Luctan (De Dea Syria, 16) was

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. ys)

the female one ; and the old Heraclitan principle that fire is the origin of all
things, rudely conceived by the popular imagination, is manifestly that which
in this god identifies the fervour of the vine juice, the brewst of the sun, with
the fervour of the generative process. The fact that the worship of Dionysus
was not native in Greece, but travelled from the East, naturally led to the
representation of this god as a wonderful conqueror, in the fashion of SEsosTRIS
and ALEXANDER the Great; from which analogy, coupled with his preaching
the gospel of wine, Bryanr and other speculators have been eager to find in
him a perverted Noau; but the application of the principle of EUHEMERUS in
this case evidently rests on too slender a foundation to afford any grounds for a
scientific interpretation.

_ XLI. Aphrodite is that goddess in whose case Mr GLapstone’s favourite
idea of Phoenician influence on the Greek Pantheon has long been recognised
as the most certain (HErRop. i. 105; Pausan. i. 14, 6). The recognition of this
Pheenician element, however, does by no means imply that the existence of an
original Hellenic impersonation of the passion of love, and the seductions of
personal beauty, should be denied. On the contrary, the female deity whom
the Phoenicians were seen worshipping in their factories on the coasts of the
Mediterranean, would most probably be accepted by the ancient Pelasgic tribes
chiefly because they found in her attributes a striking identity with their own
native Aphrodite.

XLII. Pheenician influence is also undoubtedly to be acknowledged in the
very complex and composite mythology connected with the name of Heracles.
But the person of Heracles, as we find him in Homer, exhibits nothing beyond
the exaggerated traits of a stout and muscular humanity in combat with fate
and circumstance, and the wild beasts of the forest—a plain Hellenic counter-
part, in fact, to the Hebrew Samson, of whose historical reality, to a mind not
violently possessed by German theories, there cannot be the slightest reason
to doubt. The exaggerations connected with his story are the natural and
necessary effects of the excited popular imagination brought to bear on such
a character; but these exaggerations, taken at their highest, are exhibited
on a very small platform in Homer, and present a very modest array of achieve-
ments compared with the multiform mass of myth that afterwards accumulated
round this representative Greek hero. The principle of growth, of such
luxuriant vitality in popular myths, has been obviously at work here; and the
sort of omnipresence latterly attributed to this wandering queller of monsters
is most readily explained from the influence of the Phcenician factories in the
Mediterranean, in whose Melcarth the Greeks delighted to recognise their
own stout son of Jove and Semele. And if this Tyrian Hercules, as Phoenician

56 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS,

scholars incline to believe (MoEVERS, vol. i. p. 385), really was a sun god, the
twelve labours of Hercules will, of course, only be the symbolical expression
for the progress of the Titan sun through the twelve months of the solar year.
This the ancients themselves, in the Orphic theology, at least distinctly recognised.

XLIV. In Bellerophon the Germans find a favourite example of their
theory, that all the heroes of the so-called heroic age are the degraded gods of
an early elemental worship. How this theory is worked out in the present case
it may be instructive to consider. The winged steed, of course, brings you at
once into the region of the sun. Then you turn up Eustaruius’ commentary on
the well-known episode of the Corinthian hero in the sixth book of the Iliad
(v. 181), and you find there that there was an old Greek word €hdepos, used by
CALLIMACHUS, which is equivalent to xaxdés or bad; but bad things are black
things; therefore, with the help of the digamma, transmuting €)\¢pos into
BedXepos, we arrive at the conclusion that BeddepoddvTns means the slayer of
darkness, and, of course, can be nothing but the light, or the sun. Bellerophon
is thus, by a dexterous etymological feat, already a solar god in full panoply ;
and when, in addition to this, we find that the worship of the sun was much
practised at Corinth, the native place of the hero, and that he died in Lycia, a
country famous for its devotion to the same deity, the case for a degraded
“HXuos seems to be satisfactorily made out. But, on the other hand, the oldest
version of the story in Homer has no hint of the winged horse; and for the
rest, looks in every trait as much like a purely human history of those early
Greek times as the story of Str Cotumsa shows like a real legend of a real
Catholic apostle in early Christian times. We shall, therefore, in my opinion,
more wisely say that the airy flight of the grandson of Corinthian Sisyphus on
his winged Pegasus, is only the imaginative painting out of a real human journey
made from such real and natural causes as those which Homer details ; and, if
the winged horse has anything to do with the worship of the sun at Corinth, it
is more reasonable to suppose that such a blazon should have been added for
the glorification of a real great man, than that all the great men of early Corinth
should have been clean swept from the popular memory to make way for an
unmeaning Pantheon of degraded and forgotten gods.

XLV. Descending lower down into the region of what has the aspect, not
of metamorphic theology, but of plain human fact, we may take the names of
Achilles and Theseus as examples of how far the German school is inclined
to carry its peculiar tactics of finding nothing in all early tradition but theolo-
gical ideas and symbols. As to Achilles, the favourite notion with most German
writers is that this hero is a water god,—a notion founded on nothing that I can
see, save on the etymological analogy of Achelous, the happy coimcidence of

WITH SPECIAL REFERENCE TO GREEK MYTHOLOGY. 57

Peleus with the Greek name for mud (7y)és), and the fact that the mother of
the hero was a sea-goddess ; and on this notion ForcHHAMMER, I believe, or some
one of the erudite fancymongers beyond the Rhine, constructed a theory that
the Iliad is really a great geological poem, in which water power is represented
by Achilles, and land power by Hector (from eyo, to hold, restrain, keep back) !
This is really too bad. If a man in Thurso, to take a modern example, named
WaATERS—and it is a characteristic name in that quarter, were to marry a woman
called Loco—a well-known name in Sutherland—and a daughter, the offspring
of this marriage, should join herself in wedlock to an English gentleman named
RIVERS, no sane person could see in this conjunction of congruous etymologies
anything but one of those curious coincidences which amuse a newspaper reader
for a minute, and then are forgotten. Why, then, we ask, should the occurrence
of water, and mud, and a sea-nymph, among the family names of an old Thessa-
lian throne, be supposed to possess any more profound significance, even on the
supposition that the etymologies are certain, which they certainly are not ?
And accordingly, we find this favourite water theory discarded by the Germans
themselves, the moment it does not suit the theory of the interpreter. To Max
MU.uerR Achilles can be nothing but a solar god; for his imagination, fired
with sunlight from the flaming east, can see nothing in the stout battles of
Greeks and Trojans in the Iliad but the grand struggle between the powers of
light and darkness. Of the probability of this theory I have sought in vain for
the shadow of a proof. If Helen of Troy, whose name can obviously be identi-
fied with brightness (oéAas oedjvy), must on this account take her place with
her brothers, as a sidereal phenomenon (sic Fratres Helen, LUCIDA SIDERA), this
does seem to me an exceeding weak foundation for the transformation of the
whole topographical and traditional heroes of the Iliad into a meteoric
spectacle.

If, according to the views set forth in this paper, there is no scientific
ground for raising Achilles into the category of gods, whether aquarian or
solar, much stronger are the reasons which induce us, with unsophisticated
old PLuTARcH, to see in Theseus no myth, but a great historical reality. Ifthe
principle be once accepted, that a single miraculous fact or incredible story con-
nected in the popular imagination with a great popular name, shall deprive him
simpliciter of all claim to a historical existence, we shall make strange havoc, I
fear, with some of the most brilliant and the most instructive pages of national
record. There is no need of believing all the wonderful stories that Athenian
reverence and wonder accumulated round the name of Theseus, as little as there
is of believing all the silly miracles that the Lausiac history narrates of the
Egyptian ascetics; but there is certainly as little wisdom in roundly denying
the historical germ to which, in all such cases, these accretions were attached.

I have thus pointed out, in a rapid and succinct way, what seem to be the

VOL. XXVI. PART I. P

58 PROFESSOR BLACKIE ON INTERPRETATION OF POPULAR MYTHS.

leading principles on which a sound and safe interpretation of early popular
myths must proceed. I have kept myself purposely within the bounds of what
appears to me sober statement, not being eager for the glory of adventure in
this nebulous field; and if I shall seem to have achieved a very small thing when
I keep myself within these bounds, I have at least kept myself clear of non-
sense, which in mythological science is as common as sunk rocks in the Shetland
seas. To Max Mutter, and other Sanscrit scholars, I hope I shall always be
grateful for any happy illustrations which they may supply of the general
character of Aryan myths, and of occasional coincidences of the Hellenic mode
of imagining with the Indian ; and I think the somewhat cold and unimaginative
race of English scholars are under no small obligations to him for having taught
them to recognise poetical significance and religious value in some legends,
which passed in their nomenclature for silly fables or worthless facts ; but I
profess to have been unable to derive any sure clue from the far East to the
most difficult questions of Greek mythology ; nor do I expect that, when every
obsolete word in the Rigveda shall have been thoroughly sifted and shaken,
a single ray of intelligible light will thence flow on the Athena of the Parthenon,
or the Hermes of the Cyllenian slopes. I believe that in the region of mytho-
logy they will ultimately be found to be the wisest, who are at present content
to know the least; that while some mythological fables are too trifling to
deserve interpretation, others are too tangled to admit of it ; and that the man
who, at the present day, shall attempt to interpret the Greek gods from the
transliteration of Sanscrit or Hebrew words, will be found, like Ixion, to have
embraced a cloud for a goddess, and to have fathered a magnificent lie from
the fruitful womb of his own conceit. There is no more dangerous passion than
that which an ingenious mind conceives for the fine fancies which it begets.

(EPI)

III.—On the Extension of Brouncker’s Method to the Comparison of Several
Magmtudes. By Epwarp Sane, Esq.

(Read 7th February 1870.)

The discovery of those numbers which shall, either truly or approximately,
represent the ratio of two magnitudes, necessarily attracted the attention of the
earliest cultivators of exact science. The definition of the equality of ratios
given in Evcuip’s compilation clearly exposes the nature of the process used in
his time. This process consisted in repeating each of the two magnitudes until
some multiple of the one agreed perfectly or nearly with a multiple of the
other ; the numbers of the repetitions, taken in inverse order, represented the
ratio. Thus, if the proposed magnitudes were two straight lines, Euctip would
have opened two pairs of compasses, one to each distance, and, beginning at
some point in an indefinite straight line, he would step the two distances along,
bringing up that which lagged behind, until he obtained an exact or a close
coincidence.

He seems to have assumed that, in the case of incommensurable magnitudes,
the further continuation of the process must give still closer approximations ;
but we do not find any indication of a knowledge of the fact that, in the course
of that continuation, we shall certainly come upon coincidences still more close
than any which we have already obtained.

This process for finding the numerical expression for a ratio is Inconvenient
from its bulkiness ; it is also unnatural, for the mind, in comparing two unequal
magnitudes, is rather inclined to regard them as made up each of so many
measures, than to consider how many times the one must be augmented in order
that the result may be a multiple of the other; it prefers the direct to the
inverse comparison.

Lord BrounckeEr’s method of continued fractions enables us with great
rapidity and within the compass of the magnitudes themselves, to determine
directly their ratio. It is one of the great landmarks in the progress of the
science of numbers.

By one or two slight improvements in the mode of calculation, the chain or
continued fraction became a ready tool in the hands of arithmeticians. It placed
in a clear light the whole doctrine of indeterminate equations of the first degree,
leaving scarcely anything further to be desired in this branch of the Diophantine
analysis.

VOL. XXVI. PART I. Q

60 MR EDWARD SANG ON THE EXTENSION OF BROUNCKER’S METHOD

On applying BrounckeEr’s method to two incommensurable quantities of the
second degree, it was found that the denominators eventually came to be
repeated or circulated indefinitely; and LAGRANGE showed that while every cir-
culating chain-fraction was known to represent the root of a quadratic equation,
the roots of all such equations were developable in such a fraction. Hence the
conclusion was drawn that, the root of no equation of a higher order can possibly
be represented by a circulating chain-fraction.

Although, to the mind of BrounckeEr, the continued fraction presented the
readiest way of expounding his idea, it is not essential thereto ; a much clearer
view of the true nature of the process may be obtained without it. The opera-
tion consists, essentially, in deducting, as often as possible, the less from the
greater ; the remainder again from the preceding subtrahend, and so on; in
keeping note of the numbers of the subtractions ; and in computing from these
numbers the value of the magnitudes in terms of the ultimate subtrahend. The
chain-fraction is merely one way of representing the final computation. By
stopping at the first denominator, then at the second, afterwards at the third,
and so on, we obtain a series of fractions alternately too great and too small,
but approaching rapidly to each other and to the true expression for the ratio.
Now this series may be deduced directly from the equations representing the
various subtractions ; wherefore, in our subsequent investigations, we may put
the idea of the chain-fraction entirely aside, without thereby changing the in-
trinsic character of the Brounckerian process.

On examining the two series converging to the two roots of a quadratic
equation, I observed that the circulating quotients are the same for both, but that
their order is inverted. This observation led me to a singular law, which some
years ago I submitted to the Society. It is this, that if we continue the forma-
tion of the series for one root beyond the non-circulating quotients, obliterate
these and the fractions adjoming them, and then, using only the circulators,
compute the series backwards, we shall obtain the other root of the equation;
so that both roots are given by a, so to speak, two-headed progression.

The periodical recurrence of the quotients enables us to approximate as
closely as may be desired to the roots of equations of the second order, with
very little labour; and a kind of regret accompanied the conviction that the
same facilities cannot be obtained for equations of higher degrees. On con-
sidering the arguments on which this conviction rested, it appeared to me that
the whole circumstances of the case had not been taken into account; one, and
a most influential one, had been concealed under the notation employed, that is,
under the scheme of continued fractions. If we assume any two fractions to
take the place of two contiguous terms in a Brounckerian progression, and
operate upon these in the usual way, that is, by adding to a multiple of each
member of the second fraction the corresponding member of the preceding ; and

TO THE COMPARISON OF SEVERAL MAGNITUDES. 61

if we continue this operation, using always the same multiplier, or a circulating
set of multipliers, the fractions so resulting converge to the root of a quadratic
equation. If we should assume ¢iee fractions, and combine fixed multiples of
their members, so as to form a progression of the third order, as we may call it,
to what value do the terms of this progression converge ?

In a paper read by me some time ago to the Society, it was shown that the
convergence in this case is toward the root of a cubic equation; and that the
same arrangement may be extended to the still higher orders ; as examples of -
the application of this method, two cases may be cited.

. : ft pep A :
If we begin with the two fractions 9»; and form a progression by adding

to the double of each member of the last, the corresponding member of the
preceding, we form the well-known progression

which converges toward the ratio of the diagonal to the side of a square.
If, beginning with the same pair, we form a progression by taking the sums
of the members of the last and of the penult, we obtain
11,2
OF eed?
which converges toward the ratio of the diagonal of a regular pentagon to its
side. In this case, the numerator of one fraction becomes the denominator of
the succeeding, so that it is unnecessary to write both progressions. These
are familiar examples of quadratic roots.
Let us now assume ‘three terms, 0, 0, 1, and continue a progression by
adding to the double of the last term the difference between the two previous
ones, thus—

0, 0, 1, 2, 5, 11, 25, 56, 126, 283, 636, 1429, 3211, 7215, 16212, 36428,
81853, 188922, &c.,

and we obtain an approximation to the ratio of the long diagonal to the side of
a regular heptagon. Thus, if the side of the heptagon be 283, its longest
diagonal is almost exactly 636.

Or again if, assuming the same three terms 0, 0,1, we form a series by
deducting the antepenult from the triple of the last term, thus—

0, 0, 1, 3, 9, 26, 75, 216, 622, 1791, 5157, 14849, 42756, 123111, 354484, &c.,

we obtain an approximation to the ratio of the long diagonal of an enneagon to
its side.
I have shown that, in progressions of this kind, that is, where the numerator

62 MR EDWARD SANG ON THE EXTENSION OF BROUNCKER’S METHOD

of the one fraction becomes the denominator of the other, the approximation is
toward that root of the equation which is farthest from zero; and that if the
progression be carried backwards, the approximation is then toward the root
nearest to zero.

These remarks may suffice to show that this branch of the theory of numbers
promises to yield important results. Now, the whole doctrine of quadratic
recurrence sprung from the comparison of two magnitudes ; and so the com-
parison of three magnitudes must be the true foundation on which to build
the doctrine of cubic recurrence. I propose, therefore, in the present paper, to
discuss the elementary operation by which the ratios of three incommensurable
magnitudes may be approximately ascertained.

Let there be three homogeneous quantities, A, B, C, arranged in the order
of their magnitudes, and let it be proposed, if possible, to find their common
measure.

By repeatedly subtracting the second B from the greatest A, we obtain a
remainder less than B; this remainder may or may not be greater than C; if it
be greater, we may take C from it until we obtain a remainder D less than the
least of the three proposed quantities. In this way we have an equation of
the form

A=p,B+4q4,0C+D,

in which p cannot be zero, while g may.
Treating now the three quantities B, C, D, in the same way we have a new
equation

B=p,C+¢g,D+E,

and we may proceed in this way until there be no remainder, or until the
remainder be so small as to pass the limits of exactitude demanded by the
nature of the case in hand.

We are now able, by means of successive substitutions, to obtain values of
A, B, C, in terms of the ultimate remainders ; and our first business is to devise
some convenient arrangement for this purpose.

For the sake of giving greater generality to our investigations, let us put
the successive equations in the form

A= pRB gC Sr, Dw,
Bp, 0 Pg Per hs
C=p,D+q,E+7, F,

Or 7, RE Go 4 ta

in which 7,,7,, . . . . have been written for the unit of the usual process.

TO THE COMPARISON OF SEVERAL MAGNITUDES. 63
By successive substitutions we arrive ata value of A, in terms of R, S, T,
which may be conveniently represented by the equation,
A=@n.R+Mm.S+wm.T,

and our business becomes to discover the law of formation of the functions ¢n,
On, Wn.
Continuing the operation one step further, we have
‘ fee eos Oni Le a1. Ux;
and substituting this value for R in the preceding equation,
A = {Pno1- 92 + Onk St {Gn4i. 90 + yn} T 4+ 7441. 9nU,

wherefore the law of successive formation is contained in the three equations—

g(n +1) = Poyi1- 92 + On, : ; (ty;
O(n + 1) = GQr4i-9n + Wn, ; (2),
win + 1) = 7r,¢1- On, . (3).

Now these forms hold good for every value of n, wherefore
yn =7,.9(n — 1) ,
and consequently
O(n +1) = G41. 9n+7,.9¢(n —1) ,
whence
On = qn. ¢(n —1) +7,-1.9(n — 2) ,
so that the equations (1), (2), (3) become

g(m + 1) = Pri. 9M + G,.G(m — 1) + 7,_1. O(n — 2), (1),
O(n +1) =drai- PP +7. G(n —1),~ . ; ; : (2),
Win - L) = Tar. 90, . ; (3).

By means of these formule we can construct the series of functions ¢n
independently of the others, and thence we can readily deduce the progression
6n; as for the third progression wn it is, in the usual case of 7 = 1, a mere
transcript of the progression ¢2 moved one step back. The arrangement of the
work is very simple, and may be best studied from a numerical example.

The arrangement of the intervals in music has to follow the natural sub-
division of a vibrating column, and so must be made, primarily, to suit the
ratios 1:2, 1:3, 1:5, and their compounds. If, then, it be proposed to tune a
musical instrument so as to permit of transposition from one key to another, the
ratio represented by the smallest interval on it must be contained exactly, or
very nearly, in each of these three ratios. Therefore the arrangement of the
gamut on an instrument of equable temperament must be obtained by a com-

VOL. XXVI. PART I. R

64 MR EDWARD SANG ON THE EXTENSION OF BROUNCKER’S METHOD

parison of the logarithms of the three prime numbers 5, 3, and 2. These
logarithms are incommensurable, and so it is impossible to tune a keyed instru-
ment perfectly. The comparison of these three logarithms furnishes a con-
venient instance of the application of our method.

Putting, for shortness’ sake, A = log 5, B = log 3, C = log 2, we obtain the
following equations :—

69897 00048 = A= 1B+0C04D
ATID NAIA, = B= CAE a ee
30102. 99956 = C.= 1.D'+ OE +F
22184. 8/496 = D= LE + 04 256
17609 _12590 = E= 2F +0G+H
798) 12460 = =] Gs ee Se
4576 74905 = G =. 20-0 ioe ie
1772: 87669, = i=. 1 Oe
1569 4968S = J. = AK 2
1029. 99566 = Ki = 5:14 0M. +N
203 37784 = L = 1M+5.N+P

132 74750 = M= 10.N+0P+Q
13 10644=N= 2P4+1Q+R

5 09810 = P
1 68303 = Q
1 22790=R;

and, in order to compute from these the successive approximations, we may
write the three sets of coefficients, p, g, 7, in three horizontal lines, as in the
subjoined scheme :—
Mie Gian as Wi I Ves RS MR er
0.0 04,0 2 D0 eee Oe 0
tS es ee eee insta ES ile eee)

1) of 2. & 6 19 24 630, 217 6 (24T Sb2b 2b,

r
q
P
Ae dy i Ae 87 9 (28 Vso a4 1S “353 S16t es8
B
C tt 1 8P 4 al2, plo, 19s WlS7,. 152) 2004 itash

Having written wnit beneath the first p, to serve as the beginning of the
series A, we multiply that unit by p, to get 1, the second value of A,, which
value we write beneath p,. We now take the products A, p,, A, ¢,, the sum of
which gives us A, = 1. Thereafter we take the sum of A, p,, A, g,, A, 7,, to
obtain A, = 2. In this example the first five g’s happen to be zeroes, and so
the middle terms of the expressions are awanting; the middle term is first
found in the expression for A,, whichis A,.p, + A,.g, + A,.7, =18+7+ 3;

TO THE COMPARISON OF SEVERAL MAGNITUDES. 65

and again we have it in the value of A,,, which is A,,.p,, + A,-g + Ag-7 =
220 + 70 + 28.

The series for B is formed exactly in the same way, only that its commence-
ment is delayed one step ; in other words, B, is held as zero, and B, is made wnt.

Similarly for C, C,, Cz are held as zeroes, and C, is made wnit. We may
in the same way find series for D, E, F, and so on.

The eighth set of values give 28, 19, 12 as nearly proportional to the
logarithms of 5, 3, and 2. Assuming these as sufficiently near for the purposes
of musicians, we must divide the interval corresponding to the ratio 1: 2, called
by them an octave, into 12 parts, to which the name semitone is given. In this
way, counting in semitones, we have log 2 = 12, log 3 = 19, log 5 = 28;

5
whence log : =7, log 7 =4, and so on; whence the arrangement of the
gamut is at once obtained. According to these values, we should have
log : = 2, and log y= 2, wherefore the degree of precision obtained by these

numbers is not sufficient to discriminate between the tone major and the tone
minor.

In order to obtain a closer approximation, we must proceed further along
the series. Now it is important to keep to the nomenclature and arrangement
of semitones; wherefore we search among the series C for some member
divisible by 12; no one of those above given is so divisible, and therefore we
look for some compound of two of them which may be a multiple of 12. Thus
C,, + C,, = 156, so that, still counting in semitones, we have log 5 = <= =

247 156

27 = , log 8 = 75 = 19, log 2= 7g =12. From these values the logarithm

of the tone major represented by the ratio : is still 2, but that of the tone minor

ORS all"
represented by > is 17g; in the same way the corrected values of the other

musical intervals may be obtained.

By putting A, B, C to represent the periodic times of three astronomical
phenomena, we may ascertain the intervals between their simultaneous recur-
rence. Thus, if we put A for the time of revolution of the moon’s node, B and
C for the earth’s and moon’s periodic times, we shall obtain directly the law of
recurrence of eclipses.

If we take three contiguous sets of values, and thence compute the succeed-
ing set, we obtain

Ne ea Ansys Le = Pn+e: An +e a Osean DX ge
ee eps) bts — Paae -Drag 4 -Gnai+ Pati + 7» B,
Cc. Cardi Cra Cius ee Ones Sa pean Orere fay Cars

66 MR EDWARD SANG ON THE EXTENSION OF BROUNCKER’S METHOD

eliminating p and g from these three equations, we obtain, omitting the sub-
scribed , for shortness,

r, {A, B, C, — A, B, C, + B, A, G — B, A, OC, + C, A, B, — C, A, By =
{A, B, C, — A, B, C, + B, A, C, — B, A, C, + C, A, B, — C, A, B,}.
Now the first of these quantities within the ties is the determinant obtained

from the coefficients

yaaa ONE
Be ee ie
Cr oy ee

while the second is the determinant from the next three sets of values ; or,
using CAYLEY’s notation

Ae Ane aA A, eee
| Beh Bech By hella. - eerie
en RTS lye 7 Gy ec eel

Now, in the usual operation, and when the three magnitudes are incommensur-
able, 7 is unit all along ; wherefore the determinant from nine contiguous values
never changes. But at the beginning this determinant is obviously unit, and
thus we have the ordinary well-known theorem for the usual progression in
reference to two magnitudes extended to the case of three ; that is to say, the
value of the determinant is + 1 all along. In the case of two magnitudes, the
value is alternately + 1 and —1; whereas, in the case of three magnitudes,
the sign is preserved.

The above statement holds good in the case of ncommensurable quantities ;
but when there is a common measure the quotient 7 may disappear toward the
end of the operation, and then all the subsequent determinants become zero.

Tf, in the case of three commensurables, we complete the calculation, as in
the following example in which the three primes 99137, 30763, and 3229 are
compared, a remarkable yet obvious law is seen.

| |
iE 1 1 1 1 1 vat
2 4 | 0 0) 0 0 2 0 q
| 3 9 8 2 1 1 3 11 3 pull
] 3 M29 245 493 522 767 2794 32790 99137 A |
fee! 9 76 153 162 238 867 10175 30763 B.-}
1 8 16 i 25 91 1068 3229 C
1 2 2 3 11 129 390 D
1 1 1 4 47 142 E
| 1 1 3 36 109 F
1 3 35 106 G
| i 1a 33 Hi
1 3 il
| 1 K
| |

TO THE COMPARISON OF SEVERAL MAGNITUDES. 67

If we compare the last three values of A, which are in this case
A,,, A,, A,; we observe that the order of the quotients must necessarily be,
A, =P, A, + 9, A; + 7, A,; that is to say, the computation must take the
form

it 1 il 1 1 1
(0) 2 0 0) 0 0 4 2
3 Ty 3 i 1 2 8 i) 3
1 3 | wo 106 109 142 390 3229 30763 99137 a
Th a 35 36 47 129 1068 10175 32790
il 3 3 4 11 91 867 2794 Cc
il il il 3 25 238 767 d
1 i 2 iy 162 522 e
1 2 16 153 493 if
al 8 76 245 g
1 9 29 h
1 3 Z
1 k

Hence, if the values of g were written above and between the contigu-
ous values of p, and similarly with those of 7, as in the subjoined scheme,
the computation carried from left to right leads to the ultimate values of
A, B, C; when carried from right to left it leads to those of A,, A,,,
and so on; but in each case it gives the same aggregate group of numbers ;
with a difference merely in position; and hence, whenever the numbers

| i 1 1 1 1 1 1 |

| 2 4 0 0 0 0 2 0

| 3 9 8 2 1 1 3 11 3
On (i 5%» N,G, +.» , &., are arranged symmetrically, the series
of values A,, A,, A, .. . . is identic with A, B, C, read inversely.

The continuation of the same process to the case of a greater number of
magnitudes is so obvious as to stand in no need of farther illustration. The
application of this process to problems involving the higher powers of numbers
may be expected, as the Brounckerian process has already done for squares, to
throw considerable light upon that difficult branch of the Theory of Numbers.

VOL. XXVI. PART I. S

(576977)

IV.—On Gireen’s and other Allied Theorems. By Professor Tatr.
(Received April 29th, read May 16th, 1870.)

I was originally attracted to the study of Quaternions by Sir W. R.
Hamitton’s ingeniously devised and most valuable operator

od ;
Vat7 tJ

d d

dy a5 kz ,

to which he called special attention (Lectures on Quaternions, § 620) on account
of its promise of usefulness in physical applications. But I soon found that
in order that its full power may be applied, in general investigations, it is
necessary that we should have processes of definite integration, of the kinds
required in physics, applicable to quaternion symbols and not merely to scalar
variables. I often consulted Hamitton about this want, and he promised to
endeavour to supply it at some future time. I fancy that shortly before his
death he must have in some way supplied it, though he certainly did not print,
nor does he appear even to have written, anything on the subject. In one of
the last letters I received from him, he said that he intended to conclude the
final chapter of his Elements, which is devoted to physical applications, by
some sections on the use of the operator mentioned above. That chapter
remains unfinished, and as HAmiILTon rarely wrote down the steps of even a
complex train of mathematical reasoning until he had mentally completed it, it
is to be feared that this portion of his investigations is entirely lost. So far as
the analytical aspect of Quaternions is concerned, this loss is very serious
indeed, for there can be little doubt that Hamriron’s solution would have been
of immense value from the purely mathematical point of view.

I have recently succeeded to a certain extent, by a simple, though not very
direct, process, in supplying the want—so far at least as to enable me to use
quaternions in inquiries connected with potentials—and have thus arrived at
very simple proofs of GREEN’s celebrated theorem and various allied results,
some of which appear to be new and valuable. The quaternion calculus can,
in consequence, be applied without loss of its enormous special advantages to
various general theories, such as Attractions, Spherical Harmonics, Fluid
Motion, &c., &c. Curiously enough, I find that I had almost arrived at one of
the general theorems given in the present paper so long ago as 1860 (Quaternion
Investigation of the Potential of a Closed Circuit, Quarterly Math. Journal), but

VOL. XXVI. PART I. T

70 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

though I then gave a special case I did not see that a very slight modification
of my work would have enabled me to generalise it. I was then seeking to
derive from my formule the well-known physical result, and not thinking of
extending the calculus itself.

Even the little advance that I have made in the present paper has enabled
me to see, with a thoroughness of comprehension which I had despaired of
attaining (at least by Cartesian processes), the mutual relationship of the many
singular properties of the great class of analytical and physical magnitudes
which satisfy what is usually known as LApLAceE’s equation. This is, of course,
solely due to the simplicity and expressiveness of quaternions in general.

1. In what follows we have constantly to deal with integrals extended
over a closed surface, compared with others taken through the space enclosed by
such a surface ; or with integrals over a limited surface, compared with others
taken round its bounding curve. The notation employed is as follows. If Q
per unit of length, of surface, or of volume, at the point z y z, Q being any
quaternion, be the quantity to be summed, these sums will be denoted by

J{Qds and S//fQds,

when comparing integrals over a closed surface with others through the
enclosed space ; and by

J{Qds and /QTdp,

when comparing integrals over an unclosed surface with others round its
boundary. No ambiguity is likely to arise from the double use of

Sf Qds ,
for its meaning In any case will be obvious from the integral with which it is
compared.

2. I have already shown (Proc. RS.E., April 28th 1862,) that, if o be the
vector displacement of a point originally situated at
p= tae + jy + kz,

then

S.Vo

expresses the increase of density of aggregation of the poimts of the system
caused by the displacement. (See Appendix to this paper.)

3. Suppose, now, space to be uniformly filled with points, and a closed
surface 2 to be drawn, through which the points can freely move when
displaced. :

Then it is clear that the increase of number of points within the space 2,
caused by a displacement, may be obtained by either of two processes—by
taking account of the increase of density at all points within 2, or by estimating

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. 71

the excess of those which pass inwards through the surface over those which
pass outwards. These are the principles usually employed (for a mere element
of volume) in forming the so-called “ Equation of Continuity.”

Let v be the normal to 2 at the point p, drawn outwards, then we have at
once (by equating the two different expressions of the same quantity above
explained) the equation

S{[S.Vods =f/f8.cUv ds,

which is our fundamental equation so long as we deal with triple integrals.

4. Asa first and very simple example of its use, suppose o to represent
the vector force exerted upon a unit particle at p (of ordinary matter, electricity,
or magnetism) by any distribution of attracting matter, electricity, or magnetism
partly outside, partly inside 2. Then, if P be the potential at p,

cies aT
and if 7 be the density of the attracting matter, &c., at p,
Vo=\V7P-= 4Aar
by Potsson’s extension of LAPLACE’s equation.
_ Substituting in the fundamental equation, we have

4a fffrds = 4rnM = //S.VPUvds,

where M denotes the whole quantity of matter, &c., inside >. This is a well-
known theorem.

5. Let P and P, be any scalar functions of p, we can of course find the
distribution of matter, &c., requisite to make either of them the potential at p ;
for, if the necessary densities be 7 and 7, respectively, we have as before

VE — Arr ) V7P — Arr, .
Now

Ware — EVE. foe Ve
and

Ve Pe EN Pr eeePay Pi) 20 VPP; .
But, by the fundamental theorem,
MV .PPids =/f8.(V.PP,) Uv ds = /fS8.(PVP, + P,VP)Uv ds.
Substituting the above value of V?. PP,, this becomes
Jf BAPVP, + P,VP)Uv ds = //f(PV’P, + P,V?P)ds + 2fffS.VPVPyds.
But, obviously, we have also by the fundamental theorem

Jf S(PVP, — P,VP)Uvds = 7 (PV°P, — P,V?P)ds,

72 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

and the two latter equations give

SfS.VPVP,ds = —fffP.V?Pds + /f/P,S.VPU> ds ,
= — fff PV?P,ds + //PS.VP,Uvds ,

which are the common forms of GREEN’s Theorem. Sir W. THomson’s extension
of it follows at once from the same proof.

6. If P, be a many-valued function, but VP, single-valued, and if 2 be a
multiply-connected* space, the above expressions require a modification which
was first shown to be necessary by HELMHOLTZ, and first supplied by THomson.
For simplicity, suppose 2 to be doubly-connected (as a ring or endless rod,
whether knotted or not). Then if it be cut through by a surface s, it will be-
come simply-connected, but the surface-integrals have to be increased by terms
depending upon the portions thus added to the whole surface. In the first form
of GREEN’s Theorem, just given, the only term altered is the last: and it is
obvious that if p, be the increase of P, after a complete circuit of the ring, the
portion to be added to the right hand side of the equation is

DS[S.VPUvds

taken over the cutting surface only. Similar modifications are easily seen to be
produced by each additional complexity in the space 2.

7. The immediate consequences of GREEN’s theorem are well known, so that
I take only one instance.

Let P and P, be the potentials of one and the same distribution of matter,
and let none of it be within =. Then we have

Sti (VP)?ds = /fPS.VPUp ds,

so that if VP is zero all over the surface of &, it is zero all through the interior,
i.e., the potential is constant inside 2. If P be the velocity-potential in the
irrotational motion of an incompressible fluid, this equation shows that there
can be no such motion of the fluid unless there is a normal motion at some part
of the bounding surface, so long at least as = is stmply-connected.

Again, if 2 is an equipotential surface,

HI(TP as = P/fSVPUrds = Pi /fV- Pas

by the fundamental theorem. But there is by hypothesis no matter inside 2,
so this shows that the potential is constant throughout the interior. Thus there
can be no equipotential surface, not including some of the attracting matter,
within which the potential can change. Thus it cannot have a maximum or
minimum value at points unoccupied by matter.

* Called by Hetmnonrz, after Riemann, mehrfach zusammenhdngend. In translating Hevu-

HOLTZ’s paper (Phil. Mag. 1867) I used the above as an English equivalent. Sir W. THomson in his
great paper on Vortex Motion (Trans. R.S.E. 1868) uses the expression “ multiply-continuous.”

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. 73

8. If, in the fundamental theorem, we suppose

=e
which imposes the condition that
Se Va = 0),

i.é., that the o displacement is effected without condensation, it becomes
/{[S.VrUvds =fffS.V'rds = 0.

Suppose any closed curve to be traced on the surface &, dividing it into two
parts. This equation shows that the surface-integral is the same for both parts,
the difference of sign being due to the fact that the normal is drawn in opposite
directions on the two parts. Hence we see that, with the above limitation of
the value of o, the double integral is the same for all surfaces bounded by a
given closed curve. It must therefore be expressible by a single integral taken
round the curve. The value of this integral will presently be determined.
9. The theorem of § (4) may be written

S{V'? Pads = f/f s.UvV Pds = LS (UvV) Pads.
From this we conclude at once that if
eis Ue FE, eee 5

(which may, of course, represent any vector whatever) we have

{WV cds = f{S(UrV) ads ,
Wag te,

Mifrds = ffS(UvV~)r ds.

This gives us the means of representing, by a surface-integral, a vector-integral
taken through a definite space. We have already seen how to do the same for
a scalar-integral—so that we can now express in this way, subject, however, to
an ambiguity presently to be mentioned, the general integral

STL qas

where g is any quaternion whatever. It is evident that it is only in certain
classes of cases that we can expect a perfectly definite expression of such a
volume-integral in terms of a surface-integral.

10. In the above formula for a vector-integral there may present itself an
ambiguity introduced by the inverse operation

V-1
to which we must devote a few words. The assumption

Vo=T

Gah

VOL, XVI. PART I.

74 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

is tantamount to saying that, as the constituents of o are the potentials of
certain distributions of matter, &c., those of 7 are the corresponding densities
each multiplied by 47.

If, therefore, 7 be given throughout the space enclosed by 2, o is given by
this equation so far only as it depends upon the distribution within >, and must
be completed by an arbitrary vector depending on three potentials of mutually
independent distributions exterior to 2.

But, if o be given, 7 is perfectly definite ; and as

Vo=j=V 7,

the value of V~ is also completely defined. These remarks must be carefully
attended to in using the theorem above: since they involve as particular cases
of their application many curious theorems in Fluid Motion, &c. To these,
however, I shall not further allude here, as I propose to make them the subject
of a separate communication to the Society.

11. We now come to relations between the results of integration extended
over a non-closed surface and round its boundary.

Let o be any vector function of the position of a point. The line-integral
whose value we seek as a fundamental theorem is

SS. adr,

where 7 is the vector of any point in a small closed curve, drawn from a point
within it, and in its plane.
Let o, be the value of o at the origin of 7, then

o=0,—dS(tV)o,
(Proc. R.S.E., 28th April 1862; see also Appendix to this paper), so that

S8.odr =f8.(o, — S(tV)o,)dz.
But
fdr =0,

because the curve is closed; and (Tair on Electro-Dynamics, § 13, Quarterly
Math. Journal, Jan. 1860) we have generally

SS.1V8.0,d7 = 48.V(cS0,7 — o,f V.7d7) .
Here the integrated part vanishes for a closed circuit, and
£/V.7tdz = ds Uv,

where ds is the area of the small closed curve, and U> is a unit-vector perpen-
dicular to its plane. Hence

f8.o,d7 = 8.Vo,U»v.ds.

PROFESSOR TAIT ON GREEN'S AND OTHER ALLIED THEOREMS. 75

Now, any finite portion of a surface may be broken up into small elements such
as we have just treated, and the sign only of the integral along each portion of
a bounding curve is changed when we go round it in the opposite direction.
Hence, just as Ampére did with electric currents, substituting for a finite closed
circuit a network of an infinite number of infinitely small ones, in each con-
tiguous pair of which the common boundary is described by equal currents in
opposite directions, we have for a finite unclosed surface

J8.cdp =f[/8.VcUv.ds.

There is no difficulty in extending this result to cases in which the bounding
curve consists of detached ovals, or possesses multiple points. This theorem
seems to have been first given by Tuomson (THomson & Tart’s “ Natural
Philosophy,” § 190 (7); THomson on Vortex Motion, Trans. R.S.E., 1868-9,
§ 60 (q)), where it has the form

S(adx + Bdy + ydz) = ffds (7 ot) i m (“1 +n pa Ne
It solves the problem suggested by the result of § 8 above.

12. If o represent the vector force acting on a particle of matter at
p, — S.odp represents the work done while the particle is displaced along dp,
so that the single integral

JS8.oadp

of last section, taken with a negative sign, represents the work done during a
complete cycle. When this integral vanishes it is evident that, if the path be
divided into any two parts, the work spent during the particle’s motion through
one part is equal to that gained in the other. Hence the system of forces must
be conservative, 7.¢., must do the same amount of work for all paths having the
same extremities.

But the equivalent double integral must also vanish. Hence a conservative

system is such that
J{ds8S.VoUv=0,

whatever be the form of the finite portion of surface of which ds is an element.
Hence, as Vo has a fixed value at each point of space, while Uv may be altered

at will, we must have
WieNiGa=— 0,
or
Vo = scalar.

If we call X, Y, Z the component forces parallel to rectangular axes, this
extremely simple equation is equivalent to the well-known conditions

Pg Mav aig caine, dia dk
Oe DOr IRON" GE. «de. -

76 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.
Returning to the quaternion form, as far less complex, we see that

Vo = scalar = 477, suppose,
implies that
cee,
where P is a scalar such that
WP = 457r+

that is, Pis the potential of a distribution of matter, magnetism, or statical
electricity, of volume-density 7. ,
Hence, for a non-closed path, under conservative forces

—f/S.cdp = —/SS.VPdp
= SfdzP =/dP (see Appendiz)
= FE er se ,
depending solely on the values of P at the extremities of the path.
13. A Vector theorem, which is of great use, and which corresponds to the
Scalar theorem of § 11, may easily be obtained. Thus, with the notation already

employed,
/V.odr =f[V(o — S(tV)o,)dz,

= —SfS(rV)V.o,dr.
Now
V(V.VV. cdr)o, = — S(tV)V.0,d7 — S(drV) Veo, ,
and

ad(S(tV)Vo,7) = S(rtV)V.o,dr + S(drV)Vo,7 .
Subtracting, and omitting the term which is the same at both limits, we have
SV .odr = V.(V.UrV) ands.
Extended as above to any closed curve, this takes at once the form
SV.cdp =f{[dsV.(V.UvV)c .

Of course, in many cases of the attempted representation of a quaternion
surface-integral by another taken round its bounding curve, we are met by
ambiguities as in the case of the space-integral (§ 9): but their origin, both
analytically and physically, is in general obvious.

14. If P be any scalar function of p, we have (by the process of § 11, above)

S Pdr =f(Py — S(7V)P, az
=— {S.7VP,. dr.

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. (iri

But
V.VV.rdr = dzS.1V — S.drV ,
and
A(rStV) = drS.7V + 7S.drV.
These give
[Pdr =— 4(rSrtV — V.VrdrV)P, = dsV.UrVP,.

Hence, for a closed curve of any form, we have

fPdp = /fdsV.UrVP,,

from which the theorems of §§ 11, 13 may easily be deduced.

15. The above are but a few of the simpler of an immense host of theorems
which any one with some knowledge of quaternions may easily work out for
himself, by developing a little farther, or applying to other combinations, the
processes just explained. I shall, therefore, give no more of them until I have
an opportunity of, at the same time, showing their ready applicability and great
value in physical investigations.

Appendix, added June 3d 1870.

16. At the instance of Prof. KELLAND, to whom this paper was referred,
T append a slight sketch of some of the properties of the operator V, of which
so much use has been made in the foregoing paragraphs. Most of the results
now to be given have been already published by myself, but the mode in which
they were formerly deduced has been abandoned for one more purely quaternionic.

17. It may perhaps be useful to commence with a different form of definition
of the operator V, as we shall thus, if we desire it, entirely avoid the use of
ordinary Cartesian co-ordinates. For this purpose we write

Suey = as,

where « is any unit-vector, the meaning of the right hand operator (neglecting
its sign) being the rate of change of the function to which it is applied per unit
of length in the direction of the unit-vector a. If a be not a unit-vector we
may treat it as a vector-velocity, and then the right hand operator means the
rate of change per unit of time due to the change of position.

Let a, 8, y be any rectangular system of unit-vectors, then by a fundamental
quaternion transformation

V =— aSaV — BSBV — ySyV = ad. + Bde + yd,

which is identical with Hamitron’s form given above. (Lectures, § 620.)

18. This mode of viewing the subject enables us to see at once that the effect
VOL. XXVI. PART I. x

78 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

of applying V to any scalar function of the position of a point is to give its vector
of most rapid increase. Hence, when it is applied to a potential w, we have

Vu = vector-force at p.

If u be a velocity-potential, we obtain the velocity of the fluid element at p;
and if w be the temperature of a conducting solid we obtain the flux of heat.
Finally, whatever series of surfaces is represented by

p= ,

the vector Vw is the normal at the point p, and its length is inversely as the
normal distance at that point between two consecutive surfaces of the series.
Hence it is evident that

S.dpVu =— du,
or, as it may be written,
—S.dpV =d;

the left hand member therefore expresses total differentiation in virtue of any
arbitrary, but small, displacement dp.

19. To interpret the operator V.aV let us apply it to a potential function w.
Then we easily see that «w may be taken under the vector sign, and the

expression
V(aV)u = V.aVu

denotes the vector-couple due to the force at p about a point whose relative
vector Is a.
Again, if o be any vector function of p, we have by ordinary quaternion

operations
V(aV).¢ = S.aVVo + aSVo — VSac.

The meaning of the third term (in which it is of course understood that V
operates on o alone) is obvious from what precedes. It remains that we
explain the other terms.

20. These involve the very important quantities (not operators such as the
expressions we have been hitherto considering),

S.Vo and V.Voc ,

which occur very frequently in the preceding paper. There we looked upon o
as the displacement, or as the velocity, of a point situated at p. Let us now
consider the group of points situated near to that at p, as the quantities to be
interpreted have reference to the deformation of the group.

21. Let 7 be the vector of one of the group relative to that situated at p.

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. 79

Then after a small interval of time ¢, the actual co-ordinates become
p+ to
pt7+t(o — S(rtV)o)

by the definition of Vin § 17. Hence, if ¢ be the linear and vector function
representing the deformation of the group, we have

g7 = 7 —W(rV)o

and

The farther solution is rendered very simple by the fact that we may assume ¢
to be so small that its square and higher powers may be neglected.
If ¢g be the function conjugate to ¢, we have
@7r =7t—UtVSr70.

Hence
CeCe) GG)

=T- 3[ SV)o + VSro | — 5V.1VVo

The first three terms form a self-conjugate lmear and vector function of 7, which
we may denote for a moment by a7. Hence

¢T = aT — 5 V.tVVo,
or, omitting ? as above,
¢tT = oT — 5 V.atVVo ;
Hence the deformation may be decomposed into—1st, the pure strain a, 2d, the

rotation

SVVo.

Thus the vector-axis of rotation of the group is
4VVo.

If we were content to avail ourselves of the ordinary results of Cartesian
investigations, we might at once have reached this conclusion by noticing that

d€ dn 1 (WER eae we
Wa Gaeewana)* G-%
and remembering the formulee of Stokrs and HELMHOLTZ.
22. In the same way, as

we recognise the cubical compression of the group of points considered. It
would be easy to give this a more strictly quaternionic form by employing the
definition of §17. But, working with quaternions, we ought to obtain all our
results by their help alone; so that we proceed to prove the above result by

30 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

finding the volume of the ellipsoid into which an originally spherical group of
points has been distorted in time ¢.
For this purpose, we refer again to the equation of deformation

gr =7—W(rV)c,

and form the cubic in ¢ according to HAMILTON’s exquisite process. We easily
obtain, remembering that 2? is to be neglected,*

0 = @ — (3 —iSVo)¢’ + (3 — 28 Va) g— (1 —t8Va),
or
0 = (g—1)?(¢—1 + #8Vo).

The roots of this equation are the ratios of the diameters of the ellipsoid whose
directions are unchanged to that of the sphere. Hence the volume is increased

by the factor
1—tSVo,

from which the truth of the preceding statement is manifest.

23. As the process in last section depends essentially on the use of a non-
conjugate vector function, with which the reader is less likely to be acquainted
than with the more usually employed forms, I add another investigation.

Let

w= ¢r=7T—WW(rV)oc.
Then
T= Ono — a + iS(oV)o.

Hence since if, before distortion, the group formed a sphere of radius 1, we

have
fy ryeemaal bie

* Thus, in Hamroy’s notation, , #, v being any three non-coplanar vectors, and m, My, My the
coefficients of the cubic,

—mS. AWW = S.¢ rg ug'v
= 8.(A — tV8ro)(u — tVSuc)(v — tVSvc)
= 8.0 — ?VSro)(Vew — tVpVSvo + fVvVSyc)
= S.Auwy — t[S.uvVSro + S.rAVSuo + S.rAWVSve]
= S.rAwy — .S.wrV + wS.rrV + vS.AuVo
= S.Auv — tS.AwWSVo.

mS.hpuv = S.rAG'ugv + S.uPver + S.vge'rAg'w

= S.A(Vuv — tVpVSvo + tVvVSyc) + &e.
= S.Auwy — tS. AwVSve — tS. vrAVSue + &e.
= 38.Auy — 2tSVoS.rpv.

—mS.dyuv = S.Aweyv + S.uvg'r + S.rrg'w
= S.Auy — t8.rAWVSve 4+ Ke.
= 3S.Amv — tSVoS.rpv.

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. 81
the equation of the ellipsoid is

T(@ + iS(wV)o) =i |
or

a’ + 2t8aVSao =—1.
This may be written

S.aya = S.a(@ + tVSao + iS(@V)o) =—1,

where x is now self-conjugate.
Hamitton has shown that the reciprocal of the product of the squares of
the semiaxes is

—S.xXIXE
whatever rectangular system of unit-vectors is denoted by 7, 7, 4.
Substituting the value of y, we have
—S.(i + WVSio + tS(iV)c)(j + &e.)(A + &e.)
=—S.(i + tVSic + tS(iV)o)( + 2tiSVo — tS(iV)o —tVSic)
= 1 + 20SVo.

The ratio of volumes of the ellipsoid and sphere is therefore, as before,

= = 1- Vo.
Via -

24. Before concluding I may append a generalised form of GrEEN’s Theorem,
which is obviously fitted to be of use in quaternion investigations. If we put

7=i1P + 7P’ + kP’,
we easily see by the equations at the end of § 5 that

SL S(VP,.V) rds = —SfffP,V?rds + f/f P,S (Uv. V) rds ,
SH fffsV?P ds + /f7S.VP_Uv.ds .
As a particular case, let
1 = Sap
so that
VP, = ia + JSja + kSka =— a,
VAP 05
we have
S{[S(aV) rds =/[fSapV7rds — ffSapS (UvV) rds ,
=f To.aUv as .

VOL. XXVI. PART I. N

82 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

Any constant may be added to the value of P,. The additional terms thus
introduced must vanish. This gives, as in § 9,

Sf Vi rds = ff 8S (UvV) rds .
As another verification, suppose 7 constant, and we have
jfs:aUvds= 0,

which is obviously true. Interesting results are obtained by treating this by
the processes of §§ 8, 11.
25. From one of the theorems above—viz.,

ST[S(aV )rds = f{{7S.aUp ds ,
we have by the formula of § 17
Sf Vids =ff Uv.rds,

a considerable extension of the fundamental theorem of § 3, which is, in fact,
only its scalar part. It might have been obtained, however, as the reader will
easily see, by a much more direct process. The vector part

SL NV rds =f/fVUv.7 ds ,

as we see by the meaning of VVz in § 21, is of great importance in physical
applications, especially in connection with Electricity and with Fluid Motion.

When
c=IVEP.

where P is is a scalar, the left hand member vanishes, and the value of the right
hand member limited to a non-closed surface is then found as in § 14.
26. Again, let

P= p,
which gives
VP,.=— 25;
V?P, =6

We have
— 2/fS(pV) tds = — f/f p’V?rds + f/f p?S(UvV) rds

= — 6ff/frds — 2ff78.pUvds.

Now if the constituents of 7 be homogeneous functions of p of the m'" degree, we

have for any one of them
S.pVE=— n€,

so that under these circumstances

(n + 3) ///rds =—S[7S.pUvds.

PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS. 83

Of this a particular case is

(n + 3)///&ds =—SfES.pUvds,

which suggests many curious theorems.
27. As a verification of it, let the closed surface > which determines the
limits of the integrations be itself
Eé=C,
which, of course, subjects the form of € to further limitations.
The right hand member is obviously equal to

3C_x<-yol. of >,

because — S.pUv is the perpendicular from the origin to the tangent plane at p
to the element ds. The left hand side may be broken up into a set of shells
bounded by surfaces whose equations are

ies Ore

where ¢ varies from 0 to 1. [This follows from the assumption that € is homo-
geneous.| The volume of the surface corresponding to any value of ¢ is obviously

Cu evOleotle:

Hence
ds = 8e’de x vol. of >,

so that the left-hand member of the equation above becomes
1
(n + IE 3Ce"t?de x vol. of 2 = 3C x vol. of 2,
' 0

and the proposition is proved.
28. A very interesting case is when

1
- c= Tp

in which case n = — 3, and our equation appears to become
@n291 32 0S ia
ie ‘Tp? ;

It is obvious, however, that there is an infinite element on the left hand,
when Tp = 0, #.¢., when the origin lies inside 2; and it is easy to see that the
correct result is a simple case of the well-known equation of § 4. In fact, the
expression on the right denotes, as is evident, the whole spherical opening sub-
tended at the origin by 2. Its value is therefore 0 if the origin be without %,
and 47 if within—2 being supposed to be simply-connected.

29. As a final example let us suppose in § 26 that € is a Spherical Harmonic.

84 PROFESSOR TAIT ON GREEN’S AND OTHER ALLIED THEOREMS.

Then, in addition to the condition of homogeneity there given, we have the
condition .

VE =0,
and the general equation of the section referred to gives

Qn [ff Eds = ff p’S.UrVEds,

so that, with the help of the final equation of § 26 we have for any closed sur-
face whatever
J[S.Uv(2npé + n + 3p°VE)ds = 0.

This integral, whose value is obviously the same for all surfaces bounded by
a given closed curve, can be reduced to the form

4n+6

Mf (Tp)"**8. U0 (9, - —ae-)as,
(Tp)"**

where q, is any quaternion which satisfies the condition
Vo Os

This is susceptible of various remarkable transformations, both as a double and
as a single integral. But this digression might be indefinitely extended, and
perhaps has already gone too far.

30. The essential basis of the whole of this theory is the great invention of
HAmILton, by which it is made possible to represent as a vector-operator the
square root of LAPLACE’s operator

d? d? a?

dy" diz?’

which has not yet been done by any but quaternion symbols, at least in a sym-
metrical, easily intelligible, and practically useful form.

It is rash to make any definite assertions on such matters, especially when
a writer of such extraordinary fertility, knowledge, and power as Sir W. R.
HAMILTON is concerned, but to the best of my knowledge the greater part of
the results given above is my own. Hamiiron’s treatment of V, so far as I am
aware of its having been published, will be found in Proc. R._A., 1846 and
1854, (in the latter of which there is a very curious and interesting proof of
Dupty’s Theorem,) and in his Lectures on Quaternions, § 620. My own is to
be found in Quarterly Math. Journal, October 1860; Proc. R.S.E., 1861-2,
1862-3 ; and Elementary Treatise on Quaternions, S§ 317, 319, 364, &c., 418,
421-8, Ex. 24 to Chap. IX. and 10 to Chap. XI.

( 85 )

V.—On the Heat Developed in the Combination of Acids and Bases. Second
Memoir. By THomas Anprews, M.D., F.R.S., Hon. F.R.S.E., Vice-
President of Queen’s College, Belfast.

(Read 6th June 1870.)

In a paper communicated to the Royal Irish Academy in 1841, I gave an
account of a large number of experiments on the heat disengaged when acids and
bases, taken in the state of dilute solution, enter into combination, and when
bases, insoluble in water, are dissolved in dilute acids. The following general
eonclusions or laws were deduced from those experiments :—

Law 1.—The heat developed in the union of acids and bases is determined
by the base and not by the acid, the same base producing, when combined with
an equivalent of different acids, nearly the same quantity of heat ; but different
bases, different quantities.

Law 2.—When a neutral is converted into an acid salt, by combining with
one or more atoms of acid, no change of temperature occurs.

Law 3.—When a neutral is converted into a basic salt, by combining with
an additional proportion of base, the combination is accompanied with the
evolution of heat.*

Three years later I laid before the Royal Society of London the results of
an experimental investigation of the heat developed when one base is substi-
tuted for another in chemical compounds. ‘The law deduced from this inquiry
is implicitly involved in the foregoing, of which it may indeed be regarded as a
necessary consequence. It was enunciated in the following terms :—

Law 4.—When one base displaces another from any of its neutral combina-
tions, the heat evolved or abstracted is always the same, whatever the acid
element may be, provided the bases are the same. t

Finally, the law of metallic substitutions, first announced in the “ Philo-
sophical Magazine” for August 1844, was thus stated in a paper published in
the “ Philosophical Transactions” for 1848.

Law 5.—When an equivalent of one and the same metal replaces another
in a solution of any of its salts of the same order, the heat developed is always
the same ; but a change in either of the metals produces a different development
of heat.

In 1845 a paper appeared by Grauam on the heat disengaged in combina-
tions, the second part of which refers to the -heat produced when hydrate of

* Transactions of the Royal Irish Academy, vol. xix. p. 228,
t Philosophical Transactions for 1844, p. 21.

VOL. XXVI. PART I. Z

86 DR ANDREWS ON THE HEAT DEVELOPED IN THE

potash is neutralised by different acids.* The results arrived at by this distin-
guished chemist exhibit a close agreement with those contained in my first
communication to the Royal Irish Academy.

The concluding part of the elaborate memoir of MM. Favre and SILBEr-
MANN on the heat disengaged in chemical actions is chiefly devoted to the same
subject. A large number of experiments are described, which are nearly a
repetition of those I had previously published. Their results bear a general
resemblance to those given by myself in 1841; but they widely differ in the
details. The authors of this able memoir fully recognise the accuracy of my
fourth law, which asserts the equality of thermal effect when one base is sub-
stituted for another. ‘ M. ANpREws,” they observe, “ avait en effet établi que,
quelque soit acide d’un sel, la quantité de chaleur dégagée par la substitution
d’une base 4 une autre pour former un nouveau sel est la méme, lorsque l’on
considére les deux mémes bases.” +

In a preceding paragraph of the same memoir, the authors object to what
they conceive to be my first law, and state that it is not in accordance with the
results of their investigations. As the question is one of some importance, I
may perhaps be permitted to quote the passage in the original language. ‘“‘ Ses
conclusions, savoir: que la chaleur dégagée par l’équivalent d’une méme base
combinée aux divers acides est la méme, ne s’accordent pas avec les résultats
de nos recherches, et ne nous paraissent pas pouvoir étre admises.” No doubt,
through inadvertence, MM. Favre and SILBERMANN have here given an inaccurate
statement of my first law. It did not declare that precisely the same amount
of heat is disengaged by all the acids m combining with the same base, but
that the heat is determined by the base, “the same base producing, when
combined with an equivalent of different acids, nearly the same quantity of
heat.” A comparison of the results of MM. Favre and SILBERMANN with those
in my original memoir will show that I had fully recognised and described the
deviations from the other acids, exhibited, on the one hand, in excess, by
the sulphuric acid, and on the other, in deficiency, by the tartaric, citric,
and succinic acids. ‘If we refer,” I remarked, in the original memoir of
1841, “to the first, second, and fourth tables, as being the most exten-
sive, from the large number of soluble compounds formed by potash, soda,
and ammonia, it will be observed that the sulphuric acid developes from 0°8 to
nearly 1° more than the mean heat given by the other acids; while the tartaric,
citric, and succinic acids fall from 0°:4 to 0°55 short of the same. A minute
investigation of the influence of the disturbing sources of heat will no doubt
discover the causes of these discrepancies. The high numbers for sulphuric

* Memoirs of the Chemical Society, vol. ii. p. 51.
+ Annales de Chimie et de Physique 3®™¢ sete xxxvii. p. 497 (1853).

COMBINATION OF ACIDS AND BASES. 87

acid are probably connected with that acid’s well known property of developing
much heat when combining with successive atoms of water. All the other acids
develope nearly the same amount of heat in combining with the same base, the
greatest divergences from the mean quantity being, in the case of potash,
+ 0°24, and — 0°13; in that of soda, + 0°26, and — 0°14; and in that of
ammonia, + 0°17 and — 0°:05. These differences are almost within the limits
of the errors of experiment.”*

But although there is a superficial agreement between my original results
and those of MM. Favre and SILBERMANN, they will be found, when examined
closely, to differ widely in detail, and on points of great importance. I had
found that the oxalic acid disengages almost exactly the same amount of heat
in combining with the soluble bases as the hydrochloric, nitric, and many other
mineral acids, and this observation I have always regarded as one of the main
foundations of Law 1. MM. Favre and SILBerMANN, on the contrary,
have inferred from their experiments that “the following organic acids—the
oxalic, formic, valeric, and citric—disengage sensibly the same quantity of heat,
but it is less (plus faible) than that given by the foregoing mineral acids ”—
among which they enumerate the nitric and hydrochloric. According to my
experiments, no distinction of this kind can be admitted between acids derived
from the mineral and organic kingdom, inasmuch as the oxalic acid developes
at least as much heat in combining with the bases as the hydrochloric, nitric,
and several other strong mineral acids.

The experiments to be described in this paper were made some years ago,
but their publication has been deferred from accidental circumstances. I have,
however, recently repeated a few of the more important of them, with a
slightly modified form of apparatus. The solutions were taken in so dilute a
state that the heat disengaged never exceeded 3°5C. A standard solution of
sulphuric acid was prepared and carefully analysed, by precipitating a given
weight with a soluble salt of barium, and weighing the sulphate of barium. The
strength of the alkaline solutions was adjusted with great care by means of this
standard acid. Thesame solution of each alkali was employed in all the experi-
ments, and the quantity used in each experiment was determined by careful
weighing. The acid solution was of such a strength that, after being mixed with
the alkali, an excess of two or three per cent. of acid was present. The alkaline
solution was contained in a light glass vessel, in which a large platinum crucible
holding the acid was carefully floated. By giving a rapid rotation, by means of
alight stirrer, to the acid solution in the platinum crucible, a perfect equilibrium
of temperature was soon established between the two liquids. The initial tem-
perature of the solutions was usually about 1°°5 below that of the air, and the
final temperature of the mixture about 1°°5 above it. The corrections for the

* Transactions of the Royal Irish Academy, vol. xix. p. 240.

88 DR ANDREWS ON THE HEAT DEVELOPED IN THE

heating and cooling action of the surrounding medium were determined with
great care. The mechanical process of adding the acid to the alkaline solution
produced no change of temperature, and as the heat disengaged in the com-
bination raised the liquid almost instantly to the maximum temperature, the
whole correction required was for cooling. The first temperature was read one
minute after the addition of the acid to the alkaline solution, the mixture being
stirred during the whole of that time. If 6 represents the correction, and e the
excess of temperature above the air in centigrade degrees, the value of 5 will be
given by the following expression :—

=e xO O12.

Asa proof of the accuracy of the method of mixture adopted in this inquiry,
I may mention that, being desirous to know whether the dilute acids em-
ployed in these experiments produced any change of temperature when mixed
with water, I made the experiment with nitric acid by the method just described,
substituting water for the alkaline solution, with the unexpected result of a fall
of 0°01. On varying the conditions of the observation, so as to obtain a larger
effect, it was ascertained not only that a diminution of temperature had actually
occurred, but that the observed fall represented approximately its true amount.
When hydrochloric acid of equivalent strength was diluted to the same extent,
an elevation of temperature of 0°05 was produced.

The accuracy of experiments of this kind, where the whole thermal effect
observed amounts only to 2° or 3°, depends greatly on the thermometer employed.
Unless its indications are perfectly trustworthy in every part of the scale, the
labour of the inquirer will only end in disappointment. I have therefore taken
every precaution to secure this important object. The tube of the thermometer
was calibrated and divided with care, according to an arbitrary scale, by means
of a dividing instrument contrived for the purpose, and provided with a short
screw of great accuracy made by TroucuTon & Simms. The divisions, etched
finely on the glass, corresponded to about 0°05 C., and the readings could be
made with certainty to less than 0°01. The division of the scale, corresponding
to 0°, was determined from time to time in the usual way ; and another point,
about 30° C., was fixed by comparison with four other thermometers similarly
constructed, whose scales extended from the freezing to the boiling point of water.
The readings of these four instruments, when reduced to degrees, rarely differed
from each other within the limits to which they could be read, or 0°02. The
reservoir of the thermometer used in these experiments was 75 millimetres
long, and, when immersed in the liquid, occupied nearly its entire depth.

As some uncertainty always exists with regard to the thermal equivalent of
glass vessels, | made two sets of comparative experiments—one with a thickly
varnished copper vessel, and the other with a vessel of platinum. The mean

COMBINATION OF ACIDS AND BASES. 89

result of these experiments coincided almost exactly with the result obtained
when the glass vessel was employed.

The weight of the glass vessel which contained the alkaline solution was 58
grammes, and corresponded thermally to 11:4 grammes of the solutions formed.
The thermal equivalent of the reservoir of the thermometer and of the stirrer
was 0:9 grammes. The alkaline solution weighed 160 grammes, and contained
the equivalent of 1°738 grammes of SO;. The acid solution weighed 42:5
grammes. Hence the entire thermal value of the apparatus, in terms of the
solution, formed, was—

Solution, . ; ‘ : . , 20%
Glass vessel, 1
Thermometer and stirrer,

-214°8 grammes.

A correction (additive) of 345 was made to the direct readings for the
mercury in the stem of thermometer. The results are given to thousandths of
a degree, but this apparent minuteness is due to the reduction of the indica-
tions of the arbitrary scale to degrees.

In the following detailed statement of the experimental results, Inc. is the
increment of temperature observed, corrected for the mercury in stem, and 6 is
the correction for cooling.

Potash and Sulphuric Acid.

Inc. 3°°358 3°°356 3°'366
6 ‘010 024 021
3°°368 3°:380 3°°387
Mean increment corrected, 3°°378
Potash and Nitric Acid.
Inc. 2°:971 2°:976 BEV TEN
6 ‘018 ‘019 ADIL %
2°-989 2°-995 9°:994.
Mean increment corrected, 2°:993

Potash and Hydrochloric Acid.

Ine. 3°:004 3°°002 3°:005
0) ‘017 TOMES) OILY
3°:021 S020 ae ee 3-022

Mean increment corrected, 3° 021

Potash and Oxalic Acid.

Ine. 3°°036 3°°048 3°:040
t) 017 ‘017 016
3°°053 3°:065 3°:056

Mean increment corrected, 3°°058

MOE. <XVI, PART J, DR

90 DR ANDREWS ON THE HEAT DEVELOPED IN THE '

Potash and Acetic Acid.

Inc. 27835 2°-846
6 016 ‘007
2°°851 2°°853

Mean increment corrected, 2°°852

Potash and Tartaric Acid.

Ine. 2707 ret ay o-730
6 ‘014 014 013
PA 1 ater 743

Mean increment corrected, 9°-739

Soda and Sulphuric Acid.

Ine. 37392 3°335
t) 025 024
3°°347 3°°359

Mean increment corrected, 3%353

Soda and Nitric Acid.

Inc, 2°-914 2°-919
t) 012 012
2°-926 2-931
Mean increment corrected, 2°°929

Soda and Hydrochloric Acid.

Inc. 2°-963

F) 019

2°:982
Increment corrected, 2°-982

Soda and Oxalic Acid.

Ine. 3°:029 3°013
0) “O19 020
3°°048 3°°033

Mean increment corrected, 3°-040

Soda and Acetic Acid.

Inc. 2°°816 2°-812
é ‘017 018
2°°833 2°°830

Mean increment corrected, 2°°831

COMBINATION OF ACIDS AND BASES.

Soda and Tartaric Acid.

Ine. 2°:693 2°'693
6 ‘019 “O)ILa)
Dele 2°-708

Mean increment corrected,

2°-710

Ammonia and Sulphuric Acid.

Ine. 2°°967 2°-959
ny) ‘017 ‘010
2°°984 2°969

Mean increment corrected,

2°'976

Ammonia and Nitric Acid.

Ine. 2°56
f) “010
2°'566

Mean increment corrected,

27551
"015

2°-566
2°-566

Ammonia and Hydrochloric Acid.

Tne. 9°'609 9°°607
6 ‘O15 “015
9°°624 Page)

Mean increment corrected,

Ammonia and Oxalic Acid.

Tne. 2°°635 2°°630
f) ‘015 016
2°°650 2°646

Mean increment corrected,

2°-648

Ammonia and Acetic Acid.

Ine. 2°'469 2482
6 017 016
2°-486 2°498

Mean increment corrected,

2°°499

Ammonia and Tartaric Acid.

Inc. 2°:365 2°°354
6 SON 016
2°°382 2°:370

Mean increment corrected,

2°:376

92 DR ANDREWS ON THE HEAT DEVELOPED IN THE

In the following table I have collected the foregoing results, arranging the
acids in the order of their thermal action.

Acid, Potash. Soda. Ammonia.
Sulphuric Acid, 3°°378 3°353 2°-976
Oxalic Acid, . c é 3°°058 3°:040 2-648
Hydrochloric Acid, ; ; 3°-021 2°°982 2°-623
Nitric Acid, : : F 2°-993 2°-929 2°-566
Acetic Acid, . : : 2°°852 2°°832 2°-4.92
Tartaric Acid, . , : 2 toe 2-710 2°:376

It is interesting to observe how closely the results in the three vertical
columns agree relatively with one another. The acids follow in the same order
under each base, and even the differences in the amount of heat disengaged by
the several acids in combining with the different bases approximate in many
cases closely to one another. Thus the heat given out when the sulphuric acid
combines with potash exceeds that given out when the oxalic acid combines
with the same base by 0°320, the corresponding differences in the case of
soda and ammonia being 0°°313 and 0°328. If, in like manner, we compare the
differences between the heat disengaged by the acetic and tartaric acids, we fall
upon the numbers 0°°120, 0°:122, and 0°:116. Even in the case of the oxalic,
hydrochloric, and nitric acids, which disengage so nearly the same amount of
heat, the same order is observed with the three bases. It must be particularly
remarked that the oxalic acid disengages from 0°:022 to 0°:058 more heat in
combining with these bases than the hydrochloric acid, and from 0°-065 to
0°:111 more than the nitric acid. The conclusion of MM. Favre and SILBER-
MANN, that the organic acids (oxalic, formic, acetic, &c.) disengage sensibly less
heat than the mineral acids, is thus entirely disproved ; and the original results
recorded in my work of 1841, according to which the oxalic acid disengages at
least as much heat as the nitric, phosphoric, arsenic, hydrochloric, hydriodie,
boracic, and other mineral acids (with the exception of the sulphuric acid) are
fully confirmed. The tartaric, citric, and succinic acids, it is true (as was also
shown in the same work), give out about ;;th less heat than the average of the
other acids ; but the acetic and formic acids fall scarcely 4th below the mean,
and the oxalic acid is always above it. These results, in all their main features,
are fully corroborated by the experiments recorded in this paper, which were
performed with a more perfect apparatus and a more exact. thermometer
than I had at my command in my earlier investigations. A reference to the
same paper will show that, while acids, differing so widely from one another as
the oxalic, phosphoric, arsenic, nitric, hydrochloric, and boracic acids, scarcely
present any sensible difference in the quantities of heat which they disengage in
combining with the bases ; and while of the other acids examined the sulphuric
acid (and probably also the sulphurous acid) presents an extreme deviation of

COMBINATION OF ACIDS AND BASES. 93

about 4th above the mean, and the tartaric acid group a deviation of about 35th
below it ; the bases, on the contrary (and the subsequent researches of FAVRE and
SILBERMANN have confirmed this result), differ altogether in thermal power from
one another. Thus equivalents of the oxides of magnesium and of silver give out
4°-1 and 1°'8 of heat respectively in combining with nitric acid, the former oxide
having therefore 2°3 times the thermal power of the latter. Yet, as is well
known, both these bases fully saturate the acid, and the resulting solutions are
even neutral to test paper. For these reasons, I have no doubt whatever that
the first law, as enunciated in 1841, is the expression of a true physical law,
and that in the combination of acids and bases in presence of water the heat
disengaged is determined by the base and not by the acid. It is true that in
this, as in similar physical inquiries, experimental results cannot immediately
be obtained free from complication or disturbing influences. The same remark
applies to the experimental proof of the great law discovered by Dutone and
Petit, which connects the specific heats and atomic weights of the elementary
bodies, and also to that of the remarkable relations discovered by Kopp between
the composition and boiling points of many organic liquids. We have already
seen an illustration of one of these disturbing influences, in the fact that dilute
nitric acid, when mixed with water, gives a slight fall of temperature, hydro-
- chloric acid, a rise; and the differences of specific heat in the solutions formed
will to a small extent modify the results. But the cause of the higher thermal
power of sulphuric acid I have not been able to discover, and future researches
must decide whether it depends upon some disturbing cause, or (which is less
probable) upon its possessing an exceptionally high thermal power. One
condition is, however, essential, or Law 1 will not apply. The acid and base
must be capable of combining when brought into contact, and of forming a
stable compound. In the paper so often referred to, I showed that hydro-
cyanic acid and potash, which fail to fulfil this condition, do not disengage the
normal amount of heat when mixed; and the same observation will doubtless
be found to apply to a large number of metallic oxides, which form unstable
compounds with, and imperfectly neutralise, the bases.

As regards the experimental proofs of the other laws, even those of the
fourth law, the truth of which is admitted by MM. Favre and SILBERMANN,
they are only approximative ; and here also we meet occasionally with peculiar
and unexpected results. Thus a slight fall of temperature occurs, as Hess
showed long ago, in the conversion of the neutral sulphate of potash into the
acid salt ; and I found, as indeed might have been expected from their alkaline
reaction, that in the conversion of the ordinary phosphates and arseniates into
super salts, a disengagement of heat occurs, amounting to about one-seventh of
that disengaged in the formation of the salts themselves. In other cases results,

at first view startling and apparently anomalous, will be found to be strictly in
VOL. XXVI. PART I. 2B

94 DR ANDREWS ON THE HEAT DEVELOPED IN THE

accordance with the general principles already laid down. In the formation of
double salts there is no disengagement of heat—a principle announced in
1841, and which ought perhaps to be enunciated as a distinct law, although it
is implicitly involved in Law 2. Again, if tribasic phosphoric acid or arsenic
acid is added in fractional portions to a solution of potash till the subsalts are
formed, the heat disengaged on each addition of acid corresponds to the amount
of acid added ; but after this pomt has been reached, the disengagement of heat
follows a different law. The pyrophosphoric acid, on the other hand, behaves
in the same way as the nitric and most other acids, when added in successive
portions to solutions of potash or soda; equal increments of heat being evolved
for equal additions of acid, till the pyrophosphate of potash or soda is formed.*

APPENDIX.

In the following tables I have given the results described in this communi-
cation and those of 1841 in a form which admits of comparison with one
another, and with those of MM. Favre and SILBERMANN. I have also added a
few determinations recently made by M. Tuomsen of Copenhagen.t+ It will be
seen that the original experiments of 1841 exhibit, on the whole, a fair agree-—
ment with those now communicated to the Society. From the small scale on
which they were performed (the whole weight of the solutions after mixture
being less than 30 grammes), the imperfect form of the apparatus, and the
uncertainty of the thermometric indications, I have indeed been surprised to
find them so near the truth. The results of MM. Favre and Sm~tBpeRMANN do
not exhibit the precision which might have been expected from the high char-
acter of those experimentalists, and from the accuracy of other parts of their |
great work. The mercurial calorimeter employed by them appears to have
been little adapted to its purpose; but after making due allowance for its im-
perfections, I am at a loss to account for the serious errors into which they have
fallen. M. THOMSEN’s experiments have evidently been made with care, and
his results agree comparatively with my own ; but the absolute amount of heat
obtained by him falls far short of what I have found. It is indeed much easier
to obtain results relatively than absolutely correct. The numbers given in this
paper will, I believe, be found rarely to differ relatively more than 34th from
the truth, but they may hereafter require a small correction in respect to their
absolute value. That correction can, however, be scarcely more than jth of
the whole amount; and I have little doubt that the number, for example,

* Transactions of the Royal Irish Academy, vol. xix. pp. 245-248. The observations of GraHAM
confirm the statement that no heat is evolved in the formation of any double salt. Memoirs of the
Chemical Society, vol. 1. p. 83.

+ PoccEnporrr’s Annalen, cxxxviii. p. 78.

COMBINATION OF ACIDS AND BASES. 95

given by THOMSEN to express the heat disengaged in the combination of soda
with nitric acid will prove to be as far below the true number as that given by
MM. Favre and SILBERMANN is above it.

TABLE [.—Potash.

Acid ANDREWS, FAVRE and ANDREWS,

: 1841. SILBERMANN. 1870.

- |

|
Sulphuric, . ; é 16330 16083 16701
Nitric, , : 4 15076 15510 14800
Hydrochloric, . ; 14634 15656 14940
Oxalic, : : : 1V4A771 14156 15124
Acetic, 5 : ‘ 14257 13973 13805
Martaric, ; : 13612 13425 13508

TABLE I¥.—Soda.

: ANDREWS, Favre and ANDREWS, |
ae 1841. sipenitane 1870. THOMSEN.
|
Sulphuric, : 16483 15810 16580 15689
Nitric, ; : 14288 15283 14480 13617
Hydrochloric, . 14926 15128 14744 13740
Oxalie, : . 14796 13752 | 15032 oon
Acetic, i : 14046 13600 14000
Tartaric, 13135 13651 | 13400
TasLE I1I].—Ammonia.
iMeid ANDREWS, Favre and ANDREWS,
; 1841. SILBERMANN. 1870.
Sulphuric, . : eel 14135 14690 14710
Nitric, . 3 | 12440 13676 12683
Hydrochloric, . : 12440 13536 12964
Oxalic, : ; : 12684 ial 13088
Acetic, : : cael T2195 | 12649 12316
Tartaric, . Bee Me ll 11400 aa 11744 |

(99717)

V1I.—The Genetic Succession of Zooids in the Hydroida. By Professor
ALLMAN.

(Read 16th May 1870.)

Though most of the terms employed in the following paper have already
become part of the language of science, some definitions may be here given with
the view of rendering the subject more intelligible.

The Zooids are the more or less individualised members of which the hydroid
colony is composed.

The Hydranth is the proper nutritive zooid.

The Blastostyle is a columnar zooid destined not for nutrition, but for the
origination of sexual buds.

The Blastocheme is a medusiform zooid which gives origin to generative
elements, not immediately, but through the intervention of special sexual
buds.

The G'onophore is the ultimate generative zooid, that which ¢mmediately
produces the generative elements. It may be either medusiform or sacciform.
The Trophosome is the entire assemblage of nutritive zooids in a colony.
The Gonosome is the entire assemblage of generative zooids in a colony.

From all the facts which the study of the Hydroida has made apparent, we
may regard it as certain that however long zooidal multiplication may continue,
this is not sufficient for the perpetuation of the species, but that a period must
at last come in the life of the hydroid when by an act of true sexual reproduc-
tion, new individuals are produced for the indefinite extension of the species
through time.

This truth finds its expression in STEENSTRUP’S famous law of ‘“ Alternation
of Generations,’—a law which, though not very correctly enunciated by its
framer, may be regarded when properly expounded as a statement of the fact,
that in certain animals every act of embryonal development is followed by
one or more acts of zooidal development, which invariably conduct us to an
ovum in which embryonal development followed by zooidal development again
occurs, and the entire series becomes thus repeated.

Now the various series expressing this alternation of sexual with non-
sexual development, exhibit among the Hydroida different degrees of complica-
tion, which will be more easily understood if we attempt to present them in
the somewhat technical shape of formule.

VOL. XXVI. PART I. 2c

Nicolae PROFESSOR ALLMAN ON THE GENETIC SUCCESSION

Let ¢ be the trophosome, and g the gonosome, then

Le FG KGB OU BAN eae otecens &e.,

S—_ Oe —

will be the general expression for the genetic succession in the life of the
hydroid, the sign + indicating succession by zooidal development, and x by
embryonal.

It is very seldom, however, that the trophosome consists of only a single
zooid. Such rare instances are presented by corymorpha (fig. 1), and by cer-
tain allied forms, whose trophosomes never become developed into a colony of
mutually dependent hydranths, and I believe it better to regard the hydrorhizal
fibres here as elsewhere in the light of mere extensions of the hydrorhizal or
fixed end of the colony, rather than in that of proper zooids—a view supported
by their mode of development in the primordial hydranth. In almost every
other case, the hydranths composing the trophosome become greatly multiplied
by budding.

Still less tendency is there in the gonosome to present an absolutely simple
condition. Indeed, the gonosome is perhaps never limited in its normal state
to a single zooid, and we frequently find hundreds and even thousands of zooids
entering into the composition of this portion of the hydroid colony,

But the zooids of which the colony is thus composed may not only be
numerous, but may also vary in form. ‘Those indeed which constitute the
trophosome are always of a different form from those of the gonosome. In the
trophosome it is rare to find any other form of zooid than that of the proper
hydranth. In Hydractinia, however, there is associated with the ordinary
hydranths the peculiarly modified ones, whose spiral form confers upon the
trophosome of this genus one of its most striking features, while the nemato-
phores of the Plumularide can scarcely be regarded otherwise than as special
zooids whose morphological differentiation from the other zooids of the colony
is carried to a maximum. .

In the gonosome, on the other hand, the usual condition is that of variety
of form among its component zooids; and it is quite common to find in one
and the same gonosome, three different kinds of zooids, each with its special
form among the associated zooids, and its special duty in the generative
functions of the hydroid.

While the type of heteromorphism, or variety of form, among the zooids is
fixed for every species, the polymerism, or simple multiplication of the com-
ponent zooids, is indefinite, and varies with the age, perfection of nutrition, &c.,
of the individual.

If we specialise the general expression already given (I.), so as to make it

OF ZOOIDS IN THE HYDROIDA.

directly applicable to particular cases of heteromorphic succession in the life of
the hydroid, we shall obtain the following formule, where / is used for the
hydranth, b/s for blastostyle, b/ch for blastocheme, and gph for gonophore—

(fig. 2.)

II. Hy n IOI A NIE Kn sith ioaen go Seite ag cale'ses eis os &¢e., Corymorpha. (fig. 1.)
III. Z h+bls+gph x h+ bls + gph X .ccccccceieereees &c., Dicoryne.
ra eae ns
s
IV. & 5h + bls + blch + gph x h + bls + blch + gph x ...&c., Campanularia. (fig. 3.)
g eS TH
S

These formule present three types of heteromorphism. In II. the hetero-

morphism is binary, in III. ternary, in IV. quaternary.

\h ,

a \\)}

/
SI

Fig. 1.—Diagram of Corymorpha. Fig. 2.—Diagram of Dicoryne. Fig. 3.—Diagram of Campanularia

A, the entire colony composed of aaaa, the trophosome, consisting A, portion of the entire colony ; wa.
trophosome and gonosome ; aaa, of numerous zooids; bc, the the trophosome ; bc, the gonosome ;
the trophosome, consisting of a gonosome, consisting of blasto- b, blastostyle ; cc, blastochemes. B,
solitary zooid ; b, the gonosome, style, b, and gonophores, c. a blastocheme become free and
consisting of numerous zooids. mature, and carrying within its bell
B, a single zooid (gonophore) of special zooids, which are the ulti-
the trophosome become free and mate sexual buds or gonophores.
mature,

But the hydranth may and does in almost every instance—either directly
or through the medium of the common basis or hydrophyton—repeat itself

100 PROFESSOR ALLMAN ON THE GENETIC SUCCESSION

indefinitely by budding (fig. 2) before the time arrives when an element of the
gonosome is to be budded off; and a series of homomorphic zooids may thus
introduce themselves into the heteromorphic succession, as expressed in the
following formulee—

V. hththt......&¢. +blst+gph x h+h+h+......&c.+blst+gph x......86¢.
SS eee SSS

where the hydranth becomes indefinitely repeated in the formula of ternary
heteromorphism (III.) given above; and the same will apply to each of the other
two types of heteromorphism.

Now, in all these cases, the succession from the primordial nutritive zooid to
the ultimate generative zooid, or gonophore, admits of being expressed in a
continuous line; but one or more of the zooids of the trophosome may emit
buds which will diverge from the direct line of succession, and which may
then either form the starting-point for another similar line of succession, or
may be destitute of all power of continuing the succession of the zooids. Thus,
(figs. 4 and 7) the primordial hydranth, or any of those derived from it, may
repeat itself by a bud which will diverge from the direct line, produce other
zooids by gemmation, and thus start off a new series, as expressed in the
following formula :—

(+h+th+th+t...... &e. + bls + gph §
NE OM hee: Capa, ah ki Gabet bike anne a ee
7) t

And this state of things may also repeat itself indefinitely, giving rise to an
indefinite number of collateral series diverging from one another, and from the
primary axis of succession.

As already said, however, the diverging zooid may have no power of con-
tinuing the succession. Thus, the spiral hydranth of Hydractinia is not inter-
calated in the direct succession of zooids. It is a diverging zooid, like that
which starts off the collateral series in formula VI., but one which here never
gives rise to buds, and is therefore incapable of either continuing or originating
a new succession.* .

The following formula, where #’ is the spiral hydranth, will express the
place and power of this zooid in Hydractinia :—

: x :..&¢.

: tbls Oph. % PRR &e.
ue | treme 4H

The case expressed in the formule given above is the simple one, where only
the last hydranth in the succession of buds composing a period is supposed to

* The bifurcation occasionally observed in the spiral hydranth of Hydractinia is evidently
abnormal, and cannot be regarded as invalidating the above statement.

OF ZOOIDS IN THE HYDROIDA. 101

give origin to a bud of the gonosome. But any other hydranth in the succes-
sion may just as well bud off a member of the gonosome, which may thus
form a collateral gonosomal axis. This, indeed, is by far the most usual case,
and is what is actually represented in the diagrams (see figs. 2, 4, 7). The
axis, however, thus produced will be necessarily definite, and will contrast
in this respect with the indefinitely extended axis of the trophosome, while it
will differ from the diverging bud, h’ in formula VIIL., by the fact of its having
the power of repeating the colony by sexual reproduction, while h’ has no
power of reproduction, either sexual or non-sexual.

This condition may be expressed by the following formula, in which not
only the last hydranth of the period gives off a bud of the gonosome, but the
primordial hydranth emits a collateral gonosomal axis :—

+h+h+h+...... &e. + bls + gph )
Rall, | ee xb} |} x......€e,

Besides the particular cases now given, certain other modifications of the
plan of gemmation will at once occur to any one who has made the Hydroida
a subject of study. Those here adduced, however, will serve to convey an
adequate idea of the essential features in hydroid gemmation.

It is thus, by the combination of heteromorphic and homomorphic multipli-
cation, and of direct and diverging series indefinitely repeated, that the animal
attains to the condition of those wonderful complex colonies which impress
themselves so strongly on the mind of the observer.

So also the gonosome may present not only a heteromorphic but a homo-
morphic multiplication of zooids. In no case, however, so far as I am aware,
does any zooid of the gonosome repeat itself by homomorphic gemmation, except
in some comparatively rare instances of budding in the medusa; for though
the homomorphic repetition of zooids may be in the gonosome as in the tropho-
some, carried to a great extent, it is almost always the result of budding from
a zooid of a different form. Thus the blastostyle never emits buds destined to
repeat its own form, and this form, however frequently repeated in the gono-
some, is always budded off from the hydranthal element in the trophosome,
its own buds, however numerous, being always heteromorphic with itself.

In the formule now given, one fact is obvious, namely, that the groups
included between every two acts of embryonal development are exactly similar
to one another in the nature and succession of their heteromorphic elements; in
other words, that the life series of the hydroid may be represented by definite
groups of zooids exactly repeated after each generative act.* We are indebted
to Huxtey for having assigned to our conception of the biological individual its

_ * The mere xwmber of zooids in two or more of these groups may of course vary, depending as
this does on the accident of abundant or deficient nutrition and the like.

VOL. XXVI. PART I. 2D

102 PROFESSOR ALLMAN ON THE GENETIC SUCCESSION

proper limits, when he defined it as “the total result of the development of a
single ovum,” and compared the definite groups of zooids which constitute the
life series of animals presenting the phenomenon of “alternation of genera-
tions” to the single organisms known as the individuals, which make up the
species in other animals. These groups form the periods of the series ; the
period repeats itself by true generation, and this repetition continues itself
indefinitely, like a circulating decimal, so as to represent the indefinitely
extended life of the species, while the life of the individual—in its technical
sense as the component of the species—is expressed by each period singly.

—————

Fig. 4.—Diagram of Laomedea.

«wad, hydranths belonging to the primary or direct line of succession ; a’a’a’‘a’, hydranths belonging
to a secondary or diverging line of succession ; b, blastostyle of the primary line of succession, bearing
gonophores, and surrounded by a gonangium ; D’, blastostyle with gonophores and gonangium of the

diverging line.

It is a universal law in the succession of zooids, that no retrogression ever
takes place in the series. In other words, no bud ever becomes developed into
a zooid which is of a different form from the budder, and has at the same time ~
preceded it in the line of succesion. Thus, true hydranths are never emitted
either by blastostyle, blastocheme, or gonophore ; and to this law the peculiar
gemminate hydriform bodies which are found on the summit of the female
blastostyle in certain species of Halecium form no exception ; for though closely
resembling true hydranths, they appear to have a different signification, con-_

OF ZOOIDS IN THE HYDROIDA. 105

tributing probably in some way as yet unknown to the generative functions of
the hydroid, while they have no power of continuing the succession in a direct
or collateral line like the proper hydranths of the trophosome.

The hydranth normally continues the axis in the hydroid colony, just as the
leaf-bud in the plant continues the vegetable axis ; the gonophore, on the other
hand, has no power of continuing the axis, and constitutes the terminal zooid
in each period of the series, just as the flower-bud stops the elongation of the
axis in the plant. This analogy, however, must not be pushed too far, for while
the hydranths and gonophores are simple zooids, the leaf-buds and flower-buds
are complex associations of the corresponding element of individuality in the
plant.

The normal order of succession of the buds in the trophosome is from the
proximal or fixed to the distal or free end of the hydrosoma, so that the older
buds are met with towards the base or hydrorhizal end of the main stem and
branches, the younger ones towards the summit. In the gonosome, on the
other hand, the order of succession is sometimes towards the distal, sometimes
towards the proximal end of the axis. In the calyptoblastic genera, represented
by campanularian, sertularian, and allied forms, the order of succession of the
sporosacs or blastochemes is invariably from the distal towards the proximal
extremity of the blastostyle on which in these genera they are always borne.
When a blastostyle is present in the gymnoblastic or tubularian genera, the
gonophores succeed one another, sometimes from the proximal towards the
distal end (Hydractinia echinata), sometimes from the distal towards the
proximal (Dicoryne conferta). In Tubularia their succession is from the distal
towards the proximal end of the common peduncle, which is more or less
developed in the various species of this genus ; and the same order of succession
occurs in Corymorpha.

Where no special gonosomal axis is developed, the succession is usually
from the proximal to the distal extremity of the branch (Bougainvillia, Perigoni-
mus), thus corresponding to that of the zooids of the trophosome. Sometimes,
however (Syncoryne, Gemmaria), it is from the distal to the proximal.

We have thus, then, in the gonosome of the Hydrozda, as in the inflorescence
of plants, both a centripetal and a centrifugal order of development. It is
possible, however, that irregularities may occur, and that a new bud may be
abnormally emitted at the distal side of a centrifugal series, or at the proxi-
mal side of a centripetal one, so as to disturb in individual cases the normal
sequence of the zooids.

Some further points admitting of comparison with the inflorescence of plants
may be noticed in the gonosome of such hydroids as possess a special gonosomal
axis. In Yubularia indivisa (fig. 5), and in the male colonies of Tubularia
larynx, the gonophores are—like the flowers of a raceme—carried on short

104 PROFESSOR ALLMAN ON THE GENETIC SUCCESSION

pedicels along the sides of a long common peduncle, which springs from the
body of the hydranth. Their order of development, however, is centrifugal, or

Ny
, a
ZW)
ZA a

Fig. 5.—Diagram of Tubularia indivisa. Fig. 6.—Diagram of Tubularia laryna (Female).

aa, hydranth on its stalk; b, shortly stalked gono- aa, a hydranth on its stalk; 6, gonophores at-
phores borne on a common peduncle, and increasing tached by short stalks to a common branched
in maturity from the proximal to the distal extremity peduncle, and increasing in maturity from the
of the peduncle.

proximal to the distal extremities of the
branches.

from the distal to the proximal extremity of the peduncle, so that the whole
group may be compared to a reversed raceme. In the female colonies of Tubu-
laria laryne (fig. 6), and in Corymorpha nutans, the pedicels become branched

Fig. 7.—Diagram of Hudendriwm.
aaaada, hydranthal zooids of the direct line of succession ; a’a’a’, hydranthal zooids of a diverging line ;
b b, suppressed hydranthal zooid, bearing gonophores, which are disposed in an unbelliform group.

with a similar order of development, which thus gives us the compound re-
versed raceme or cyme. ;

In certain proliferous meduse, the buds are borne on the manubrum with

OF ZOOIDS IN THE HYDROIDA. 105

a centripetal order of development, thus giving us, according as the buds are
sessile or pedunculated, the true spike, or the true raceme.

The reversed spike, or spike with a centrifugal development, shows itself in
such forms as Dicoryne conferta (fig. 2, bc); while in Campanularia (fig. 3),
Laomedea (fig. 4, 6b’), Obelia, and other calyptoblastic forms, we have a reversed
spike surrounded by the gonangial sheath ; and were it not for the centrifugal
development of the generative buds upon the blastostyle, and the complete
closure of the gonangium, strongly recalling the spadix with its spathe in the
inflorescence of an araceous plant.

In Ludendrium the male gonophores are disposed in an umbel (fig. 7, 0) with
the axis, in some cases prolonged beyond it, while in others there is little or
no extension of the axis beyond the depressed portion which carries the gono-
phores. Though we cannot here recognise any difference in the order of
development among the gonophores composing the umbel, we are justified
in assuming this order to be as in the true umbel—a centripetal one ; for in the
female colonies of most species of this genus, such as Hudendrium ramosum,

Fig. 8. EHOmO:
A blastostyle of Hydractimia, carrying its gono- A hydranth of Clava with its gonophores surround-
phores, which increase in maturity toward s ile _ ing it in globular clusters.
proximal or attached end.

the gonophores are separated from distance to distance upon the stem imme-
diately below the hydranth ; and here their order of development is plainly seen
to be centripetal.

In Hydractinia echinata (fig. 8) we have the closely approximated gonophores
sessile on a blastostyle, and the development centripetal, as in the true spike,
while the axis extends beyond it as a naked prolongation, reminding us of the
naked prolongation of the spadix in certain Aracee.

In Clava squamata, and in Clava multicornis, the gonophores form dense

VOL, XXVI. PART I. 2E

106 PROFESSOR ALLMAN ON THE GENETIC SUCCESSION.

clusters, surrounding the hydranth in a sort of verticil (fig. 9). Each cluster
consists of sessile gonophores, borne on a greatly depressed common peduncle,
and thus recalling the form of inflorescence known as a capitulum. The order
of development, however, appears to be centrifugal, instead of being, as in the
true capitulum, centripetal, and would therefore, perhaps, more truly suggest
a comparison with the depressed cyme which constitutes the axillary inflores-
ence in many Labiate.

In the comparison just instituted between the gonsome of the Hydroida
and the inflorescence of plants, it will be noticed, that whenever in the
Hydroida the generative buds are borne upon a special gonsomal axis, like
the flowers in an inflorescence, the order of succession is far more frequently a
centrifugal than a centripetal one. In the calyptoblastic forms, indeed, it is
always centrifugal. This is exactly the opposite of what prevails in plants ;
for here the centripetal forms of inflorescence greatly exceed the centrifugal
ones.

We must be careful, however, not to assign to the resemblances which may
be noticed more importance than they are justly entitled to. But yet, after
setting aside such as are merely superficial and accidental, many still remain
which have their origin in certain deep-seated properties, and may be referred
to the common phenomenon of gemmation, which by agamic multiplication in
the animal as well as in the plant, gives rise to colonies whose members in each
case, mutually dependent on one another, continue to be organically associated
into definitely arranged and determinate groups.

ee a da a

(0742)

VI1.—Influence of the Vagus upon the Vascular System. By WitL1AM RUTHER-
ForD, M.D., F.R.S.E., Professor of Physiology, King’s College, London.

(Received, April 1869. Read, 3d May 1869.)*

The innervation of the vascular system is a subject which has engrossed the
attention of physiologists ever since the days of GALEN. Yet, notwithstanding
the number of distinguished observers who have contributed to our knowledge
of this difficult topic, there are still many points of the greatest importance
which are enveloped in the deepest obscurity, and not a few regarding which
opinions are much at variance.

During the past three years I have been more or less engaged in prosecuting
an inquiry, the chief object of which, at the outset, was to ascertain as pre-
cisely as possible the influence which the pneumogastric nerve exerts over the
heart. But, as the investigation proceeded, various ideas started forth which
led me to inquire into the influence which the vagus exerts over certain
vascular territories, more especially the blood-vessels of the stomach. This
line of research, although intricate and difficult to pursue, has nevertheless led
to important results, and has enabled me to throw some light upon the manner
in which the tissues rule over the blood-vessels which minister to their nutrition.

I need not, however, further anticipate here what is fully expounded in the
following pages ; but, before proceeding further, I desire to express my deep
obligations to many of my pupils for the valuable assistance which they afforded
me in the performance of the experiments. My thanks are especially due to
Mr Harninc, Mr Apam, Mr ALLEYNE, Mr Hamitton, and Mr Spence, without
whose skilful co-operation my kymographic experiments must have lacked
much of the precision which they happily possess.

INNERVATION OF THE HEART.

That the heart possesses within itself the conditions necessary for its
rhythmical movement is a theory which was advanced by GALEN, and is now
believed by all physiologists.

The peculiar nervous arrangements essential for the rhythmical movement
are—as REMAK points out—ganglia situated in various parts of the organ.

* An Abstract of this paper was printed in the Proceedings of the above date. Urgent duties
prevented me from preparing the paper in an extended form for the Transactions of 1869. By the
permission of the Council its publication has therefore been delayed for a year.

VOle XXVil, PART I: ; Dar

108 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

It has long been known that the movements of the heart may be influenced
by nerves connecting it with the cerebro-spinal axis. It is unnecessary, however,
that I should enter into a full historical account of this subject, inasmuch as this
has already been given at great length by Von Bezoup.* I need, therefore, only
say that it is now perfectly ascertained that the nerves which convey influences
between the cerebro-spinal axis and heart are branches of the sympathetic and
vagus. The sympathetic filaments take origin in the brain and medulla
oblongata, pass through the cervical portion of the spinal cord, the last cervical
and first dorsal sympathetic ganglia, and from thence to the heart (M. and E.
Cyon).t These nerves convey to the cardiac organ influences which accelerate
its action. Von Bezotp{ thought he had proved that they are continually
prompting the heart to move ; he having observed that on dividing the cervical
portion of the spinal cord—wherein these nerves are contained—that the heart
beats more slowly than it does previous to the injury. He, however, omitted
to take into account the fact, that on dividing the cervical portion of the spinal
cord nearly all the blood-vessels of the body are paralysed, and that the lowered
blood-pressure which results therefrom may be the cause of the slower action
of the heart which follows the lesion. The brothers Cyon found, that although
retardation of the pulse follows division of the spinal cord, no such change is
usually observed if the cardiac motor nerves coming from the last cervical and
first dorsal ganglia are divided, although these same nerves are cut across when
the cervical portion of the spinal cord is divided. We have, therefore, no
reason whatever for supposing that these nerves are continually in action, but,
on the contrary, the evidence advanced by the brothers Cyon is entirely
opposed to such an idea.

It has also been maintained by Von BeEzoxp§ and others, that cardiac motor
nerves are to be found in the trunk of the cervical sympathetic nerve. With
regard to this matter, I have performed many experiments on rabbits, and have
invariably failed to observe any excitement of the heart follow stimulation
of this nerve unless the irritant (electricity) was transmitted through the nerve
close to the inferior cervical ganglion. In that case accelerated cardiac action
often followed the irritation ; but such result is no proof that the trunk of the
cervical sympathetic contains motor nerves for the heart, seeing that the irritant
was applied to the nerve near enough to the inferior cervical ganglion to throw
into action the cardiac motor nerves derived from the spinal cord. I therefore
agree with Lupwic and WEINMANN,|| in considering the cervical sympathetic as
not at all a cardiac nerve.

* Von Brzoup, Untersuchungen iiber die Innervation des Herzens, 1* und 2* Abtheilung.
Leipsic, 1863.

+ M. and E. Cron, Retcnerr and Du Bors Reymonn’s Archivs, 1867, p. 389.

t Lib. cit. 2° Abtheilung, pp. 230 and 257. § Lib. cit. 1° Abtheilung, p. 147.
|| Lupwie’s Lehrbuch der Physiologie, ii‘* Band, p. 178.

UPON THE VASCULAR SYSTEM. | 109

The heart is connected with the vagus by a superior and an inferior branch.
The former, in the rabbit, leaves the vagus with the superior laryngeal nerve,
or it may be somewhat below the origin of the latter ; it courses down the back
in close proximity to the sympathetic, joins one or two branches of the inferior
cervical ganglion with which it proceeds to the heart. In dogs this nerve is
bound up with the trunk of the vagus and cervical sympathetic in one common
trunk; in cats it is joied to the sympathetic. The function of this nerve
was discovered by Lupwie and Cyon.* It is a vaso-inhibitory and also an
excitocardio-inhibitory nerve; that is to say, when it acts it dilates vessels,
and it also excites the filaments of the vagus (inferior cardiac branch) which
inhibit the heart’s movements. The influences which travel through the nerve
start from the heart and pass to the medulla oblongata, there to inhibit the
nerve-cells in the medulla connected with the motor nerves for the abdominal
blood-vessels, and also to excite the nerve-cells in the medulla connected
with the cardio-inhibitory fibres of the vagus. This nerve was named by
the discoverers of its function “ Nervus Depressor,” because it lowers the
blood-pressure ; this it does by diminishing the work done by two great portions
of the vascular system—the heart—and abdominal blood-vessels. The in-
fluences which travel through the nerve pass towards the medulla, probably
their only starting-point is in the heart; but with the cause which determines
the action of the nerve we are totally unacquainted. Its discoverers always
failed to find it in action; that is to say, they never saw the blood-pressure
rise when the nerve was divided. This of course was a very unsatisfactory
circumstance, not a little calculated to cast grave doubts as to the real function
of the nerve having been discovered at all. I am glad to say, however, that in
the course of experiments hereafter to be detailed, I succeeded in finding this
nerve in action on several occasions (see Experiments XLI., XLIV., XLVL.,
LI.) The nerve certainly acts in the manner indicated by Lupwic and Cyon ;
but my experiments do not enable me to state what are the causes of its being
thrown into action.

The inferior cardiac branch of the vagus usually arises with the inferior
laryngeal nerve, and from this origin it proceeds to the heart, where, according
to BEALE,t it joins the cells of the cardiac ganglia. {

* Sachs. Acad. Bericht, 1866, p. 307.

{ Philosoph. Trans. 1863, p. 562, and fig. 41.

{ From physiological evidence it is generally believed that the cardio-motor nerves (sympathetic)

are also connected with the ganglia in the heart. The termination of the depressor nerves within the
heart is quite unknown.

110 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

FUNCTION OF THE INFERIOR CARDIAC BRANCH OF THE VAGUS. Js it motor as
well as inhibitory ?

Effect of Stimulating the Nerve.

In 1845, the brothers WEBER* made the well-known observation that, on
irritating the vagi, or those portions of the central nervous system from which
they spring, that the heart beats more slowly, and may even come to a stand-
still in a state of relaxation. From this observation they concluded that the
vagus exercises an inhibitory power over the heart’s action. The accuracy of
the experiment has ever been beyond dispute, but the explanation, though now
accepted by nearly all—if not by all—physiologists, has nevertheless been
opposed by such distinguished investigators as Scuirr,t Mo.escuort,{ and
ListEr.§. These authorities, while admitting that powerful stimulation of the
vagus arrests cardiac action, maintained that gentle stimulation quickens it.
They, therefore, concluded that the vagus is really a motor nerve of the heart,
and that the arrest of cardiac action which follows powerful irritation of the
nerve, is due to exhaustion of the latter.

As it is unnecessary to slay the slain, I need not adduce the arguments
necessary to show how fallacious is the method of reasoning upon which these
authors have hinged their conclusion,—that has already been ably done by
PFLUGER,|| Von Bezoup,{ and others. I will only make a single remark,
namely this,—were it true that while powerful stimulation slows the heart,
weak stimulation quickens it, the conclusion that both effects must necessarily
be due to the influence of the stimulant upon the same fibres of the vagus is by
no means warranted. It seems to me that the only legitimate explanation
which ScutrrF and others could have given of their facts, is that the excitement
of the heart due to stimulation of the lower end of the vagus—after its section
in the neck—results either from general excitement of the animal, or from the
presence of cardiac motor nerves in the vagus—2z addition to those which
inhibit the heart’s movements; a weaker stimulus being necessary for the —
former than is required for the latter.

It is settled beyond all dispute that the inferior cardiac branch of the vagus
contains fibres which inhibit the heart. The experiments hitherto performed

* Omodei Annali Universali di Medicina, vol. exvi. p. 225, November 1845.

+ Experimentelle Untersuchungen iiber die Nerven des Herzens. Archiv. fiir Physiolog. Heil-
kunde 8‘* Jahrgang.

t Wiener Med. Wochenschrift, 25ter Mai 1861.

§ Proc. Roy. Soe. vol. ix. p. 367.

|| RetcHert’s and Du Bois Reymonnp’s Archivs, 1859, p. 13.

q Lib. cit. Erste Abtheilung.

UPON THE VASCULAR SYSTEM. 111

do not, however, seem to me to conclusively show that cardiac motor fibres
are absent from this nerve. ScHIFF’s statement is that when the vagi are
divided in the necks of rabbits, and the lower end of one or both nerves
very gently stimulated, the heart’s action is quickened. He has further said
that it is difficult to hit upon the precise amount of stimulation which will effect
this. Although other observers of undoubted reputation for skilful experi-
mentation have failed to obtain this result, the above statement is nevertheless
positive evidence which cannot be discarded unless the negative evidence be
very strong. It seemed to me that it was possible to investigate this matter in
a manner more exact and reliable than that adopted by previous experimenters ;
accordingly, I performed a number of experiments in 1866-67, in the following
manner :—In frogs and rabbits I exposed the vagi in the neck, and then opened
the trachea and larynx anteriorly,—in order that asphyxia and consequent ex-
citement might be prevented, and also to enable me to see movements of the
arytenoid cartilages. JI then divided the vagi on a level with the thyroid
cartilage. I always stimulated the nerve by induced currents obtained from
Du Bois Reymonp’s induction machine. The electrodes consisted of clean
copper wire ; the battery of one of DAntELL’s cells. On stimulating the lower
end of the vagus, I always watched the corresponding arytenoid cartilage as
well as the heart. The movement of the former served as a strict test for the
proper application of the electrodes, in short—for the proper stimulation of the
nerve. The observations were begun by ascertaining the strength of current
necessary to affect. the recurrent laryngeal filaments in the vagus—so that
movement of the arytenoid cartilage ensued. Having ascertained this, I made
the current still weaker, and then began the observations on the heart. As is
well known, the strength of the induced currents obtained from Dv Bors Rey-
MOND’S machine depends on the distance between the primary and secondary
coils. The strength is inversely as the distance. Seeing that I began with
very weak currents—that is, with the secondary far removed from the primary
coil—and, being anxious to test the effect of all currents intermediate between
the very weak ones at the commencement and those strong enough to retard
the pulse, I always increased the strength of the current while the nerve was
being stimulated, and the effect upon the heart observed. As long as the
stimulus was not strong enough to slow the heart, the nerve was usually
stimulated for about half a minute. Whenever the animal struggled, the obser-
vation was at once abandoned, and repeated when all excitement had subsided,
—the effect of struggling being to increase the cardiac movement. In the case
of rabbits, the cardiac pulsations were counted with the aid of a stethoscope,
the number being taken previous to and during the stimulation of the nerve.

VOL. XXVI. PART I. 2G

112 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

EXPERIMENTS ON RABBITS.

EXPERIMENT I.—8tH AvcGust 1866.—StronG Rappit. TRACHEA AND LARYNX OPENED.
BotH VAGI DIVIDED IN THE Neck. Dv Bois ReyMonpd’s INDUCTION MACHINE WITH ONE
DANIELL’S ELEMENT EMPLOYED AS THE STIMULATING AGENT.

; : Pulse in 10”,
: Distance in Millimetres of Stake of Larval

Tame, Py Eo Before Irritation of | During Irritation of Muscles, :

Vagus. Vagus.
Cardiac end of
left Vagus,

4:22’ 740 a a0 Contraction.
23’ 800 51 50 Rest.
24’ 800—750 52 52 of
iy 750—730 52, 52 52, 52 Contraction.
26’ 730—700 51, 52 52 As
277 700—670 51 51 3
28’ 670—640 52, 52 52, 52 95
30’ 640—620 yl ay. D2 OL js
il 620—630 Diep O Pail 51, 50, 50 a
Bie 630—640 50, 51 51, 50 se

38’ 640—600 52, 51 50, 51 A
40’ 600—550 51, 50 Syke yay =
41’ 550—500 50, 50 51, 50 55
44’ 500—450 50 50 3
45’ 450—400 50 49 5
48’ 400—350 52, 50 50 E.
Ley 350—300 50, 50 50, 48, 49 3
5A! 300—250 48, 49, 50 49, 50 35
| 59’ 250 50, 49 49, 49 3

pole 250—230 48, 50 33, 29 5
ay 250 48, 48 48, 48 a
8’ 240 48, 49 THe), TS, as

EXPERIMENT II.—Strone Youne RABBIT. TRACHEA AND LARYNX OPENED. BOTH VAGI
DIVIDED IN THE NEcK. 1 DANIELL. :

: ot Rips Pulse in 10”.
; Distance in Millimetres of State of Laryngeal
ee PEE ee Beccary Before Irritation of | During Irritation of Muscles.
; Vagus. Vagus.
Cardiac end of
right Vagus.
10°15’ 550 ss re Contraction.
ate 700 45 45 Rest.
“9) 700—650 44 45 55
val 650—600 46, 45 46, 46 Fs
23’ 600—550 45 45 Contraction.
28° 550—450 46 46, 46 B
30’ 450—400 46 47
3.2’ 450—400 46, 46 46, 46 5
34’ 400—3506 46 46, 45, 46 3
36 350—300 45 45, 44
celtel’ 300—250 44 45, 44 5

Time.

11 o'clock.
. 5/
. 8’
10’

EXPERIMENT III.—Srtrrone Op Rassit,

Time,

UPON THE VASCULAR SYSTEM.

EXPERIMENT Il —continued.

Distance in Millimetres of
Primary from Secondary
Coil.

Pulse in 10’,

Before Irritation of | During Irritation of

115

State of Laryngeal
Muscles.

Vagus. Vagus.
250—220 42 42
220—200 43, 44 44, 44
200—180 42 42
180—170 43 43
170—160 42 42
160—150 43 43
150—140 42 42
140—130 42 38
130—100 42 Stoppage.

Contraction.

TRACHEA AND LARYNX OPENED.

DIVIDED IN THE NgcK. 1 DANIELL.

Distance in Millimetres of
Primary from Secondary
Coil.

Pulse in 10”,

Before Irritation of
Vagus.

5 o'clock.

700
700—650
630
630— 600
600—560
560—460
460—400
400—350
350—300
300—250
250—220
220—200

700
700—650
650—600
600—580
580—500
500—450
450—400
400—350
350—300
300—250
250—200
200—185

150

100

During Irritation of
Right Vagus.

Cardiac end of
left vagus,

BotuH VAGI

State of Laryngeal
Muscles.

Rest.

”?
Contraction.

Rest.

”

bB)
Contraction.

114 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

I might give the results of nine other experiments similar to the foregoing,
but these so thoroughly agree with the above that I think it unnecessary to
detail them. I shall, however, have occasion to refer to some of their results in
the sequel.

EXPERIMENTS ON FROGS.

EXPERIMENT XIII.—Strone Froc. Boru Vaci Divipep. LARYNX LAID OPEN ANTERIORLY.
HEART EXPOSED. PERICARDIUM UNOPENED. 1 DANIELL.

: : ere Pulse in Half a Minute.
; Distance in Millimetres of State of Laryngeal
Tee: a! Ee en any. Before Irritation of | During Irritation of Muscles.
Vagus. Vagus.
Right vagus.
Le A 800 16 16 Rest.
14’ 800—750 16 16, 16 =
ie 750—700 16 16 -
aliG 700—650 16 16 53
2.2’ 650—600 16 16 s
‘26’ 600—550 16 19 Contraction.
30’ 560 16 16, 16 Rest.
34’ 550 16 15, 16 Contraction.
soit, 550—500 16, 16 16, 16 3
“43” 500—450 16 16,16 <,» -
“48” 450—400 16 16, 16 5
"53! 400—350 15 15, 15 5
56’ 350—300 15 16, 15 MS
"59! 300—250 15 15, 15 +
10°4’ 250—200 14 14,13 %
“OE 200 14 14, 12, 12 3
14’ 200—170 14 12, 12, 10 =
20’ 170—140 15, 14 14, 12,11

or 140—100 15 8, Arrest. RS

The left vagus was then irritated, but only with a view to ascertain what was
the feeblest current necessary to produce movement in the larynx, and also the
weakest current which could arrest the heart’s movements. A current at 530
mm. was the weakest which threw the left recurrent laryngeal nerve fibres
into action, while the weakest which sufficed to arrest the heart was one at
170 mm. Further observations on the left vagus were not undertaken, seeing
that the heart’s action had become irregular.

UPON THE VASCULAR SYSTEM. 115

EXPERIMENT XIV.—Strone Froc. Boru Vaci Divipep. LARNYX OPENED. HEART
EXpPosED. PERICARDIUM InTAcT. 1 DANIELL.

= —— |

: ; ate Pulse in Half a Minute.
: Distance in Millimetres of State of Laryngeal |
Tne Faunary ae pocondany Before Irritation of | During Irritation of Muscles.
; Vagus. the Vagus.
Left vagus.
4-14" 700 17 iy Rest.
LG 700—650 17 16 5
19’ 650—600 16 16 9n |
2, 600—580 16 16 Contraction.
25’ 580—550 16 16, 16 »
30’ 550—500 16 NGsG o
34’ 500—480 16, 16 16, 16 ar
39' 480—450 16, 16 16 ‘
“44! 450—440 L617 Wf. We ”
48’ 440—420 75 tte Wi, WG ie
ay 420—400 a, Way Wis ING ay
NEA 400—380 Wi 16 54
+59’ 380—360 16 16 ”
54’ 360—340 16 16, 16 ”
8’ 340—320 16, 16 16, 16 3
pillar 320—300 16 16, 16 .
NY? 300—280 16, 16 16, 16 oS
Sil 280—260 16 16 43
“26° 260—250 16 16 8
30’ 240 16 15 a
“Be 220 16 15 <5
*35" 200 16 16 09
aon. 180 15 16 ms; |
4.9’ 160 17 13 “0 |
44’ 140 17 13 3 |
‘48’ 120 18 12 0 |
lie | 100 21 Arrest. A |

It is unnecessary for me to give the results of other two experiments upon
frogs, seeing that they are precisely similar to the above. The experiments al-
ready detailed amply suffice to show the method of experimentation adopted in
the inquiry. I am at a loss to conceive a mode of research better calculated to
yield accurate results. The stimulation of the recurrent laryngeal fibres of the
vagus served as an index of the effect of the irritant upon the very nerve sup-
posed to contain motor fibres for the heart, and enabled me to judge whether
or not the vagus was being properly stimulated. Hence the fact that I never
observed quickening of the heart’s action follow stimulation of the nerve although
negative in its nature, is, nevertheless, I venture to think, exceedingly reliable
on account of the method of procedure adopted. It may be well, however, for
me to repeat, that I never registered the heart’s pulsations while the animal was

restless ; had I done so, I might have shown that accelerated cardiac action
VOL, XXVI. PART I. 2H

116 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

often follows stimulation of the vagus, but such observations must obviously
have been utterly fallacious, seeing that violent movements invariably excite the
heart’s action. With a view to explain how Scuirr and his supporters obtained
their results, EckHARD* has hinted that possibly the irritating current may have
been sent through the vagus so low in the neck that it affected other nerves in
addition to the vagus. Certainly EckHarp’s conjecture is sufficient to serve as
an explanation of the above, but whether or not it be the true explanation, I
cannot say, inasmuch as I did not see Scuirr and his supporters perform their
experiments.

A striking fact is very clearly brought out by the above mode of experi-
menting—viz., that a very much stronger iritant is necessary so to stimulate
the cardio-inhibitory fibres of the vagus, that the heart’s action may be re-
tarded, than is required to stimulate the recurrent laryngeal fibres, so that the
laryngeal muscles may be thrown into action. The following table demonstrates
this fact :—

TABLE I.—SuHow1nc CoMPaRATIVE STRENGTH OF THE STIMULI NECESSARY TO THROW THE _
INFERIOR LARYNGEAL AND THE INFERIOR CARDIAC NERVES INTO ACTION.

Vagus Divided in Neck, Lower End Stimulated.
Distance in Millimetres of Primary from Secondary Coil indicating
No. of Experiment. Nature of Animal. lee crear arg
(A.) To throw the Laryngeal (B.) To Inhibit the Heart’s
Muscles into Action. Action.
I Rabbit. 740 240
TE, i. 550 130
BET: 7 630 Right Vagus. 210
Do. cake 590 Left - 190
Vi. oa 600 240
Vi #3 575 | 220
VI. Rs 650 250
WE - 520 170
VAL . 610 260
IX. 660 200
X. 5 630 210
aT; 5 580 185
20h, 5 620 200
xennE Frog. 550 200
XaiVe s 580 160
Ove es 620 200
XVI. e 550 200

It may be seen, from the above table, that the strength of current necessary
to stimulate the inferior laryngeal and inferior cardiac filaments in the trunk of
the vagus, differed in different cases. The cause of this is probably threefold :
1st, The strength of the electrical current was not absolutely constant ; 2d, The

* Experimental Physiologie des Nervensystems, 1867, p. 201.

UPON THE VASCULAR SYSTEM. aay

degree of sensibility varies in ‘different animals; 3d, The preparation of the
nerve cannot, of course, be conducted so that precisely the same amount of
injury is inflicted upon it in different cases. When I first elicited the difference
between the results given in column A. and those in column B., it occurred to
me that possibly the inferior laryngeal nerve fibres are more excitable than
those of other motor nerves. But afew experiments on rabbits and frogs satis-
fied me that such is not the case. It is to the last degree unlikely that the
inferior laryngeal nerve is more excitable than the inferior cardiac nerve, and,
therefore, I think, we must look to the peripheral terminations of the two
nerves for the explanation of the facts above given. An ordinary motor nerve
may be supposed to encounter little—if any—opposition when it acts upon the
muscular plasm, but the inhibitory nerve has to act on a nervous apparatus
in which there are counter-influences constantly at work. Only a powerful inhi-
bitory influence can hold these in check, and indeed so powerful are these
promptings to motion within the heart, that stimulation, however strong, of the
inhibitory nerve, cannot keep the heart quite still for more than a few seconds.

Having, in the above manner,” entirely failed to find any acceleration of the
heart follow stimulation of the vagus, another method of experimentation
suggested itself to my mind. It has been shown by Borkrn that atropia
paralyses the cardio-inhibitory fibres of the vagus, that is to say, it so affects
them or their terminations in the heart, that when they are stimulated the
frequency of the pulse is no longer diminished.t I determined to produce this
paralysis, and then see whether or not acceleration of the heart’s action followed
irritation of the lower end of the vagus.

EXPERIMENT X VII.—In a rabbit I divided both vagi in the neck, and stimu-
lated the lower end of the right vagus by a powerful current (Secondary 40 mm.
distant from primary coil of mduction machine. One Daniell). The heart’s
action was arrested. I then injected ten milligrammes of atropia sulphate into
the jugular vein. When two minutes had elapsed, I stimulated the same nerve
with a current of the same strength. The heart’s action, instead of being
arrested as before, was slightly accelerated. Before stimulation of the nerve,
the pulse in 20” numbered 90—during stimulation it numbered 96. After an
interval of four minutes, I stimulated the nerve again with the same current.
Before stimulation the pulse was 104 in 20’—during stimulation it rose to
112. After a further lapse of time, I excited the lower end of the left vagus in
the same manner, but no acceleration of the pulse ensued. The acceleration of

* Should the reader at any time have occasion to repeat the above experiments, he will require to
observe the arytenoid cartilages very narrowly, in order to detect the finest movements which may re-
sult from irritation of the vagus. The animal should be arranged so that the light may be reflected
from the inner surface of the arytenoid cartilage. The slightest movement of the glittering mucous
surface can then be readily detected.

+ VircHuow’s Archivs, Band xxiv., 1862, p. 89.

118 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

the pulse in this case took place when no signs of excitement were exhibited
by the animal, it therefore seemed to indicate that the right vagus, at any rate,
contains efferent cardio-motor nerves* which are not paralysed by atropia
sulphate. The following experiment led me to abandon this idea :—

ExperImeNnT X VIII.—In a rabbit I divided the trunk of the left vagus in ~
the neck, dissected down upon the subclavian artery, and divided the inferior
cardiac branch of the right vagus. I then severed the trunk of the vagus on
the right side of the neck and irritated its lower end with an induced current
(secondary coil 80 mm. from primary coil. One Daniell). The heart’s action
was accelerated. Before stimulation the heart gives 92 beats in 20’—during
stimulation the number rose to 100. On repetition of the above a similar result
was obtained. The acceleration in this case could not possibly be due to the |
action of any motor nerves contained in the inferior cardiac branch of the vagus,
for that branch had been divided on the same side as that on which the trunk
of the vagus was irritated. I, therefore, concluded that in this and in the pre-
ceding experiment, the acceleration of the heart’s action was probably due to a
reflex excitement of the heart resulting from spasm of the laryngeal and —
cesophageal muscles, as well as the greatly increased movement of the stomach
and intestines which follows powerful stimulation of the lower end of the vagus.

I have, therefore, entirely failed to find any evidence to the effect that the
inferior cardiac branch of the vagus contains any cardio-motor fibres in addition
to those which are cardio-inhibitory in their action.

EFFECT UPON THE VASCULAR SYSTEM OF SECTION OF THE VAGI IN THE NECK.

It is well known that division of both vagi in the cervical region is—in the
case of mammals at any rate—usually followed by accelerated cardiac action
and increase of the arterial blood-pressure.

(a.) Cause of the accelerated Cardiac Action.

Rerpt ascribed it to “the struggles and terror of the animal produced by
division of the nerves.” Undoubtedly this is to some extent true, but accelera-
tion of the heart may be observed after division of the vagi during complete —
narcotism produced by opium. Brown-Stquarp{ thought that the excitement
of the heart is due to accumulation of carbonic acid in the blood ;—it being well
known that division of the vagi is usually followed by a slower rate of respira- |

* Since the above was read I have experimented still further with regard to this point. The
experiments which I have performed on rabbits and cats have convinced me thoroughly that the vagus
does not contain “accelerator” fibres for the heart, and that any acceleration of the heart which may
be observed when the lower end of the vagus is stimulated after atropia-poisoning is not due to a direct
action of the vagus upon the heart.

t Physiological Researches, 1848, p. 132.

+ Jl. de la Physiologie, v. p. 656,

UPON THE VASCULAR SYSTEM. 119

tion. Considering that this distinguished physiologist long ago pointed out the
irritating effects upon certain nervous centres, which result from accumulation
of this substance in the blood, it is not surprising that he should have advanced
the above theory. The following experiment shows, however, that acceleration
of the pulse may follow section of the vagi although a hyperoxygenated con-
dition of the blood be maintained before and after the section.

EXPERIMENT XIX.—In a strong rabbit I exposed the vagi, introduced a
canula into the trachea, and then by means of a special apparatus maintained
artificial respiration with such rapidity, that the respiration could be completely
stopped for twenty seconds without slowing of the heart ensuing. It was,
therefore, certain that a hyperoxygenated state of the blood had been fairly
produced.* While care was taken to maintain the artificial respiration at the
same rate, I divided the vagi and watched the results for some time after.
They arethe following :—

Time. Pulse in 15”.
5:12’ Previous to Hyperoxygenation 60 61 59
Palo i division of Vagi oo 60) 58
sh TAS Vagi divided

paar 65 64 64
Aol 66 64 65

The above facts show that a quickened action of the pulse may follow section
of the vagi although the slightest approach to asphyxia is prevented. Further,
recent researches by Voir and RavuBert prove, that until the pulmonary textures
undergo inflammation the increased depth of the respirations after division of
the vagi entirely compensates for their diminished frequency, so that the amount
of oxygen and carbonic acid in the blood undergoes no change.

It is now generally believed that the acceleration of the pulse after division
of the vagi is due to escape of the heart from the restraining influence of these
nerves, and seeing that the acceleration very frequently follows the above-
mentioned lesion, it is inferred therefrom that the cardio-inhibitory fibres of the
vagi are in almost constant action. It occurred to me that if this explanation be
the true one, and the only one, we should expect to find no acceleration of the heart
follow division of the vagi after their cardio-inhibitory fibres have been paralysed
by such a substance as atropia. Accordingly, I performed a number of experi-
ments with a view to test this point; but as these bear equally upon the
following question, I shall briefly allude to it before proceeding further.

* Usually within three seconds after the respiration of a rabbit is arrested the heart comes almost
to a stand-still. This is due to irritation of cardio-inhibitory nerves by the asphyxiated condition of
the blood.

+ Centralblatt. 1868. No. 47.

VOL. XXVI. PART I. P|

120 DR RUTHERFORD ON THE INFLUENGE-OF THE VAGUS

(b.) Cause of the Increased Blood-Pressure.

During the operation of the cardio-inhibitory nerves, the work done by the
heart is diminished (see fig. 2). The rise in the blood pressure, which com-
monly follows section of the vagi, is therefore ascribed by all to increased force
and frequency of the heart’s contractions. If this be the only cause of the rise
in the blood-pressure, then we ought to find that it undergoes no exaltation on
division of the vagi during a paralysed state of their cardio-inhibitory fibres.

The following experiments were undertaken with a view to furnish an
answer to this question. Do accelerated cardiac action and increased blood-
pressure follow division of the vagi—during paralysis of their cardio-inhibitory
fibres—produced by such a substance as atropia sulphate ?—

EXPERIMENTS SHOWING THE EFFECT UPON THE BLOOD-PRESSURE AND FREQUENCY
OF THE PULSE WHICH SOMETIMES FOLLOWS SECTION OF THE VAGI IN ANIMALS
WHERE THE CARDIO-INHIBITORY NERVES ARE PARALYSED BY SULPHATE OF
ATROPIA,.

EXPERIMENT XX.—A SMmAzLu TERRIER DoG. CANULA IN CAROTID ARTERY.
TRACHEA OPEN.

Time. Pulse in 15”. — er He oe wes General Notes.
11°44’ 16 4:5
47’ 16 4:5
30” 0°67 milligramme atropia sulphate
injected into vein.
48’ 30” 40 4°65
50’ 30” 29 4:2
52’ 22 4-4
30” 04 milligramme atropia sulphate
injected into vein.
53’ 35 4:2
5A’ 29 4
55’ 30” 25 3°9
56’ Right vagus divided in the neck.
ue a0 23 4
58307 Left vagus divided in the neck.
59 56 5°9
45” 50 59
Wee, 42 6-4
8’ 43 6°45
30” Distal end of left vagus irritated
by a strong induced current, but
r no effect was produced on the
heart’s action, clearly showing
that the cardio-inhibitory nerves
were completely paralysed.

* Tn all these experiments Lupwie’s Mercurial Kymograph was used.

UPON THE VASCULAR SYSTEM. 121

In the above, notwithstanding the paralysis of the inhibitory nerves of the
heart, section of the vagi was followed by a most distinct increase in the fre-
quency of the heart’s contractions, and a rise in the blood-pressure.

EXPERIMENT XXI.—A MIDDLE-SIZED CoLuiEe Doc.

Time. Pulse in 15”.
13" 29
0’
10’ 3)
30” 35
ale
i 36
es 33
153 32
30”
eG’ 35
20° 30"
DH 39
93° 39
28’ 40
29’

TRACHEA OPEN.

CANULA IN CAROTID ARTERY.

Mean Pressure in inches

General Notes.

of Hg.
59
0°67 milligramme atropia sulphate
injected into vein.
5°6
53
0°4 milligramme atrophia sulphate |
injected into vein.
52
56
53 |
Left vagus divided. |
5'8
Right vagus divided.
6°6
68
6°8

Cardio-inhibitory nerves proved to |
be paralysed.

EXPERIMENT XXII.—A Spanre, DoG, FIVE MONTHS OLD. CANULA IN CAROTID ARTERY,

TRACHEA OPEN.

Mean Pressure in inches

Time. Pulse in 15’. of Hg.
11:19’ 27 3°6

20’ 30” 26 4°6

oe 26 4°05
30”

Q4’ 30 4°05

Pile 38 4:05
30”

ey B04 66 4:3

30’ 60 4°]

Bie BO? 60 34)

35’ 50 ay)

40’ ays) 4°]

4]’

49’ 60 An3
30”

45/ 68 51

47’

General Notes.

0°67 milligramme atropia sulphate
injected into vein.

0-4 milligramme atropia sulphate
injected into vein.

Left vagus divided.
Right vagus divided.

The cardio-inhibitory nerves were
found to be completely paralysed.

122 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS.

EXPERIMENT XXIII.—OLD SpanreL Doc. CANULA IN CAROTID ARTERY. TRACHEA OPEN.

Time. Pulse in 15”. — ism aye General Notes.
4,28’ 25 51
30” 2 milligrammes atropie sulph. in-
jected into vein.
29’ 46 vA
34’ Clot in canula. Apparatus cleaned.
| 49’ 42 4°9
43’ 38 4:3 Hitherto the respiration has been
rapid and irregular.
48 40 : 4:7 Animal sobbing.
49’ 20 minims Tincture Opii given.
30” 40 61
52’ Clot in canula. Apparatus cleaned.
54 33 4°6
Doe 0°13 milligramme atropie sulph.
given.
56’ 30” 34 4°6
59’ Left vagus divided.
5 o'clock, Clot. Apparatus cleaned.
3’ 30° —! 4°9
4’ Right vagus divided.
| 5 30" 41 51
| 6’ 30° 40 5°61
| os 43 5:2
12’ 44 a4
13’ Cardio-inhibitory nerves found to
| be completely paralysed.

These experiments show that in dogs division of the vagi in the neck may
be followed by accelerated cardiac action and increased blood-pressure, although
the cardio-inhibitory nerves be paralysed. It is clear, therefore, that physiolo-
gists generally are mistaken in supposing that the acceleration of the pulse
which commonly follows division of the vagi, is entirely due to the heart’s being
liberated from its controlling nerves. A portion—perhaps, in some cases, the
whole—of that acceleration may be due to division of other filaments than those
which retard the heart’s action. I shall not, at present, enter ito a discussion
of the causes of the increased blood-pressure observed in these experiments.
The reason for this will be readily perceived when my remarks upon other
experiments performed on dogs have been perused. (See page 137.)

The consideration of these changes which follow division of the vagi will be
resumed after we have inquired into the present state of our knowledge regard-
ing the

INNERVATION OF BLOOD- VESSELS.

By the investigations of BERNARD and Brown-SEQuARD, it has been estab-

UPON THE VASCULAR SYSTEM. 125

lished that the contractile elements of the blood-vessels are supplied by motor
nerve filaments derived from the sympathetic. Diminution in the calibre of the
blood-vessels is produced by these nerves. According to Lupwic and Turry,
the general centre for the vasomotor nerves is situated in the medulla oblongata.
This cerebro-spinal centre is more or less constantly in action, whereby vessels
are usually kept in a semi-contracted state. The amount of contraction in the
vessels—in other words, the degree of activity of the cells in the vasomotor centre
—may be increased or diminished by certain nerves which convey influences to
the medulla. BERNARD* was the first to show, by experiment, that vessels may
be dilated by the irritation of certain nerves. He found that when he divided
the auricular nerves in rabbits, and excited their central ends, the vessels of the
ear of the same side became turgid. Slight contraction preceded the dilatation.
Lovént has confirmed BERNARD’s observation, and has shown that dilatation of
vessels in the rabbit’s leg follows irritation of its afferent nerves ; in short, that
dilatation of the vessels of a part may be produced by influences transmitted
through the afferent nerves of that part to the cerebro-spinal vasomotor centre.
Like BernarD, he found that transient contraction generally precedes the dila-
tation of the vessels so induced. The most remarkable instance of a nerve
capable of dilating vessels is to be found in the superior cardiac branch of the
vagus already alluded to. When this nerve is divided and its cranial end
stimulated, dilatation of abdominal blood-vessels takes place without any previous
contraction, such as commonly results when a mixed nerve such as the sciatic
or the trunk of the vagus is stimulated. In all the above cases the vascular
dilatation succeeds stimulation of the central ends of the divided nerves ; that is
to say, the peripheral end of the cranial portion of the divided nerve. Two
facts, however, have been discovered which are opposed to the idea that the
motor centre for all the blood-vessels of the body les in the medulla oblongata;
one concerns the submaxillary ganglion, the other, the ganglia upon the nervi
erigentes of the penis. It is well known that if the chorda tympani nerve be
divided, and its peripheral end stimulated, dilatation of the blood-vessels in the
submaxillary gland is the result. In like manner, as recently shown by
EcxuArp{ and LoveEn§, when the nervi erigentes are divided in the dog and the
peripheral portions stimulated, erection of the penis results, principally from the
dilatation of vessels induced by the irritation. On these nerves there are many
ganglionic corpuscles ; and the most feasible explanation of this vascular dilata-
tion in the case of the submaxillary gland and penis is, that the ganglionic cells
existing in connection with these structures, are in part, at any rate, vasomotor
cells, and correspond to the ganglia in the heart. These three groups of ganglia

* Ji. de la Physiologie, 1862, p. 416. { Beitrage. Giessen, 1863.
+ Lovin, Ludwig’s Arbciten, 1866, p. 1. § Lib. cit. p. 18.
VOL. XXVI. PART I. 2K

124 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

forming three peripheric motor centres connected with the vascular system, the
only ones as yet known. But it is interesting to observe that the submaxillary
and cardiac ganglia are obviously very directly connected with the medulla
oblongata by motor and inhibitory nerves, and although nothing can be stated
with precision regarding the central connections of the ganglia presiding over
the vessels of the penis, they have, nevertheless, probably an intimate con-
nection, like the others, with the medulla. The dilatation of vessels which
results from the action of these vaso-inhibitory nerves is, as regards the vessel,
passive ; it is due to the elasticity and blood-pressure being no longer opposed
by the action of the contractile elements of the vascular wall, these having been
brought to rest by a cessation of action in the vasomotor nerve apparatus.

The vasomotor nerves may have their action increased as well as diminished
by the action of other nerves. The contraction of vessels in distant parts by
the sudden application of cold to it may be a small extent of skin. The remark-
able increase in the blood-pressure which follows stimulation of the central end
of the superior laryngeal nerve (AUBERT and Roever),* the contraction of
vessels and increase of blood-pressure which usually follows the gentle stimula-
tion of the central ends of mixed nerves, are some of the facts which support
this idea; such nerves may be very appropriately termed excito-vasomotor
nerves. These nerves appear all to pass inwards to the vasomotor centre in
the medulla oblongata.

Other facts might be mentioned, but I may briefly say that every advance in
our knowledge of this question only tends more and more convincingly to show
that the innervation of the contractile elements of the blood-vessels is similar to
that of the cardiac muscular fibres. These contractile elements are directly
supplied by motor nerve fibres ; and the evolution of energy in the cells con-
nected with the latter may be diminished by one set of nerve fibres—cardio and
vaso-inhibitory—and increased by another—excito-cardio and _ excito-vaso-
motor.

The idea commonly prevails that when a part becomes the seat of active
nutritive change, its blood-vessels undergo dilatation by reason of the increased
attraction for blood manifested by the tissues. The vis & fronté is supposed to
become so powerful that it can overcome the contraction of the arterial walls,
and thereby produce dilatation. It struck me that the vascular dilatation in
such a case is possibly the result of an imfluence transmitted by the tissue
through its vaso-inhibitory nerves. The only author who has come near to this
idea is LovEn. A considerable time after the above had presented itself to my
mind, he published the excellent memoirt to which reference has already been
made. He showed that the blood-vessels of a part may be dilated by artificial

* Centralblatt, 1868, p. 578. + Lib. cit.

UPON THE VASCULAR SYSTEM. 125

stimulation of the afferent nerves of that part. There, however, he stopped.
He has advanced no theory regarding the bearings of this fact upon our con-
ceptions of the mode in which vascular dilatation in a part commonly takes
place, nor has he thrown out any suggestion as to the agent by which these
nerves are normally brought into play. I believe that the experiments which
I have yet to detail will be found to very decidedly advance our knowledge
regarding this matter.

The vasomotor nerves for the blood-vessels of the stomach are contained in
the splanchnic nerves. The vaso-inhibitory and excito-vasomotor nerves of
that organ appear to me to be for the most part, if not entirely, contained in
the pneumogastric nerves. If it be true—as I imagine—that when the gastric
blood-vessels undergo dilatation, vaso-inhibitory nerves are brought into play,
we should—if these nerves be contained in the vagi—expect to find that if the
vagi be divided during dilatation of gastric blood-vessels, the vessels will undergo
contraction, and we should desire to see that stimu-
lation of the upper end of the divided nerve is able
to produce dilatation of vessels. In order that the
sequel may be better understood, I would refer the | mo.
reader to the following diagram representing the
stomach, s; the vagus, v; the splanchnic nerve, sp ;
the medulla oblongata, m. o.; and the spinal chord, VA |
s.c. The arrow near the vagus indicates the direc- ds
tion in which vaso-inhibitory and vaso-excito motor - a
influences travel through the vagus to affect the vaso-

motor centre in the medulla oblongata—while the dS | / v| |
See thee /
\ /
\ /
See ee

Fig. 1.—Diagram showing innerva-
tion of Gastric Blood-vessels.

arrow near the splanchnicus indicates the direction
in which vasomotor influences travel through that
nerve to the gastric blood-vessels.

The following experiments were undertaken to gtomach, s Sree @ondeec ite:
ascertain whether or not one can obtain evidence Seamer Rees OS Mages:
of the passage of vaso-inhibitory influences through
the vagus during the dilatation of the blood-vessels of the stomach which takes
place during digestion, and also to ascertain the effect of irritating the vagus
upon the gastric blood-vessels :—

126 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

INFLUENCE OF THE VAGI UPON THE BLOOD-VESSELS OF THE STOMACH.

Effect of Division and Irritation of the Nerves.
(a.) During Digestion.

EXPERIMENT X XIV.—Rabbit two months old. Three hours after food was
given the vagi were exposed, and the trachea opened in order to prevent
asphyxia and consequent struggling. The abdomen was opened, and the
stomach found to be largely distended with food, and its outer surface very
vascular. Intestines were moderately vascular. The stomach was then opened
by an incision extending from right to left along its anterior surface, and its
contents partially everted. The lining membrane was of a dusky red hue. The
vagi were divided in the neck four minutes after the exposure of the gastric
mucous membrane. Pallor of the membrane followed immediately upon the divi-
sion of the nerves, and remained during the time occupied by the rest of the
experiment—forty-five minutes. The superior cardiac branches of the vagi
(depressor nerves) were then divided ; the result was a slight increase in the
pallor of the mucous membrane.

The effects of irritating the vagi were now attended to. The irritant used
was Farapic Electricity from Du Bors REymonn’s machine, with 1 SMEE’s cell.

ine eee Sonic con anne Nerve Stimulated. Pobre Sena Fer
o* in Millimetres. j 5
10’ 230 Upper end left vagus. No evident change.
12’ 180 Upper end left vagus. Became redder.
Si 10 milligrammes atropiz sulph.* given to paralyse cardio-inhibitory fibres
of vagus.
| 15’ 180 Lower end left vagus. No evident change.
| 2,14 120 Upper end right vagus. Pallor followed by slight
| redness.
25° 80 Upper end right vagus. Pallor.
34’ i) Lower end right vagus. No evident change.
44’ 70 Lower end left vagus. No evident change.

EXPERIMENT XX V.—Full-grown strong rabbit fed two and a half hours
before the experiment was begun. Abdomen opened ; stomach and intestines
very vascular. Division of the vagi was followed by decidedly diminished
vascularity of the outer surface of the stomach which was in this case unopened.
Owing to an interruption the experiment was not carried beyond this point.

* Although this substance paralyses the cardio-inhibitory, it does not paralyse the vaso-inhibitory
nerves.

UPON THE VASCULAR SYSTEM. 127

EXPERIMENT X X VI.—Strong full-grown rabbit. Fed two hours before the
experiment. The cavity of the stomach was not opened. When the superior
cardiac branches* of the vagi were divided, no evident change resulted in the
vascularity of the stomach or intestines. On dividing the vagi the whole outer
surface of the stomach became paler ; but no such change was observed in the
mesentery or intestines. The vascular change in the stomach was permanent.

The upper ends of both vagi were then repeatedly stimulated, with variable
results. Sometimes the surface of the stomach became paler, at other times
redder. Irritation of the lower ends of the nerves produced no effect.

_ Experiment X X VII.—Rabbit ; fed an hour before the vagi were divided.
The outer surface of the stomach was not apparently so vascular as in the three
former cases. Division of the vagi produced no evident change on the gastric
vessels.

EXPERIMENT XX VIII.—A cat fed on milk an hour previous to division of
the vagi. On section of these nerves the vascularity of the outer surface of the
stomach instantly became greatly diminished, and remained so.

(b.) Section of the Vagi during Fasting.

EXPERIMENT X XT X.—Full-grown rabbit which had fasted for twelve hours.
Outer surface of stomach pale. Division of depressor nerves produced no
evident change in gastric or intestinal vessels. Division of both vagi likewise
produced no evident change in gastric or intestinal vessels. Irritation of upper
end of right vagus (1 Smeg, secondary 200 mm. distant from primary coil)
caused slight reddening of outer surface of stomach, no change on intestinal!
vascularity. This observation was repeated with a stronger current (secondary
coil at 150). A slight increase in the pallor of the stomach was the immediate
result, but this yielded during the continuance of the irritation to distinct red-
dening of the gastric wall. The irritation was continued for twenty seconds.
Anesthesia was then produced by means of chloroform in order to get rid of
the effects of the irritant upon the sensory nerve centres. The upper end of
the right vagus was then stimulated (secondary coil at 100). Slight but distinct
increase in the gastric vascularity at once ensued. Irritation of the upper end
of the left nerve yielded the same result. Irritation of the lower ends of the
nerves caused no change.

EXPERIMENT XX X.—Full-grown rabbit which had fasted for fourteen hours.
Division of the nervi depressores and vagi produced no evident change on the
vascularity of the outer wall of the stomach or of the intestines, which both
before and after the division were but slightly vascular.

* These nerves produce dilatation of abdominal blood-vessels.
VOL. XXVI. PART I. 2 L

128 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

ee eee Seeeline Goll of Eetuction Nerve Stimulated. Result as regards Gastric
gi. Machined. Vascularity.
mm
ve 200 Upper end of right vagus | Slight reddening.
irritated for 30”.
12’ 150 Upper end of right vagus | Pallor succeeded by well-
irritated for 30”. marked redness.
135’ 120 Upper end of right vagus | The same result.
| | irritated for 15”.
Chloroform was now given until complete anesthesia resulted.
20’ 120 Upper end of right vagus | Slight increase of pallor suc-
irritated for 30”. ceeded by well-marked
redness.
24’ 80 Upper end of right vagus | The same result.
irritated for 30”.

EXPERIMENT XX XI.—Cat which had fasted for sixteen hours. The blood-
vessels of the outer surface of the stomach were small and contracted. Division
of the nervi depressores and vagi produced no apparent change in the vascu-
larity of the stomach or intestines. Irritation of the upper end of the vagus
caused tolerably distinct blushing of the gastric wall.

The general result of the experiments just given is, that section of the vagi
produces no change in the gastric blood-vessels if these be not in a dilated con-
dition, such, ¢.g., as obtains durig digestion, while division of these nerves
during dilatation of the gastric vessels is generally followed by marked and
permanent contraction of these vessels. It is true that this result did not occur
in one (experiment xxvii) of the five experiments in which the nerves were
divided during digestion. In that case, however, it was quite evident that the
gastric vessels were not so dilated as they usually are; but, of course, such a
statement is not without fallacy, seeing that every case cannot be identical as
regards the vascular dilatation that obtains during digestion. I am not, there-
fore, prepared to give any decided opinion regarding the results of experiment
xxvii ; but it seems clear that the general result of the effects of division of the
vagi supports the idea that, during digestion, vaso-inhibitory influences pass in
a centripetal direction through the vagi. The effects of irritating the cut ends
of the nerves were various. It is certain that no evident change in the vascu-
larity was ever produced by stimulating the lower ends of the nerves, so we
may safely say that the influences which pass through the nerves to control the
gastric vessels certainly do not pass in a centrifugal direction. When the upper
ends of the nerves were subjected to sufficiently powerful stimulation, pallor of
the gastric wall sometimes followed, at other times blushing; frequently the
blushing succeeded the pallor, and sometimes no perceptible effect resulted.

UPON THE VASCULAR SYSTEM. 129

These results receive a feasible explanation by the supposition that the vagus,
like other mixed nerves, contain fibres which excite, and those which inhibit
contraction of the vessels. Because the vagus is a mixed nerve, the results
of its division must obviously be more trustworthy than the results of its
stimulation. We cannot suppose that while during digestion influences pass
from the stomach through the vagi to inhibit the gastric vessels, there are
also influences travelling from the same source which produce an opposite
effect ; and, therefore, we may expect that when we divide these nerves
during the dilatation of vessels which obtains during digestion, we shall simply
stop the transit of those vaso-inhibitory influences from the stomach, hence
the division of such nerves is a much simpler case than artificial stimulation,
seeing that during such stimulation we must throw into play fibres whose
functions are antagonistic.

Seeing that the experiments just given show what are the evident changes
in the gastric vessels that follow stimulation of the vagi, it is convenient to give
here results of experiments which show the effect of this stimulation upon the
arterial blood-pressure.

EFFECT UPON THE ARTERIAL BLOOD-PRESSURE WHICH FOLLOWS STIMULATION OF
THE VAGUS AFTER ITS SECTION IN THE CERVICAL REGION.

(a.) Stemulation of the Lower End of the Nerve.

If the lower end of the vagus be stimulated by a sufficiently powerful cur-
rent, the heart’s action is retarded, the work done by that organ is diminished,
and in consequence the arterial blood-
pressure falls. The following tracing by
Lupwic’s Kymograph from the carotid
artery of a rabbit affords a good illustra-
tion of the above fact. The tracing must
be read from right to left. The vertical
lines have been added to show when the
vagus was at rest and when it was stimu- See nee | eI
lated. The portion of the tracing hetween i
the two lines shows the influence of the cardio-inhibitory fibres of the vagus
upon the heart.

The above trace shows the influence of the vagus upon the heart, but it con-
tains no indication as to whether or not the vagus contains vasomotor fibres.
This fact can only be ascertained by stimulating the nerve after its influence
over the heart has been got rid of. I accordingly paralysed the cardio-inhibi-
tory fibres by sulphate of atropia, and then stimulated the lower end of the
nerve as before. The results were various. If the animal were not paralysed

Fig. 2.

150 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

by curara, a slight rise in the blood-pressure frequently resulted from the
stimulation. Such a result is shown in the following tracing from a rabbit.

Fig. 3.

Rapesco

During— Before—
Stimulation of the Vagus.*

A change similar to the above is; however, by no means the rule even in the
same animal. The following tracing shows no change in the blood-pressure
during stimulation of the vagus, although it was taken shortly after the fore-
going from the same animal, and although the lower end of the same vagus was
stimulated by a current of the same strength.

Fig. 4.

snot ne

During— Before—
Stimulation of the Vagus.

This tracing represents what I always found when the atropised vagus was
stimulated in cases where the influence of extraneous movements upon the
blood-pressure had been got rid of by means of curara paralysis. We may,
therefore, say, that the vagus certainly contains no vasomotor-nerve fibres
which act in a centrifugal direction, for if it did, stimulation. of the nerve
after palsy of its cardio-inhibitory fibres would always raise the blood-pressure
whether curara be given or not.

(b.) Stimulation of the Upper End of the Nerve.

Already much has been written with regard to the changes in the blood- —
pressure which result from stimulating the upper end of the vagus after it has
been cut across in the cervical region. According to DRESCHFELD+ such stimu-
* Jn reading tracings taken by such an instrument as Lupwie’s Kymograph, it is necessary to
remember that the vertical variations in the mercurial column are always the double of what the tracing

indicates, because the tracing shows the movements of a column of mercury in a U-shaped tube.
+ Von Brzoup’s Untersuchungen, 1867, p. 326.

UPON THE VASCULAR SYSTEM. 131

lation always raises the blood-pressure, but if the cerebrum be removed,
or if it be paralysed by morphia, vagus stimulation always lowers the
pressure.

This is simply untrue. Stimulation of the nerve may increase or lower the
‘pressure whether morphia narcotism be induced or not. Since I performed
my experiments on this subject, KowaLewsky and ADAMUK,* AUBERT and
RoEvert have published the results of their researches regarding this question,
and I am glad to say that these exactly agree with what I had previously found.
Seeing that these authors have already published results similar to mine, I
need not do more than briefly say, that when the upper end of the vagus is
stimulated, the respiration is very apt, more especially in rabbits, to come to a
stand-still. Asa result of this, carbonic acid accumulates and oxygen diminishes
in the blood, thereby bringing about a condition of that fluid which acts as an
irritant to the vasomotor centre in the medulla, and increases the tonicity of
the blood-vessels so that the blood-pressure is raised. This source of fallacy
must be guarded against by using artificial respiration. Struggling, too, is apt
to result from stimulation of the upper end of the vagus, to guard against which
we may narcotise the animal by means of opium, or may produce paralysis by
curara. When we give opium or curara and then stimulate the nerve, a rise in
the blood-pressure is not so frequently observed as when the nerve is stimulated
before these poisons are administered; obviously because extraneous convulsive
movements have been got rid of. However, whether we give these toxic agents
or not, stimulation of the upper end of the vagus in rabbits and cats (where the
depressor nerve is a separate branch) may be followed by increase or by
diminution of the blood-pressure, most frequently the latter. I have often ob-
served that in the same animal a rise or fall of the blood-pressure may be
obtained by using for the production of the latter a more powerful stimulus than
that which may have been found sufficient to produce the former. The explana-
tion of this seems to be, that in the case of the vessels, as in that of the heart,
a weaker stimulus suffices to throw the excito-motor nerves into action than is
necessary to cause the inhibitory nerves to produce their effect. The following
tracings illustrate the results of stimulating the upper end of the vagus. They
must be read from left to right.

The fibres in the vagus, then, which influence 0b/ocd-vessels, all convey in-
fluences towards the medulla oblongata, and these fibres appear to be both
vaso-inhibitory and excito-vasomotor, the former causing dilatation of blood-
vessels and consequent lowering of the blood-pressure (fig. 5), the latter caus-
ing contraction of blood-vessels and consequent increase of the blood-pressure
(fig. 6). Doubtless the influences which travel through these two kinds of

* Centralblatt, 1868, p. 546. + Ibid. p. 578.
VOL; K&XVI. PART I. 2M

1382 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

nerve-fibres start from the parts to which the vagus is distributed, principally,
therefore, from the stomach and lungs. |

Fig. 5.—Fall in blood-pressure on irritating upper end of vagus in a rabbit (both vagi divided), to which 10 milli-
grammes of atropia sulphate had been given. (Secondary 30 mm. distant from primary coil. One Daniell.)

Before— During—
Stimulation of the Vagus.

_ Fig. 6.—Rise in blood-pressure on irritating upper end of vagus in the same rabbit as that from which the fore-
going tracing was taken. (Secondary 120 mm. distant from primary coil. One Daniell.)

Before— During—
Stimulation of the Vagus,

Having seen what are the effects upon gastric blood-vessels of division and
stimulation of the vagi, and also the changes which the latter gives rise to as
regards the blood-pressure, we shall now consider the effect of division of the
nerves as regards the blood-pressure. The experiments on each class of animals
are divided into two groups : 1st, those showing the effect of dividing the nerves
during digestion, and those showing the effect of this during fasting. The
main object of the whole being to ascertain whether or not during the former,
vaso-inhibitory influences are transmitted from the stomach through the vagi to
diminish the action of the vasomotor nerves ruling over the gastric blood-
vessels, and thereby to bring about dilatation of these. While this is the main
point of the experiments, they at the same time furnish data which serve to
explain the acceleration of the pulse which sometimes follows division of the
vagi after paralysis of their cardio-inhibitory fibres (see page 122),

UPON THE VASCULAR SYSTEM. 133

EXPERIMENTS SHOWING THE EFFECT OF DIVISION OF THE VAGI UPON THE BLOopD-
PRESSURE AND FREQUENCY OF THE PULSE IN ANIMALS DURING DIGESTION,
AND DURING FASTING.

A. EXPERIMENTS ON DOGS.
(a.) During Digestion.
Cardio-inhibitory Nerves paralysed by Atropia.

EXPERIMENT XXXII.—RETRIEVER Puppy ABOUT THREE MONTHS OLD, FED THREE HOURS
BEFORE THE EXPERIMENT. CANULA IN CAROTID ARTERY. TRACHEA OPEN.

Time. Pulse in 15”. MSE Eewue HA Tae General Notes.
4°51’ 34 4°5
59’ 30” 3 milligrammes atrop. sulph. in-
jected into vein.
ESO elias 46 47
5’ 46 4:7
12’ Both vagi divided.
16’ 52 Ds
21’ 50 61
27’ 50 59
33’ 54 6:3
34’ Cardio-inhibitory nerves found to
be completely paralysed.

Result.—Increase of pressure and acceleration of pulse after division of vagi.

EXPERIMENT XXXIII—A Terrier Doc, rep aT 1.30 P.M. CANULA IN CAROTID
ARTERY. TRACHEA OPEN.

Time. Pulse in 15”. Micon Eressuie un mches General Notes.
of Hg.
332) 20" 30 5.7
50” 30 57
37’ 1 milligramme atrop. sulph. injected
into vein.
5 ial Fa 56 6°6
40’ 62 6:7
45° 56 6°7 Vagi not paralysed.
55 56 6°2
50’ 1 milligramme atrop. sulphate given.
20” 56 6:2
53’ 50” Both vagi divided.
55’ 62 74
35, 60 1G
58’ 55” . 54 7
DiC leeya Cardio-inhibitory nerves found to
° be quite paralysed.
|

Result,— Permanent increase of pressure, and temporary acceleration of
pulse after division of vagi.

134 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

EXPERIMENT XXXIV.—A StroneG RETRIEVER DoG, FED TWO HOURS BEFORE THE
EXPERIMENT. CANULA IN CAROTID. TRACHEA OPEN.

Time. Pulse in 15”. ee General Notes.
of Hg.
12°14’ 32 6
es 32 6
23° 2 milligrammes atrop. sulph. in-
jected into vein.
25° 28 4°8
27 30 i)
| 31’ 307 30 54 Vagi not quite paralysed.
32’ 1 milligramme atrop. sulph. given.
30” 42 4°]
35’ 36 4°8
36’ 32 4°8
38’ 30” 35 5
43’ Both vagi divided.
50’ 38 6:4
52’ 35 68 |
saa 34 67
4’ Cardio-inhibitory nerves still para-
lysed.

Result.—Permanent increase of pressure, and temporary acceleration of
pulse after division of vagi.

EXPERIMENT XXXV.—SMALL Doc FED AT ONE 0o’CLOCK. CANULA IN CAROTID.
TRACHEA OPEN.

Time. Pulse in 15” te oe Pree, ee General Notes.
5°49’ 30 5
50° 2 milligrammes atropia sulphate |
injected into vein.
40” 70 5°8
53” 66 54
54’ 10” 66 5 Both vagi divided.
Ai
56’ 64 a1
59’ 60 uD
6 20 56 6°4
5! 56 6°5
10’ 53 6°6

Result.—Increase of pressure after division of vagi. The frequency of the
pulse was diminished; but it is doubtful whether or not a similar decrease
would not have taken place had the vagi remained intact. When the pulse un-
dergoes a great increase in frequency on the administration of atropia, as in the
present instance, a steady decrease almost always sets in shortly afterwards.

ee ee

UPON THE VASCULAR SYSTEM. 135

(b.) During Fasting.

1. Cardio-inhibitory Nerves Paralysed by Atropia.

EXPERIMENT XXXVI—A Srrone RetrrieveER DoG WHICH HAD FASTED FOR SEVENTEEN
Hours. CANULA IN FEMORAL ARTERY. TRACHEA OPEN. 3 MILLIGRAMMES ATROPIA
SULPHATE INJECTED INTO VEIN AT 10°10 A.M.

Time. Pulse in 10”. wien prea HGS General Notes.
10°15’ 33 6
16’ 33 6
18’ 33 6°4
20’ Both vagi divided.
23’ 30” 34 6°4
45” 33 6°6
24’ 10” 33 6°3
40” 34 6
26’ 33 6°4
Zul 32 63
28’ 34 6
30” Cardio-inhibitory nerves found to
be completely paralysed.

Resulit.—Division of vagi, followed by no change in blood-pressure, or fre-
quency of pulse.

EXPERIMENT XXXVII.—TERRIER WHICH HAD FASTED FOR EIGHTEEN Hours. CANULA IN
CAROTID ARTERY, TRACHEA OPEN.

Time. Pulse in 15”. sian Browne Sug General Notes.
11:46’ 30 5
51’ 31 583
52’ 2 milligrammes atropia sulphate
injected into vein.
57’ 50 5° Both vagi divided.
58’
59’ 39” 49 54
12s 2’ 48 55
8’ 46 5-2 Cardio-inhibitory nerves _ ascer-
8’ 30” tained to be paralysed.

Result.—Division of vagi, followed by no change in blood-pressure or fre-
quency of pulse.
VOL. XXVI. PART I. 2N

156 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

EXPERIMENT XXXVIII.—Strone RETRIEVER DoG WHICH HAD FASTED FOR SEVENTEEN
Hours. TRACHEA OPEN. CANULA IN FEMORAL ARTERY.

Time. Pulse in 10”. BE Presi ES General Notes.
12:23’ 28 53 Animal sobbing.
25’ 20” 24 54 Animal sobbing.
40” 3 milligrammes atropia sulphate
injected into vein.
26° 50” 28 54 The animal is now quiet.
49’ 20” 27 5:2
ji! 45? Both vagi divided.
56° 10” 22 59
58’ 22 6
1 1406 20 5°6
6 22 6
16’ 22 59
1 Cardio-inhibitory nerves still para-
lysed.

Result.—Division of vagi, followed by increased blood-pressure and dimin-
ished frequency of the pulse.

TABLE IJ.—GENERAL RESULTS OF THE FOREGOING EXPERIMENTS ON Doas.

No. of Experiment. Vagi divided during Blood-Pressure. Pulse.
XXXII. Digestion. Increased. Accelerated.

XXXIITI. 5 55 Unaltered.
XXXTV, ” ” ”
XXXYV. ” ” ”
XXXVI. Fasting. Unaltered. re
XXXVII. 2 “5 3

XXXVIII. Slightly increased. Retarded.

Tn all these experiments the cardio-inhibitory nerves were paralysed previous to the division
of the vagi.

When these experiments were performed, I was too much influenced by the
fact that Lupwic and Cyon* had always failed to find the depressor nerve |
in action; I therefore fancied that although it is impossible to divide the vagi
in dogs without at the same time dividing the nervi-depressores, such experi-
ments might nevertheless serve to show whether or not the gastric and vaso-
inhibitory fibres of the vagi are thrown into action during digestion. But, on
several occasions in experimenting on rabbits and cats, I have, as before stated
(see page 109), had the good fortune to find the nervi-depressores in action—as
shown by the rise in the blood-pressure which followed their section. J am

therefore convinced that the depressor branches of the vagus are by no
* Lab, Cit.

UPON THE VASCULAR SYSTEM. 137

means so inactive as their discoverers have concluded, from the small number
of experiments performed by them. Seeing, therefore, that these nerves are
not unfrequently in action, and seeing that in the dog, as above stated, the
trunks of the vagi cannot be divided without at the same time cutting across
the depressor nerves, I am compelled to admit that, in the group of experiments
on dogs just given, the increased blood-pressure which followed section of the
vagi may have resulted from section of the depressor nerves, and from these
only. The question at issue must therefore be decided by experiments on cats
and rabbits, seeing that in these animals the superior cardiac branch (depressor
nerve) leaves the vagus high in the neck, and can therefore be divided sepa-
rately from the trunk of the latter.* If, however, the results of the foregoing
experiments on dogs were due to the action of the depressor nerves, then we
should require to adopt the conclusion that these nerves act during digestion,
and are inactive during fasting. Such a conclusion is opposed by the results of
experiments on rabbits and cats, which are yet to be detailed. There is no
evidence whatever that the depressor nerve acts more during digestion than
during fasting; and therefore, after all, these experiments on dogs really do
support the idea that during digestion vaso-inhibitory fibres, distinct from the
“ depressor” nerve fibres, in the vagus are thrown in action.

The following experiments on rabbits and cats are, however, entirely free
from the source of fallacy that obtains in the case of dogs, because in the
former animals the nervi-depressores can be divided without the vagi being at
the same time implicated.

B. EXPERIMENTS ON RABBITS AND CATS.
(a.) During Digestion.
Cardio~inhibitory Nerves paralysed by Atropia.

EXPERIMENT XXXIX.—Stroneé RABBIT FED AT 2 P.M. CANULA IN CAROTID ARTERY.
TRACHEA OPEN.

Time. Pulse in 10”. Mean a inches Conerall Notes:
S05 @ 44 4
59’ 10 milligrammes atropie sulph.
: injected into vein.
30” 44 4°5
AL" Both depressors divided.
30” 40 44
2! Both vagi divided.
ay 39 55
4’ 15” 38 53
6’ 20” 38 5°2
10’ 39 5°3

* The reader will now understand why remarks on the blood-pressure were omitted from the
commentary on the first group of experiments on dogs (see page 122).

138 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS
Result.—Division of vagi followed by increased pressure, but no change in

frequency of pulse.

EXPERIMENT XL—A Strone RABBIT, WHICH HAD FASTED FOR TWELVE Hours, wAs FED
AT 1:30 P.M. CANULA IN CAROTID. TRACHEA OPEN.

Time. Pulse in 10’, weed eae “es General Notes.
3 15! 38 3°5
6 38 3°4
8’ 39 3°55 Both depressors divided.
ay 10 milligrammes atropia sulphate
injected into vein.
107-207 42 3°5
oF 20" Right vagus divided.
12’ 20° 41 3°7
14’ Left vagus divided.
1G? 46 4°2
19,07 45 4:4
laa ta 44 4°3
22! Cardio-inhibitory nerves still para-
lysed.

Result.—Division of vagi followed by increased pressure and acceleration of
pulse.

EXPERIMENT XLI.—A Strone RABBIT FED AT 1 P.M. CANULA IN CAROTID ARTERY.
TRACHEA OPEN.

Time. Pulse in 10’. ee of He in inches General Notes.

3°19’ 42 3°3

20’ 43 34

24’ 10 milligrammes atropia sulphate

injected into vein.

3°25’ 52 35

26’ 10” Both depressors divided.

27’ 40” 47 4°]

287 30” 45 4

29' 25” Right vagus divided.

31’ 46 4:2

32) Left vagus divided.

34’ 50 4°8

35’ 52 4°7

387.15” 51 4°8

40’ 35” 50 4:9

Result.—Increase of pressure and acceleration of pulse following division of
vag.

UPON THE VASCULAR SYSTEM. 139

EXPERIMENT XLII.—Strone Raspit, FED AT 10 A.M. CANULA IN CAROTID ARTERY.

TRACHEA OPEN.

Time. Pulse in 10’.
eyes 2! 36
(a e 37
Ix
30” 50
Gt” 46
imeobe 44
Sao” 43
9”
TO’ 25” 42
11’ 40” 4]
13’
Ae 10” 40
15’ 20”
ig 30° 50
18h 15” 51
40”
20° 10° 49
22% 47
94’ 5” 47

Mean Pressure in inches

Hg.

4°8
4:7
4°8

General Notes.

9 milligrammes atropiz sulph. in-
jected into vein.

Both depressors divided.

Right vagus divided.

Left vagus divided.

Cardio-inhibitory nerves proved to
be paralysed.

Result.—Division of vagi, followed by increased pressure and accelerated

pulse.

EXPERIMENT XLIII.—Strone Rassit, FED AT 10.30 A.M. CANULA IN CAROTID ARTERY.

Time. Pulse in 10”.
1:15’ 4]
16) 10" 42
ne 4]
1387s 5
55” 58
20’ 54
DK tei 52
93’ 15” 53
94’ 5”
2 a0" by)
267 5 50
Pid
28’ 30” 54
30’ 55
34’ 54
ives tae 55

VOL. XXVI. PART I.

TRACHEA OPEN.

Mean Pressure in inches
Hg.

General Notes.

10 milligrammes atropia sulphate
injected into vein.

Both depressors divided.

Both vagi divided.

140 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

Result.—Division of vagi, followed by decided rise in pressure and slight
acceleration of pulse.

EXPERIMENT XLIV.—StrRonG RABBIT FED AT 1.5 P.M. CANULA IN CAROTID.
TRACHEA OPEN.

Time. Pulse in 10”. a aie SHS General Notes.
2:56" 34 32
Sie oe 34 31 |
58’ 10 milligrammes atropia sulphate
given.
59’ 30” 58 3-4
3° 2! 56 34
Ae ats 54 3°3
on LOG Both vagi divided.
6’ 207 52 4:5
7 40” 50 4:4
10’ 45 4:6
12’ 44 4'5
T3e1l0? Both depressors divided.
50” 45 4°9
15’ 44 4°8
19’ 45” 42 4°8

Result.—Division of vagi, followed by increase of pressure and retardation of
pulse. Division of nervi-depressores, followed by increased pressure, but by
no change in pulse.

The rise in the pressure observed in this group of experiments was certainly
not due to any voluntary muscular movements on the part of the animal. In
all cases, none of the recorded observations were made during struggling or
other violent movements, unless it is so stated in the General Notes. In order,
however, to satisfy all that this increase of blood-pressure is really due to a
change in the state of the vascular system not dependent upon extraneous
muscles, I performed the following experiment, in which, in addition to the
atropine, I gave curara, in order to paralyse all voluntary movement.

UPON THE VASCULAR SYSTEM. 141

EXPERIMENT XLV.—Strone Rappit Fep at 11 A.M. CANULA IN CAROTID ARTERY.

TRACHEA OPEN.

Time. Pulse in 10”.
1:28’

Bey 46

37’ 44

44’

47’ 44

48° 30” 41

49°

5 42

ba 10”

53’ 43

30”

54! 20” 42

55’ 40” 4]

Bt 40

59’ 10” 43
2° 2! 40

| Mean Pressure in inches

of Hg.

General Notes.

4°2
4°]

4:2
4:3
4°]

| Left vagus divided.

8 milligrammes atropia sulphate
injected into vein.

2 milligrammes curara injected into
vein.

Both depressors divided.

Right vagus divided.

Cardio-inhibitory nerves still para-
lysed.

Result.— Division of vagi followed by rise in blood-pressure, but by no change
The animal having been paralysed by curara as well as
by atropine, this rise in pressure cannot be ascribed to anything but a change
within the vascular system independent of extraneous muscular movements.

in frequency of pulse.

EXPERIMENT XLVI.—A FUuLL-Sizep Cat FEp at 2.20 p.m. CANULA IN CAROTID.

TRACHEA OPEN.

Time.

Pulse in 10”.

Mean Pressure in inches

General Notes.

of Hg.
4°40’ 30 4:5
45’ 5 milligrammes atrop.sulph. injected
into vein.
30” 38 4
47’ Both depressors and cervical sym-
pathetics divided.*
48’ 36 5'1
49’ 30” 36 51
52 Both vagi divided.
53’ 39 6-4
58’ 39 63
pills 0” 38 6°2
4 39 6°4
5’ 30° Cardio-inhibitory nerves still para-
lysed.

“ In the cat the depressor nerve usually joins the trunk of the sympathetic soon after leaving

the vagus in the upper part of the neck.

and depressor.

It is, therefore, most convenient to divide both sympathetic

142 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS.

Result.—Division of vagi and depressors followed by increased blood-
pressure ; but no change in the frequency of the pulse.

(b.) During Fasting.
EXPERIMENT XLVII.—Strone RABBIT WHICH HAD FASTED FOR EIGHTEEN Hours. CANULA
IN CAROTID ARTERY. ‘TRACHEA OPEN.

Time. Pulse in 10”. a ee — General Notes.
12:28’ 38 +
30’ 38 4 ;
32’ 8 milligrammes atropia sulphate
injected into vein.
35’ 57 3°6
36’ 55 4:2
37’ Both nervi-depressores divided.
30” 54 4°3
38" Right vagus divided.
45” 53 4:2
40’ 51 4°]
41’ Left vagus divided.
49’ 50 4
44’ 49 3°9
45’ 50 3°9

Result.—Division of vagi followed by no change in pressure or pulse.

EXPERIMENT XLVIIJ.—Rassit WHICH HAD FASTED FOR EIGHT Hours AND a HALF.
CANULA IN CAROTID ARTERY. TRACHEA OPEN.

Time. | Pulse in 15”. a ah a HEELES General Notes.
4°54! 48 38
56’ 50 4
5:10’ 9 milligrammes atropia sulphate
injected into vein.
13 74 38
iN! 76 3°6
18’ 74 3°5
30” Both depressors divided.
20’ 75 . 3°5
Di Both vagi divided.
27’ del 34
30° 74 3° 5
32’ 70 3°6 :
34’10” 68 34

Result.—Division of vagi followed by no change in pressure or pulse.

UPON THE VASCULAR SYSTEM. 143

EXPERIMENT XLIX.—StroncG RABBIT WHICH HAD FASTED FOR NINE HOURS. CANULA IN
CAROTID ARTERY. TRACHEA OPEN.

Time. Pulse in 10”, Me EE “ras General Notes.
3°58’ 40 3°9
4° 4 4] 4°2
aM 10 milligrammes atropia sulphate
injected into vein.
i 15” 54 37
8’ 20” 52 3°9
9°30" Both depressors divided.
als LY 51 3°8
13! Both vagi divided.
14’ 45” 58 3°7
16’ 20” 49 3°8
19” 48 38

Result—No change in pressure or pulse followed division of vagi.

EXPERIMENT L.—Srtrone@ OLD RABBIT WHICH HAD FASTED FOR SIXTEEN HOURS. CANULA IN
CAROTID ARTERY. TRACHEA OPEN.

Time. Pulse in 10”. see ee Tee Benes General Notes.
11:30’ 25” 48 4:5
32’ 47 4°4
33’ - 10 milligrammes atropia sulphate
given.
34’ 49 4°5
| 36’ 18” | 48 4°6
2 ian | Both depressors divided.
Se ese 47 4°8
41’ Right vagus divided.
42! 35” 48 | 4:7
45’ | Left vagus divided.
48’ 20° 47 4°8
50! 48 4:7
i as 48 4:5
57’ 46 4°6

Result.—Division of vagi followed by no change in pressure or pulse.

WOE SOV TEAU Sie ae

ho
ia]

144

DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

EXPERIMENT LI.—Strone RaBBIt WHICH HAD FASTED FOR EIGHTEEN HOURS. CANULA IN
CAROTID ARTERY. TRACHEA OPEN.
: 4
Time. Pulse in 10”. Mean oe noe inches General Notes.
AG 44 4'5
8’ 8 milligrammes atropia sulphate
given.
9°30” 46 4:6
Wal? 48557 AT 45 |
12’ Both depressors divided.
1 a7 50 58
1a 85? 49 56
16’ Blood coagulated. Canula cleaned.
20’ 45 5:2 |
Dil’ DO’ | Both vagi divided.
Daly 46 5)
28' 45 52
30’ 46 53

Result.—Section of vagi followed by no change in pulse or pressure.

Sec-

tion of superior cardiac branches of vagi followed by increased pressure and
temporary acceleration of pulse.

EXPERIMENT LIJ.—Stronc RABBIT WHICH HAD FASTED FOR TWELVE HOURS. CANULA IN

CAROTID ARTERY. ‘TRACHEA OPEN.

Time Pulse in 10”. oe EaP nef He ae General Notes.
3°35" 40 3°8
36’ 8 milligrammes atropia sulphate
given.
30” 50 3°5
38’ if 50 3°7
40’ Both depressors divided.
45” 49 3°6
43/ 50 37
44’ Both vagi divided.
45° 54 4°] |
46’ 20” 52 3°9
49’ 10” 50 37
53” 48 38

Result.—Division of vagi followed by a transient rise in pressure and pulse.
This was probably due to excitement, seeing that it soon disappeared.

UPON THE VASCULAR SYSTEM. 145

EXPERIMENT LILI—A Larce Strone CAT WHICH HAD FASTED FOR THIRTEEN HOURS.
CANULA IN CAROTID ARTERY. ‘TRACHEA OPEN.

Time. Pulse in 10”. Mean pressure-in inches General Notes.
of Hg.
9°20’ 32 | 3°6
22/ 33 3°5
23’ 5 milligrammes atropia sulphate
given.
30” 40 37
25’ 4] 3°8
28’ Both depressors and cervical sym- |
pathetics divided.*
29° 30” 40 37
32’ Both vagi divided.
33 25” 38 | 38
3D 15” 39 | 3°9
38’ 37 | 3°8
40’ 5” 38 3°9 |

Result.—Division of vagi followed by no noteworthy change in pulse or
pressure.

The general result of the foregoing experiments on rabbits and cats may be
learned from the following table.

TABLE II].—GENERAL RESULTS OF THE FOREGOING EXPERIMENTS ON RABBITS AND CATS.

No. of Experiment. Nature of Animal. | Vagi divided during Blood- Pressure. Pulse.
XL. Rabbit. Digestion. Increased. Accelerated.
XLI. ” ” ” oe)
XLII. ” » >, »
XLII ‘ ‘ . 3
XLIV. 5 3 5D Retarded.
XXXTX. os bs Bs Unchanged.
XLV. » ” ” ” |
XLVI. Cat. *s i a
XLVII. Rabbit. Fasting. Unchanged. 7
XLVIII ” ” ” 39
XLIX. ” ” 92 99
L. ” ” ” tP)
LIL. ” ” 2) te}
LIT. ” ” ” oP)
LIT. | Cat. PR a nA

Tn all these experiments the cardio-inhibitory nerves were paralysed previous to the division
of the vagi.

* See note to Experiment XLVI.

146 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS

These experiments (XX XIX.-LIII. inclusive) show,— ;

1st. That in rabbits division of the vagi may be followed by increased blood-
pressure and accelerated pulse, although the cardio-inhibitory nerves are
paralysed, and therefore totally inactive before the division of the nerves.

2d. In experiments XX XIX., XLV., XLVI. the pulse was unaffected by
the section: in experiment XLIV. it was retarded, and, notwithstanding, the
blood-pressure was increased in all these cases. A rise in the blood-pressure
following division of vagi may therefore be quite independent of the heart.
_ 3d. The blood-pressure was unaffected when the vagi were divided during
fasting, while it rose after their section during digestion. When we remember
that when the vagi are divided during digestion a permanent blanching of the
stomach takes place (see p. 126), it is evident that the rise in pressure in the
experiments under consideration must be ascribed to contraction of gastric
vessels chiefly if not entirely. It therefore appears that the increased blood-
tension which frequently follows division of the vagi cannot—as has hitherto
been supposed—be wholly ascribed to increased action of the heart liberated
from its inhibitory nerves (see p. 120). Instead of depending only on one
factor it really depends on two. Cessation in the action, Ist, of cardio-inhibi-
tory ; and 2d, of vaso-inhibitory fibres of the vagi.

4th. The evidence afforded by these experiments and those previously given
(see page 126), shows that during digestion inhibitory influences pass from the
stomach through the vagi to paralyse those vasomotor cells in the medulla
which preside over the gastric blood-vessels. But during fasting, when the
gastric blood-vessels are in a contracted state, both the vaso-inhibitory and
excito-vasomotor fibres of the vagi are at rest. (Were the latter in an active
state, a fall in the blood-pressure would follow division of the vagi during
fasting). It appears, therefore, that although the vaso-inhibitory fibres of the
vagus play an important part in dilating the gastric blood-vessels, the rdle
assigned to the gastric excito-vasomotor fibres of the vagi is as yet unknown.
Vasomotor nerve-cells appear, like their homologues the cells of the cardiac
ganglia, to be continually evolving energy. By reason of this, they would
constantly keep the blood-vessels in at least a semi-contracted state, were it not

that their power of generating energy may be controlled by inhibitory nerves. |

These nerves appear to be brought into play by the tissues of a part when it
demands a greater influx of blood, but when it has no such demand, it does not
appear that the excito-vasomotor nerves are brought into action to increase the
evolution of force in the vasomotor nerve-cells, but it seems that in this case the
tissues simply cease to excite vaso-inhibitory nerves. I am, therefore, inclined
to think that these excito-vasomotor nerves discharge their functions on occasions
much more extraordinary than those on which the vaso-inhibitory fibres operate ;
but what those are must be left for future research to determine.

lin 2 iis

UPON THE VASCULAR SYSTEM. 147

5th. Although in all the experiments the pulse remained unchanged when
the pressure underwent no alteration after division of the vagi, it was variously
affected when the pressure was increased. In four cases (see Table III.) it was
accelerated, in three it remained unchanged, while in one it was retarded.
Seeing that the acceleration in these cases and in those previously given (see
page 120) took place when the cardio-inhibitory nerves had been paralysed
previous to the section of the vagi, it is certain that it could not be due to
escape of the heart from control. To what cause, then, shall we ascribe it? I
can think of none other than a direct influence of the increased blood-pressure
upon the lmmg membrane of the heart. It is now generally agreed that—as
Lupwie and Tuiry* pointed out—7/ the vagi have been previously divided—
that is, if the cardio-inhibitory nerves are not in operation, increased blood-
pressure usually accelerates and very rarely retards the pulse.t The retardation
is commonly the result of an extraordinary increase of the pressure. My expe-
riments on this question have convinced me of the truth of the above, but I have,
moreover, frequently noticed that a considerable rise in the blood-pressure may
take place without causing any change in the rapidity of the heart’s action. As
these results are all illustrated in Table III., I therefore think that in the first
five experiments there recorded, the acceleration and retardation of the pulse
were due to the increased blood-pressure. Whether this be or be not the true
explanation, it is certain that the acceleration of the pulse which so frequently
follows section of the vagi, is not, as is generally supposed, dependent merely on
escape of the heart from the influence of its controlling nerves, but depends on at
least another cause—and that probably is—a rise in the blood-pressure. Seeing
that such is the case, the amount of acceleration of the pulse which may follow
division of the vagi, cannot any longer serve as an accurate index to show the
extent to which the cardio-inhibitory fibres of the vagi may be in action previous
to their section, indeed we have as yet no accurate test by means of which this
may be ascertaimed. It has, indeed, been stated by Von Bezoxp, that a trust-
worthy test is to be found im the action of atropia. Most of my experiments
show that when sulphate of atropia is administered previous to division of the
vagi, a varying degree of acceleration of the pulse almost always ensues. It
has been said by the above-named author{ that this acceleration is entirely due
to palsy of the cardio-inhibitory nerves—the heart simply attaining the speed
which it would maintain but for the inhibitory action of these nerves. Von
BeEzoLp came to this conclusion from observing that atropia never accelerates

* Wiener, Sitz. Berichte, 1864, Band 49.

+ Much confusion has been produced by certain authors discussing the influence of the blood-
pressure upon the cardiac movements without distinguishing between the influence of the pressure before
and after section of the cardio-inhibitory nerves.

¢{ Von Bezourp. Untersuchungen aus dem physiologischen Laboratorium. Wiirzburg, 1867,
Erstes Heft.

VOL. XXVI. PART I. 2Q

148 DR RUTHERFORD ON THE INFLUENCE OF THE VAGUS, ETC.

the pulse if it be given after division of the vagi. Unhappily this is untrue. I
have more than once noticed that if atropia sulphate be administered to rabbits
and dogs after division of the vagi, a decided acceleration of the pulse was the
result. For example—

ExpERIMENT LIV.—I divided the vagi of a rabbit, and after waiting five
minutes, I counted the pulse and found it 228 in the minute. I then very
slowly injected 50 milligrammes of atropia sulphate dissolved in one cubic centi-
metre of water into the jugular vein. The speed of the pulse at once rose to
258 ina minute. Ten minutes afterwards the pulse was 240 in the minute. I
then gave another dose of 25 milligrammes atroepia sulphate dissolved in half a
cubic centimetre of water. The pulse very rapidly rose to 270. The blood-
pressure was slightly diminished by these doses. It therefore appears that
atropia may stimulate the cardio-motor nerve apparatus as well as paralyse the
cardio-inhibitory nerves, and so we cannot trust this substance to indicate the
times of action and inaction of the cardio-inhibitory nerves.

Much that is obscure yet remains in connection with the innervation of
vascular system, but I venture to think that the researches detailed in the
foregoing communication clear away not a little fallacious dross from this
matter, while they likewise fill up some important blanks, and thereby render
more complete our knowledge of this complicated and recondite subject.

/ Trans. Roy Soc Vol XXVI_

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(149 )

VIII.—On the Old River Terraces of the Earn and Teith, viewed in connection with
certain Proofs of the Antiquity of Man. By the Rev. THomas Brown,
F-RS.E. (Plate IV.)

(Read 3d January 1870.)
Introductory.

No subject of modern scientific inquiry is more important than the series of
deposits in which geology comes in contact with the period of human history.
This must be my apology for some of the seemingly trivial details contained in
the following paper. When these observations were begun, nothing could be
further from my thoughts than any reference to the antiquity of man. But I
shall perhaps best introduce the subject by simply narrating the way in which
I was led forward step by step, till the whole inquiry assumed the form in which
it is here presented.

In the autumn of 1863, I spent some weeks at Bridge of Earn, on the estuary
of the Tay, and noticed, as every one must, the carse lands lying along the river
Earn, from which they rise by a steep escarpment, running on a dead level back
to the base of the hills. They were deposited, our recent geological authorities*
say, at a time when the land stood lower and the sea higher than now, and
are the estuarine mud of that former period. I had no idea of questioning this
opinion, or of examining the deposit, but in my walks I was struck by the
marked absence of marine fossils. Long ranges of sections were beautifully
laid open, and the absence of marine organisms seemed so remarkable that I
was led to make a closer examination. In the deposit I found there were two
divisions, a lower and a higher, separated by a bed of peat about a foot in

* It may be enough to refer to a series of papers from 1860 to 1866 in the Journal of the Geological
Society of London, by Mr Jamizson, of Ellon, forming one of the most valuable contributions made of
late years to Scottish geology, and one frequently quoted and relied on by Sir C. Lysty. I give two
quotations :—

“The land sank again until the sea in most places reached a height of from 30 to 40 feet above
the present tide-mark. . . .. The clays and beds of silt forming the carses of the Forth, Tay, and
other rivers were accumulated.”—1860. Vol. xvi. p. 371.

“A depression now took place... . . In the valley of the Tay and Forth this old coast-line
was 25 or 30 feet above the present, but on the coast of Aberdeenshire, not more than 8 or 10. The
old estuarine beds or carses of the Forth, Tay, and other rivers were formed, together with correspond-
ing shingle beaches and caves along the coast.”—1865. Vol. xx. p. 195. In this paper the deposits
of the Earn are specially described.

VOL Xvi. PART I, OR

150 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

thickness, which ran for miles through the sections. Taking the order of
succession as it usually occurs, we find the following series :—Immediately
beneath the surface soil (sometimes 3 to 4 feet), there is the carse clay from
9 to 103 feet in thickness, grey-coloured, tenacious, unlaminated, intermingled
with sand towards the base. Underlying this is a stratum of peat, the materials
of which seem to have been drifted from some distance, and one remarkable
thing is that the leaves, &c., which form this peat are found passing up into the
clay, plentifully intermixed with it at first, but getting less abundant as you
ascend. The clay and peat are in this way so associated that one might
almost view them as forming a single deposit. The portion of the series which
underlies the peat consists of laminated clay with partings of sand, and
laminated sands with partings of clay, going down under the surface of the
river. This lower series is unconformable to the overlying peat and clay, and
occasionally the former surface is seen to have been denuded, and the peat
and clay are found filling up the hollows. Some miles further up, near the
railway station at Forgandenny, the sandy layers are found to predominate,
with small gravel intermixed, and the whole has been consolidated into a
tolerably compact sandstone conglomerate, two yards of which are exposed at
the base of the cliff underlying the peat. In regard to the peat itself and the
immediately overlying clay, it is found everywhere to contain wood, marsh
plants, such as the Arundo phragmites, hazel-nuts, mosses, &c. At one point

I found a series of leaves—willow, plane, &c.—in a singular state of pre-.

servation, spread out between lamine of clay, displayed as in a herbarium,
and this continued layer after layer for a yard above the peat. The hazel-
nuts which occur in the peat are of a large size, and still show something of
the shining brown colour which belongs to them. There are occasional speci-
mens Of beetles also, the elytra of which retain much of their brilliancy. My
examination of these deposits was by no means complete, but their general
character seemed sufficiently obvious. What was to be said in favour of
their marine or estuarine origin I really could not tell. No single trace of any
marine organism would turn up. For miles and miles the deposit was
laid open, but examine it where you might, all the remains were fresh-
water or land. The evidence was indeed to a great extent negative, and I
was not willing to come to any definite conclusion, but everything seemed
to indicate that these beds were merely a river formation. They rise about
27 feet above the present level of the stream. If only we could suppose a
time when the river floods had, like those of the Nile, the power of rising
27 feet, how natural and how easy the explanation of the whole phenomena
would be.

Next autumn (1864) I went to Crieff, further up the Earn. Even on approach-
ing the town, looking through the windows of the railway carriage, I was struck

See ee ee ee ee

OF THE EARN AND TEITH. 151

by the resemblance of the high banks lying along the river to those already
observed at Bridge of Earn. The general aspect of these terraces is well shown
in the sketch here given, where a represents the level of the present banks of
the river, 6 an intermediate terrace on the opposite side, and ¢ the high level
terrace. This last was obviously a formation: similar to what I had seen the
previous year. It lay more than 100 feet higher above the sea than that at
Bridge of Earn. The only agent holding the same relation to it in both
localities was the river, along which lay the steep escarpment and level surface,
telling in each case the same story. And again the question presented itself,
was not this simply a river formation in both cases? might not the sea have as

ante, fe aes
a

Sketch 1.—Near Crieff.

little to do with that deposit at the Bridge of Earn as in the neighbourhood of
Crieff? I had long felt, however, that much time and attention would be
required for the satisfactory examination of these river deposits. Men had too
often been content to take a bit here and a bit there from different river-courses,
without the continuous examination of any one in particular. If reliable results
were to be reached, it seemed that some one of our rivers must be fixed on,
followed from the hills to the sea, and made to tell its story from end to end.
This was the more apparent on comparing the deposit at Crieff with that at
Bridge of Earn. But such an examination demanded more time than was then
at my disposal.

The succeeding season I spent some weeks at Comrie, still higher up the
Karn, and there the same deposits again presented themselves in a form which
seemed still more to deserve investigation in connection with the carse lands
at the mouth of the river.

I had gone one day to the foot of Glenartney, near Cultibregan, where the
Ruchil flows from the hills down on the plain of Dalginross. Looking up the

152 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

valley there appeared three terraces, as represented in sketch 2.* Along the
river side is the lowest terrace, a, about six feet above the water, being the
height of the present banks. Further back, and rising above it, is the second
terrace, b, about sixteen feet higher than the first, or twenty-two feet above
the water; and still further back is the third, running along the sides of
the valley, its level being, at the point where I measured it, about 57 feet above
the bed of the stream. In this last the line of escarpment has been somewhat
broken by denudation, but the continuity of the terrace itself is obvious at a
glance. The whole of these levels consist of gravels and sands with clay mn

Sketch 2.—Near the Foot of Glenartney.

different proportions. Near the point ¢, the highest terrace was well laid open,
and showed the following structure, beginning at the surface :-—

Feet. Inches.

1. Gravel with clay, the pebbles lying on their flat sides, 2 0
2. Pan, : : : : ' 0 1
3. Gravel, sandy above, coarser beneath, : : 2 3
4, Fine brown sand, in layers, . : : 0 8
5. Fine gravel with sand, é depth unknown.

This was evidently the work of running water, and the question again arose
whether it had not been deposited by the river at some period when its floods
ran much higher than at present, and whether that threefold system of
terraces might not be found to throw light on the whole of these old river
deposits. On going across to the Turrit, where it comes out from the hills

* For this series of sketches I am indebted to a young friend, Mr W. B. Murray, an art-student
of our Edinburgh School, who has been very successful in his rendering of the scenes. Along the
Earn it has, in three or four cases, been necessary to suppose the woods thinned, in order to show —
the real form of the ground, but this has been done as sparingly as possible. On the Teith there was
less need for this except in Sketch 12, and even on the Earn all the finest examples of the terraces,
such as those in Sketches 6 and 7, are given exactly as they appear in nature.

OF THE EARN AND TEITH. 1535

below Ochtertyre, I found the same three levels in still more striking propor-
tions, and it at once became a question how far they could be continuously
traced along these river valleys.

Beginning at the foot of Glenartney there could be no doubt as to the lowest
level forming the present banks of the river. It passes downwards and spreads
out into extensive meadows. And equally marked was the extension of the
second terrace, the steep escarpment of which goes sweeping for miles, forming
a great irregular triangle from Cultibregan to Lennoch, the level flatness of the
surface being not less remarkable than that of the Carse lands at Bridge of Earn.
The highest terrace, however, is often discontinued, especially along the right
bank of the river ; but away to the north, beginning at Coneyhill, portions of it
may be seen at Tomperran, Lawers, and especially at Monzievaird. It soon
became apparent that these terraces were a good deal interrupted, appearing
and disappearing by turns, while at intervals the threefold system is in full
preservation. But these interruptions are not to be wondered at, when we
think of the denuding agencies to which the deposits were exposed. ‘The
loose sands and gravels, of which they were composed, were just the materials
most liable to be washed away, and their position on the sloping sides of the
valleys was precisely that on which the denuding agencies would most power-
fully act. It is plain also, that the rains and floods of the old time were much
more powerful than now, and, if we picture them to ourselves, rising to a height
and acting with a force to which nothing at the present day can be compared,
it is little wonder that the terraces have in many places been removed, and in
others greatly worn down and obscured.

Making fair allowance for all this, it became a question. whether anything
like a continuous chain of these deposits could be traced along the course of
the valley. During the autumn of 1865, and again in 1866, I had some weeks
of leisure on my hands, and I thought something might be done to ascertain
the point. Taking the Ordnance Survey map in my hand, I filled in as I went
along the results of my observations, using a separate colour to mark each of
the three levels. At all points of importance I endeavoured to ascertain, from
actual sections, the internal structure of the terraces, for there is always a risk
of mistaking for a terrace what is really due to the rocky structure of the
country. I took also a series of measurements showing the height of the
deposits above the river course ; but being alone I had to content myself with
only approximate results. Thus, I followed the Earn, from where it leaves the
loch to where it meets the tide, through many pleasant days, amidst scenes of
quiet river-side beauty, which I shall not soon forget. The results are given
im the accompanying map (Plate IV.) That I have succeeded in all cases
im reading the deposits aright, or in tracing their boundaries, is, I am afraid,
more than I can hope for. I offer it merely as an approximative eye-sketch,

WOLAXx<X VI. PART I. | eS

154 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

sufficient, I believe, to show the general course of these deposits. Their outline
on the side furthest from the river is not attempted.

Having thus examined the Earn, it seemed desirable to test these views in
some different district; and next summer (1867), accordingly, I went to
Callander, on the banks of the Teith, the chief stream in the basin of the Forth.
From the details about to be given it will be seen that the same terrace system
is developed along the Teith, if possible, more strikingly than I had seen it
on the Earn. Other occupations made it impossible for me at once to follow
up the subject, but, having during last autumn verified the leading points, I
shall now endeavour to state the results.

Origin of the Terraces.

The great point of interest is the question as to how these terraces were
formed, and I go into the discussion of this the more willingly, because it will
lead me to describe the way in which these deposits occur, and will show their
continuity along the different valleys.

One explanation ascribes their formation to the sea at a time when the
land was to a great extent submerged, and when our river courses were fiords.
These terraces, it is said, are the old shores, against which the tides rose and ~
fell. Great prominence has been given to this view by various writers, and
especially by Dr Ropert CuHAmsBers in his work on “ Ancient Sea Margins,”
part of which refers to the Tay and its tributaries.* There is one difficulty,
however, in the way of this opinion, from the utter absence of marine fossils.
Even where the most delicate leaves of land plants are beautifully preserved we
can find no trace of the sea. Another difficulty lies in the impossibility of con-
ceiving how the threefold terrace system could have been formed by marine
action. The sea can lay down only one line of beach atatime. Take the
valley, sloping upwards for 240 feet from Bridge of Earn to the foot of Glen-
artney,—suppose it once filled by the waters of the sea, and that they gradually
retired, leaving, as they went, the highest terrace, how is the second terrace
to be formed? Will you let down the land, reintroduce the sea, and bring it
again to the foot of Glenartney? But what would become of the highest
terrace, in the meantime, all down the valley, at Kinkell for example? Exposed
to tides and waves, must it not have been swept away? There is yet another
difficulty, not less fatal, to which we shall immediately refer.

Some of our leading geologists, rejecting this view, have held these terraces
to be the margins of-ancient lakes. The flow of the waters, it is said, had been
barred, and our valleys had become the beds of old lakes. From time to

* Nowhere, perhaps, is this opinion more ingeniously stated and defended than in a series of
papers by the late Mr Cuartes Macuaren. See his Select Writings recently published, vol. i.
pp. 186-201. It is from the valley of the Tay that he takes his examples.

OF THE EARN AND TEITH. 155

time the barriers had been lowered, and as the waters fell these terraces are
the old lake margins showing the different heights at which the waters once
stood.*

One fatal objection which applies equally to this view and to the theory
of their marine origin, is that these terraces lie up and down the valley not
horizontally, but according as the bed of the stream rises and falls. The
parallel roads of Glenroy are an example of how it would have been if they had
been formed either by the sea or by the standing waters of a lake. In Glen
Roy they lie on a horizontal level, keeping their own height without regard to
the bottom of the valley. On the Earn, however, the case is reversed ; the
terraces follow the inclination of the river bed ascending as it ascends towards
the hills, descending as it descends towards the sea. Take the intermediate
terrace, for example, on which lies the Roman Camp south of Comrie. Beginning
at a point above Cultibregan we can trace it as it spreads out and goes down
to Lennoch three and a half miles below. To the eye it seems to lie on a dead
level; and yet, as shown by the Ordnance Survey, it has a decided incline,
being 71 feet higher at Cultibregan than it is at Lennoch, while its height
above the river bed is nearly the same. The river course appears to have
descended about 68 feet, so that the two have nearly kept pace with each other,
and the same thing is found all along the valley. The terraces descend as the
river descends from where they leave the hills to where they meet the tide.

This is of course decisive, but the true nature of these deposits can only be
fully understood when one follows them continuously from point to point along
the whole river valley. There are localities, it must be confessed, where to an
ordinary spectator the theory of lake margins would suggest itself as exceed-
ingly probable. Near Strowan, for example, under the hill on which stands
the monument to Sir Davip Batrp, the river passes through a gorge, and looking
up along its course you see the terrace-like deposits lming the wide open valley
on either side, one of them bearing on its surface the church of Monzievaird.
Would not one naturally say that the gorge had once been barred and a lake
formed, of which these are the old margins? But the fallacy of this is seen

* As this paper deals only with a question of local geology, I do not refer to any writers except
those who have treated of the two rivers to which these researches are confined.

Mr Mitne-Home applies this explanation of Lake Margins to the Terraces of the Teith, Trans.
Roy. Soc. of Edin., vol. xvi. p. 416.

Dr Fiemine supposes a lake to have occupied these river valleys, Lithol. of Edin. p. 76, 1859.

I may refer also to a discussion before the Geol. Society of Edin., 19th March 1867, the report
of which appeared at the time. Only two theories, the Lacustrine and Marine, found support.

Mr CHarues Nicorson, M.A., B.Sc., read a paper on the Surface Formations of the Tay at Perth,
describing the Terraces, and advocating the view that they are of Lacustrine origin.

The President Dr Paes, Mr Coynn, C.E., and others, gave “ their opinion on these Terraces in
opposition to Mr Nicotson’s Lacustrine Theory maintaining their marine origin, Dr Pacz instancing
the minute examination by Dr R. Cuampers of the old sea margins, and Mr Coyne giving his
opinion from minute measurements and personal observations.”

156 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

when you pass through the gorge to the lower side of the supposed barrier, and
find portions of the same terraces on the farm of Trowan, thus holding on their
course utterly disregarding the supposed limits of the lake. Another instance,
to which, as will be seen, some importance attaches, occurs on the Turrit, and
especially on the side valley that goes up towards Monzie. In the angle

wi Se
aah

=
—————SS S

T
—

RPI
Ni RR Tile, 47
= = = ay ne 7, 1 par
SS == 2 WA, Ss yy =x, |
Tes a RRS SW Nine orm Se Gir

Sketch 3.—On the Shaggie above its junction with the Turrit.

between the two we have the three terraces strikingly developed as represented
in sketch 3, where, however, a row of bushes and trees are supposed to be
removed, in order that the form of the ground may be seen. -

Se t | 5,

: es \ \ ..
F 5 yo Y ra f Iw Ve

’ oe ian aX % S ‘ SS Bie) a,
ma f he Piero SI Ve SAN INS : BEN = ;

ae -~ M=
seas me 7 Yj

Sketch 4—On the Shaggie above its junction with the Turrit.

Supposing the spectator to cross the bridge shown in sketch 3, and ascend
the second terrace }, then turn and look down the stream, he would have
before him the view given in sketch 4.

OF THE EARN AND TEITH. 157

Here we have in front the narrow ravine between the spectator and the
point R through which the stream passes ; and we might have argued that the
glen had once been closed, a lake formed, and that these terraces are the
former margins. But following the course of the deposits we find the terrace
¢ passing into the ravine and continuing especially along the right bank. And

lage ay ven
eT ee
a ue q- *

Sketch 5.—Dalvreck Bridge on the 'Turrit.

what is more decisive, if you go to the other end and look across and up the
_glen where the water comes out from under the Bridge of Dalvreck, the three
terraces come out distinctly as here given in sketch 5. On the one bank the

SS = es
y

AUS MEU LIP II PPP

Sketch 6.—Dalpatrick.

second level > holds its course at a similar height above the river bed, while
in the other bank the highest level ¢ passes* out into the open going round

* There is a beautiful section showing that it consists of finely laminated sands with a little
gravel and clay.

VOlm xO, PART T. Dy Tt

158 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

over the grounds of Ochtertyre; and when one finds these deposits in the
same position above the ravine and below it, it seems vain to ascribe them to
some lake formed by the barrage of the river.

Tracing the course of the Earn from Crieff to Dalpatrick, we again reach a
point above the old castle of Innerpeffray where the channel is narrowed between
high grounds. Standing near Easter Dalpatrick and looking upwards, we have the
scene as represented in sketch 6. The present banks and meadows, a, are sur-
mounted by the second terrace b, and that by the higher level ¢, all in full
preservation.* Again, the idea might be suggested of a bar above Innerpeffray
forming a lake, but again the explanation is forbidden by the continuous course
of the deposits, and more especially by their appearance when the narrow
portion of the river has been passed, and the banks again spread out into a
wide open valley. This takes place immediately below Kinkell, where the
threefold terrace system is very remarkable, as shown in sketch 7.

~ oo
ss ~
ee Sy

Sketch 7.—Kinkell Bridge, leoking up.

A section of terrace ¢, on the eastern side of the Machany, is laid open by
the cutting of the road, and is given in Pl. IV., fig. 1. The details are—
a, Humus.
b, Carse clay, grey, unlaminated tenacious.

c. Laminated clay with partings of sand. Lamine a half inch in thickness.

The series of deposits at this point are specially important, because there is no
position further down the valley where it is possible to suppose that a barrier
could have ever been thrown across. These are not lake margins.

The lower portions of the river, as it passes Dunning, Forteviot, &c., were
examined somewhat more rapidly. The threefold system of terraces seems to
have been less distinctly preserved. A point, however, is given in sketch 8, —

* Immediately beyond the farm-house the railway gains the summit of terrace c, and the view
shown in sketch 1 is seen looking up the valley.

OF THE EARN AND TEITH. 159

some distance below the bridge of Dalreoch, where the three levels are present,
the second especially being well exposed.* They may be seen also distinctly at
Forgandenny, from the railway crossing, looking across towards Boatmill.
But all through these lower portions of the river course the higher and middle
terraces show a tendency to coalesce, forming, with the present meadows, only

Sketch 8.—Near Dunning.

two levels. It was the higher of these which first attracted my attention at
Bridge of Earn. By comparing the map with the series of views, the reader
will have some idea of how continuously these terraces pervade the whole course
of the river valley of Strathearn from the mountains to the sea.

Sketch 9.—Loch Lubnaig, looking up.

Turning now to the river Teith, we have first to notice the shore deposits of
Loch Lubnaig, along the western side of which they may be seen running

* The highest terrace c is made too prominent in the sketch.

160 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

under the shadow of Ben Ledi. First, there is the present shore of the lake,
above which rises the second terrace to the height of nearly fourteen feet, and
this in its turn is surmounted by the third, about thirty-seven feet over the
water. Their position is well given in sketch 9. The point seen in the
distance is perhaps the spot where the highest terrace is best displayed, and.
is given more fully from the upper side in sketch 10. When I first saw it in
1867 its form was that of the dotted line, a shelf projecting and running at its
own level along the mountain side. Now it has been cut up by the railway,
which has been carried for a considerable distance through these terraces,
showing many remarkable examples of drifted gravels and stratified sands,
with, in some instances, underlying boulder-clay.

Sketch 10.—Loch Lubnaig, looking down.

These deposits on Loch Lubnaig would seem to point to a time when the
waters of the lake stood permanently higher than now. One of our best geo-
logists, Mr Minne Home, has advocated this view, placing the barrier which
held back the waters, at the pass of Leny; and it would have been difficult
to resist this opinion, but for the circumstance that when we get below the pass
we find the same terraces quietly falling into their"places, and resuming their
course as before. Sketch 11 shows their form when they leave the narrow por-_
tion of the valley, going off towards Callander on the left bank—a similar appear-
ance being presented as they sweep round towards Loch Vennachar on the right.
The height of the terrace levels is almost identically the same with those on
the shores of Loch Lubnaig, and there is the closest resemblance in their inter-

OF THE EARN AND TEITH. 161

nal structure. If it be said that these in sketch 11 are the margins of a lower
lake, there is, first, the difficulty of accounting for their beg so exactly the
same height above the water, and then there is the fact of their continuity
down past all supposed barriers at Gart or elsewhere. Leaving the pass of
Leny, and going on towards Callander, we find it is for the most part on these
terrace levels that the new west end villas are built ; and when the railway was
being cut in 1867, it was striking to observe the close resemblance which the
fine grey sands, with their false beddings, and the coarse gravels, bore,
both in their structure and relative positions, to those seen in the sections on
Loch Lubnaig.*

a W sw 75,
ey Nx
i; SS INN os

SSi S
NS

Sketch 11.—Looking from below the Pass of Leny towards Callander.

Further to the east it would be easy to give from different points of the
river course examples of the threefold levels ; but it may be interesting to take
one from its great feeder, the Keltie, so well known as forming the Falls of
Bracklin. It is seen (sketch 12) a little above its junction with the Teith,
and the view will serve to show that the same system found on the main stream

pervades also the tributaries. The upper level presents itself in two stages,
and to this fact we shall afterwards refer.

The succeeding portion of the Teith, down as far as Doune, shows a con-
tinual succession of the same deposits. The village of Dalvaich especially lies
in the midst of a series of these terraces, deserving a far more careful examina-
tion than it was in my power to give them. They may be followed down
through the grounds of Lanrick Castle, and come out well at the fine old
churchyard of Kilmadock. But one of the most striking examples either on
the Earn or Teith is that given in sketch 13, above Deanston, on the opposite

* To illustrate this, two sections are given in Plate IV. Fig. 2 is from the terrace on the
shores of Loch Lubnaig ; fig. 3 is from the railway cutting at Callander. In both the fine grey lami-
nated sands, with their false beddings, are seen to have been denuded in a remarkable way, and are
overlaid by coarse gravels.

VOL. XXVI. PART I. 2U

162 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

side of the river, near the farm-house of Clarkton. The highest terrace there
shown passes on to the railway station, and has a great part of the town of
Doune built on it. It is the same also which may be traced across the river
into the grounds of Blair-Drummond, where, in the view immediately opposite

= = 2 ————————————————— |

//, i m_,
an Ae: Hl fe = Hau
Phim, Ane fi? Ui, ie ( a =
it \ Ay i, iil Meas A
SHAAN EN .
Hi Z /

(

RN OE eet rey
— AAUP ri PMN GATE ‘Ar at De
4 LVEDD? 7 . a
GA inc, pe On IO ee
See SF _ OPER NO SB a gee lg Wg eRe

i ——

Sketch 12.—On the Keltie, near Cambusmore.

Doune Castle, the three levels are distinctly seen ; and here a point is reached
of great importance in regard to these terraces. The Carse of Stirling, lying

Sketch 13.—On the Teith above Deanston.

between the Forth and Teith, begins to spread out its flat level, and the highest
terrace gradually descends and coalesces with the second, just what we saw
take place on the Earn, when approaching the Carse lands in the neighbour-
hood of the sea; and yet all the way some traces of the threefold system may
here and there be found. Even at Kildean, near Stirling, and well within tide

OF THE EARN AND TEITH. 163

mark, thisis shown. The highest bank c is only twenty-four feet above the river,
while the lowest @ is six; but the opposite side shows an intermediate level 6
of some twelve or fourteen feet. The whole are thus on a very inferior scale.
Perhaps it may be thought that this diminution is due to the greater width of the
valley. But a still stronger reason, I believe, is the comparative weakness of
the current arising from the lower gradient of the incline along these portions
of the river course. To this I shall again refer.

The details thus given make it plain that we cannot have recourse to the
lake theory, for the terraces are not horizontal, but slope with the valley, and
in the case of both rivers we reach a point below which any barrage is incon-
ceivable, and yet these deposits hold on their course.

But there are certain additional circumstances to which I ask attention, and
which I was led to notice only as the result of having continuously examined
the whole course of these rivers.

Sketch 14.—Kildean on the Forth.

First, there is the difference between the upper Earn and its feeder, the
Ruchil. There are three streams which meet at Comrie, where the Earn,
coming straight from its loch, is jomed by the Lednoch from the north, and the
Ruchil from Glenartney in the south. Both these feeders show the terraces,
those on the Ruchil being, as we have seen, specially remarkable. On search-
ing for them along the upper Earn I was struck by the difference. The present
banks of the stream are unusually low, and above these there is a second
terrace, some 12 or 14 feet over the river ; and this was all I could make out.
Some banks which I took for the highest terrace I found were due to the rock
structure of the country, and some gravel deposits near Aberuchil belonged
evidently not to the Earn, but to a mountain stream which comes from the
south. The general appearance of these low terraces along this part of the
river will be seen from sketch 15, which the reader is requested to compare
with sketch 2, when the difference of scale will be obvious. The threefold

164 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

terrace system of the Ruchil is feebly represented by the 12 or 14 feet
banks which are continued along the Earn, past Dunira, and on to the loch.
In seeking the explanation of this difference a remarkable circumstance was —
brought out. The floods which come down the Earn at the present day are
quite feeble compared with those of its tributaries. The Ruchil is remarkable
for the suddenness and strength of its floods, while the flow of the Earn is
quiet and equable. Now, the height of the o/d terraces on the two streams
exactly corresponds with this. The coincidence is striking, and can have only
one meaning. Along the stream, where the floods are still powerful, the
old terraces are powerfully developed. Along the stream, where the floods are
feeble, the old terraces are feebly developed. The only possible conclusion is,

ia . a ™~ Le») Wise Se a +
A wl opt om mS a “F a as |
a me Res, All De. “ <a —
|

eats a . ii me ie 4

Sketch 15.—On the Upper Earn.

that it was by the floods of these rivers that the old terraces were really built
up in a former age, and that their flooding power was then in proportion to
what it is at present.

A second circumstance of the same kind is seen when we compare Loch Earn
with Loch Lubnaig. Along the shores of the latter, as we have already shown,
there are well marked terraces, and on Loch Earn also a similar deposit is
present, but in far less proportions, spreading out especially towards the bottom
of the lake, where in September 1869 I found it 12 to 14 feet above the water.
This is a weak representative of the 37 feet terrace of Loch Lubnaig. But
the remarkable thing is that it almost exactly corresponds to the proportionate —
rise and fall of the water in the two lochs at present, as caused by a flood on
the one hand, and a drought on the other. Both lakes, I was told, were at the —
lowest ebb at which they had been seen for years. In the case of Loch Earn,

OF THE EARN AND TEITH. 165

the water was between 3 and 4 feet below the highest water-mark that could
be found ; while on Loch Lubnaig the difference was about 8 feet. How much
of this was due to the absolute depth of the water, and how much to the action
of the wind, I could not, of course, say. It was the western shore of the loch
in both cases where I took the measurements. But the striking thing is to
observe the closeness with which the results correspond with the proportions of
the old terraces. In the loch, where the floods and winds of the present day
raise the waters 3 to 4 feet, you have the old terrace about 14 feet high. In the
loch, where the waters at present are raised some 8 feet, you have the old
terrace lying 37 feet high. It is hardly possible to resist the inference that
these old terraces are due simply to the greater flooding power of some former
epoch.

A third fact which came out was, that these old terraces vary in height just
as the present banks of the stream vary at different parts of the river course, and
in something like the same proportions. Usually the present banks and haughs
of the Earn are 5 to 6 feet above the stream, but in some places we find them only
3, and in other cases they rise to about 10, as near the Bridge of Strowan.
The difference is due to the form of the valley, and still more, I believe,
to the force of the current. Now, there is precisely the same kind of
variation in the levels of the old terraces. As the present banks and haughs
may be anything from 3 to 10 feet, so the second terrace varies from 16
to 24, and the highest from 35 to 60. The cases where extremes occur
are rare; but this general truth must be recognised, that as the present banks
vary in height with the locality, so do the ancient terraces.

Connected with this, however, there is one further circumstance which
deserves to be noted ; the height of the old terraces varies with the incline of the
river bed. Where the incline is greatest, there, of course, the current ran
strongest, and there the terraces are highest. When the gradient is low, the

terraces get low. It is difficult, indeed, to bring out the exact truth on this
point, for it is necessary to make allowance for the varying width of the valley ;
but in comparing the different parts of the river, there is seen to be a distinct
proportion between the steepness of the incline and the height of the deposits.
Thus, from the foot of Glenartney to Kinkell, the distance in a straight line is
about ten miles, and there the descent of the river is nearly 200 feet. From
Kinkell to Bridge of Earn is more than eleven miles, and there the descent is
about fifty feet. Now, it is along all the upper section, where the current ran
strong, that the terraces rise high ; and along all the lower portion, where the
current ran slow, the terraces subside. Precisely the same thing is seen on the
Teith. Above Doune the descent is rapid, and the current strong. Below
Doune all is flat, and the current gets slow, and it is in the upper portion that
the terraces are raised high, while below Doune their height markedly diminishes.
VOL. XXVI. PART I. 2X

166 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

Here, again, the facts point to these river currents as the agent which built
up the terraces. In estimating the facts, however, care must be taken to
include some considerable length of the river course. A short reach, where
the water runs level and slow in the midst of rapids, may be the very place
where the deposits most accumulate. But take any considerable distance, and
it will be found that in proportion as the gradient is steep* these old terraces
rise in height, and as the gradient gets low the terraces diminish.

If we could only suppose a time when the river had the power of rising in
flood to the requisite height, the whole phenomena would admit of the most
easy and natural explanation. The simple key to the whole would be the
principle, that gust as the river deals now with its present channel and present
banks, so it dealt in the old time with those high lying terraces.t

Putting together then the whole of these facts, we can now see what this
threefold system of terraces means. It is simply a record of the different levels
at which the most powerful river floods stood at different periods of the past.
The highest of these lmes of deposit is evidently the oldest, a gravelly and
sandy terrace which runs along our valleys at a height of from 35 to 60 feet.
Then we find a descending scale as the floods grew less and less powerful,
subsiding towards the present state of things. Rather more than half way
down the scale there is an intermediate terrace about 16 to 24 feet above the
river, and forming an outstanding feature of these deposits. It seems to
indicate that when the descending waters reached that stage a pause of con-
siderable duration took place, during which the action of the highest floods
went on at that level. Subsequently there was a descent from the middle
terrace to the present banks. Between each of the stages indeed, there are
intermediate lines of deposit occurring here and there in different localities,
and putting them all together it would be possible to construct a whole series
of graduations by which the highest terrace would be found to descend to the
second, and the second to the lowest. Still there can be no doubt that the
three terraces form the prominent feature of these deposits.

The general result thus seems to be, that along the sides of these river
valleys, we read the history of various ages during which the floods gradually
ran at a lower and lower level, and in that record there are three great lmes —
which stand out from the rest as indicating each some considerable period —
during which the waters remained stationary, till at last the intervals were all
passed over and the present state of things was reached.

* This does not apply when one follows the stream up among the mountains, where for the most
part the terraces are absent. Is it that denudation has swept them wholly away? or is it that during
the epoch of these old floods there were glaciers still lingering in the upper portions of the river-valleys ?

+ More than twenty years ago, Mr Mitne-Homm described the terraces above Perth as haughs
or river flats, but he seems to connect their formation with the bursting of lakes. See “Trans. Roy. —
Soc. Ed.,” vol. xvi. p. 418. I do not refer to other districts. “

OF THE EARN AND TEITH. 167

Their Geological Position and Age.

In regard to the time when these terraces were formed it is difficult to
pronounce with confidence, but there are certain indications which deserve
attention.

Near Comrie we find some antiquarian remains which go a good way back
into the past. The site of the Roman Camp is close to the village, and a little
further to the east there is what the Government Surveyors have laid down as
a small roundel or Druidical structure, a circle raised above the surrounding
ground, in the middle of which there once stood a rude and apparently
unsculptured monolith, now prostrate. These Roman and Druidical remains
are all on the expanse of the second terrace formerly referred to. So far back
then as they carry us the intermediate terrace had been already formed.

Leaving archeology and appealing to the methods of the geologist, it is
clear that these deposits have been laid down subsequently to the glacial epoch
in Scotland, for no glacier can have touched the valleys since the terraces were
deposited. We have traced them on Loch Lubnaig up to a height of 400 feet
above the sea, and their state of preservation makes it plain that up to that
level at least no glacier nor icecake has since their formation grazed hill or
valley.

It is equally plain, and for the same reason, that the sea had finally retreated
from the land. Some minor change of level there may have been about the
Carse of Stirling, but already the sea must have finally left the valleys free for
the action of the river floods.

There seems indeed to be good ground for believing that a series of peculiar
deposits is interposed between the oldest of these terraces and the glacial
epoch. In working back, and trying to make out stratigraphically the place of

the highest terrace, we are in contact with a set of gravels, &c., which in the
present state of our knowledge are particularly obscure. I refer to a series of
mounds or hillocks sometimes round or sinuous, sometimes drawn out as long
lines in the form of escars or kames, but invariably when laid open showing
that they have been deposited by water in a state of disturbance. - Occasionally
they come down into the valleys, but for the most part they stretch away out
over the higher grounds. Examples are to be seen along the course of the
Earn, but they are still more striking on the Teith, and especially near the
village of Dalvaich. In sketch 12 it will be observed that the highest terrace ¢
is shown in two stages. Following the oldest and highest of these stages it
_ appears to connect itself with the deposits in question, which, a little to the
west of Dalvaich and north of the turnpike road, begin to spread out and go far
up over the face of the country. One of these lines of kames is given in sketch
16 showing another and higher ridge of the same kind behind it. The end

168 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

view of the first of these lines is seen in sketch 17, showing the form which
these deposits assume. The question as to how they were formed is one of the

Sketch 16.—Kames west of Dalvaich.

most difficult in geology. That they are not due properly to the action of
flooded rivers is plain, for they may be traced up and out over the level face
of the country, and are nowhere more fully developed than towards the water- —

must have been feeblest. It would be natural to suppose that they were

OF THE EARN AND TEITH. 169

submarine banks belonging to the time when Scotland was submerged, but
this seems disproved by the fact, that wherever such submarine banks occur,
they swarm with all kinds of marine life, while in regard to any trace of such
life, these kames are invariably an utter blank. But, indeed, similar difficulties
‘attend all the theories hitherto suggested. Nor ought this to surprise us.
We do not know what it is for a country once incased in ice as Greenland
now is, to have the ice-sheet lifted off or melted from the face of hill and plain.
Into what forms the subjacent materials of gravel and sand would be thrown—
what would be the modes of operation of the forces let loose, it is difficult to
conjecture. No example of such a process has been witnessed, and yet it is
certain that Scotland passed through it. It is little wonder if among its results
there should be some residual phenomena for which it is difficult to account.
Among these it would seem we must place the kames or escars, and the gravel
mounds associated with them. At all events, their position appears to lie
between the period of arctic climate and the time of that series of terraces
which this paper describes, and which were, it is probable, built up out of the
materials furnished by these pre-existing gravel deposits.

In deciding the geological position of the terraces, however, we must not
forget the fossils of the peat and the associated carse clays referred to at the
beginning of this paper. For several miles above Bridge of Earn these remains
occur in abundance, but evidently they have been drifted from some distance,
brought down by the current, and they show what the flora of Stathearn had
been at the time when the peat and carse clay were deposited. In regard to
the extension of the carse clay itself, it can be traced up as far as Kinkell,
where its grey colour and fine unlaminated structure were quite distinct,* form-
ing part of terrace c,as shown in sketch 7. Above Kinkell, the same terrace runs
on, but the place of the clay is taken to a great extent by sands and gravels,
and these materials get on the whole coarser the further you go up the stream.
All this is easily explained. The coarser the material, the less easily is it
moved forward by the current, while clay in the form of mud is floated to the
furthest distance. The highest terrace, therefore, which consists at first of
gravel and sand, with a little clay, presents through all the lower reaches of the
river little else than large sections of the finest carse clay. These carse clays,
and the underlying peat near Bridge of Earn, form, as we have seen, properly
only one deposit, and the result would seem to be that these fossil leaves and
hazel nuts, &c., give us the flora which grew along the valley at the time when
the oldest of these terraces were formed.t

* See Fig 1, Pl. iv. :

+ There seems good ground for holding that the peat beds of the Earn belong to the time when
the land stood comparatively high. But when Mr Jamieson makes the land again sink, and brings

in the sea in order to deposit the estuarine mud of the carse, not only does the fossil evidence go
against this, but there is the decisive fact already pointed out, that the peat and the carse clay are so

VOU. XXXVI. PART I. OX

170 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

If this be so, it would appear that the glacial epoch must, to a great extent
at least, have passed away. Its close had been marked by the formation of
the kames, those ridges of gravel whose strange forms meet us on our
uplands and over the face of the country. Then there came a more genial
time, shown by the large size of the hazel nuts, when the present Flora was
established. And then it was that those vast floods seem to have flowed forth,
which filled our valleys, and left their record in these highest terraces. What
was done during the Moray floods may have been done of old on a still greater
scale, and these high-lying deposits may be the proof of it.

Antiquity of Man.

The views thus far stated must be judged of on their own merits, apart from
any question as to the antiquity of man. The advocates of extreme opinions
on this subject have relied to a great extent on geological evidence, and some
of their strongest arguments have been derived from the flint implements of the —
Somme in France, and the Brixham Cave in Devonshire. It is held that
wrought weapons, the work of man, are found along with the remains of extinct
mammalia, and occur in such a way as to show that man had been their co-
temporary. If this were all, however, the argument would have little force, for
the inference would be perfectly open, either that the human period must be
carried further back, or that the time of these extinct mammalia must be
brought further down. Were we mistaken as to the duration of man—must
it be carried much further back among these extinct animals? or were we
mistaken about these extinct animals—do they come down into the human
period? Men would lean to one or other alternative according to their pre-
possessions. Other circumstances had therefore to be appealed to, and, in fact,
the stress of the argument has come to rest on the position of the deposits in
which these remains occur. Those on the Somme were examined by one of —
our best observers, Mr Prestwicu, who reports that the oldest beds containing
these fossils lie along the valley, at the height of about 80 feet above the river
course. The time when these were deposited was the time when man and —
the mammoth lived together. Since then the river has worn down the valley,
cutting through rock, &c. some 80 feet, and the human period must be carried
back through the ages which can rationally be supposed needful for this opera-
tion. He refuses to admit “hundreds of thousands of years,” but if his view
be taken, the period must be very long. The argument in the case of the
Brixham cave is similar. The remains of human art and of these extinct
animals are found together in a deposit which must have been carried by
running water into its present position. But the entrance to the cave is in the
associated as to form properly one deposit. At whatever time the one was formed, the other was also.

The conclusion to which all the evidence seems to point is, that the whole system of these river
terraces was formed at the time when the land was elevated above its present level.

OF THE EARN AND TEITH. 17h

side of a valley 60 feet above the present bed of the stream, and we are told
that the human period must be carried back through the long ages needful for
wearing down the rocky floor of the valley. These are, I believe, among the
strongest arguments from geology.

But now, if the analogy of our Scottish rivers may be trusted, there seems
fair ground for asking whether such arguments have not been carried too far.

1. First, it is plain that previously to the time when these high-level terraces
were deposited along our river courses, the rocky structure of our Scottish
valleys had been hollowed out as deep as they are now. In proof of this, it is
enough to refer to the fact that the boulder-clay which belongs to an ante-
cedent period is found occasionally forming the floor of the valley over which
the streams flow.* If, then, the formation of the valleys of France and Devon-
shire were analogous to ours in Scotland, their rocky structure was excavated
not after, but before, the deposition of the high-level gravels.

2. Secondly, The floods which piled up these old high-level deposits seem to
have had the power of doing so at a time when the river-bed was cleared of
all other materials, and stood at as low a level as the present streams.

This might be shown by various examples along the Earn; but we take a

Sketch 18.—Below the Castle of Monzie, 1866.

single case from the valley of Monzie. Sketch 18 shows a spot where the
stream has cut into the highest terrace,t laying it open from the base up to a
height of 50 feet, and showing that it is composed of stratified gravels, sands,

* Seen in the valley of the Turrit, for example, above Crieff, and also in that of Monzie, in both of
which it underlies the high-level gravels. My attention has been called to the fact that this view had
been brought forward in the Memoirs of the Government Survey on Berwickshire, p. 50. 1863.

} This forms a continuation of terrace ¢ shown in Sketch 4 on the left bank of the stream. It
lies a short way further up the valley.

172 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

&c., the work of running water flowing down the valley. Eighty yards further
up, the same bank (sketch 19) is laid open up to its whole height of 120 feet,
where, in one of the gullies, innumerable alterations of the same strata are
shown. ‘The last 20 feet* at the top are particularly well seen, as also are the
50 feet at the base in Sketch 18.

Now that this terrace was formed by river floods is shown by its connection
with similar deposits further down the stream. Those seen in Sketch 1 must

Sketch 19.—Below the Castle of Monzie.

have been a river formation from the slope of their surface down the stream,
and not only is their structure identical with that at Monzie, but—allowing for
the effects of partial denudation—they may be traced running on till the one
series actually meets and passes into the other, forming two stages of terrace ¢,
as in Sketch 12.

But if these deposits were formed by river floods, a single glance at such a
section as that in Sketch 18 will satisfy every geologist that the running water
began its work at the bottom, down on the level of the present river bed.
When the first stratum was laid down, the running water had found the valley
cleared out down as low as it is now. Explain that matter how we may, these
old floods must have-had the power of doing this ; beginning at the level where ,
the stream runs now, they could form a terrace, piling stratum above stratum,
up to the height of 120 feet, as far, indeed as the water was able to rise. _

It is plain, therefore, that if the analogy of our Scottish rivers will apply to

* See Plate iv. fig. 4, showing the arrangement of the gravels and sands forming the upper portion —
of the terrace. ]

OF THE EARN AND TEITH. 173

those of France and England, the long period required for the supposed lower-
ing of the river-bed is got rid of. Ifthe stream which flows through the valley
at Brixham could do what has been done by the burn in the valley of Monzie ;
or.if the Somme could do what the Teith has done along its course, any amount
of high-level gravels might have been piled up, or carried into caves after the
valleys had been hollowed out, and the bed of the stream brought down to
where it now is.

But are we right in supposing that the analogy of our Scottish rivers will
apply to those of England and France? The presumption certainly is that
their formation was analogous.* But we have more than mere presumption.
Mr Tyter, an English geologist, has been investigating the case of the Somme,
and finds evidence of a pluvial period—a time of high floods sufficient to
account for the high-level gravels. The body of facts which he has brought
together deserve careful consideration, and it will certainly be matter of deep
interest if the deposits of the Somme are found to record the same story which
we have been reading along the Earn and Teith.t

_ I have no wish to push this argument beyond what is perfectly fair. These
terrace-like deposits form a subject which has been too little investigated. It
may turn out that they reveal to us a period of river floods much greater in
volume than men are generally prepared as yet to admit. And it may also be
that those inferences which have been drawn as to the duration of the human
period may be very seriously affected. It would surely be safe to have more
complete examination before judgment is given. These deposits belong to what
in geology is a very recent period indeed, and all I would ask is that the facts
be more fully investigated, lest in arguing for extreme views as to the antiquity
of the race men be found importing fallacious elements into the calculation.

But how shall we account for the volume of water necessary to produce
such floods? Looking to the width of some of our valleys, and to the height
of these deposits, is it not difficult to believe in the existence of such torrents 4
In dealing with this question care must be taken not to exaggerate the diffi-

* An attempt has been made to deny this on the ground that Scotland was submerged during the
glacial epoch, while Picardy and Devonshire were not—the object being to show that the French
valleys were excavated at a later period than the Scottish. But this argument can hardly be urged by
those who hold that the formation of valleys is due mainly to subaerial forces and hardly at all to
marine action. If the difference between the two countries be that France not having been submerged
was continuously acted on by these eroding agencies, while Scotland was withdrawn from them by
being buried beneath the sea, how will that prove the French valleys to be of later formation than the
| Scottish? So far as that difference goes it should surely prove the reverse.

+ “On the Amiens Gravel,’ by AtForpD Tytmr, Esq., F.G.S., “ Journal Geol. Soc. Lond.” vol. xxiv.
p. 103. In one respect Mr Tyuer’s reading is different. He regards the lower terrace, the loess, as
the bank of the ancient river when in its ordinary state, and the higher terrace as its bank when in
| flood—referring both to the same period. This differs materially from the view which I have been led
to take, namely, that each terrace is the highest flood-mark of its own time, just as the present banks
and haughs are related to the floods of the present time.

VOL. XXVI. PART I. De,

174 REV. THOMAS BROWN ON THE OLD RIVER TERRACES

culty. Our rivers, as we see them at present, flow along a comparatively
narrow channel, the greater part of the river valley being usually occupied by
level meadows. When a flood comes, it is only the narrow channel that needs
to be filled; and then, should the water rise but a very little over the brim, it
will spread out like a sea on either hand. But, after all, it is the banks and
meadows which fill the space of the valley. Except in the central channel the
sheet of water may be comparatively shallow. And so in that old time the
river would have only its central channel lined by banks proportionally higher.
The flood would be needed to fill the river bed, flowing perhaps over the brim
and out over the surface. In this way these old terraces would be formed just
as the present meadows are. The volume of water needed was by no means
what would have been required to fill the valley if it had been empty. It would
be enough if the confined river bed were filled to overflow.

But, if things were on such a scale that the river channel was lined with
banks 50 or 60 feet in height, where was the water to come from which could
rise to such a height? One explanation has been sought for in the melting of
the ice and snow as the glacial epoch passed away. At present, when the ice
and snow melt in northern latitudes, the arctic rivers rise annually from 40 to
50 feet. This of itself would go a long way to solve the problem. Besides,
there was more than the annual melting which takes place at present under —
ordinary conditions. The fact that the glacial epoch was passing away, must be
taken into account. Ifthese terraces may be taken as a record of the time when
the great icy covering was melting off the face of the land, and Scotland was
passing from the rigour of an arctic climate to its present condition, the currents
which filled our valleys may have been increased to an extent which it is diffi-
cult to estimate. Swollen lake and flooded river may have risen to a height
sufficient to meet all the conditions of the problem we are considering.

Another explanation which has been suggested, is the existence of a period
of great rain-fall—a “ pluvial epoch,” as it has been named by Mr TYLer. —
This may have arisen either from the quantity of rain having been increased, or —
from the rain-fall having been concentrated—a greater amount falling im a
given time. Some idea of these floods of the old time may be got from the
account of the Moray floods, as given so admirably by Sir Toomas Dick LAUDER.
They occurred in 1829, and were owing to a fall of rain to the amount of —
33 inches having taken place in twenty-four hours. In regard to the height
to which, on that occasion the water rose, the writer mentions having himself
seen a man wade into the water and capture a salmon on the haughs 50 feet
above the usual level of the Findhorn, pursuing the fish with his umbrella and —
driving it ashore. The violence and velocity of the currents he describes
in striking terms. “It was scarcely possible to follow with the eye the trees
and wreck which floated on its surface. The force was as much more than

OF THE EARN AND TEITH. 175

that of a raging ocean as gunpowder ignited within the confined tube of a
cannon is more terribly powerful than the same material when suffered to
explode on the open ground.”* It is no wonder if, with such force at work,
there should be strange tales to tell of the results of denudation. Instances
of farms, where six, eight, or ten acres were eroded and swept away, are so
common as hardly to deserve notice. At Mains of Orton, on the Spey, when
the proprietor, Mr Warton Durr, came, after the flood, to examine his farm,
he found he must make a new bargain with the tenant, and deduct some 50
or 60 acres which were gone. At Braemoray, the whole low land was
annihilated, and the green slopes of the hill converted into naked precipices.
At Relugas, the pleasure-ground and lawn were swallowed up, and in their
place that river might be seen raging for 300 yards along the brink of a red
alluvial precipice 50 feet high. At Dalrachney, the river Aven attacked a
wooded bank from which it carried off a mass of not less than 90,000 cubic
yards, leaving a sandy precipice 90 feet high. At Tillyglens, on the Dorbach
part of the farm, an acre in extent was carried off bodily before the eyes of the
farmer ; and, as he looked at it sailing away, he observed another half acre
detach itself from the hillside and descend some 60 feet into the valley, carrying
a grove of trees on its surface.

But if the flood could thus tear down, in the same proportion it could build
up, often leaving its deposits where they were little welcome. On one of the
farms of Captain Macponatp of Coulnakyle, consisting of 200 acres, 150 were
ruined by a deposit of sand and gravel to the depth of 3 feet. At the mansion
house of Ballindalloch, the garden was covered by sand to such an extent that
only the tops of the apple trees were seen rising through it, presenting a strange
appearance still laden with fruit. Yet more remarkable was the height of the
deposits at the Mill of Logie, near Relugas. The flood completely filled with
sand the lower story of the mill rising 284 feet above the ordinary level of the

river. These examples are instructive ; but, in order to appreciate the subject,

the whole volume should be studied, showing the marvellous power of such
torrents, both in denuding and in building up. If we suppose, that from what-
ever cause,t there had occurred in the old times a series of torrents, surpassing
the Moray floods as these latter surpassed the ordinary summer floods of

* Page 101.

t+ It has been suggested by Mr Bucuan of the Scottish Meteorological Society, that if the bed of
the sea round our coast were elevated, and especially in the direction of Greenland the effect on
the climate would be greatly to increase the river floods. Now already, on stratigraphical grounds, we
have been led to the conclusion that it was precisely at that period of elevation that our high river
terraces were formed. (See note, page 169.) The coincidence is remarkable. The whole strati-
graphical evidence makes it probable that these high gravels were deposited just at the time when
meteorology teaches us to expect that the river floods would be much beyond the present ; and if even
in the present state of things there could be such results as the Moray floods have to show, we may be
prepared for the still more striking effects of that former age.

176 REV. THOMAS BROWN ON THE OLD RIVER TERRACES.

our rivers, nothing could be easier than to explain the whole phenomena of
these terraces with their high-level gravels. |

There need be little difficulty as to where the materials would be found
which were to form the great masses of these sandy and gravelly deposits. The
escars and the associated mounds sufficiently show what immense accumu-
lations of such materials had been provided under the action of ice, and perhaps
of the under-ice rivers of the glacial period. If exposed to the action of torrents
on a somewhat greater scale than those of the Moray floods, such materials
would soon be disposed of. No great lapse of time need to be supposed. If
the whole of the above results in Morayshire were produced in about twenty-
four hours, it would be difficult to say what might not be done by a single
century of such inundations.

Note.—In this paper attention has been called to the absence of marine fossils from the terraces
at Bridge of Earn and elsewhere. If such fossils should occur, it would be important to inquire
whether they belonged to the time when the terraces were formed. Sometimes portions of antecedent
deposits are overlaid or enclosed by the materials of the terraces—portions of rock in situ, for example,
or of boulder clay. In the same way there might be found portions of those marine shell clays which
belong to a previous period. ;

Cea)

IX.—On Spectra formed by the passage of Polarised Light through Double
Refracting Crystals. By Francis Deas, M.A., LL.B., F.R.S.E.

(Read, 6th June 1870.)

It is familiarly known as one of the commonest experiments in optics that
when a beam of polarised light is passed through a thin film of mica or selenite,
and subsequently analysed either by reflection or by double refraction, two
colours are seen complementary to one another, and alternating with one another
at each 90° of a revolution of the analysing plate or prism.

It might be expected that the coloured light thus obtained would, if thrown
into the form of a spectrum by means of dispersion prisms, exhibit some
peculiarities, and such is the case as will be seen from the following experi-
ments :—

To make the experiments intelligible, it may be well in the first place to say
a few words about the instrument employed, and the method of using it.

Any spectrum microscope ought to answer the purpose provided that in
addition to the spectroscopic arrangement a pair of NIcoL’s prisms can be
attached, one below the stage and the other over the eye piece. Both should
be capable of being rotated, and it tends much to facility of working as well as
to exactness of result that both the polarising and the analysing prism should
carry graduated heads, so that their axes may readily be turned to any re-
quired degree of inclination to one another.

The instrument I employed was a large SmirH and Beck. The spectro-
scopic arrangement consists of an adjustable slit attached to the under part of
the substage below the achromatic condenser, and a set of direct vision prisms
which are inserted in the body of the instrument immediately above the object
glass.

By proper focusing, an image of the slit is thus formed by the achromatic
condenser in the focus of the object glass, and a fine spectrum obtained filling
the whole field.

This arrangement, it will be seen, differs considerably from the spectrum
microscope in common use in which the dispersion prisms are placed close to
the observer’s eye, the slit being in the focus of the eye lens. The former
arrangement has this manifest advantage, that owing to the distance of the
prisms from the eye, the spectrum fills the whole field ; also, that the apparent
breadth of the spectrum can be varied at pleasure by a change of the magnify-

VOL. XXVI. PART I. 3A

178 FRANCIS DEAS ON SPECTRA FORMED BY THE PASSAGE OF

ing power employed. Each form of arrangement has, however, its advantages
as well as disadvantages, which it would be out of place to discuss here.*
The polarising part of my apparatus consists of two NIco.’s prisms, for one
of which, when desired, a double image prism can be substituted.
The polarising prism is carried on the substage. It is inserted just above
the slit in a short tube in which it can be freely turned by a graduated head.
The analysing prism is placed in the usual way—in a cap over the eye piece.
The film of selenite to be examined having first been mounted in balsam
between two thin glasses is placed on the stage of the microscope like an
ordinary object.
It is a great convenience in this class of experiments to have the stage of the
microscope not only capable of rotation in the optical axis of the instrument,
but graduated. f
By this means we can at any time, without displacing the film under exami-
nation, adjust its neutral axes at any required angle to the plane of polarisation.
With regard to the mounting of the selenite films for examination the
following method will be found convenient :—Make in the turning lathe several
wooden disks about two inches diameter and one-eighth of an inch thick.
Through the centre of each a hole must then be bored of about half an inch
diameter. A small portion round the hole is then scooped out so as to form a
cup, and in this the selenite is placed and secured with sealing-wax.
The axes of the selenites are then determined and marked on the rims of
the disks.t In this way any two or more selenites can be used in combination
with their axes set at any required angle to one another.
It remains only to trace the course of a beam of light in passing through the
foregoing combination. First the ray, having been reflected from the mirror,
passes through the slit. It is then polarised by the first Nicox’s prism, after
which it passes through the lenses of the achromatic condenser, and appears as
an image of the slit in the focus of the object glass. Having passed through —
the selenite and the object glass, the ray enters the dispersion prisms and is
drawn out into a spectrum. This is magnified by the eye piece through which
the ray, having passed, is lastly analysed by the second Nicov’s prism.
The loss of light from the number of the above media is not so great as”
might be supposed, still an intense source of light is desirable for satisfactory
results. A good artificial light placed close to the mirror will be found the best.
In diffused day-light rays are apt to enter the object glass by reflection from
the brass work without first passing through the polariser, by which the beauty
of the spectrum is impaired. .
* A similar form of instrument will be found described in the “Quarterly Journal of Science”
for October 1869. i

+ The graduated rotatory stage above mentioned, and which is supplied by Surra and Beck,
affords a ready means of doing this.

POLARISED LIGHT THROUGH DOUBLE REFRACTING CRYSTALS. 179

To understand the bearing of the experiments, it is necessary to keep in
view the different effects of a doubly refracting film upon polarised light, accord-
ing to the position of its axes, with respect to the planes of polarisation.

Suppose we take a film of selenite, such as those commonly sold as an
adjunct to the polarising microscope, giving, as its two colours, a pinkish red
and its complementary green. Such a film will, if examined between two
Nico1’s prisms, act on the light according to the following laws :—

1st. When a neutral axis of the film is in the plane of primitive polarisation,
the film will exercise no influence on the light ; if, therefore, the prisms are set
with their axes perpendicular the field will remain dark, if the prisms have
their axes parallel the field will contain only white light.

2d. If the prisms are placed with their axes perpendicular, and the film
is made to rotate, there will be four points of darkness at each quarter of a
revolution, viz., when an axis of the film is in the plane of polarisation, and
between these four points, the same colour (say green) will occur.

3d. If the prisms are set with their axes parallel, and the selenite is rotated,
the field will be white at the four points where it was previously dark, and of
the complementary colour (red) between each of these four points.

4th. If the selenite is fixed with its neutral axis inclined 45° to the plane of
primitive polarisation, and the analyser made to rotate, the field will be alter-
nately red and green in the four quadrants.

5th. The colours are always of maximum brightness when the axes of the
prisms are perpendicular or parallel, and the axes of the selenite inclined 45° to
the plane of polarisation.

Suppose, now, we repeat the above experiments, using the polarising spectrum
microscope above described, and let us call the point in the revolution of the
selenite at which either of its axes is in the plane of primitive polarisation, the
zero point, from which the number of degrees through which it is turned are
measured.

Let the prisms be set with their axes perpendicular to one another, and the
selenite rotated on the stage. The spectrum will, of course, vanish at the four
zero points. Between these points, however, remarkable phenomena occur. A

person unacquainted with the true nature of the colours of polarisation, and

proceeding on the analogy of homogeneous light, might expect to get a spectrum
consisting only of green rays, seeing that that is the colour of the field when the
Spectrum arrangement is removed. This, however, is not the case, and the
result very beautifully illustrates to the eye what is well known theoretically to
be the true nature of these colours. What we obtain is a continuous spectrum
consisting of all the prismatic colours, in greater or less intensity, with the
striking peculiarity that there is a well-marked dark band in the red, similar in
appearance to the well known absorption bands which many substances pro-

130 FRANCIS DEAS ON SPECTRA FORMED BY THE PASSAGE OF

duce in the spectrum, only blacker and better defined than these are ever
seen.

The following is the mode in which the band makes its appearance. As the
zero point is passed, the light first makes its appearance in the green of the
spectrum, from which point, as the selenite is rotated, the light opens out in
both directions. When the light reaches the red, the black band makes its
appearance, and attains a maximum blackness when the selenite is at 45°, viz.,
when, without the use of the dispersion prisms, the field would contain green
light of maximum brightness. When this point of revolution is passed, the
band again fades, the spectrum becomes obscured at each end, the darkness
creeping in towards the green, till at 90° the spectrum has again vanished. The
same phenomena recur at each quarter of a revolution.

Let the Nicou’s prisms now be set with their axes parallel, and the same
selenite rotated on the stage as before. The result is what we should be led to
expect from the last experiment. At the zero points the selenite exercises no
influence, and we have a continuous ordinary spectrum. As a zero point is
passed, however, a dark band makes its appearance, but this time in the green
rays. The band is at first faint and nebulous, but becomes blacker and sharper
as the stage is rotated, till at 45° it attains its maximum. The spectrum in this
experiment never vanishes, but is apparently quite continuous throughout, save
for the appearance of the black band.

Lastly, let the selenite be fixed at 45° from the zero point, and the analyser
rotated. We have now a combination of the two previous experiments. The
band in the red appears alternately with the band in the green at each quarter
revolution, the former being at its maximum when the axes of the prisms are
perpendicular, the latter when these axes are parallel.*

The above are the appearances which present themselves in the case of most
films of selenite of a medium thickness. In some cases, however, two, or even
three, black bands occur simultaneously, these being always followed by as
many complementary bands, when the analyser is turned through 90°. The
number of bands can generally be multiplied by usmg two or more films in
combination, and the appearances can be still further varied by changing the

degree of inclination of the axes of the two films to one another. If the two

films are placed with their similar axes coincident, we obtain, of course, the
spectrum appropriate to a film equal to the sum of the thicknesses of the two
films, while, if dissimilar axes are superposed, the spectrum is that due to the
difference of the same. I have two films which, when properly combined, give
no less than six well-marked bands simultaneously.

* Tf the axis of the selenite makes a greater or less angle than 45° with the plane of polarisation,

the result is that while the same band still recurs after 180° of a revolution of the analyser, the com- ~

plementary band is no longer separated from it by 90°, but by a greater or less angle.

POLARISED LIGHT THROUGH DOUBLE REFRACTING CRYSTALS. 181

But the most striking of the phenomena presented by films which give more
than a single band, remains still to be noticed, viz., the motion of the bands along
the length of the spectrum. This can generally be easily seen by using two films
in combination, and properly adjusting their axes. |

The following may be taken as an illustration of this experiment, of which
many varieties may be made. Suppose the two films are so adjusted as to give
two black bands, one in the orange and one in the blue, which we may call a
and 0 respectively. On rotating the analyser, each band is seen to divide into
two halves. The right hand half of band a runs along the spectrum, and unites
with the left hand half of band 6, which advances to meet it, the two coalescing
into a single band in the green. At the same time that this has been going on,
two entirely new bands have made their appearance. These seem to originate
respectively beyond the visible rays at each end of the spectrum, and to advance
in opposite directions till they are met respectively by the left hand half of the
original band @ and the right hand half of the original band 6. The result is,
that when the analyser has been turned through 90°, we have a spectrum with
three black bands, one in the extreme red, one in the green, and one in the
indigo.

Continuing still further to turn the analyser the above phenomena are re-
versed. ach of the three bands splits into two, moving in the reverse of their
former directions, until when 180° is reached the original spectrum with its two
bands recurs.

A curious variety of this experiment occurs when a circularly polarising film
is interposed between the analyser and the film producing the bands. The
nature of the movements of the bands is now entirely changed, the order of
motion being all in the same direction, and the bands appearing to chase one
another along the length of the spectrum, making their appearance at one end
and disappearing at the other. To produce this effect, the “ band-producing”
film should be set with its neutral axis at 45°, and the circularly polarising film
superposed with its neutral axis in the plane of primitive polarisation. If the
axis of either film is turned through 90°, the motion of the bands is reversed ;
7.é., f the bands formerly moved from left to right, they now move from right
to left. Ifthe two films are both placed with their axes at 45° to the plane of
polarisation, the only effect of the circularly polarising film is to alter the posi-
tion of all the bands by a corresponding amount (¢.e., to increase or diminish
their refrangibility) without affecting the nature of their motions.*

A very pleasing and beautiful variety of the foregoing experiments may be
obtained by using a double image prism as the analyser instead of the Nicot’s

* The effect of circularly polarising the light before it passes through the selenite, is simply that
the occurrence of the bands is irrespective of the inclination of the axis of the selenite to the plane
of primitive polarisation, and depends solely on the position of the analyser.

VOL. XXVI. PART I. 2B

182 FRANCIS DEAS ON THE SPECTRA FORMED BY THE PASSAGE OF

prism. Two spectra formed respectively by the ordinary and extraordinary
ray are thus obtained, which by rotating the double image prism may be made
to lie parallel to one another, or be partially superposed at pleasure, while by
turning the polarising prism the spectra can be made of any desired relative
intensity. Suppose that we adjust the two prisms with their axes at right —
angles, and interpose the selenite used in the first experiment, which gave a
band alternately in the red and in the green, we get now two spectra parallel
to one another, the band in the red of the one occurring simultaneously with
the band in the green of the other. The two bands are thus seen to be strictly
complementary, for the band in the red of the one spectrum appears, attains its
maximum, and vanishes simultaneously with the similar changes of the band in
the green of the other spectrum. This coimcident appearance of the bands,
moreover, is independent of the inclination of the axis of the selenite to the
plane of polarisation, the only effect of a change in which is to increase or
diminish the maximum intensity of both bands alike, a result which, as has
been noticed, does not hold with regard to the alternation of the two bands in
the same spectrum.*

When the two prisms are placed with their axes parallel, so that the two —
images of the slit are seen alongside one another, and consequently the two —
spectra partially superposed and different colours mixed, the appearance of the
bands is very striking. A band occurring in either spectrum is no longer black,
but of the colour of that part of the other spectrum which coincides with it.
The appearance is, in fact, as if a stripe had been cut out of the one spectrum
through which the colour of the other spectrum is seen, while on either side of
the band we have in striking contrast the colours due to the compounding of the
different parts of the two spectra.

The beauty of the effect depends of course greatly on the extent to which
the double image prism separates the two images. It should be so cut that the
compound colours caused by the overlapping of the spectra shall be as different
as possible from either of their constituent colours. The selenite should then
be set at 45°, so as to make the spectra of equal and the bands of maximum
intensity.

With films which give numerous bands the effect is very beautiful, and may
be still further enhanced by rotating the polariser, when the bands will shift
their position, at the same time changing their colours.

Experiments with Sections of Double Refracting Crystals giving Coloured Rings.

The coloured rings produced when polarised light is transmitted through a
double refracting crystal cut perpendicularly to its axis, have always been

* See Note on p. 180.

POLARISED LIGHT THROUGH DOUBLE REFRACTING CRYSTALS. 185

admitted to be among the most beautiful of the phenomena which the science
of optics can produce.

When homogenous light is used it is well-known that the rings assume
entirely the colour of the light used, the spaces between the coloured rings
being black.

The splendour of the phenomena, however, obtained by the use either of
ordinary or of homogenous light, is incomparably inferior to that displayed by
projecting the rings against the spectrum. The spectrum microscope is admir-
ably suited for this exhibition.

The method I adopted was simply to place the section of the crystal imme-
diately over the eye lens of the microscope, and between it and the analysing
prism.

The rings are thus seen of every colour in the spectrum, alternating with
jet black rings between each, those in the red being the broadest; and the
breadth of the rings gradually diminishing to the most refrangible end of the
spectrum.

It is impossible to give any satisfactory idea of the appearances by mere
description, and no little skill or labour would be required to paint any
adequate representation of the effects seen in some of the following combinations.

Take, as an example, a section of a crystal of sugar which gives a very fine
system of rings. I have counted easily.as many as forty-five when projected
against the spectrum. This crystal is one of those which gives in polarised
light two black brushes, not a black cross like Iceland spar. When the NIcot’s
prisms are at right angles the brushes are at their maximum intensity, and the
spectrum with its series of rings is seen to be cut in two by the jet black
brushes. When the analyser is turned through 90° the brushes which would
now, if seen by ordinary polarised light, be white, are of every colour in the
spectrum acording to the part of it they fall upon, and shaded off at their sides
by a nebulous haze of colour through which the black rings are visible.

In intermediate positions of the analyser the brushes become entirely nebu-
lous, so that the rings can be seen through their whole extent. In this position
of matters the circle appears divided into four quadrants, and the rings are
distinctly seen to be dislocated so to speak, i.e, the rings in the alternate
quadrants are pushed out so that each coloured ring in the one quadrant is con-
tinuous with a black ring in the next.

This effect is still better seen by circularly polarising the light before its
passage through the crystal. The effect of this is a curious one. Instead of
the circle being divided into four alternate quadrants, it is now divided into
two semicircles, the rings in the one being alternate with those in the other.
The semicircles are separated by two narrow coloured brushes which revolve
with the analyser, and seem as if they swept out the black rings in the one

184 FRANCIS DEAS ON THE SPECTRA FORMED BY THE PASSAGE OF

segment to be replaced by the coloured rings of the next. If we again circu-
larly polarise the light by interposing a second circularly polarising plate between
the crystal and the analyser, the brushes entirely disappear, and both the black
and the coloured rings are continuous throughout, forming perfect circles.

When the analyser is rotated through 90°, the centre of the system which
was formerly black is now coloured, and, at the same time, all the black rings
have exchanged places with the coloured rings, the change being effected by a
lateral displacement in opposite directions of the two halves of the circle.

If, for the second circularly polarising film, we substitute a film of a different
thickness, the rings assume curiously distorted forms. With one film which I
used the rings became ellipses, with another they all united so as to forma
circular helix, which appeared to unwind like a screw as the analyser was
turned.

The appearances produced by using different crystals are, of course, similar
mutatis mutandis.

By circularly polarising the light before and after its passage through a
crystal of nitre, the brushes are wiped out, and the lemniscates are beautifully
seen, unbroken throughout.

When a crystal of Iceland spar is used, and the Nicow’s prisms set with their
axes inclined 45° we get eight segments, of which the four light segments look
as if they stood out in relief against the dark segments, while the sections of
the black rings, especially near the centre of the system, look more like straight —
lines than circular arcs, and form a system of octagons. 4

The effect upon the rings, produced by placing on the stage a film of selenite
in the position in which it should give the black bands previously described, —
is a strange one. Instead of a black band occurring, the coloured ring
belonging to that partof the spectrum is seen to split into two. It sends off
a branch as it were from its lower part, which shoots across the adjoiing
black ring, and joins itself with the lower part of the next coloured ring. :
This last ring then in turn sends off a branch from its middle part, which in
like manner unites with a third ring, which in turn does the same to its neigh-
bour. All this takes place within the space which should be occupied by the ~
black band. 3

The beauty of these last experiments, wonderful as it is, may be still further”
enhanced by the use of a double image prism as the analyser. The result is”
analogous to that obtained with the same arrangement in the case of the selenite
previously described. We now get not only two spectra but two systems of rings —
which, by superposing the spectra, may be made to interlace with one another. —
Wherever a black ring of the one spectrum crosses a black ring of the other, —
the intersection is of course still black. Where a coloured ring of the one
system crosses a black ring of the other, it retains its original colour ; but if a

POLARISED LIGHT THROUGH DOUBLE REFRACTING CRYSTALS. 185

coloured ring crosses a coloured ring, the intersection is of the resultant colour
of the two combined.

Still more complex figures are got by employing two or more crystals in
combination.

Indeed, there is no end to the variety of exquisite beauty, both in colour and
in pattern, which a little ingenuity may produce. Pigments would be almost
as helpless as words in representing many of these. The appearance produced
by a single crystal with a double image prism as analyser, may be not imaptly
compared to a tesselated pavement of every colour made for a fairy palace,
while that produced by combining two crystals may be said to resemble a suit
of chain armour wrought for a fairy king in jewels of which no two are of the
same hue.

Addition to the above Paper. By J. CLerK Maxwe tt, LL.D., F.RSS. L. & E.

In Mr Deas’ paper a number of interesting experiments are described, in
which, by means of a spectroscopic microscope fitted with polarising and analys-
ing prisms, the true nature of the phenomena observed by Brewster, Bior,
and others, in plates of selenite, &c., is made exceedingly intelligible to the
understanding, while, at the same time, the eye is satiated with new forms of
splendour.

The subject is one to which the attention of experimenters is not so strongly
directed as it was fifty years ago; and therefore it is desirable that the remark-
ably simple methods of observation here described, and the perfection with which
the phenomena may be seen by means of modern instruments, should be more
generally known.

In the text, the paper appears purely descriptive, without any theoretical
application, and the esthetic beauty of the phenomena might be assumed to be
the object of the experiments. But the carefulness of the selection of the
experiments and the faithfulness of the description make me think that the
author himself looked at what he saw in the light of the theory of double
refraction and the interference of light. I, therefore, think that a simple state-
ment of the relation of the visible things here described to the results of theory
would greatly increase the value of the paper; for in scientific education the
identification of what is observed with what is deduced from theory is of more
value than either the process of observation or the process of deduction.

This might be done as follows :—

Begin with the plane polarised light, the equations of motion of which are

@=c cosnt ge.

Now let it pass through a plate of crystal of which the axis is inclined a to
VOL, XXVI. PART I. 3 C

186 FRANCIS DEAS ON THE SPECTRA FORMED BY THE PASSAGE OF

the axis of x; and let this crystal produce a retardation whose phase is p in
the light polarised in the plane of the axis

parallel to axis x’ = ccosacos (nt + p)
perpendicular to axis y’ = csinacosnt.

Next, let the light fall on an analyser in a plane inclined 8 to the axis of
the crystal. The analysed light will be

x” = c cosa cosB cos (nt + p) + ¢ Sina sinB cos nt.
The intensity of this light will be
c {cos 2a cos?B + sin?a sin?B — 2 sina cosa sin B cos B cos p}

orge {1 + cos 2a cos 2B — sin 2a sin 28 cosp }.

We may represent this whole process geometrically as follows :—

Let OCO’ represent the original polarised
light, OCA the angle between the plane of
polarisation and the axis of the crystal.
The light is resolved into ACA’ and DCD’.
Now, let a semicircle be drawn with radius
OA, and let OAp = =p be the phase of retar-
dation ; draw pT perpendicular to AO, and
draw an ellipse with centre C and touching
AO in T and also the other sides of the
parallelogram. This ellipse is the path of the
light emergent from the crystal. Now let BCB’ be the plane of the analyser.
Draw Td T’0’ tangents to the ellipse perpendicular to BB’, then 0C0’ repre
the amplitude of the emergent light.

The result of the process may be made still simpler thus :

Draw CO = ¢, in the plane of polarisation, CA parallel to
the axis of the crystal, and CB parallel to the analyser.
Draw OA perpendicular to CA, AB to CB, and OD to CB,
then CB=c cosacos B, and BD=csinasin B; make DBP=p,
the phase of retardation,and BP=BD. Then CP represents
the amplitude of the emergent light.

The emergent light will be either a maximum or a
minimum when p = 0° or nz.

The minimum will be zero, or blackness, only in the following cases,

1. When a + B=5

and p = 0 or 2nz.

oe wily

2, Whena—B=5 and p= (2n+ 1)z.

POLARISED LIGHT THROUGH DOUBLE REFRACTING CRYSTALS. 187

3. When a = 0 and Bas.
4, When a = 5 and = 0.

To compare our results with the experiments, we observe that for a given
thickness of the crystal p is a function of the kind of light, so that in passing
from one end of the spectrum to the other the value of p increases (or dimi-
nishes) in a continuous manner. When the film is thick, p will make several
entire revolutions within the spectrum. When it is thin, there will be only one
or two, or a fraction of a revolution. Take the case of a thick film, then there

will be a certain set of black bands when 8 = 5 —a. We may call these No. 1.
For these p = 2nz.

When 6B = : +a there will be another set of black bands, No. 2, inter-
mediate in position to No. 1. For these p = (2 + 1) z.

When 8 = 0 or 5 the system of bands vanishes.

When 6 =—a the black bands of No. 1 become bright and of maximum
intensity.

When £8 =a the black bands of No. 2 become bright and of maximum
intensity.

When oa = : all these phenomena are at their greatest distinctness.

In turning the analyser there is simply a dissolution of one system into the
other, without motion of the system of bands in the case of a single plate of
crystal. But if we place the crystal with its axis inclined 45° to the plane of
primitive polarisation, and place above this a film of retardation . with its axis
parallel to the original polarisation, then we have as before for the light emerg-
ing from the first crystal,

B= e708 (nt +p) y= c+. cos nt.

r/2
Resolving these rays in the direction of the axis of the second film, we have
ef = 5¢ (cos (nt + p) + cos nt)
= se (cos (nt + p) — cos nt),

and since w” is retarded : it becomes

—— x (sin (nt + p) — sin nt),

188 ON THE SPECTRA FORMED BY THE PASSAGE OF POLARISED LIGHT, ETC.

y” remaining the same. We may put these values into the form

pe i ma
i ecos (nt + p) cost

W P) gin 2
y = ccos (nt + 4) sing.

This shows that after emerging from the circular polarising film the ray is

plane-polarised, that the plane of polarisation inclined : p to that of primitive

polarisation.

If the emergent light is analysed by a dispersion prism, and a Nicot’s prism
inclined 8 to the plane of primitive polarisation, there will be black bands
(perfectly black) for all colours for which

p = 2B or 2B + 2nz7,

and as the prism is turned these bands will march forwards in a regular manner
across the spectrum.

This very beautiful experiment, in which the phenomena of rotatory polarisa-—
tion are imitated, is not so well known as it deserves to be. One form of it is
due, I believe, to Brot, and another to WHEATSTONE, but the arrangement here
described is by far the most convenient.

When the second plate is thick, then for some points of the spectrum its’
retardation is (27+4) 7. At these points the bands will move forwards when
the analyser is turned. At an intermediate set of points the retardation is
(2n—4)7. At these points the bands will appear to move backwards. At
intermediate points the retardation is m7. At these points the bands will not
move, but will become deeper or fainter. I suppose this to be the explana- |
tion of the experiment described at p. 181, but the arrangement of the films
is not very precisely described.

The experiments with the rings in crystals are very well described, and
must be beautiful, but are not so instructive to a beginner as those with the
selenite plates. Those, however, who have made out the meaning of the expe-
riments first described have a good right to regale themselves with gorgeous
entanglements of colour.

isiso.e)

X.—On the Oxidation Products of Picoline. By James Dewar, F.RS.E.,
~ Chemical Demonstrator in the University of Edinburgh, and Lecturer
on Chemistry at the Edinburgh Veterinary College.

(Read 6th June 1870.)

The combined researches of ANDERSON and WILLIAMS on the basic compounds
contained in coal tar have led to the discovery of two well-defined series of
organic bases, called respectively the Pyridine and Chinoline series, the mem-
bers of both of which possess the properties of nitrile bases. The isomerism
between the pyridine and the aniline series of bases excited considerable
interest at the time of its discovery. The subsequent researches of WILLIAMS
on the products of the distillation of chinchonine led to the discovery of bases
having the same composition as the members of the pyridine and chinoline
_ series of coal tar. When first discovered they were supposed to be identical.
Since that time, however, a careful examination and comparison of the tar
series of bases with the chinchonine series has led Mr WILLIAMs to the interest-
ing discovery that the two lutidines, as also the two chinolines, are in reality
not identical, but isomeric. This introduces a greater complexity into the
study of the constitution of these compounds.

I began this investigation in the summer of 1867 under the able direction
of Professor Auc. KEKULE, in the University of Ghent. At that time the
whole of the then known facts regarding the properties of these bases had been
accumulated by ANDERSON and WILLIAMS, and by Perkin, who had obtained
pyridine from a naphthaline derivative. Save by this latter method the only
process of preparation known was destructive distillation. All the attempts
that had been made to elucidate the internal constitution and relationship of
these bases had failed to yield positive results, their extreme stability in
presence of the most powerful reagents presenting a barrier to investigation.
In 1869 Professor ADoLPHE BAEYER made the brilliant syntheses of picoline
through the action of tri-brom-allyl and of acryl aldehyd, respectively, on
ammonia; and through the action of higher aldehyd homologues has shown the
reaction to be general. Thus, by the synthetical labours of BAEYER, we have
acquired for the first time a definite knowledge regarding the mode of forma-
tion and constitution of this class of organic compounds.

Having formerly employed permanganate of potassium in the oxidation of
phenol (see Proc. Roy. Soc. Edin. Session 1866-67, p. 82), I naturally attempted
the oxidation of these bases by the same agent; and I have since found that
‘VOL. XXVI. PART I. 3D

190 MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE.

A. W. Hormann had successfully employed it to oxidise chinoline into ammonia
and oxalic acid. Finding the members of the pyridine series to be easily
attacked by this reagent, I commenced a careful examination of the products
of the oxidation of picoline, with the object of learning something regarding its
internal structure. A preliminary note on the results obtained was communi-
cated to the British Association at its Norwich meeting, 1868 (see Report Brit.
Assoc. 1868).

I am indebted to my friend Dr Ronatps of Bonnington for a liberal supply
of a quantity of bases that he had carefully prepared, with the object of insti-
tuting an investigation into these compounds himself. The crude bases placed
in my hands had been repeatedly fractionated on a large scale, and the indi-
vidual fractions were thus tolerably pure to begin with. Finding they contained
traces of pyrrol and hydrocarbons, I redissolved in acid the fraction boiling
from 130° to 160° C., and subsequently treated it in the way recommended by
ANDERSON and WILLIAMs to purify these bodies. The mixture of bases thus
obtained was subjected to a series of careful fractional distillations. With the
object of effecting the best possible separation, [ employed the method recom-
mended by WARREN, and found it admirably suited to effect a comparatively
easy separation, so far as fractional distillation can be made to yield a pure
product. The bases were transferred to a retort connected with an ascending
spiral of copper tube enclosed in a paraffine bath of large dimension, the tem-
perature of which was continuously equalised by constant stirring. Five suc-
cessive fractionations by this method gave a separation as complete as was
necessary for the object I had in view. From the laborious researches of
ANDERSON and WILLIAMS, we know that these bases for a large range of tem-
perature have the same composition, and that perfectly pure products can be
obtained only through the fractional precipitation of the platinum alts.
Although I did not make an exhaustive separation by WARREN’s method, the
fractionation was so effective that, by comparing the temperature of the boiling
vapour in the retort with the temperature of the paraffine bath throughout the
whole course of a distillation, a difference of 10° C. at starting gradually
increased to 20°. No fraction that I obtained, separated in this process by a
difference of 2° C. in the imtermediate condenser, had a perfectly constant
boiling point. On different occasions fractions separated in this way, boiling
between 130° and 140° C., have been employed in the following experiments.
Analyses of the platinum salts of portions boiling between those temperatures
showed the fractions to be substantially picoline, with a possible trace of
lutidine.

Picoline, as is well known, resists oxidation by the most powerful agents
adapted for this purpose, nitric and chromic acids being without visible action,
even at high temperatures. Of all oxidising agents, permanganic acid, or its

MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE. 191

potassium salt, seems to be the most powerful. Having on a former occasion
employed this reagent to effect the oxidation of phenol alcohol, I naturally
investigated the action effected on these nitrile bases. I found that the whole
of the members of this series of compounds could be readily oxidised by the
use of this substance. The higher members of the series were oxidised more
readily than the lower, but in all cases it was effected with ease and rapidity.
The following is a description of the apparatus used in the experiments on the
oxidation preducts of these substances and of the mode of conducting the
operation. A flask, about three litres in capacity, was connected by a wide
tube with a large reversed Lizpie’s condenser, so as to effect a rapid condensa-
tion of volatile products and their immediate return to the field of chemical
action. The flask having been placed on a sand bath, 150 grms. potassium
permanganate, 14 litres water, and 25 grms. picoline were introduced, and the
whole heated to near the boiling point. The reaction began suddenly, with
great evolution of heat, necessitating the removal of the source of external heat.
The reduction of the permanganate was completed in half an hour. After the
contents of the flask had cooled, the oxide of manganese was separated by
filtration from the strongly alkaline liquid, and washed repeatedly with boiling
water. The alkaline liquid was then transferred to a flask, and the basic sub-
stances distilled off. The residual alkaline liquid was then concentrated by
evaporation to 200 c.c. and 300 c.c. and dilute sulphuric acid (containing 70 per
cent. H,SO,) added to it. After standing for some time this acid liquid became
thick from a deposit of long white crystalline needles of a complex of new
acids. In different experiments the relative proportions of the reacting sub-
stances were considerably varied, but the yield of the new acids in every case
was small, a large portion of the picoline having been completely oxidised, while
some of it had remained unacted upon. This must always be the case, as a
large quantity of the original base, from the violence of the reaction, was driven
away from the flask, and, when condensed, it fell back into a boiling liquid of
increasing alkalinity, in which the base was comparatively insoluble. After
separating, by filtration, the new crystalline acid referred to, the filtrate was
transferred to a retort, and the volatile acids distilled off.

GENERAL RESULTS OF OxIDATION.— When the dilute alkaline fluid was taken
immediately after the permanganate was exhausted, it was found to contain
carbonate of potassium. When neutralized with hydrochloric acid and chloride
of calcium added, a white precipitate of oxalate of calcium was obtained. The
oxalate was mixed with a small quantity of some higher acid, probably malonic,
| as the per-centage of lime found was 36:3, oxalate containing 38°3 per cent.
The quantity of volatile acid produced by the reaction was small. The
presence of nitric and acetic acids, however, was readily proved. The surplus
base remaining after the oxidation operation was transformed into the double

192 MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE.

platinum salt, and a fractional crystallisation made. The platinum was deter-
mined in these different salts as follows :—
1st Crystallisation—0°177 grm. pt-salt gave 0'077 grm. pt=43°7 per cent.
2d 5 —0°6910 ,, - 02102 ,, ==83°95 per cent.
The first sample agreed in composition with the double chloride of platinum
and ammonium ; the second, with that of a mixture of the double platinum salts
of pyridine and picoline. There was produced by the reaction, therefore,
carbonic, nitric, oxalic, acetic, and a complex of new acids, ammonia, and _pro-
bably a small quantity of pyridine. The relative proportion of the products
obtained depended on the quantity of the oxidising agent used, the volume of
the solution, rapidity of the action, and the quantity of bases present in the field
of action. . 7
DI-CARBO-PYRIDENIC Acip C,H,N | COs The crystalline acid substance ‘
separated by adding excess of sulphuric acid to the concentrated alkaline fluid,
after standing overnight was collected on a filter, and repeatedly crystallised
from hot water (in which it was easily soluble) until free from oxalic acid and
potassium salts. When first separated from the acid solution it appeared in —
long needles ; but after several recrystallisations from water it was obtained in
the form of perfectly colourless plates, resembling naphthaline. (The crystallo-
graphic constants of this acid have not yet been determined.) The crystals |
did not contain any water of hydration. The following are the results of the
analyses of the acid and its silver salt :-—

Acid. “a
Weight of acid taken, . : F : : ‘ ; 5 0°2195 grm.
Carbonic anhydride produced ' , : : : : : 00685 _,,
Water produced, ; 0°4040 _,,
Nitrogen found by Gorries’s method to beat to the CO, produce the
proportionate vol. of, : : ¥ 5 : ~~ Loo Ta

Calculated centesimally these figures give—
Sample. C,H;NO,.

Carbon, } : : : : : ‘ ; : 50:19 50°29
Hydrogen, . : , : : : : , ‘ : 3°47 2°99
Nitrogen, ‘ ; : : : : : ; : — —_—

Silver Salt.*
As with the acid, the salt was dried at 100° C.

Is Il.
Weight of salt taken, . : : : ; 0°8766 grm. 0:5456 grm,
Carbonic anhydride produced : , : A 0°6985 _,, 0°4518 _,,
Water produced, — : ; 3 : : 0°0554 _,, 0'0602 ,,
Weight of salt taken, . ; , , 4 ‘ 0°6980 grm. 0°3951 grm.
Silver obtained, . é : ; 2 : : ILI yy 06525 ,,

* J, and II. were samples of silver salts obtained from different experiments. I. was got by doubl
decomposition of the sodium salt ; II. from the ammonium salt.

MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE. 193

Calculated centessimally these results give—

i Il. C,H,NAg,0,.
Carbon, ; é : ; : ‘ : 21°73 22°58 22°04
Hydrogen, . : ‘ 5 : . ; ; 0°70 1:22 0°78
Silver, . : a : , ; : ; ; 56°60 56°78 56°69

In order to determine the equivalent of the acid, I took 0°5392 grm. of acid
dried at 100° C. and titrated with pure caustic soda solution, every 1°813 c.c.
containing 23 merm. of sodium ; 11°7 c.c. of soda neutralised the acid taken. The
point of saturation was well marked. The equivalent of the acid, from this de-
termination, was, therefore, 83°55 ; as determined by analysis of the silver salt it
was 83°5. The atomic weight of the acid taken as C,H,NO, was exactly double the
equivalent found, which, if true, would necessarily involve the acid being bibasic.
In order to determine the basicity of the acid, the ammonium salts were the
only combinations that I specially examined. 0°4739 grm. of the acid carefully
treated with excess of ammonia, and dried at 100° C., increased in weight by
00481 grm., gain = 10°13 per cent. ; gain on the acid ammonium salt of fore-
going formula = 10°17 per cent. The neutral ammonium salt was extremely
soluble in water, whereas the acid salt was much less soluble, and could readily
be obtained in the form of fine silky needles when the solution was evaporated.
From the above data there can be little doubt that the acid was bibasic, bearing
the same relation to pyridine that phthalic acid doestobenzol. The acid melted
at a temperature of about 210° C., frothed, evolved a small quantity of carbonic
anhydride, and emitted the readily recognisable smell of these bases. It was
easily decomposed when heated with soda-lime, evolving a basic substance, no
doubt pyridine. The mercury, copper, cadmium, and zinc salts were all readily
soluble in water. The barium and calcium salts were also soluble, and were
obtained by adding the respective chlorides to the neutral sodium or ammonium
salt ; they crystallised in minute prismatic needles. The silver salt of this acid
was specially characteristic. On the addition of nitrate of silver to a solution of
the acid or its neutral ammonium salt, a white gelatinous precipitate immediately
separated out—it was insoluble in boiling water, and was not visibly affected by

exposure to light. The insolubility of this salt would enable us more readily to

separate the acid from the other products of the oxidation reaction than the
process first quoted in this paper.

Along with the acid just described, as separated by the process detailed,
there was found associated with it another acid substance having a very much
higher atomic weight. The crystalline mass, obtained by the addition of sul-
phuric acid to the alkaline liquid from the oxidation operation, purified by solu-
tion in and recrystallisation from alcohol, and dried over sulphuric acid by means
of an air-pump, had an equivalent weight of 121. The crystals were hydrated,
and lost 4°9 per cent. of their weight when dried at 100° C. The sodium

VOL. XXVI. PART I. 3 E

194 MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE.

salt of this mixture gave, on the addition of nitrate of silver, a gelatinous preci-
pitate agreeing in appearance and composition with that got from dicarbopyri-
denic acid ; it contained 56°6 per cent. of silver. The equivalent of this mix-
ture, as determined by the composition of the ammonium salt dried at 100° C.,
was 336; 0:8335 grm. of acid mixture dried at 100° C., and treated with
ammonia, increased in weight by 0:0422 grm. This acid substance, therefore,
was clearly a mixture of dicarbopyridenic acid with some acid of a very much
higher atomic weight. It remained solid when heated to 220° C., at which
temperature dicarbopyridenic acid readily melted, and was much less soluble
in water than the latter.

The difficulty and expense of obtaining these oxidation products in any
quantity, prevented me from making the exhaustive investigation I would have
liked. .

The formation of dicarbopyridenic acid by the oxidation of a mixture of
picoline and lutidine, whether obtained from lutidine alone or by the complete
_ destruction of picoline, is quite analogous to the formation of phthalic or teraph- —
thalic acid, by the oxidation of the homologues of benzol, or by the complete —
destruction of benzol itself, as shown by Cartus; the only difference being that
Carius employed the lowest member of the benzol series, whereas picoline is the
second known member of the basic series. The production of the same acid from
pyridine itself would in no wise influence speculation regarding the constitution
of the higher members of the series. For the present, we may consider pyridine
as the nucleus from which all the other members of the series are derived. —
Although such a supposition must be considered purely hypothetical, in reality —
it is a great advantage to classify by analogy, relatively to other series, disjointed
groups of organic compounds. The two series of bases, viz., the coal-tar and
the chinchonine, bear the same empirical relation to pyridine that benzol does
to its homologues and to naphthaline.

Benzol. Toluol. Naphthaline.
C,H, C,H, C,H,
CAB; Cpt,
Pyridine. Picoline. Chinoline.
Oris lels| C,H,N Cn
CH, C,H,

Now, although it has not yet been proven that lutidine and chinoline have a
similar formative relation to pyridine that tolnol and naphthaline have to
benzol, still it is by. no means an improbable analogy. The isomerism in the
pyridine series, so far as is known, commences with the third member, lutidine,
as found by WILLIAMS on comparing the chinchonine lutidine with the coal-tar
lutidine ; whereas the chinoline obtained from either source differs essentially
in chemical characters. If we consider picoline as in all likelihood methyl-

MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE. 195

pyridine, then the « and # lutidines may, in all probability, be represented as
dimethyl-and ethyl-pyridine respectively, and we would expect the dimethyl-
pyridine to give directly dicarbopyridenic acid, or an isomer, on oxidation.
Pyridine may be written graphically as benzol in which nitrogen functions in
place of the triatomic residue CH”’, and thus may be represented as a closed
chain,—

G36. 7
lala ape)

And, considering the stability and mode of formation of these bases it is not at
all improbable that they may not be produced by the simultaneous action of
acetylene and its derivatives on hydrocyanic acid; thus as three molecules of
acetylene condense and form benzol, so may two molecules of acetylene, and
one of hydrocyanic acid, condense and produce pyridine.

There is a large class of substances that bear the same relation to the mona-
mines that dicarbopyridenic acid does to pyridine, with this difference, that the
best known are all monobasic instead of being bibasic acids. Thus glycocol,
alamine, leucine, and their homologues, may be looked upon as the monocarbo-
acids of the ethylamines, in which the carboxyl radicle is united directly to the
carbon, the isomeric carbamic acids (or urethanes, as they are called) being the
derivatives in which the carboxy] is united directly to the nitrogen,—

Ammonia. Methylamine. Glycocol. Methyl-carbamic Acid.
H CH, CH,CO,H CH,

NEL NH NH NH
H H H CO,H

Analogous derivatives are obtained from the aromatic ammonias. A class of
derivatives similar to the above, must necessarily be derivable from the diamines.
In the case of the nitrile bases or triamines, only derivatives could be obtained
analogous to glycocol and its homologues. Bodies of a like constitution to
glycocol readily break up into carbonic anhydride, and the corresponding
ammonia ; the reverse transformation has not yet been effected. The amido-
bibasic acids bear the same relation to the monamines that dicarbopyridenic
acid does to pyridine, with the exception of the difference in the constitution of
the closed nitrile nucleus. No bibasic acid, other than pyridenic, has been
| discovered. A strictly analogous compound, in the monobasic series, is the
acid carbo-pyrrolic, C,H,NO,, which bears a similar relation to pyrrol that the
amido-mono-carbon acids do to the ammonias, only that the pyrrol, although a
| nitrogenous body, is not, strictly speaking, a nitrile base. But the close analogy

196 MR JAMES DEWAR ON THE OXIDATION PRODUCTS OF PICOLINE.

between pyrrol and true nitrile bases, they being simultaneously produced in
the majority of reactions, would lead us to expect a like class of derivatives
being obtainable from pyridine.

Husner (Ann. Chem. u. Pharm., vol. 141) has described the oxidation
products of nicotine, got by the action of sulphuric acid and bichromate of
potassium. In that memoir he describes an acid so obtained, having the
formula C,H,NO,. This acid is identical in composition with mono-carbopyri-
denic acid. The base nicotine itself differs from dipyridine by only four hydro-

Nicotine. Dipyridine.
C,H,N C,HN
C,H,N Gea

gens; and as the nucleus of nicotine is a nitrile nucleus, it is not at all impro-
bable that this acid may be a member of the series to which dicarbopyridenic
belongs, so that nicotine may be similarly constituted to ANDERsoN’s poly-
merised. bases.

The stability of these bases to the majority of reagents (especially the
primary member of the group, pyridine) would predispose us to look upon it as
analogous to benzol, and to suppose that the atoms are symmetrically grouped.
The syntheses of BaryER support this view, and there is not any reason why
we may not have as many stable derivatives from this nucleus as from benzol. I
have already pointed out the analogy between the chinoline series and the
pyridine series; and in a short time I hope to be able to publish details support-
ing the theoretical relations above given.

In the meantime, the following analogies may be pointed out between
benzol- and nitrile-derivatives ; thus—

Benzol. Naphthaline. Anthracine. Pyridine. Chinoline.
C,H, C,H, C,H, C,H, C,H,N
CRE (opie E CoH, NCH Grlels
C.Be C,H, G7 3 C,H, CPE:

Indol, from its general characteristics, evidently belongs to the pyrrol series,
the following showing, in all probability, the relative structure of indol and
pyrrol :-—

Indol. Pyrrol.
C,H, C,H,
NE. NREL

C,H, C,H,

According to this hypothesis indol is simply benzol-pyrrol.

NEM

pT pW] suey Te T }1ap atquosdteqy gr

VF. M‘ Farlane, lath” Edin?

J B. Abercrombie del

RS eg ee es

W.H. MS Farlane Lith? Edin™

JBA& WT del

. i Pa le ee td

WH. M°‘Farlane, Lit

1B. Abercrombie delt

(ets)

XI.—An Account of the Great Finner Whale (Balenoptera Sibbaldii) stranded
at Longniddry. Part I. The Soft Parts. By Wm. Turner, M.B. (Lond.),
Professor of Anatomy in the University of Edinburgh. (Plates V., VI.,
VII., VIIL)

(Received November, 1870.)* \

CONTENTS.

PAGE PAGE
Introduction, ; ‘ : : . 197 Baleen, . : ; ; : 5 BY
External Form and Dimensions, . . 199 Organs of Alimentation, : ‘ n 222,
Colour, . : ‘ : : é . 202 Organs of Circulation, . : , . 227
Fetus and Membranes, . ‘5 ; 5 PAO Organs of Respiration, . : ; 5 Pets
Skin and Blubber, F : : . 209 Genito-urinary Organs, . i : . 240
Mammary Gland, . : : : + wall Comparison with other Finners, . . 242

On the 3d November 1869, a huge Finner whale was stranded on the beach
at Gosford Bay, Longniddry, Firth of Forth.

Most of the large Fin whales which have been examined by British and
Continental anatomists have been found floating dead on the surface of the sea,
and have then been towed ashore by their captors. But, from the account
which was given in the Edinburgh daily newspapers, it would appear that, for
some days previously, this animal had been recognised by the fishermen, swim-
ming to and fro in the Firth. On the morning of the 3d it was seen from the shore,
blowing with great violence from its nostrils, flapping its huge tail, and obviously
struggling to disengage itself from the rocks and shoals, amidst which an un-
usually high tide had permitted it to wander. Shots were fired at it, and, from
the wounds produced, blood poured forth which tinged the surrounding waves.
As the tide receded, the animal was fairly stranded ; and, after some vigorous
but ineffectual attempts to disengage itself from its position, it slowly died.
The animal lay some yards above low-water mark, so that for several hours each
day it could be examined, and photographs taken from various points of view.

Under the powers conferred by Act of Parliament, the carcase was taken
_ possession of by the receiver of wrecks for the Board of Trade and sold by public
auction. It was purchased by Mr Joun Tarr, Oil Merchant, Kirkcaldy, for
L.120. After lying for a fortnight on the beach at Longniddry, a strong rope

* A preliminary account of this animal, illustrated by a number of specimens, photographs, and
drawings, was read to the Society on the 20th December 1869, and an abstract of this communication
was printed in the Proceedings of that date. By permission of the Council I have been allowed to
supplement the preliminary notice with additional observations, and to extend it in a form for the
Transactions of the Society.

VOL. XXVI. PART I. 3 F

198 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

was secured around the root of the tail, and, when afloat at high water, it was
towed by a powerful steamer to Kirkcaldy, a town on the opposite shore of the
Firth, distant about ten miles.

It was flensed on the beach, immediately to the east of Kirkcaldy harbour ;
and, as this could only be done at low water, the process of removing the
blubber, taking out the fat within the abdomen, cutting off the baleen and flesh,
disarticulating and removing the bones, occupied several men for nearly a —
month.

As one of the largest sized whalebone whales comes so very seldom within
comparatively easy access of a great city, the opportunity was taken by crowds ~
of persons to inspect the huge creature, not only as it lay on the beach at
Longniddry, but whilst the process of cutting up was going on at Kirkcaldy.

As the classification and structure of the larger Cetacea possess many in-
teresting points for investigation, I gladly availed myself of the presence of this
rare visitor to devote such time as I could spare, in the midst of the work of
the University session, to its examination.

The colour, general form, and dimensions of the animal were observed when
the whale was lying on the shore at Longniddry. The observations on the in-
ternal structure were made as it was being cut up at Kirkcaldy, or on speci-
mens which were brought over to the Anatomical Museum of the University,
and submitted there to a more careful examination than could have been con-
ducted on the sea beach.

The distance from Edinburgh at which the whale was lying, during the
flensing, rendering a journey by rail and steamer necessary at each visit, the
exposed position of the animal on the sea beach below high-water mark making
access to it practicable only at low water, the great bulk of the creature, the
difficulty of getting at the internal parts owing to the size of the cavities, the
greasy, slippery condition of all the surroundings, and the impediments offered
to handling or removing the viscera on account of their magnitude and weight, —
have made the examination of this whale a very laborious task. For these
reasons, aS well as from the putrid state into which the carcase passed, the
extremely offensive gases generated by so huge a mass of putrifying flesh, and
the great heat evolved by its decomposition, it was impossible to study many of
the structures to which I should have wished to have devoted my attention.
In many respects, therefore, I regret to say that my description will necessarily —
be incomplete and fragmentary.

In conducting the examination, I was most ably assisted by the thoroughly
cordial and, I may say, enthusiastic, co-operation of my assistant, Mr STIRLING, —
and my pupils, Mr Mitten Coucurrey and Mr James Foutis, to whom I take
this opportunity of expressing my thanks for the important aid which they ren-
dered. To Mr Joun Tarr of Kirkcaldy I and my assistants are indebted for —

STRANDED AT LONGNIDDRY. 199

permission to examine the parts as they were exposed during the flensing, and
to remove such specimens as could conveniently be taken away.

External form and dimensions.—The whale was a female. When I first
saw the animal on Gosford beach, it was lying with its head pointing inland,
and it rested on the right side of the belly, chest, and right lower jaw. The
middle line of the belly was in contact with the ground, and the under surface
of its horizontal tail lay on the shingle. The head, owing to its great weight,
had fallen over to the right, so that it overhung the right lower jaw, and per-
mitted the whole length of the inner surface of the left half of the lower jaw,
and a large part of the dorsum of the tongue to be seen, together with the outer
edges of the baleen plates on the left side (Plate V. fig. 1). ;

The length of the animal, measured with a graduated tape-line along the
curve of the middle line of the back from the tip of the lower jaw to the end of
the tail, was 78 feet 9 inches. The girth of the body immediately behind the
flipper was estimated at 45 feet, dimensions which it preserved almost as far
back as the extent of the abdominal plicz, behind which it tapered off rapidly
to the tail. Its girth in line with the anal orifice was 28 feet, whilst around the
root of the tail it was only 7 feet 9 inches. In front of the flipper the girth
was considerable, as far forward as the swell or greatest projection of the lower
jaw, but in front of this it tapered off to the symphysis. The lower jaw arched
outwards and forwards with a wide sweep from the angles of the mouth ; then the
two halves converging met at the symphysis and formed there a keel-like ridge.

_ The tip of the lower jaw projected 145 foot beyond the tip of the upper jaw.
_ The inner surface of the lower jaw was bevelled off close to its upper border,
so as to admit the edge of the upper jaw within it. The length from the angle
to the tip of the mouth, along the upper curved border of the lower jaw, was
21 feet 8 inches, and 17 feet 4 inches im a straight line.

The dorsum of the upper jaw was not arched in the antero-posterior direc-
| tion as in the Balena mysticetus. It sloped gently upwards and backwards to
the blow-holes, from which a low but readily recognised median ridge passed
forwards on the beak, gradually subsiding some distance behind its tip. On
each side of this ridge was a shallow concavity. Immediately in front of the
blow-hole the ridge bifurcated, and the forks passed backwards for several
inches enclosing the nostrils, and then subsided. The outer borders of the
| upper jaw were not straight, but extended forward almost parallel to each other
from the angle of the mouth for some distance in a gentle curve, and then con-
verging in front formed a somewhat pointed tip. Their rounded palatal edges
fitted within the arch of the lower jaw. The transverse diameter of the upper
jaw over its dorsum between the angles of the mouth, was 13 feet 3 inches.

From the blow-holes the outline of the back curved upwards and backwards,
| it was uniformly smooth and rounded, and for a considerable distance presented

200 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

no dorsal mesial ridge. But somewhat in front of the posterior fourth of the
back a ridge appeared, which culminated in the dorsal fin. Unfortunately the
height of this fin could not be taken, as the summit had been cut away before I
saw the animal. It was triangular in form, its anterior border convex, its
posterior border falcate, whilst its apex had obviously projected upwards and
backwards. A line drawn from its posterior border vertically down the side of
the whale reached the ventral mesial line some distance behind the anus. From
the tip of the lower jaw to the anterior border of the dorsal fin was 59 feet 3
inches. Behind the dorsal fin the sides of the animal sloped rapidly down-
wards to the ventral surface, so that both the dorsal and ventral mesial lines
were clearly marked, and the sides tapered off backwards to the tail. .

The lobes of the tail curved outwards and backwards from the terminal
part of the sides of the animal; a rounded interlobular median notch marked
the termination of the caudal spine, and separated the two lobes from each
other. The anterior border of each lobe was rounded, and convex from root to
tip, the posterior was sharp, and concave from root to tip; the tip was pointed
and the surfaces flattened. The greatest girth of one of the tail lobes was 5
feet 8 inches, whilst the distance between the tips of the two lobes was some-
what more than 16 feet.*

The ventral surface of the throat, and the sides and ventral surface of the
chest and belly, were marked by numerous longitudinal ridges and furrows.
Some extended as far forward as the symphysis of the lower jaw, others to the
angle of the mouth ; some mounted as high as the root of the flipper, and even
above its posterior border. These folds terminated at their hinder ends with
ereat regularity along a line, which commencing some distance behind the root
of the flipper sloped obliquely downwards and backwards to the ventral sur-
face. The ventral folds were consequently the longest, one about the middle of
the belly measured 45 feet. The number of these folds on each side of the
ventral mesial line it was difficult exactly to determine, on account of the posi-
tion in which the whale was lying, but at least thirty appeared to be present,
though as a ridge occasionally bifurcated or gave off a branch, and as, after some
time, its forks blended with adjacent ridges, the number necessarily varied in
different localities. When I first saw the animal the furrows separating the
ridges were not more than from 4 to 3 an inch broad, whilst the ridges them-
selves were in many places 4 inches in breadth, but as the body began to swell
by the formation of gas from decomposition, the furrows were opened up, be-
came wider and shallower, and the ridges underwent a corresponding diminu-
tion in breadth. At the same time a considerable change took place in the
contour of the body in the thoracic and abdominal regions, which presented

* The extreme ends, probably one foot from each lobe, had unfortunately been cut away before _
the measurement was taken.

STRANDED AT LONGNIDDRY. 201

a huge lateral bulging, giving a greater girth than when it first came
ashore. Close to the posterior ends of the mesial abdominal plice was a deeply
puckered scar, the umbilicus.

The flipper projected from the side of the body 31 feet 4 inches behind the
tip of the lower jaw, measured in a straight line, and 14 feet behind the angle
of the mouth. It curved outwards and backwards, terminating in a free pointed
end. Its surfaces were flattened ; its anterior border rounded and convex from
root to tip, measured 12 feet 3 inches ; its posterior border concave from root
to tip 10 feet, whilst its girth at the root was 9 feet 6 inches. The distance
between the two flippers, measured over the back, between the anterior borders
of their roots, was 18 feet 6 inches.

The slit-like entrance to the female passage was situated 22 feet in front of
the fork of the tail. Its antero-posterior diameter was 16 inches. It was
bounded laterally by elongated prominent folds of the integument, which
represented the labia majora, and were indented by longitudinal furrows.
In front of the aperture was a rounded elevation representing the mons,
which was placed 10 feet behind the longitudinal plicee on the middle of the
belly. Behind the mons was a deeply depressed part of the imtegument,
immediately posterior to which was a thick clitoris, triangular in its outline.
Its length was 6 inches, the breadth at the root 4 inches (Plate VI. fig. 6). The
clitoris curved backwards, and overlapped the external orifice of the urethra,
which orifice was surrounded by a well-marked fold of mucous membrane.
Both on its superficial and deep aspects it presented a rugose appearance.
On each side of the root of the clitoris a projecting fold lying between the
labia majora passed backwards, external to the urinary meatus. These two
folds formed the labia minora; they bounded the vestibule, and their inner
surfaces, as well as the floor of the vestibule, possessed a number of complex
ridge-like elevations of the mucous membrane. When this membrane was cut
through, a quantity of erectile tissue, in which were many large veins, was
seen. Eight inches on each side of the female passage was a funnel-shaped
elevation of the integument, at the summit of which a circular aperture, which
readily admitted the tips of the fingers into a fossa about 4 inches deep, was
seen. Projecting from the bottom of this fossa, but not through the circular
aperture at its summit, was a large nipple about 3 inches long, which possessed
an orifice at its free end—the termination of the great lacteal duct—into which
the forefinger could be passed. A number of pedunculated papille were situated
at the summit of the nipple around this orifice (fig. 7).

Thirteen inches behind the female passage was the orifice of the anus,
which was small and contracted, but could easily be dilated so as to admit
the hand. The integument immediately around the orifice was rugose, and in
the neighbourhood both of the intestinal and genital openings the skin was

VOL. XXVI. PART I. 3G

202 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

indented by several longitudinal furrows (fig. 8). A well-marked sphincter was
observed beneath the integument around the anus. The mucous membrane at
the anal end of the rectum had a blackish tint.

The eye was situated immediately above the angle of the mouth, from which
it was 1 foot 6 inches distant. The fissure between the lids lay antero-pos-
teriorly. The ear orifice was a narrow slit situated in a line behind the eye,
from which it was distant 3 feet 10 inches. The transverse distance over the
dorsum between the two eyes was 11 feet 5 inches, the corresponding distance
between the two ears was 13 feet 7 inches.

The blow-holes were placed in the fossa between the two subdivisions of
the dorsi-mesial ridge of the beak. Two longitudinal slits or nostrils, each
large enough to admit the extended hand, were separated by an intermediate
septum. Anteriorly the slits were only 4 inches asunder, but owing to their
divergence the posterior ends were 15 inches apart, and the transverse diameter
of the septum was correspondingly increased. The upper surface of the septum
was marked by a longitudinal mesial groove. The antero-posterior diameter
of the blow-holes was 1 foot 6 inches. From the tip of the lower jaw to the
anterior end of the blow-holes, 14 feet 9 inches. From the anterior end of
the blow-holes to the mesial notch of the tail 64 feet.

Colour.—On the dorsum of the beak and of the cranium, on the back of the
body, and for some distance down its sides, the colour was dark steel grey,
amounting in some lights almost to black. On a line with the pectoral flipper
the sides were mottled with white, and on the ventral surface irregular, and in
some cases, large patches of a silvery grey or milk whitish tint were seen. An
experienced whaling seaman, Mr Water Roppam, who had charge of the car-
case, told me that he had repeatedly seen this kind of whale in the northern seas,
and stated that, owing to the silvery hue of the belly, it was known to the whalers
by the name of “silver bottom.”* The surfaces of the clitoris and of the labia
minora were mottled with black and silvery grey tints like the skin of the belly.

The dorsal fin was steel-grey or black, except near its posterior border,
where it was a shade lighter and streaked with black lines. The anterior
margin of the lobes of the tail, its upper surface near the root and for the ante-
rior two-thirds, were black, whilst the posterior third of the same surface and
the interlobular notch were lighter in tint. The ventral folds had a light sepia
colour, and the furrows were not so dark as the ridges. The upper surface of
the flipper was steel-grey, mottled with white at the root, at the tip, along its

* In the 2d vol. of Dr Scornssy’s Account of the Arctic Regions, p. 531, it is stated, on the
authority of Captain Day, that amongst the whales pursued by the southern whale-fishers is one called
“sulphur bottom,” a species of Fin whale of great length and swiftness. Can it be that sulphur
bottom is a corruption of silver bottom? and that this whale frequents both the northern and southern.
oceans ? :

STRANDED AT LONGNIDDRY. 203

posterior or internal border, and on the under surface ; white patches were also
seen on the upper surface near the tip, and here they were streaked with black
lines running in the long axis of the flipper. White patches also extended from
the root of the flipper to the adjacent parts of the sides of the animal. The
outside of the lower jaw was black, whilst the inside was streaked with grey
and brown.

A few days after the death of the whale, the scarf skin had become loose,
and large portions of it had separated, leaving the pinkish-white cutis exposed,
and giving therefore a different colour to these parts of the integument than
they had originally possessed. This circumstance is worthy of note, and may
serve to explain appearances which have been described by some authors in
connection with the colour of the skin in specimens of fin whales which they
have examined. The surface of the skin was smooth and shining. No parasites
were found attached to it, and no hairs or bristles were observed to project
from any part of its surface.

Although the animal had reached the enormous length of nearly 80 feet,
yet it had not attained its perfect adult state. For, as the subsequent exami-
nation of the skeleton showed, the disk-like epiphyses of the thoracic and
lumbar vertebree were not yet united to the bodies of those bones. The whale,
therefore, was at the period of growth which, as Professor FLowEr has pointed
out,* may very appropriately be termed “ adolescent.”

Fetus and Membranes.—When the whale was lying on the beach at Long-
niddry, the seaman in charge told me that he believed the animal to be in calf.
On the fourth day after the operation of flensing on the beach at Kirkcaldy had
commenced, as I was watching a man taking away the blubber and muscles from

the posterior part of the side of the abdominal wall, I observed an elongated,
_ dark-coloured mass lying loose amidst the coils of intestine, almost opposite
the umbilical scar. I requested the man to hand it to me, and at once re-
cognised it to be a wreath of young baleen about 4 feet long, which had
obviously become detached from the roof of the mouth of a young animal,
and had by some means or other escaped into the abdominal cavity of the
parent. The discovery of this baleen clearly proved that the whale was in the
gravid state. We at once commenced to remove a larger portion of the abdo-
minal wall in order to obtain a view of the uterus, but before this could be accom-
plished, the rising tide compelled us to cease our operations. As this happened
on a Saturday, work could not be resumed until the Monday following, and as
my University duties prevented me from being present, the search was conducted
_ by Messrs CoucuTrey and Fovutis, who after several hours of hard work ex-
posed the head of the calf by the removal of a mass of blubber from the right

* Proc. Zoological Society, Nov. 8, 1864.

’

204 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

side of the neck of the parent animal. The head of the calf indeed was so far
forward that the tip of its beak was only 2 feet 9 inches behind the condyle of
the mother’s right mandible. An additional mass of blubber was then taken away
from the exterior of the ribs on the right side, when more of the calf was ex-
posed. It was lying obliquely between the blubber and the muscles which
covered the outer surfaces of these ribs, and the space in which it was contained
had obviously been formed by a forcible separation of the blubber from the sub-
jacent muscles ; for when the blubber was cut through, the pressure on the
calf, owing to its position between a weighty mass of blubber and the elastic
ribs, was so great that its head protruded through the incision, and even
partially tore through the superficial textures.

The lower jaw of the calf was directed towards the ventral surface of the
mother, and the left side of its body was in relation to the outer surface of her
right ribs, and its tail was directed to her abdominal cavity. After the removal
of an additional portion of blubber, the calf was extracted by my assistants, and
in the process of removal it was observed that about 5 feet from the tail the body
of the calf was so twisted on itself, that the position of the two lobes of the tail was
reversed. A large quantity of the foetal membranes lay alongside of the calf,
more especially near its caudal end; but they were torn, and had lost their bag-
like form. Some coils of the intestine were also situated beside the tail. It
is much to be regretted that the uterus could not be preserved in the course of
this examination. The huge size of the coils of the intestine, and the desire
which the men employed had to get rid, on account of the smell, of the con-
tents of the abdominal cavity, rendered it impossible to make such an exami-
nation of these viscera as was desired.

From a consideration of the position of the calf there can be no doubt that
either immediately before or after the death of the mother, the foetus had been
disconnected from its proper attachments and extruded into an artificial space
external to the abdominal cavity. The torn state of the foetal membranes and
umbilical cord, the presence of coils of the intestine in the space in which the
foetus was lying, and the loose mass of baleen in the abdominal cavity of the
mother, all point to a rupture not only of the uterus, but of the wall of her
abdomen, which had permitted the passage out of the cavity both of the foetus
and of portions of the gut.

To what cause, then, are we to ascribe the rupture and consequent displace-
ment? Some of those who examined the whale were of opinion that they had
been occasioned by a severe injury sustained by the mother prior to, or at the :
time she came ashore. But I am rather inclined to think they must have
occurred whilst she was being towed by the tail across the Firth from Long-
niddry to Kirkcaldy. For, during the two weeks she lay on the beach at the
former place, decomposition had advanced to a considerable extent, putrid

STRANDED AT LONGNIDDRY. 205

gases were disengaged, and consequent softening of the soft parts had occurred.
As the sternum is short, and only articulates with the first pair of ribs, and
as the inner ends of the remaining ribs diverge considerably from each other,
and have no strong attachments in the ventral mesial line, the great pressure of
the sea on the wall of the abdomen, as she was towed by the tail, would tend
to rupture the uterus and abdominal wall, to drive the contents of the abdomen
forwards towards the head, and to force the foetus into the position in which it
was found.

Owing to the displacement of the foetus, the dissection of this animal does
not enable me to state with certainty the normal position of the foetus in
utero in this cetacean. Very little indeed is known of the uterine position of the
foetus in this group of mammals. In a communication made to the Royal
Belgian Academy,* M. vAN BENEDEN figures the position in utero of the foetus
in Globiceps. Its head is directed to the maternal genital orifice, its body is bent,
and the tail is folded backward under the thorax, so as to lie close to its flipper.
He believes that the foetus of Balenoptera rostrata has the same position in
utero, and doubts the statement made by M. Borcx, that the young of rostrata
is born first by the tail. In the Longniddry Balenoptera, on the other hand,
the head of the foetus was directed towards the head of the mother ; and unless
we suppose that during the displacement a complete revolution in the relative
position of its caudal and cephalic ends had taken place—an occurence which,
owing to the great length of the foetus, scarcely seems possible—the uterine
direction of the young one would have been with its tail towards the maternal
genital passage.

The gravid state of the whale necessarily exercised an influence on its shape,
more especially by increasing its girth in the abdominal region—a circumstance
which should be kept in mind in comparing the drawing of this animal (fig. 1)
with those which have been given by other naturalists of the Finners which have
come under their observation.

The form of the foetus differed in several particulars from that of the mother.
Its greatest girth was around the head, from which it tapered forwards along
the beak, and backwards to the root of the tail. From the unexpanded con-
dition of the lungs, and the flaccid state of the hollow viscera of the abdomen,
the thoracic and abdominal cavities had not attained their proper girth, and the
body and caudal end of the foetus presented a peculiar, elongated, worm-like
appearance. The dorsal fin did not rise so abruptly in the foetus as in the adult,
so that it was difficult to determine its exact antero-posterior length. Its post-
erior border had a well-marked falcate curve (Plate V. fig. 2).

The foetus was a male. The penis, 11 inches long, hung pendulous.from the
ventral surface, and at each side of its root a crescentic fold of skin arched out-

* Bulletins, vol. xx. 2d series, No. 12.
VOL. XXVI. PART I. 3 H

206 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

wards (Plate VI. fig. 9). Behind each of these folds was the mouth of the
shallow nipple fossa ; the nipple was rudimentary, and concealed by the promi-
nent anterior border of the fossa. The posterior border was feeble, and here the
fossa blended with the general surface of the abdominal wall. Passing back-
wards, midway between these fosse, was a well-defined raphé reaching to the
anus. The abdominal wall was much torn in front and at the root of the
penis, and the exact attachment of the umbilical cord could scarcely be recog-
nised, but it was estimated to be connected about 18 inches in front of the
root of the penis; for the cord, though carefully divided at the time when the
foetus was removed from the mother, had been used, along with the penis, as
a convenient object to lift with by the men employed to carry the calf, and con-
sequently both they, and the part of the wall to which they were connected, had
sustained injury. The tail was subdivided into two elegantly curved horizontal
lobes (fig. 3). The sides and ventral surface showed the characteristic plicated
appearance (fig. 4). On the top of the head, 1 foot 5 inches behind the blow-
hole, was an oval patch 1 inch long by ?ths broad. It was raised somewhat above
the level of the integument. The shape of the flipper is represented in fig 5.

The colour of the integument was a warm grey, mottled here and there with
yellow. Patches of dark steel-grey pigment were observed on the back ; but
none of the light silver-grey tints, seen in the large whale, were observed on the
belly. I believe that desquamation of the cuticle had taken place very exten-
sively before the calf came into my possession.

I had anticipated that the comparatively small size of the foetus would,

by giving me greater command over the dissection, have enabled me to have ~

worked out all those points in the anatomy of this whale, which I could not
overtake in the older animal. But in many respects I was disappointed, for
the weight of the foetus, which amounted to about half a ton, and its length of
almost twenty feet, rendered it a most unwieldy object to transport to the
Anatomical Museum. Moreover, putrefaction had to some extent advanced
before I had the opportunity to examine it ; the abdominal wall was torn, and
the viscera in that cavity were so much injured, that but little definite informa-
tion respecting the stomach and intestines could be obtained. The muscles also
had undergone a remarkable kind of decomposition ; the odour exhaled from
them was peculiarly acrid and offensive, which, together with their softened
condition, rendered it impossible to make a proper study of those important
parts of the locomotory system. The bones of the skull and spine were also to
some extent displaced.

A number of measurements were taken, a table of which I subjoin ; but in
consequence of the displacement just referred to, some of the dimensions are

a ae

probably not absolutely exact, but are to be regarded as the closest approxima- —

tion which could be obtained :—

STRANDED AT LONGNIDDRY. 207

: Feet. Inches.
Length of male foetus, . : : 1) 6
From tip of lower jaw to panier end of blow Bee :
From posterior end of blow-holes to posterior border of dorsal fin,
From posterior border of dorsal fin to interlobular median notch of tail,
Antero-posterior diameter of blow-holes, .
Transverse diameter of blow-holes,
From tip of lower jaw to angle of mouth in a miele line,
From tip of lower jaw along curve to angle of mouth,
From angle of mouth to anterior border of root of flipper, P
From tip of lower jaw in a straight line to anterior border of root of flipper,
Length of flipper along anterior border,
Greatest diameter of flipper from anterior to posterior Wbddie
Girth of flipper at root,
Girth of body just behind dorsal fin,
Girth round root of tail,
Between extreme points of tail-lobes in a auaent shin
Between extreme points of tail-lobes along posterior concave perder
Greatest girth of tail-lobe, ;
From median notch of tail to anal orifice,
Transverse distance between nipple fosse,
From anal orifice to midway between nipple fosse,
From nipple fossa to fold of skin at root of penis,
Length of penis,
Vertical diameter of doheal fin,
Greatest transverse diameter of cavity of mouth,

— —
WNHNONADOTONOOGQeH WHReH DW CO

—

MP CcocOOCO CO OnwnnrkRYE wrFowmnrre PR wWoOOasw
—_
bos

A vertical line, drawn from the root of the posterior border of the dorsal fin
to the ventral mesial line, was 161 inches behind the anal orifice.

The displacement of the foetus and the torn state of the membranes did not
give me the opportunity of observing the exact relations of the latter to the foetus
and to the mucous surface of the uterus. Although several square yards
passed through my hands, yet I did not succeed in recovering the whole extent
of these important structures. Notwithstanding these deficiencies many points
of interest bearing on the placentation of the cetacea were observed.

The outer surface of the chorion had the general villous appearance which
is characteristic of the diffused form of placenta. In my first, and somewhat
hurried inspection of this membrane, I did not notice any portion which did
not possess villi. But on a second examination, made at more leisure on the
membrane preserved in spirit, I observed that a portion of the chorion was
bare. Unfortunately this had been torn across and a portion lost, so that the
proper form of the non-villous part could not be ascertained. It had appa:
rently, however, been of some extent, for the portion preserved was oblong in
form, and measured 11 inches by 3. In all probability it had formed a part of
one of the prolonged poles of the membrane.

The villi began at the edge of this bare part by a well-defined line; immediately
beyond and parallel to which the chorion was doubled on itself, so as to form a

208 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

strong marginal fold, which projected for about one inch, and was thickly studded
with villi on its surface and free edge. (Plate VII. fig. 17.) A second fold,
also covered with villi, lay close and parallel to the marginal fold. Similar
villous covered folds of the chorion, many of which were one foot and upwards .
in length, traversed the chorion in various parts of its extent ; frequently they
ran parallel to each other, and two or more were sometimes close together, but
at other times they were separated by intervals of 3, 4, or 5 inches. Usually
the greatest projection of one of these folds was about 2 inches, though some-
times it reached 3, or even 4, but towards their extremities they gradually sub-
sided to the general plane of the chorion.

Besides these elongated folds, villous covered folds of another form, but not
so numerous, projected from the surface of the chorion. They were triangular
in shape, flattened on their surfaces, and with the apex and lateral borders
free. A very characteristic specimen is répresented in Plate VII. fig. 18. Its
margin of attachment was 4 inches, whilst its diameter from this margin to the —
free apex was 54 inches. On the elongated and triangular folds, but more
especially the former, the villi were thickly studded, but on the intermediate
surface of the chorion they were more sparingly distributed, and were for the —
most part collected on minute and ridge-like elevations, which intersected each
other, and presented an irregularly reticulated appearance. The membrane
between these slight ridges was comparatively smooth and transparent. ;

The mucous surface of the uterus in the mother must have possessed
numerous depressions of considerable length and depth, into which the elongated ~
and triangular folds of the chorion would have fitted. In the special aggregation
of the villi on these folds an approach to the cotyledonary type of the placenta
found in the Ruminantia may be traced.

The opposite surface of the chorion was in relation to the placental blood- —
vessels, some of which were of considerable size ; one, which was measured, had ©
a circumference of 23 inches. Where the folds on the villous surface were well —
marked an artery coursed along and gave off many collateral branches, which —
entered into the fold to end in the villi. The chorionic vessels were surrounded
by a delicate connective tissue, which was loosely connected with the attached
surface of the amnion. Lying in this connective tissue were numerous opaque, —
white, slender threads, which differed from the small arteries in not being tor-
tuous, and in giving off their branches at very acute angles. These threads
had to the naked eye the appearance of fine nerves. When examined with the
microscope, they were found to possess an external investment of well-marked —
connective tissue, which surrounded lines of an irregular granular or semi- —
globular substance which looked like the disintegrated medullary sheaths of ~
nerve fibres. The free surface of the amnion was smooth and glistening.

Although nothing definite seems to be known of the period of gestation of

STRANDED AT LONGNIDDRY. 209

the Finners, yet from the length of the calf, and the well-developed state of its
parts, it is probable that the whale was at or about her full time. Dr Scoressy
considered that February and March were the months in which the Balena
mysticetus gave birth to her young,* but Escuricur and Rernwarpr, from obser-
vation made at the Danish whaling factories, think that it is between the end of
March and the beginning of May.t If my supposition be correct that the whale
was at her full time, then this Balenoptera gives birth to its young in the later
autumn months, and not, like the Greenland Right whale, in the spring of the year.

This view of the period of parturition of the great Finner is strengthened
by evidence which I have received from another source. In the month of
October 1869, a large female Finner, which, from information that I have ob-
tained,t I believe to be of the same species as the Longniddry whale, was
found in a creek about a quarter of a mile to the south of Hamna Voe, North-
maven, Shetland. It was dead, and floating by its side was a dead calf, which
was well developed, and bore to the mother about the same proportion as the
Longniddry animals did to each other. Alongside the calf was a quantity of
membranes, which, from the statements of the fishermen, were evidently the
foetal membranes. The calf had obviously been born about the time of the
death of the mother, and had apparently reached the full period. The maternal
mammary glands were so charged with milk that a quantity was observed to
flow out through the teats.

The capture of two of these whales in the pregnant condition within so
short a period in arms of the sea, lends support to the statement which has
more than once been made, that the Finners resort to bays and creeks for the
purpose of bringing forth their young.

Skin and Blubber.—The colour of the skin has already been described ; a
few words may, however, be said on its structure. The epidermis readily
peeled off the cutis when decomposition had begun. It was distinctly laminated
and thicker than the human cuticle. On the belly, for example, it measured
1th of an inch, and on no part indeed of the surface of the trunk was it
seen to possess a greater thickness. In this respect it contrasts strongly with
the skin of the Balena mysticetus, which in some places has the cuticle one
inch thick.§ The superficial layer could be peeled off as a thin horny stratum,

* Account of the Arctic Regions. I. 470.

+ Memoir translated for the Ray Society, p. 10.

t I am indebted for information regarding this whale in part to Mr J. Waukmr of Maryfield
House, Bressay, and in part to Mr Coucutrey. The latter gentleman has just returned from a visit to
the Shetland Isles, and when there not only collected at my request various interesting facts about this
animal, but also procured for me a number of its bones.

§ Dr Knox (Catalogue of Anatomical Preparations of the Whale, Edinburgh, 1838) points out
the thinness of the cuticle in the species of the great northern Rorqual which he dissected ; nowhere,
he says, did it exceed 3%;ths of an inch. He compares it with the B. mysticetus, and shows how
in the one there are conjoined thin cuticle and short baleen, in the other thick cuticle and long baleen.

VOL. XXVI. PART I. oa

210 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

which, when dried, had the appearance of gold-beater’s skin. The deeper
layers contained more pigment than the superficial, and in those parts of the
skin where the colour was most marked the deep surface of the cuticle had a
rich black hue. When the epidermis was removed, rows of distinct elongated
papille were seen; and in vertical sections through the entire skin the rela-
tions of these papille to the cuticle* could be studied (Plate VIII. fig. 29).
The papillz were filiform, and as a rule simple, but in some cases two or
even three papille arose by a common stem, which then subdivided. They
were comparatively long, and their apices reached therefore much nearer
to the surface of the skin than might have been supposed. In some of the
sections I observed distinctly the small arteries of the cutis giving off branches
which entered the bases of the papillae and extended for some distance within
them.

The blubber or subcutaneous tissue was composed of adipose tissue, for the
oil was contained in well-defined fat cells. These cells were supported by
bands of connective tissue, many of which possessed considerable breadth and
strength. Blood-vessels passed in some numbers through the blubber, partly
for its nutrition, and partly for the nutrition of the integuments on its
surface. The blubber varied considerably in thickness in different parts of
the subcutaneous tissue of the adolescent animal. On the sides and upper
edge of the lower jaw, it was from 10 to 16 inches. Beneath the ear-slit 8
inches ; along the ventral surface about 4 inches. On the top of the beak
and cranium 8, 12, and even 15 inches. In front of the dorsal fin from 12 to
16 inches, and behind this projection from 14 to 21 inches, which seemed to be
the maximum thickness. The thickness of the blubber at the tip of the caudal
spine was 3 inches, and at the symphysis of the lower jaw 44 inches, so that
the length of the skeleton was within 74 inches that of the entire animal. In
the foetus the blubber was very imperfectly formed ; and the thickness of the
subcutaneous tissue was almost uniform, on the belly not exceeding one inch;
and on the back scarcely reaching two inches.

In the older animal, an enormous mass of soft fat was situated within, and
formed a sort of fatty lining for the abdominal cavity. From the heat which
was disengaged by the putrefaction of the carcase, this fat was liquefied, and
ran in streams on to the shingle, where it again solidified, and was collected
into barrels.

Mr Tair estimated that he had obtained from the blubber ten tons of oil,
and from the inside fat six tons, so that the pecuniary value of the whale from

* In the Anatomical Museum of the University of Edinburgh are several specimens (161 to 164)
prepared upwards of twenty years ago by the late Professor Goopsir, one from the B. mysticetus, three
from a “ Rorqual,” probably the Balenoptera musculus, which give most illustrative views of the fili-

form papille of those animals. <

STRANDED AT LONGNIDDRY. 211

these sources alone was very considerable. He has also furnished me with an
estimate of the weight of the other portions of the carcase; from which we
may make an approximation to the weight of the entire animal. The flesh
and viscera 36 tons, the baleen and “ gum” 10 cwt., the skeleton 9 tons 10
ewt., the blood and refuse 12 tons, which, with the oil and fat, make in all
74 tons as an estimate of the weight of the entire animal.

Mammary Gland.—The position of the nipple has already been described in
the section on the external form of the animal. The gland itself was exposed
by the removal of the blubber on one side of, and for several feet anterior to,
the genital fissure. It formed an elongated body, and measured between 7 and 8
feet in its antero-posterior diameter, and of this extensive mass only 8 inches lay
behind the nipple. Its greatest transverse diameter was 20 inches, and the thick-
ness of the gland substance, which surrounded any part of the great central
duct, was more than 6 inches. Its broadest part was in the region of the nipple,
gradually tapering off to its anterior end. Its colour was a rich red; and its
subdivision into lobules by bands of connective tissue could be readily recog-
nised by the naked eye. When cut into, it was seen to be traversed along its
entire length by a central duct, which increased in size as it passed from before
backwards, and at the base of the nipple formed an enormous sinus, the trans-
verse diameter of which was about 8 inches. Numerous large ducts, into many
of which the closed hand could be passed for some distance, opened out of this
central duct, and extended into the various parts of the gland. The transverse
diameter of one of these ducts was 54 inches. The orifices of the primary ducts
opening into the great central canal, and those of the smaller ducts which opened
into the primary, were mostly oblique in their direction, and a well-marked
fold of the mucous membrane bounded one-third, and sometimes more, of the
aperture. As a general rule, the direction of these ducts was towards the
nipple, but some ran in the opposite direction. The mucous membrane which
lined the ducts and central canal was firm, and marked on its free surface by a
characteristic ridge and furrow-like appearance (Plate VI. fig. 11). These ridges
were parallel to the long axis of the duct. At the base of the nipple the great
sinus-like dilatation of the central canal suddenly narrowed to the duct within

- the nipple, which was not larger than would admit the middle finger or thumb.
_ The lobules of the gland were polygonal in shape and variable in size; some of

the larger ones had a diameter of {th inch. Sections through the lobules
examined microscopically gave very illustrative views of the structure of a com-
pound racemose gland. The clusters of acini or gland vesicles, with their con-
tained secreting cells, could be seen with great distinctness, and the arrange-
ment of the interlobular connective tissue could be traced.

In the subcutaneous tissue around the nipple and at its base, numerous

plexiform vessels were seen, so that it is probable that erectile tissue exists
: Me

212 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

in this locality. Lying outside the mammary gland was a muscle which, by its
contraction, would aid in expelling the milk along the ducts, and through the
orifice of the nipple.

The size of the secondary ducts of this gland, and the dilatation of the single
central duct into a great reservoir for the collection of the milk, have obviously
special reference to the aquatic mode of life of an animal which suckles its
young. For, as Joan Hunter long ago pointed out,* the mode in which these
animals give suck is very inconvenient for respiration, as if the mother were to
turn round so as to elevate the nipple to the surface then her nares would be
under water ; whilst, if the mother remains in her normal position, then the
nose of the calf must be under water, and the time of sucking can only be
between each respiration. It is necessary, therefore, that the gland should be
so constructed as to allow of a considerable accumulation of milk in the ducts,
which may be readily drawn off by the calf in the intervals between the respi-.
ratory acts.

Baleen.—When the lower jaw was removed by cutting through the massive
fibrous columns, which connected the condyles of this bone to the base of the
skull, and when the occipito-atloid jomt was disarticulated, the skull was turned
over on its dorsum, and a complete view of the roof of the mouth, and of the
baleen in situ was obtained. Extending from behind forward in the mesial
plane of the palate was the great central crest or keel, which was much broader
and more prominent posteriorly than anteriorly, and was covered on its
free surface by a black mucous membrane. Immediately on each side of the
base of the keel the palate was covered by a smooth and almost flat, black —
mucous membrane, and external to this again was the lateral series, or wreath,
of deep black baleen plates with their inferior free edges fringed with black —
sete.

The wreaths of baleen plates on the two sides converged as they passed for-
wards, and at the anterior part of the mouth they became continuous with each
other, as is the rule indeed in the Finner whales.t Posteriorly, where they lay
close to the entrance into the gullet, they were separated by a considerable
interval ; though here also they inclined inwards to the base of the great mesial
palatal keel. The inner edge of each wreath had a curved outline with the
concavity towards the mesial keel. The outer edge was convex, and in its
curvature closely corresponded to that of the margin of the beak itself. This
border was bounded by a raised fold, the coronary or wreath-band (Horn-_
Kranzband of RosENTHAL), and was situated one foot within the outer edge of
the beak. Where the two wreaths became continuous in front, the junction
took place seven inches within the tip of the beak.

* Structure and Economy of Whales. Phil. Trans. 1787.
+ Escnricat and Retnnarpt. I,have also seen this in two specimens of Balenoptera rostrata.

STRANDED AT LONGNIDDRY. 213

Each wreath was estimated to contain about 370 rows of plates,* and each
row consisted of several plates or blades or bristles. The rows lay transversely
and parallel, though not in straight lines, for they were somewhat curved, the
convexity forwards, the concavity backwards, and the smaller inner subsidiary
plates were arranged in an oblique manner. Intervals varying from one
half to three-eighths of an inch existed between the rows in different parts
of the series. The transverse and vertical diameters of the plates varied
considerably, not only in different. parts of the wreath, but also in each row,
for the plates diminished in size from the outer to the inner edge of the row.
At the anterior part of the mouth they were little more than coarse black
bristles, and the free part of these projected in some only half an inch, in
others one inch and a half, into the cavity of the mouth. Extending backwards
along the outer or labial part of the wreath the baleen increased in size, at first
being somewhat elongated narrow plates, and then increasing in their trans-
verse diameter at their base of attachment, until they assumed the unequally
four-sided form, with its surfaces directed forwards and backwards, of the blade
represented in Plate VI. fig. 12, which may be regarded as a very character-
istic specimen of one of the large plates of this Balwnoptera. The dimen-
sions of this plate were as follows. The transverse diameter along its base of
attachment 1 foot 6 inches ; vertical diameter, inclusive of the part imbedded in
the intermediate substance, along outer free border, 2 feet 94 inches, along
iner free border 8 inches. Length along the border fringed with sete 3 feet
3 inches. The setz varied in their length, some measuring as much as 17
inches. On the surface of the plate numerous longitudinal parallel lines, which
at its inferior edge became continuous with the sete, were observed. Transverse
rings, which sometimes were close together, at others were separated by wider
intervals, passed from one surface to the other around the outer and inner free
edges of the plate. A plate of this form and of somewhat similar dimensions
formed the external or labial blade of each transverse row in by far the greater
portion of the wreath.

Internal to this large plate the baleen, though of the same black
colour, was elongated and narrow ; the blades possessed the form represented
in Plate VI. fig. 13, their transverse diameter was not more than ;8,ths of an
inch, and their vertical diameter, inclusive of the part imbedded in the inter-
mediate substance, was in some 7, in others 6, in others 5 and 4 inches. Each
of these narrow subsidiary plates had an uniform breadth, and the sets, which
_ Were often more than 6 inches long, arose not from the sides, but only from

* Although the rows of plates were counted without difficulty in the greater part of the wreath,
yet at the posterior end, and at the front, of the mouth the exact enumeration was attended with con-
siderable difficulty, owing to the bristle-like baleen being arranged in less definite rows than were the
blades of this substance.

VOL. XXVI. PART I. o Kk

214 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

the free end. Whilst the sete: generally had the same deep black colour as the
plates, in some cases they had more of a deep soot brown tint. The baleen at
the inner end of each transverse row consisted, not of plates, but of short bristles,
similar to those already referred to at the anterior end of the series. As the
vertical diameter of the plates and the length of the setze were so much greater
in the outer than in the inner parts of each transverse row, it followed that the
lower bristle-friged aspect of each wreath arched, from without, obliquely
upwards and inwards, so that the roof of the mouth presented a considerable
concavity from side to side.

The plates were all imbedded at their attached palatal borders in a dense
semi-elastic, slate-coloured material, the intermediate substance or “gum” of
the whaling seamen. This substance varied in its thickness from its attached
to its free surface to from 1 to 4 inches in different parts of the wreath, and was
thinner along the outer and inner borders than in the intermediate portions. It
was continuous, along the inner border of the wreath, with the cuticle investing
the palatal mucous membrane, and along the outer border, with the coronary
or wreath-band already referred to. The free surface possessed an irregular
softened, water-worn appearance.

After decomposition had begun the baleen and intermediate substance,
intimately connected together, could be readily peeled off the surface of mucous
membrane from which they grew, and their mode of growth and structure could
be examined.

All anatomists know, who have studied the structure of whalebone, that,
when a blade is carefully detached from the surface of the palate, the edge or
base of attachment is cleft along the line of its transverse diameter into two
lamine. If these lamine be drawn asunder numerous holes are seen at the
bottom of the cleft, which open into tubes or canals that traverse the substance
of the plate in the vertical direction. It has been pointed out by Escuricut
and REINHARDT, that in the short baleen plates of the Rorquals or fin whales the
length of these tubes is comparatively greater than in the much longer plates
of the Greenland Right whale. In the Longniddry whale, the deep black colour
of the baleen made the plates so opaque, that the existence of the tubes could
only be surmised by the longitudinal markings visible on a surface examination,
and it was not until after sections were made in the vertical or transverse direc-
tion, that the tubes could be distinctly seen.

In vertical sections the tubes were cut longitudinally, and could be followed
for some distance (Plate VII. fig. 19). They contained a delicate, brownish-
yellow substance, which could be easily drawn out of the tube. In the part of
the plate which surrounded the tubes numerous black pigment granules were
distributed in such a manner as to give to the section the appearance of
longitudinal striation.

STRANDED AT LONGNIDDRY 215

Transverse sections of the plates, examined with low magnifying powers,
were, however, the more instructive (fig. 20). The number and size of the
tubes was by no. means uniform in the different parts of the same trans-
verse plane. Sometimes a single comparatively large tube was alone met with ;
at others two, or even a larger number, occupied the antero-posterior diameter,
and in this case the tubes were considerably smaller. The soft brownish-
yellow contents were readily recognised, and in many of the sections this sub-
stance was seen to be perforated with holes, which looked like transversely-
divided small blood-vessels.

The solid portion of the plate was spotted with black pigment, and dis-
tinctly striated. The strie ran in two different directions, and indicated a
laminated arrangement. One set of strize or lamellz surrounded, in a concentric
manner, the individual tubes, and in their arrangement might be compared
with the lamelle surrounding the Haversian canals in a transverse section of
bone. They may be called the tubular lamelle; and the tube, its contents,
and the lamellz surrounding it, might be termed a tubular system. The other
lamellee were situated on the peripheral part of the plate, and formed a sort of
envelope enclosing the tubular system of lamelle. These may be called the
peripheral or cortical lamelle ; and they formed that part of the plate which has
been called the cortical layer or “ enamel” of the whalebone. When examined
with higher powers of the microscope, the lamellz were seen to be composed of
elongated and flattened cells, each containing a distinct nucleus, and more or
less black pigment (fig. 21). These cells were obviously peculiarly modified
epithelial cells. The intervals between the outermost lamelle of adjacent
tubular systems were filled up by cells, which presented less of a flattened and
more of a fusiform or rod shape ; these cells, though interstitial in their position,
were apparently continuous with the cells of the cortical layer.

Transverse sections through the setz displayed in each a central tube or canal,
surrounded by the usual arrangement of concentric tubular lamelle (fig. 22).
The tube within the seta contained a similar soft brownish material to that
found in the tubes within the blade itself. Each seta represented, therefore,
a single tubular system,

When vertical sections through the intermediate substance, in which the
bases of the plates were imbedded, were examined with low powers of the
microscope, the deep surface attached to the palate was seen to be much more
highly charged with pigment than the more superficial parts, and so regularly
Was it disposed, that it might almost be described as a special pigmentary
layer of the structure. The deep surface had an uniform rich black colour, and
was perforated by numerous apertures, which in the vertical sections were seen
to lead into clefts which passed some distance into the intermediate substance
(fig. 23). The black pigmentary layer was prolonged along the walls of these

216 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

clefts. Under higher powers of the microscope, the intermediate substance
was seen to consist of flattened cells, epithelial in character (fig. 24), and the
black pigmentary layer was due to a special accumulation of pigmentary
granules in the deepest cells of this substance. This layer may be considered
therefore as comparable to the Rete Malpighi of the human cuticle.

The intermediate substance was intimately united to the lamine formed by
the cleavage of each plate at its base ; so close indeed was this union that it was
impossible to separate them from each other without injury to the latter. It
not unfrequently happened, in tearing away the substance from between the
plates, that a portion of the cortical layer of the adjacent part of the plate
peeled off along with it. A distinct horizontal lamination was seen on the
surface of vertical sections made through the intermediate substance.

In my further researches into the structure of the baleen, I have derived
considerable assistance from the examination which I made of the baleen of a
recently killed, lesser Pike whale, B. rostrata, about 18 feet long, which was
captured at Burntisland in September last. In this animal the plates were for
the most part white, or yellowish-white, but, when quite fresh, a distinct pink
or rosy colour was seen, more especially in that part of the blade which lay
within and next to the intermediate substance. Some days after death the
pink or rosy colour became converted into purple.

When a fresh blade was examined in a good light, the pink colour was —
found to be not on the surface, but within the substance of the plate, and
arranged in regular lines, which ran parallel to each other from the attached
border to the free border fringed with setee, and in many cases it extended even ©
into and along the latter. When a pocket lens was used in the examination,
the colour was seen to be due to a red fluid contained in the numerous tubes
which traversed the plate in its vertical diameter. Sometimes the fluid formed
an unbroken column of one, two, or three inches in length; but at others the
column was much subdivided, and reminded one of the appearance presented
by a broken-up column of mercury in a barometer tube when out of repair. In-
some of the tubes, more especially those situated near the outer and inner
edges of the plate, the red fluid was either absent, or extended only a short F
distance down the tube. Many of these tubes appeared as if subdivided +
little septa passing across their canals, not unlike the arrangement one has-
seen in the medullary part of a hair. When the baleen plate was cut across —
transversely, and forcibly squeezed between the finger and thumb, the red r
fluid oozed out of the divided tubes, and when collected on a glass slide was _
examined microscopically. Under a high power numerous circular, disk-shaped,
non-nucleated corpuscles, which possessed the optical characters of blood cor- -
puscles, were found in it (fig. 25), and along with these were three-sided pris-
matic crystals, probably the triple phosphate, and numerous actively moving

STRANDED AT LONGNIDDRY. 217

vibriones. It was clear, therefore, that the pink tint of the baleen in the Pike
whale was due to the blood* situated in the tubes which traversed its substance
in the vertical direction.

I am not aware that any explanation has previously been given of the cause
of the pink colour of the baleen in the lesser Pike whale. Indeed many
writers seem to have paid but scanty attention in their descriptions to the
existence of this tint.

Both in the Longniddry and the Pike whales the surface of the palate, from
which the baleen grew, possessed numerous transversely elongated folds of the
palatal mucous membrane (the pulp-blades of Escuricur and Reinnarp7),
corresponding in their arrangement and transverse diameter to the different
sizes of the baleen plates in the various transverse rows, and fitting into their
cleft basal edges (fig. 26). The largest of these folds in the former animal pro-
jected as much as {ths of an inch from the general palatal surface. The free
lower edge of each fold was fringed with multitudes of well-marked elongated
filiform papille, which fitted imto and indeed filled up the tubes in the
plates and setze already described. These may be called the tubular papille.
If great care was taken in stripping off the plates, the papillze could be drawn
out of the tubes, and in fig. 26 a view of a number of these structures from
the interior of the tubes of a plate of the Longniddry whale is given. The
tubular papille varied in length in this preparation, some being 3 inches long,
whilst others were considerably shorter ; but none of these papille represented
the full length of the tubes they originally occupied, as they always broke short
in the act of removal. They varied also in thickness, in correspondence with
differences in the bore of the tubes; and they were thicker at their attached
than free extremities.

Folds and papille of this character have been described with more or less ful-
ness of detail by Hunter, Ravin,t Rosenruat,t Knox,§ Owen,|| Escuricut and
REINHARDT, FLOweER,** and Matm,tt in connection with the baleen in the
different whales which they have examined ; and in the Anatomical Museum of
the University of Edinburgh are several specimens, prepared, I believe, in the
year 1843, by the late Professor Goopsir, which furnish very illustrative views
of the folds and larger papille of the baleen plates. They have been regarded
as the nidus, matrices or pulps, from, and in connection with, which the specially

* As confirmatory evidence of this fluid being blood, I may state that I requested my friend, Dr
Artuur Gamces, to apply the chemical test for blood. He found that the fluid gave with guaiacum
and peroxyde of hydrogen the characteristic greenish-blue colour of hemoglobin.

+ Ann. des Sc, Naturelles, 2 Sér. t. v.

~ Abhand. der Akad. der Wissensch, zu Berlin, 1829, p. 127. § Catalogue, op. cit.

|| Odontography, p. 312. | Ray Society’s Translation, op. cit.

** Proc. Zool. Soc., 1865, Nov. 28.
++ Monographie Illustrée du Baleinoptére, Stockholm, 1867.

VOL. XXVI. PART I. 3h 0

218 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

modified, horny, epithelial cells of the baleen plates were developed. The
vascularity of the folds and of the papillz has also been recognised by these
anatomists, but no exact description of the arrangement of the vessels has
as yet been given.

The fresh condition of the baleen in the B. rostrata led me to think that it
might be possible to inject its papille, and to obtain a more complete view of
the arrangement of their vessels than had yet been described. I accordingly
carefully detached the entire palatal mucous membrane, with its baleen wreaths,
from the upper jaw; and after introducing injecting pipes into several of the
palatine arteries, I succeeded with the aid of my assistant, Mr StrrLine, whose
skill as a minute injector is so well known, in injecting the vessels of the baleen.
But before proceeding to describe their arrangement, it will be necessary to speak
of two other groups of papille, which appear hitherto to have been overlooked by
anatomists. When the surface of the palatal mucous membrane, situated be-
tween the bases of the transverse folds, was examined with a pocket lens, it was
found to be studded with short papille, which fitted mto clefts similar to those
already described (fig. 23), as extending into the intermediate substance from
its deep attached surface. These papille we will call intermediate. Similarly,
when the sides of the transverse folds were also examined with a pocket lens, they
were seen to give origin to numerous minute papille, which passed into minute
apertures in the inner wall of each of the laminz, produced by the cleavage of

the baleen plate at its base. These laminz were continuous with the cortical —

layer of the plate to which they belonged, and their papillz may be called peri-
pheral or cortical.

In the injected preparations, the following appearances were seen in vertical
sections (fig. 27). The palatal mucous membrane was highly vascular, and the
principal vessels ran parallel to the horizontal plane? They gave origin to
smaller vessels, which were distributed to the three groups of papille. Those
which passed to the intermediate papillz, occupying the spaces in the attached
surface of the intermediate substance, did not enter the transverse folds or pulp
blades; they were very slender, but formed distinct loops (fig. 27). The vessels for
the other papillz entered the transverse folds. Those destined for the peripheral

or cortical papille formed a well-defined superficial network of small vessels,

which gave off, at intervals, capillaries which entered these papille, and formed
loops in the usual manner. The vessels for the elongated, filiform, tubular
papillee were considerably larger. Asa rule, two entered the base of each papilla,
and extended along its axis into the tube. These vessels preserved their size
for a very considerable distance down the tube, and occasionally anastomosed.
They were easily recognised by the naked eye, both in vertical and transverse

sections of the plates and sete ; and it was in them that the blood was contained —

which conferred on the baleen of B. rostrata its characteristic pink markings.

STRANDED AT LONGNIDDRY. 219

When the papillz -were carefully extracted from the tubes, and examined
with high powers of the microscope, they were seen to consist of a delicate,
wavy, connective tissue, the filaments of which lay parallel to the long axis of
the papilla. The nucleated corpuscles of the connective tissue were distinctly
recognised after the papilla had soaked some time in glycerine. On the free
surface of the papillz a very distinct layer of flattened polygonal cells, with
their borders in close contact with each other, like epithelial cells on a free
surface, was met with. These cells were soft and delicate, and were evidently
the youngest layer of epithelial cells lying next the papille, which had not
yet undergone the horny transformation. In some of the papille I saw, more
especially at their broader attached ends, elongated fibres, having a double
contour, which I believe to have been medullated nerve fibres.

The baleen of the foetus of the Longniddry whale possessed some features
of interest, to which I may now refer. Only the wreath, which was met with
early in the dissection of the mother, was preserved, for the opposite wreath,
which had also been shed from the palatal surface, was lost in the course of the
dissection. The wreath was 4 feet long, and 33 inches in its greatest transverse
diameter. The anterior end had been broken away, and lost, but the posterior
end was flattened, and terminated in an obtuse angle. Notwithstanding the
loss of its most anterior portion, as many as 335 transverse rows were counted
in the wreath, and they were slightly curved with the convexity forwards.
Owing to the comparative thinness of the intermediate substance, the interval
between any two adjacent transverse rows was not more than th of an inch.
Here, as in the adult, the outer or labial plate in each transverse row was by far
the largest ; indeed, those internal to it were little more than short bristles in the
fetus. In the greater part of the wreath seven, eight, or sometimes nine plates
or bristles were counted in each transverse row. Towards the anterior end
only five were counted; but posteriorly, where the external plate, like those
internal to it, consisted of a mere bristle-—the number of bristles in the row
‘had increased to about thirty, and at the same time the rows increased very
materially in their obliquity. Quite at the posterior end the bristles were so
feeble as scarcely to be visible.

In the foetal wreath I recognised not only the transverse arrangement just
described, but also a distinct antero-posterior or longitudinal arrangement of
the baleen. The outer longitudinal row was formed by the series of large plates,
whilst those internal consisted of the bristle-like baleen. The number of longi-
tudinal rows varied, however, in different parts of the wreath, where eons
occurred in the number of elements in the transverse rows.

The baleen had not the rich black colour so characteristic of the plates in
the older animal. The plates were dark grey, intermingled with black. The
sete were light grey, and the intermediate substance had a similar tint. The

220 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

substance bore a greater proportionate thickness to the vertical diameter of the
entire plate than in the older animal. In one of the largest unequally four-
sided plates, whilst the greatest vertical diameter was 24 inches, the padding
at its thickest part was 1-6 inch; but at the inner and outer border of the plate
it was only 0°8 inch. The greatest transverse diameter of this plate at its
attached border was 2°3 inches. The longest sete projecting from the free
lower border of the plate measured 14 inch. The foetal baleen plates had a
distinctly fibrous appearance, and, from the thinness of the cortex, could be
readily torn along the vertical diameter into numerous fine parallel horny
fibres, which in each plate corresponded in number to the sete, and consisted of
the tubular systems, with their contained papille. The openings into the
tubes were visible in the cleft between the basal laminz of attachment of the
plate. No transverse rings, such as have been described in the older animal,
were seen on the surface of the foetal baleen plates, a circumstance which adds
to the probability of the view entertained by Escuricut and Reruarpt, that
the rings indicate a periodical change in the formation of the cortical part of
the blade. When transverse sections through a plate were examined micro-
scopically, the tubes, the tubular lamelle, and the peripheral lamellz were
seen, but on a much smaller scale; the peripheral lamelle especially being
thinner, and not so distinct as in the older animal, so that the entire plate
was consequently much thinner. The intermediate substance readily tore up
in the vertical direction, and the torn surface was longitudinally streaked, to all
appearance, in conformity with the development of its epidermal cells, in con-
nection with the basal papille. Numerous black pigment granules were scat-
tered through both the plates and intermediate substance.

The surface of the palatal mucous membrane, from which the foetal baleen
had been shed, presented folds or pulp-blades, which, in their general plan,
though with some modifications in form, agreed with those already described
on the palate of the mother. A series of transversely elongated folds corre-
sponded to, and fitted within, the clefts at the bases of attachment of the large
external plates of the transverse rows. Internal to these, owing to the baleen
having so much more of a bristle than a plate-like form, the elevations of the —
mucous surface were not transversely elongated, but had more the shape of sub-
conical papille (Plate VI. fig. 15). The corresponding surface of the baleen
wreath, instead of presenting a series of transversely, elongated, short clefts,
as in the mother, possessed polygonal pits, mostly of a regular hexagonal form i
(fig. 16), into which these sub-conical papille fitted. Towards the anterior partof
the palate, the folds were so faintly marked as to be recognised with difficulty.

As the violence which had occasioned the rupture of the uterus, and the —
displacement of the foetus, had in all probability, also, been the cause of the
separation of the baleen wreaths from the palate, the elongated tubular papillz ;

STRANDED AT LONGNIDDRY. 221

had, for the most part, been torn off the folds of the palatal mucous membrane,
and were included within the tubes of the baleen plates. In some localities,
however, some of these papille still retained their proper attachments to the
folds; and they presented an appearance which reminded one, though on a
smaller scale, of that which has already been described and figured in the older
animal.

JoHN Hunter, in his account of the mode of growth of whalebone,
pointed out very clearly that a baleen plate is formed upon a thin broad pro-
cess of a vascular substance, which fits into the hollow at the base of the
plate, and that the first part of the growth takes place on the inside of the
hollow. He was also of opinion that the cortical layer of the baleen, and the
intermediate substance arose on the surface of the vascular membrane, and
were continuous with each other. He showed their relations to hair, nails, and
other epithelial structures, and stated that the free surface of the intermediate
substance softens like the old cuticle of the sole of the foot when steeped in
water. Escuricut and Rernnarpr described epidermic cells as continually
forming, not only on the pulp-blades, but on the smooth intervals of the palatal
membrane between the blades, the cells of the latter constituting the compara- .
tively soft intermediate substance, whilst those of the former hardened into the
horn-like material of the baleen plate. The medullary or tubular portion of the
plate formed on the free lower edge of the pulp-blade, and on the numerous,
soft, elongated, filamentous papillae which fringe it, whilst the cortical layer of
the baleen plate formed on the free lateral surfaces, and inner and outer edges
of the pulp-blade, which it ensheaths.

This description by the distinguished Scandinavian anatomists is, I believe,
as far as it goes, perfectly accurate ; but the observations which have just been
_ recorded enable me to supplement it with some new and important particulars.
For, in addition to the elongated, filamentous, vascular papille of the tubes,
_ two other sets of vascular papillee have been observed—a cortical and an inter-
mediate—each of which has its appropriate epithelial investment. Hence we
may now state, that each of the three great groups of epithelial cells found in
the baleen wreath takes its rise from, and constitutes the epithelial investment
of, a distinct set of vascular papille. The cells which form the tubular lamelle,
are the cornified epithelium of the filamentous tubular papille: those which
form the peripheral or cortical lamellz are the cornified epithelium of the cor-
tical papille ; whilst the softer intermediate substance consists of the epithelial
cells, which invest the sides and summit of the intermediate papillee.

Many anatomists, in discussing the characters and morphological position
which whalebone occupies amongst the textures, have compared it with the
teeth, and have regarded it as a special modification of the dental tissue,
springing from the surface of the palate. But it seems to me, that a more exact

VOL. XXVI. PART: I. 3M

222 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

comparison may be found in the well-marked vascular folds of mucous mem-
brane, covered by epithelium, which lie transversely across the palate in the
Ruminantia. In the giraffe, for example, these folds are very strong, and they
are, moreover, fringed along the free edge with well-defined papille, which are
also covered with an epithelium. If we were to suppose these papille con-
siderably elongated, their epithelium cornified, and the whole series of papille,
springing from any single fold, bound together by a cortical, cornified, epithelial
layer, we should then have an arrangement of parts closely corresponding in
structure to that of a plate of whalebone. But the Balzenoidea are not the only
placental mammals in which a cornified epithelium is developed in connection
with papillary growths from the surface of the buccal mucous membrane. For,
as is well known, in the Carnivora, the papille on the dorsum of the tongue are
invested with a horny epithelium arranged in the form of retroverted spinules.

I am also of opinion that we must assign to the baleen a more important
function than that of the mere hair sieve or filter, with which it is most usually
compared. For structurally it is much more highly organised than hair. It is
highly vascular, and, I believe, also nervous, and can therefore play the part of a
. whole series of tactile organs, by means of which the animal would be enabled
to estimate the amount and character of the food which it receives into the
cavity of the mouth.

As Georrrey St Hitarre* and Rosert Knoxt had discovered rudiments of
the teeth in the gum of the very young foetus of the Balena mysticetus, and as
Escuricut{ had also observed them in the foetal stage both of Megaptera and —
Balenoptera, I removed the gum from the edges of both the upper and lower
jaws, with the view of examining if the rudiments of these organs still existed
in the almost fully developed foetus of the Longniddry Finner. I found in con-
nection with the periosteal surface of each gum a well-defined band, which
corresponded precisely with the margin of the jaw, and which received a number —
of arteries coming through foramina in the bones. This band, from its position,
was obviously the part in which the teeth, if present, ought to have been found.
A careful examination, however, both of the band and of the tissue on each
side, failed to discover the smallest rudiment of a tooth. Hence it follows that
in the Baleenoidea not only do the teeth not pierce the gum, but all trace even
of their rudiments disappear before the termination of foetal life.

Alimentary Organs.—Owing to the wide sweep of the lower jaw, the cavity
of the mouth was of great size, and the space included between the two halves :
of the lower jaw reminded one of a huge barge ; indeed it was no uncommon ~

* Annales du Museum. Vol. x. p. 364. ;
+ Catalogue, op. cit. p. 22. Kwox’s preparations are in the Anatomical Museum of the University”

of Edinburgh. ,
+ Die Nordischen Wallthiere, 1848.

STRANDED AT LONGNIDDRY. 223

thing, when the animal was lying on the beach, to see a number of persons
standing within the left mandible on the dorsum of the tongue as it was exposed
by the falling over of the beak to the right side. The roof of the mouth was
formed by the palate and baleen plates ; its sides corresponded to the great
antero-posterior cleft between the upper and lower jaws; its floor was formed
by the dorsum of the tongue included within the two halves of the mandible.
The dorsum of the tongue was almost flat near the front of the mouth, but
somewhat further back it presented a considerable elevation, which arose like
a hillock, and fitted within the concavity of the roof of the mouth between the
opposite wreaths of the baleen. The tongue was very compressible and elastic.
The mucous membrane on its surface was of a dark slate colour, and was at
once reflected from the dorsum at the tip and sides of the tongue to the inner
surface of the lower jaw, so that the tongue was tied to that bone, and obvi-
ously could not be protruded from the mouth. The surface of the mucous mem-
brane was firm and tough ; it was marked by ridges and furrows, which, for the
most part, were placed longitudinally, though some extended in the tfansverse
- direction.

The mouth rapidly narrowed towards the posterior buccal orifice. In the
adolescent animal the diameter of this orifice was 10 inches. The mucous
membrane was, in this locality, brownish-yellow in colour, and spotted with
patches of brown and black pigment. Numerous rounded or somewhat oblique
orifices opened on its free surface. These communicated with pits, the largest
of which formed depressions 2ths of an inch deep in the mucous membrane,
big enough to admit peas; these were obviously the mouths of gland follicles.
The upper boundary of the orifice was formed by the soft palate, which was
about an inch and a half thick, and distinct muscular fibres entered into its
construction.

In the foetus the posterior buccal orifice was much more constricted, for its
diameter was only 2 inches. It was bounded above by a broad, well-defined velum,
which extended backwards for 64 inches, and possessed a broad attachment on
each side to the pharyngeal wall, sending also a posterior pillar backwards on
each side as far as a line opposite the arytenoid cartilages (Plate VIII. fig. 30).
The greatest breadth of the soft palate was 7 inches. Its position was almost
horizontal ; mucous membrane covered its upper and lower surfaces and posterior
border, and from the latter no uvula projected. The absence of an uvula in the
lesser Pike Whale had previously been noticed by Drs Carte and MacatisTER.*
Owing to the breadth of the attachment laterally of the velum, the passage from
the mouth to the pharynx was much more in the form of a canal, which may be
termed the bucco-pharyngeal canal, than asimple opening. This canal gradually

* Philosophical Transactions, 1868, p. 232.

224 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

widened in its backward passage, for whilst only the tips of the four fingers could
be introduced into its buccal orifice, the fist could be readily passed through it
from the pharyngeal end. ‘The mucous membrane surrounding the buccal orifice
and lining the bucco-pharyngeal canal, was spotted with pigment, and with pits,
such as have been described in the same region in the older animal. The mucous
membrane was also thrown into faint transverse folds, which corresponded in
their direction with the fibres of the well defined palato-glossus muscle. The
part of the pharynx situated immediately above the velum was greatly dilated,
and measured 24 inches in circumference. It constituted the nasal subdivision
of the pharyngeal chamber. The antero-posterior diameter of the pharynx from
the posterior border of the soft palate to the commencement of the cesophagus
was 9inches. In its general form it was funnel shaped ; for whilst the transverse
diameter just behind the attachment of the velum was 74 inches, it rapidly nar-
rowed behind, where it joined the cesophagus to a tube, 12 inch in diameter.

When the interior of the pharynx was more completely exposed by a mesial
longitudinal incision, not only could the posterior buccal orifice be more clearly
seen, but the relationsof the superior laryngeal opening were exposed (Plate VIII.
fig. 31). In front of this opening was the elongated, tongue-like flexible epiglottis,
which projected forward and upward. It was invested by mucous membrane,
and from its anterior surface a well-defined hyo-epiglottidean fold of mucous
membrane passed forwards to the body of the hyoid. Projecting from the
middle of its posterior surface was a vertical rounded elevation, which obviously
corresponded to the “‘ cushion” described by CzERMAK on the back of the human
epiglottis, and which, doubtless, like that cushion, plays an important part in
the closure of the laryngeal orifice during deglutition. From each side of the
epiglottis a strong aryteno-epiglottidean fold of mucous membrane passed back-
wards to the lappet-like processes of mucous membrane which invested the
horns of the arytenoid cartilages, which formed the posterior boundary of the
orifice. These lappets were separated by a median cleft. No hood-like fold
of mucous membrane, such as Drs CARTE and MACALISTER have described in
B. rostrata, as affording protection to the orifice of the larynx during degluti-
tion, existed in this animal. The superior orifice of the larynx was large
enough in the foetus to admit both fists at the same time.

The muscular wall of the pharynx was formed of the constrictors, the fibres
of which passed from below upwards, to be attached to the superior mesial
raphé of the pharynx. The fibres of at least two pairs of constrictor muscles,
arising from the hyoid bone and thyroid cartilage, were distinctly recognised.
The muscular coat of the cesophagus was comparatively thin, and presented the
longitudinal and circular arrangement.

Numerous glands existed in the submucous coat of the pharynx. The posi-
tion of many of these was marked, more especially on its lateral and anterior

STRANDED AT LONGNIDDRY. 225

walls, by crypt-like depressions in the mucous membrane, some of which were
large enough to admit a kidney bean, others not bigger than a pea (figs. 30, 31).
These crypts were collected into groups, the best marked of which were situated
close to the junction of the anterior border of the soft palate with the anterior
wall of the pharynx.

In studying the method by which this and other whalebone whales collect
their food in their huge mouths prior to deglutition, it should be kept in mind
that they are not provided either with teeth, or with a protrusible tongue by
which to grasp the prey. It is probable that when in search of food, the animal
swims about with its mouth wide open, until a sufficient quantity of food is
collected on the dorsum of the tongue, in the space between the two halves of
the mandible, prior to being swallowed.

Though the depression of the lower jaw in the act of opening the mouth
is doubtless due to muscular action, yet, when once open, the jaw may, I
believe, remain depressed without the continued action of muscles. The huge
- fibrous columns, which pass, one on each side, from the base of the skull to the
condyles of the lower jaw, so suspend that bone, as to support it without the
need of calling into action any muscle ; for it was observed, as the animal was
floating at high water, that the lower jaw was open, and swayed gently to and
fro with the movements of the waves. To draw the jaw back prior to degluti-
tion, the temporal and other elevator muscles must be called into action ; and, as
the jaw is raised, the tongue is pressed upwards against the lower edges of the
baleen, and the water contained in the cavity of the mouth is forcibly squeezed
out between the rows of plates. The food retained in the mouth by the sieve-
like fringes of the baleen, is then forced back through the bucco-pharyngeal
canal, doubtless by the action of the tongue, into the pharynx, when the con-
strictors grasp it and force it back into the cesophagus. Here the soft palate
acts as a valve to prevent its passage upwards to the nose, and the superior
laryngeal orifice is closed by the co-aptation of the epiglottis, arytenoid cartilages,
and aryteno-epiglottidean folds of mucous membrane, so that it cannot enter the
larynx. In these respects, therefore, the mechanical arrangements for prevent-
ing the passage of the food into the respiratory passages, closely remind one
of the structures found in the corresponding locality in the human subject.
As it is also important that water should not pass from the mouth into the
pharynx whilst the animal is collecting its food ; and as the respiratory process
is performed, not by the mouth but by the nose, the contraction of the fibres of
the palato-glossal sphincter would effectually close up the bucco-pharyngeal
canal at the time when these processes were going on.

The stomach was so injured in various places by the men engaged in flensing
the animal, that little more was ascertained in connection with it, than that it
was subdivided into at least four compartments, which communicated with each

VOL. XXVI. PART I. 3N

226 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

other by valvular orifices. One of these valves was secured, and reminded one
on a large scale of the human pyloric valve, or of the valve I described and
figured some time ago* at the end of the fifth compartment of the stomach of
Gilobiocephalus svineval. The stomach was to all appearance empty.

An omentum was in connection with the stomach, which, when stretched
out, was big enough, in the mother, to cover the floor of a large room. It was
made up of fibres, composed of connective tissue, which crossed each other so
as to form a most elegant lace-work pattern, with distinct perforations in the
meshes of the net. Blood-vessels were seen in the larger bands of fibrous tissue
which traversed the net. Scarcely any adipose tissue was found in it, which is
the more remarkable, when we remember the enormous quantity of fat situated —
as a sort of inner padding for the wall of the abdominal cavity.

The intestinal canal was of great length, and by far the longer part of its
extent consisted of huge coils, of which as many as fifteen were counted, though
it is probable that a greater number existed. The hinder end of the gut, as —
it passed backward to the anus, was almost straight, and about 20 feet long. —
No accurate measurement of the length of the intestine could be taken, but it
was estimated at about 80 feet, for the various coils, as soon as they were
removed from the abdomen, were carted away to the manure heap. The
circumference of the tube was not uniform throughout, varying in different —
localities from 20 to 30 inches. Extending along the border of the intestine
at the line of reflection of the mesentery was a very remarkable looking tube
with thick walls, which exhibited an alternating series of dilatations and con-
strictions, which gave it a beaded appearance (Plate VIII. fig. 32, m). This —
tube gave off a number of branches, which ramified in the subserous areolar
coat of the gut, and formed there a complex anastomosing network. Along —
with this moniliform tube was a large vein (v), and accompanying it was a
nerve (7), considerably larger than the human pneumo-gastric, which gave off —
branches to the wall of the intestine. This nerve was obviously a large offshoot —
of the sympathetic. The intestine possessed a distinct peritoneal coat (p),
which rested on the subserous areolar tissue. The muscular coat was thick,
and the longitudinal and circular arrangement of fibres was strongly marked.
A distinct submucous coat was present. The mucous membrane was brownish- —
yellow, and thrown into strong valvule conniventes, some of which extended
two-thirds, others half round the canal of the gut. The largest valvule pro- —
jected at least one inch into the canal. Numbers of parasites were attached to —
the surface of the mucous membrane. I have not as yet had time properly to
examine them, but they are in general appearance like the Echinorynchus brevi- é
collis which Mato found in the intestine of the Balenoptera which he examined.

* Journal of Anatomy and Physiology. Vol ii. p. 73.

STRANDED AT LONGNIDDRY. 227

I can say nothing more of the anatomy of the liver than that it was subdivided
into two lobes. The pancreas was not recognised in the course of the dissection.
Organs of Circulation.—My observations on the arrangement of the heart and
blood-vessels were made chiefly on the foetus, but in several points were supple-
mented by a reference to the corresponding structures in the adolescent animal.
The heart was contained in a well-formed pericardium. In the mother it was of
enormous size ; and in the foetus it was considerably larger than the heart of an ox.
It presented externally the usual arrangement of grooves, which marked its sub-
division into four chambers, and in these grooves the coronary vessels ramified.

In the foetus the right auricle, when opened into, showed a smooth inner
surface for the most part, but the anterior wall and the interior of the appendix
had well-defined fleshy columns projecting into the cavity. In the intervals
between these columns the auricular wall was dilated, and formed a number of
pouch-like recesses. The superior cava, large enough to admit five extended
digits, opened into the anterior and external part of the cavity, and had no
valve at the orifice. The inferior cava, large enough to admit the fist, opened
into the posterior and external part of the auricle. No trace of an Eustachian
valve was seen at its mouth. The mouth of the coronary sinus readily admitted
the tips of three fingers, and opened between the inferior cava and the auriculo-
ventricular orifice, and was also without a valve.

In the interauricular septum an almost circular foramen readily admitting
five extended digits was situated. Surrounding this opening, and attached to
its edge, a loose, membranous, annular fold, formed by a duplication of the
endocardium was seen. When put on the stretch it projected into the auricle,
and the projecting border was free and pierced with large fenestre. Although
this fold was situated in the right auricle, when I opened into that cavity, yet
it could without difficulty be passed through the foramen into the left auricle.
At the attached border, again, the membrane was almost entire, and most per-
fect in its anterior, external, and posterior portions, where the depth from the
attached to the free borders was 4 inches. This membranous fold was situated
at some distance from the mouth of the inferior cava, so that it could not be
regarded as the Eustachian valve in the sense in which it is customary to use
that term. From its position it would, however, seem to have served some pur-
pose in connection with the flow of blood from one auricle to the other during
foetal life; but it is possible that, by growth both in thickness and surface, it
might, after the birth of the creature, have closed up the orifice and completed
the auricular septum. I think it probable that the structure described by Dr
Knox (Catalogue, p. 24), in the heart of a foetal mysticetus, as “a membranous
sac, the size of a full-sized thimble, presenting at the bottom a delicate reticu-
lated net-work, and projecting into the left auricle,” was similar to the annular
fold observed in this foetal Balenoptera.

228 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

A well-defined tricuspid valve was placed at the right auriculo ventricular
orifice. The cusps had the same relative position as in the human heart, and
the arrangement of the carneze columne, musculi papillares, and chorde ten-
dineze was closely similar. In the older whale one of the cusps measured 10
inches in width at its base, and the depth from base to apex was 83 inches.
Some of the chorde tendinez were 12 inches long, and the girth of one of the
largest of these, where it arose from a papillary muscle, was 24 inches. As it
subdivided before it joined the cusp, the size of its branches was very ma-
terially smaller.

The pulmonary artery arose from a distinct conus arteriosus. It ran for-
wards and to the left, and divided into two branches for the right and left
lungs. Its left branch gave origin in the foetus to a widely patent ductus
arteriosus, which joined the arch of the aorta immediately behind a spot oppo-
site the origin of the left subclavian artery (Plate VIL, z).

In the mother a strong, fibrous, rounded cord, 5 inches long, passed between
the pulmonary artery and aorta in the place of the ductus arteriosus. Its
circumference at its aortic attachment was about 6 inches, and it was some-
what thicker at its opposite extremity. When transversely divided it was seen
to be distinctly laminated, and extending along its axis was a canal readily
admitting a large sized catheter. This canal widened out into a funnel-shaped
passage at its two extremities, where it opened into the aorta and pulmonary
arteries. Hence, even in the adolescent animal the arterial duct was patent,
though, from the small size of the canal, any intermixture of blood which might
have occurred would be so small as not to affect the characters of the enormous
volume of that fluid contained in the arterial system. It is interesting also to
note that Knox found a pervious ductus arteriosus in the great Rorqual which
he examined, and Dr Muniz observed it in an adult Balenoptera musculus.*
The trunk of the pulmonary artery in the mother was 3 feet 7 inches in internal
circumference, and its coat, which was distinctly laminated, varied in thickness
from 14 inch to ?ths of an inch. The internal circumference of one of the
primary branches was 1 foot 5 inches, the thickness of its coat 3th of an inch.
The internal circumference of one of the pulmonary veins was 19 inches.

The left auricle, in the mother, had much thicker walls and a redder
colour than the right ; but in both, the appendages were large, and the fleshy
columns within them, and on the adjacent part of the auricular wall, were enor-
mously developed, one of the largest measuring 5 inches by 3, another 6 inches
by 2, and so on. The pouch-like dilatations, already referred to in the descrip-
tion of the foetal auricle, between these columns readily admitted one or both
fists. From the mode in which the columns intersected each other, they and
the pouches gave to this part of the auricle quite a cavernous character. The

* Proc. Zool. Soc., Feb. 14, 1865.

STRANDED AT LONGNIDDRY. é 229

muscular wall at the bottom of some of the pouches was often so thin as to be
translucent when held up to the light. Many of these pouches were situated
parallel and close to the auriculo-ventricular groove.

The left ventricle had thicker walls than the right, and, in connection with
its walls and auricular opening, carneze columne, musculi papillares, chord
tendinez, and a bicuspid valve were seen.

The arch of the aorta in the mother rivalled in its calibre one of the main
pipes for the supply of water to a district of a large city. The internal circum-
ference of its ascending part was 3 feet 2 inches, whilst its coat varied in thick-
ness from 14 to 14 inch. The coat was distinctly laminated, of a yellow colour,
and very elastic. A well-defined inner membrane lined it and the other parts of
the arterial and venous systems. The external circumference of the aorta in
the foetus was 10 inches. It then dilated prior to giving origin to the great
branches of the arch, and immediately beyond these vessels it diminished
materially in size as it became the posterior thoracic aorta. The external cir-
cumference of the innominate artery in the mother was 1 foot 9 inches.

The aorta arched to the left over the root of the lung (Plate VII. fig. 28).
A pair of coronary arteries (a) arose from the commencement of its ascending
part, one passing on each side of the root of the pulmonary artery. Each
coronary immediately subdivided into three branches, the largest of which
turned round its own margin of the heart in the auriculo-ventricular groove, and
supplied the corresponding auricle and ventricle. The second branch of the
right coronary entered the wall of the right auricle ; the third turned round the
root of the pulmonary artery. The second branch of the left coronary artery
descended in the anterior inter-ventricular groove; the third passed to the
substance of the left ventricle. In the mother each coronary artery was as
large as the posterior aorta of an ox.

From the anterior surface of the transverse part of the arch three large
branches arose, the brachio-cephalic, left carotid, and left subclavian (0, ¢, d).
The right branch, by far the largest, was the arteria innominata or brachio-
cephalic (a). Five inches (in the foetus) from its origin it bifurcated into a right
common carotid (e) and right subclavian (7). The right subclavian gave off,
one inch from its origin, a large branch, the right posterior thoracic (y), which
was traced into the great thoracic rete mirabile. One inch and a-half further
on the subclavian bifurcated into the axillary (4) and internal mammary (7)
arteries, the latter of which was somewhat the larger of the two, and supplied
the inferior wall of the chest. The axillary passed in front of the first rib,
immediately above the scalenus anticus muscle; but before doing so it gave off
a considerable branch which ran forwards along the side of the neck. The
axillary was traced into the flipper, and, in the dissection of the fore-arm,

VOL. XXVI. PART I. 30

230 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

branches of this artery were found lying, along with distinct nerves, in con-
nection with the flexor and extensor muscles of the digits.

The right common carotid (¢) ran forwards for 6 inches and then bifurcated. |
The branches should, I think, be regarded as the cervico-facial (4) and internal
carotid (7) arteries. The cervico-facial, much the larger, passed to the deeper
parts of the head, but gave off also a large branch to the face. The internal —
carotid was torn across; but branches arose from it which passed to a rete
mirabile inthe neck. The state of the parts prevented me from tracing out to
their termination the branches of the right common carotid artery.

The second branch of the transverse part of the arch was apparently a left
common carotid artery (c). It gave off a small branch to the side of the neck,
and then bifurcated 7 inches from its origin. The larger branch of bifurcation
was the cervico-facial (m), which divided into many branches for the head and ~
face. The smaller branch was apparently the internal carotid (7).

The third branch of the transverse part of the arch was the left subclavian
artery (d). It gave off a large branch, the left posterior thoracic (0), to the great —
thoracic rete, and then divided into the left axillary (y) and internal mammary (g)
arteries. The rete mirabile was not confined to the thoracic cavity, but ex-
tended upwards into the neck, and prolongations were traced through the inter~
vertebral foramina into the spinal canal. The large foramina at the roots of the
transverse processes of the cervical vertebrz were also occupied by considerable
masses of this highly vascular network.

The posterior thoracic aorta ran backwards, and gave off the series of inter- —
costal arteries. It then entered the abdomen and supplied the various viscera ;
but the distribution of its branches, owing to the injured state of the viscera, —
could not be followed out. It was noticed that in the foetus the hepatic artery
was as large as the human common iliac. The abdominal aorta was prolonged
backwards as the great caudal artery, which was protected by the series of
arches formed by the chevron bones. From the caudal artery, opposite the —
body of each vertebra in the foetus, two branches, which entered the middle of ©
its ventral surface, were traced into the ossifying centrum of each vertebra. .

It may not be out of place to refer to what has been stated as to the arrange- —
ment of the great arteries, which arise from the transverse part of the arch in
some of the other Cetacea, where the vessels have been carefully dissected. —
Kwox,* Escuricat,t and Carte and Macauisrer{ have all pointed out that im
the Balwnoptera rostrata, three great arteries, the brachio-cephalic, left carotid, —
and left subclavian arise from the transverse part of the arch. KwNox also states
that, in his great Rorqual, the arrangement of the vessels arising from the arch

* Catalogue, p. 18. + Die Nordischen Wallthiere, p. 104. SS
+ Philosophical Transactions, 1867, p. 245. }

STRANDED AT LONGNIDDRY. 231

followed closely that of man ; and he refers to brachio-cephalic, left carotid, and
left subclavian arteries; and Matm observed a similar disposition in his
Balenoptera. It seems, therefore, that these great arteries have a similar mode
of origin in different species of Finners. In Delphinus and Globiocephalus, how-
ever, the great arteries arise in the form of two brachio-cephalic arteries, and
the left posterior thoracic arises usually quite independently ; but as I have on
former occasions * described these arrangements, I need not in this place enter
into any further details.

It will be necessary now to give an account of the very remarkable monili-
form tube, which I have referred to in the description of the intestine of the
adolescent animal. It was found along the entire length of the mesenteric
attachment of the gut, and extended back along the rectum. It exhibited an
alternating series of dilatations and constrictions, which varied in their dimen-
sions in different parts (Plate VIII. figs. 32, 33). The dilatations were some-
times globular, at others ovoid in form, and in some cases were flattened on
their surfaces. The largest measured as much as 1 foot 6 inches in transverse
external circumference, whilst the smallest were only 8 or 9 inches. When the
dilatations were ovoid the elongation was mostly in the direction of the long
axis of the tube, in which direction the circumference of the dilatation was
therefore somewhat greater. The constrictions also varied in size, the smallest
being about 4 inches in external circumference, the largest 1 or 2 inches more.
The tube possessed very strong and dense walls, which varied in thickness in
different parts. In the larger dilatations the thickness was as much as 1+ inch,
but in the smaller not more than 4inch. The walls were white, tough, and
very resisting. Examined microscopically, the tissue which composed them
was seen to be chiefly the white fibrous, but mingled with it were elastic fibres.
The inner surface of the wall presented a corrugated appearance, owing to the
presence of a number of permanent, circular folds, wrinkles or ridges, which
passed quite around the inner surface of the tube (fig. 33). In many places
these folds were situated close together ; but elsewhere they were separated by
intervals in which the inner wall of the tube was comparatively smooth. These
ridges were in part formed of a folding of the lining membrane of the tube, and
in part of the fibrous tissue of the wall. Some of the largest of these folds
projected as much as 1 inch, or even more, into the lumen of the tube, and as
this projection was carried all round the inner wall, the lumen was necessarily
much constricted in these localities, and in the smaller divisions the bore was
sometimes reduced to a hole in the middle of the fold less than 1 inch in
diameter, whilst on each side of it the tube might perhaps dilate into a space
2, 3, or more inches in its diameter. Hence the dilated and constricted

* British and Foreign Medico-Chirurgical Review, October 1862, p. 479, and Journal of Anatomy
and Physiology. Vol. ii. p. 66,

232 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

character of the tube visible externally was an index of important internal
arrangements. Numerous branches, into which an injection was readily thrown,
arose from the moniliform tube, and passed directly into the subserous coat of
the gut. They were about the size, at their origin, of the human brachial artery,
and ran straight and parallel to each other for some distance, giving off but
few branches ; then they altered their direction, and formed, by anastomosing, a
series of arches from which numerous branches arose, which ran towards the
free margin of the intestine, again to anastomose, and give rise to still smaller
branches, which penetrated the muscular and mucous coats of the gut.

In connection with the exterior of some of the dilatations of the moniliform
tube a peculiar structure was dissected. It consisted of a number of closely
crowded lacune, varying in size from a pea to a walnut (Plate VIII. fig. 34),
separated more or less perfectly from each other by septa formed of a delicate
smooth membrane, similar to that which also lined the interior of the lacune.
The arrangement to some extent corresponded with that of a multilocular cyst,
the loculi of which communicated with each other. In one spot a distinct tube,
the size of the stem of a common tobacco pipe, was seen to open into a group of
these lacune. In some places, more or less elongated, and sometimes ovoid, bodies
of a dark brown colour, were situated immediately beneath the delicate semi-
transparent ling membrane. These bodies had the appearance of lymphatic
glands, and this view of their structure was confirmed by a microscopic examina -
tion, for, notwithstanding that the specimen had been for sometime in spirits
of wine, distinct, pale, circular, lymphoid corpuscles were seen to enter in large
numbers into the structure of these bodies. I did not succeed in tracing out
any connection between this lacunary system and the wall of the intestine,
though it is possible that the small tube, just referred to, may have proceeded
from or to the wall of the gut.

It was unfortunate that in the portions of intestine, with the moniliform
tube attached, which were sent over to the Anatomical Museum for examination,
none of the expanded part of the mesentery had been preserved. I was con-
sequently unable to trace the branches which proceeded from the proximal
surface of this tube to their origin. I have little doubt, however, but that they
were derived from the mesenteric artery.

In the foetus the intestine was, as a rule, so softened by putrefaction that it
could not be preserved. One or two coils were, however, somewhat more per-
fect, and after being hardened in strong spirits of wine, I was enabled to effect a
partial examination.

The mesenteric artery did not possess that complete series of arterial
arcades, which we are familiar with in man. It branched comparatively seldom, .
and its branches ran towards the border of the intestine. Those which arose
nearest the gut did not enter directly the intestinal wall, but passed to an

STRANDED AT LONGNIDDRY. 233

elongated structure, which lay parallel to and next its mesenteric border. This
structure occupied the position of the moniliform tube in the parent whale, but
did not possess its beaded appearance. Indications, in places, of a tube travers-
ing its long axis were seen; but in the greater part of its extent it was appa-
rently subdivided into a large number of minute spaces, so that the surface of
section had quite a cavernous aspect. From this structure numerous fine
branches arose, which passed into the subserous coat of the intestine, to be
distributed there like the branches of the moniliform tube in the parent animal.
It would seem, therefore, that in the foetus the moniliform tube is not developed
in the same precise manner as in the adolescent whale, but that a series
of inter-communicating spaces occupy the position in which it subsequently
appears. The formation of the moniliform tube, out of this lacunary system,
would be occasioned by a great increase in size of those lacunz which lie in
the same longitudinal series, and by the great hypertrophy of their originally
delicate walls. It is probable that the lacunee described on the surface of some
parts of the dilated tube in the parent (fig. 34), represented in it the original
condition of the mesenteric lacunary system of the foetus.

In the Cetacea, important arrangements, in connection with the vascular
system, exist in various parts of the body for the purpose of modifying and
equalising the force of the blood current. The great cervico-thoracic rete
mirabile, with its numerous offshoots into the spinal canal and cranial cavity, is
the arrangement which has been most carefully studied by different anatomists.
But in considering the function of this network, it is not sufficient to regard it
as merely a reservoir, or huge sponge, which contributes,.by its complex rami-
fications, to produce an enormously extended area for the reception of the blood,
when the whale dives to a great depth from the surface of the ocean. It serves,
I believe, the purpose, by minutely subdividing the arterial stream, of distribut-
ing and equalising the force of the blood current before it reaches those delicate
organs the brain and spinal cord. It may be regarded, therefore, as the teleo-
logical equivalent of the arteries in the human pia mater, of the circle of Willis,
of the tortuosities in the vertebral and internal carotid arteries, aud of the rete
mirabile in connection with the intra-cranial arteries in ruminants and in
the pig.

With what, then, are we to associate the large moniliform tube in the me-
sentery of this whale? From its beaded character it might at the first glance
be supposed to belong to the lymphatic system ; but the careful consideration
of the distribution of its branches, and of its relations to the mesenteric arteries,
have led me to the conclusion that it is a remarkable modification of the mesen-
teric arterial system, which serves the same office, for the intestine, that the rete
mirabile does for the brain and spinal cord.

The great size of the aorta and of the trunk of the mesenteric artery, the

VOL. XXVI. PART I. 3 P

234 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

paucity of the system of arterial arcades, the proximity of the intestine to the
aorta, the pressure, from the elastic recoil of the arterial wall, of the enormous
column of blood in the aorta, would seem to render some mechanical arrange-—
ments necessary, by means of which that pressure may be distributed and
regulated before the blood enters the slender arteries within the wall of the
intestine. .

The structure of the moniliform tube admirably adapts it for this purpose.
The blood flows through it on its way to the intestinal arteries, and is diffused
into the numerous dilatations or bays which bulge out from its sides. The
transverse inflexible folds on its inner wall diminish at intervals the lumen of
the tube, and where they project so far as to leave but a narrow aperture in —
the axis of the tube, they act as strictures in retarding the flow of the current.
At the same time their circular arrangement enables them to act as internal
girders, and to strengthen the walls so as to prevent over distension of the tube.* —

I have already referred to the analogy between the rete mirabile in the cetacea,
and the network in connection with the intra-cranial arteries in the pig. I may
now allude to a modification which the pig exhibits in the arrangement of its —
mesenteric arterial system. The arteries subdivide in the middle of the mesen-—
tery, and form there a compact network—a rete mirabile—from which numer- —
ous small arteries radiate outwards to the intestine.t These radiating vessels
closely correspond in appearance to those which I have described as arising
from the moniliform tube in the Longniddry whale. The Cetacea, therefore, —
present affinities to the Pachydermata, not only in the diffused character of the
placenta, but in the possession of closely allied modifications of the cerebral —
and intestinal arterial systems.

The presence of a moniliform tube, in connection with the intestine, does —
not seem to have been previously recognised in the Cetacea by anatomists.

The superior vena cava was formed by the junction of the two innominate
veins, on the right of the ascending aorta. Each innominate vein began at the
root of the neck in the form of a dilated sinus, into which the veins from the
neck, flipper and inner wall of the chest opened. The inferior cava received
a number of hepatic veins before it pierced the diaphragm. The umbilical vein
was 27 inches long in the foetus in its course from the umbilicus to the
liver.

The portal vein in the foetus had a diameter of 3 inches before it entered — .

* My colleague, the Professor of Engineering, Professor FLEEMING JENKIN, to whom I pointed out

the structure of this tube, concurs in the opinion of its function expressed in the text.
+ The mesenteric rete in the pig has long been known to anatomists—see Barotay on the —
Arteries, Edinburgh, 1812; T. J. Arrx1 in Reports of Edinburgh Meeting of British Association,
1834, p. 681 ; Owen, Comparative Anatomy of Vertebrates, vol. iii.; Gurur, Anatomie der Haus
saugethiere, Berlin, 1860. The complexity of the rete in the pig is due to the plexiform arrangement
of both the mesenteric vein and artery.

STRANDED AT LONGNIDDRY. 235

the liver. In the coil of intestine from the adolescent animal, from which
fig. 32 was taken, a vein larger than the human inferior cava, ran close and
parallel to the great moniliform artery of the intestine, and received numerous
veins, the rootlets of which took their origin within the coats of the gut. In
the foetus a vein lay along with the artery in the expanded part of the me-
-sentery.

At the upper part of the cavity of the thorax in the fcetus, close to the
apex of the pericardium, a well-defined, though small, thymus gland was found.
It was subdivided into two lobes, each of which was brown in colonr, thin,
and flattened in form, and 5 inches in length by 43 inches in its greatest
breadth. The lobes were subdivided into distinct lobules by intermediate con-
nective tissue, and they received numerous blood-vessels. In proportion to
the size of the animal the gland was obviously smaller than might have been
anticipated. The thyroid gland, supra-renal capsules and spleen were not re-
_cognised during the dissection.

Organs of Respiration.—When the cavity of the thorax was opened into, by
the removal of the inferior wall, the lungs were exposed. In the foetus each
lung was an elongated, flattened organ 2 feet 8 inches in length. It was in-
vested by a distinct and smooth pleura, and was not subdivided into lobes by
fissures. A similar absence of fissures and lobes I have also seen in the lung
of B. rostrata. The pulmonary artery, veins, and bronchus entered its substance
_ through the hilum on its mediastinal surface. When the lung was removed and
washed with a jet of water, the softened pulmonary substance broke down, and was
washed away, and the arrangement of the intra-pulmonic part of the bronchus
could be seen. This tube, as a rule, branched in a dichotomous manner, though
collateral offsets sometimes proceeded from it. It was accompanied by the pul-
monary and bronchial arteries, and by bronchial nerves of some size.

The cartilaginous framework was much more perfect than in the human
bronchus. The tube was hooped with cartilaginous, spirally arranged, ring-like
plates ; in the larger tubes usually not more than once and a-half, but in the
smaller tubes a greater number of times (fig. 35). Sometimes in these latter the
cartilage formed perfect rings, and both in them and in the larger tubes
the cartilaginous plates not unfrequently bifurcated. The branching of
the plates was always well-seen at the angle of the bifurcation of the tubes.
The plates were invested by a well-defined perichondrium. The hoop-like and
spiral coils of these cartilaginous plates have an important office in connection
with the respiratory process in this animal. They not only aid in keeping the
tubes open, but, by their elasticity, aid in the recoil of the lung during the great
expiratory effort which the whale makes in the act of blowing. The diameter
of the right bronchus in the foetus was 2 inches, that of the left 24 inches ; in
the mother one of the bronchi was 7 inches in diameter.

236 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

Three inches in the foetus, above the place of bifurcation of the trachea, that
tube gave off a supplementary bronchus, 14 inch in diameter, to the right lung,
which seems to be, as SANDIFoRT and Escuricut have pointed out, the usual
arrangement in the cetacea, the Greenland right whale being excepted.* The
trachea had three, somewhat irregularly formed, cartilaginous hoops immediately
above the bifurcation ; but from the highest of these up to the arytenoid carti-
lages, a distance of 64 inches, which corresponded to two somewhat subdivided
tracheal rings, and to the interval between the separated inferior borders and
plate-like processes of the cricoid, the cartilage was deficient inferiorly, and the
ventral wall of the wind-pipe was formed of fibrous membrane. The mucous
membrane of the trachea, more especially on the anterior wall, was marked by
numerous fine reticulated folds, the chief of which ran parallel to the long axis
of the tube. The diameter of the trachea was about 5 inches.

The cartilaginous framework of the larynx consisted of a thyroid, a cricoid,
a pair of arytenoid cartilages, and an epiglottis. The form, arrangement, and
connections of these cartilages were examined in the foetus (Plate VIIL., figs.
36, 37, 38).

The thyroid cartilage consisted of a median and two lateral portions. It
was a comparatively thin plate, and possessed two surfaces, a superior and in-
ferior, which were flattened, and two margins, an anterior and a posterior. The
median part, tongue-like in form, was bifid at its hinder border, and projected
for some distance backward ; a deep notch marked its superior border; from
this notch, to the end of the forks of the tongue-like part, the diameter was 44
inches. The lateral portion curved outwards, and was then prolonged back-
wards, as the elongated and somewhat rounded posterior cornu to be articulated
by a moveable joint with the outer surface of the cricoid. The anterior cornu
was continuous with the anterior border of the cartilage ; it was short and rudi-
mentary. The cartilage was connected to the body and great cornua of the
hyoid bone by a strong membrane, and a pair of thyro-hyoid muscles passed
between them.

The cricoid cartilage was an incomplete ring ; superiorly, it formed a thick
mass of cartilage 7 inches in its antero-posterior diameter. Its surfaces were
curved, and it turned round the sides of the wind-pipe towards its ventral
aspect, and ended in the greater part of its extent in a free rounded border.
From the hinder part of this inferior border, however, five plates, similar in
form to the cartilaginous hoops of the trachea, arose and turned round the
side of the larynx to the ventral surface; but the plates from opposite sides
did not meet in the mesial line. An interval, varying in its transverse diameter

* Die Nordischen Wallthiere, p. 148, Ray Society's translation of Memoir on Greenland Whale,
p. 103.

STRANDED AT LONGNIDDRY. Iie

from 3 to 4 inches, separated the opposite inferior margins from each other.
It was filled up by a strong fibrous membrane, which was continuous laterally
with the perichondrial investment of the cricoid and its plate-like offshoots,
anteriorly with the perichondrium investing the posterior horns of the two
arytenoid cartilages, and posteriorly with the membrane which filled up the
interval between the ventral borders of the first two cartilages of the trachea.
This membrane, which may be called the inferior crico-tracheal membrane, was
of great importance as completing the wall of the windpipe on its ventral aspect.
The posterior margin of the cricoid was comparatively narrow; the anterior
margin possessed at each lateral angle a broad surface for articulation with the
body of the arytenoid cartilage, distinct capsular and synovial membranes con-
nected the two cartilages.

Each arytenoid cartilage, irregular in form, consisted of a body and two
cornua. The body formed a thick plate of cartilage. The anterior cornu
curved upwards and forwards into the lappet-like fold of mucous membrane
behind the superior laryngeal opening. The posterior cornu curved back-
wards and inwards within the area enclosed by the sides of the cricoid ; it
almost reached the mesial plane, where a transverse fibrous ligament connected
it by the tip to its fellow. The two posterior horns formed an imperfect hoop,
invested by the mucous membrane of the larynx, which was prolonged directly
backwards and downwards to form the mucous lining of the laryngeal sac.
The free rounded border of the cricoid was connected to the posterior cornu
of the arytenoid by the inferior crico-tracheal membrane. A. crico-thyroid
muscle existed also on each side, and muscular fibres were seen to occupy the
position of the crico-arytenoidei postici and arytenoideus.

The epiglottis contained a bar of yellow fibro-cartilage, which passed back-
wards along the axis of the entire structure, to be attached to the superior
surface of the middle portion of the thyroid cartilage. In the older animal,
from which it had been removed without much injury,* the entire organ
measured 25 inches in length, whilst its breadth at the base was about 10
inches ; it was thick and massive, and rounded in form at its free end. The
fibro-cartilage was covered by mucous membrane, which was prolonged back-
ward as the aryteno-epiglottidean folds, and forward as the hyo-epiglottidean
fold. When this membrane was removed from the hinder surface of the epi-
glottis and its arytenoid connecting folds, a strong aryteno-epiglottideus muscle
was exposed, which curved upwards and inwards, decussating with its fellow
in the substance of the epiglottis, and obviously was arranged to act as a

_* The other laryngeal cartilages were so much injured during their removal from the adolescent
whale, that I was unable to examine them satisfactorily. I may, however, refer to their great size and
thickness, more especially of the cricoid and body of the arytenoid. The cartilage was traversed in
various directions by very distinct vascular canals.

VOL. XXVI. PART I. 3 Q

238 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

powerful sphincter for closing the glottis during deglutition. A deeper set
of fibres of the same muscle was exposed by the removal of the thyro-hyoid
membrane.

There were no true vocal cords passing from the thyroid to the arytenoid
cartilages, or laryngeal ventricles, but a slight fold of the mucous membrane,
extended obliquely in the antero-posterior direction within the aperture of the
glottis, on each side, a short distance below the free edge of the aryteno-
epiglottidean folds. These might, perhaps, be regarded as rudimentary false
vocal cords.

One of the most interesting structures connected with the larynx was the
great laryngeal pouch or cul-de-sac. It was 10 inches in length in the foetus, and
extended backward from the thyroid cartilage, in close relation to the ventral
surface of the inferior crico-tracheal membrane, to within 2 inches from the ‘
bifurcation of the trachea. Its outer wall was formed by a powerful muscle,
which arose from the superior surface of the median tongue and adjacent lateral
plate of the thyroid, from the inferior free border of the cricoid, and from the
body of the arytenoid. The fibres were arranged in transverse rings around
the walls of the pouch, and they formed a thick mass at its posterior end. The —
pouch was lined by a mucous membrane, which was continuous with the general
mucous lining of the larynx, by extending upwards on the inner surface of the —
bodies of the arytenoid cartilages, and by passing round the free border of the —
hoop formed by their posterior horns. The mouths of numerous large crypts
opened on the surface of this membrane. :

Owing to the peculiar arrangement of the arytenoid cartilages and the
presence of this pouch, the laryngeal chamber might be regarded as subdivided —
into three compartments. The supero-anterior which formed the glottis proper,
was bounded by the epiglottis, the aryteno-epiglottidean folds, and the anterior —
horns and bodies of the arytenoid cartilages with their investing and intermediate
mucous membrane. The posterior was bounded above and to the sides by the
cricoid cartilage, in front by the two posterior horns of the arytenoids, which
ran obliquely from above backwards and downwards; through the fissure
between these horns it communicated with the glottis, whilst behind it was con- —
tinuous with the canal of the trachea. The inferior was the laryngeal pouch
above described, which communicated directly and freely with the glottis at
the base of the epiglottis, but with the posterior chamber through the fissure —
between the arytenoid horns. This pouch is often regarded as occasioned by a —

deficiency in the ventral part of the ring of the cricoid cartilage. But from

the description of the arrangement of these parts, and from the figure 37, it
will be seen that although this plate of cartilage is defective, yet that the ring
is completed ventrally by the strong inferior crico-tracheal fibrous membrane, |
beneath which the pouch is situated. The laryngeal sac is rather to be regarded

STRANDED AT LONGNIDDRY. 239

as a diverticular prolongation of the mucous membrane between the thyroid
and cricoid cartilages, accompanied by an imperfect development of the crico-
thyroid membrane.

The air entering the lungs during inspiration would have to pass from the
glottis into the trachea through the fissure between the posterior horns of the
arytenoids ; but the air, entering the laryngeal pouch, would pass into it below
these two horns. The close approximation of these cornua would aid in the
closure of the glottis, and in the retention of the air in the lungs when the whale
has dived to a depth from the surface.

The presence of a laryngeal pouch or sac in the B. rostrata, which he dis-
sected, had not escaped the acute observation of Joun Hunter. In his
account of that animal he says,* “The arytenoid cartilage on each side sends
down a process, which passes on the inside of the cricoid, being attached to a
bag which is formed below (behind) the thyroid, and before (below) the cricoid ;
these processes cross the cavity of the larynx obliquely, making the passage at
the upper part a groove between them.” Sanpirortt then pointed out and
described its arrangements in two foetuses of Balena mysticetus. KNox ob-
served it{ not only in B. rostrata, and B. mysticetus, but in his great northern
Rorqual, and he specially directed attention to the mode in which it was
supported by the posterior horns of the arytenoid cartilages. Escuricut has
also recognised this sac not only in the foetus of B. rostrata, but in that of
the Megaptera longimana ;; and REINHARDT and he have anew carefully
described it in the Greenland Right Whale. A description, with several
figures, of the sac in B. rostrata has recently been published by Messrs CartE
and MACALISTER.|

Of these authors the last named alone discuss the probable use of this very
remarkable pouch. ‘They consider, that by the contraction of its muscular walls,
it may expel the contained air so as to augment the power of, and to sustain the
expiratory current. They suggest that it might aid in the production or modu-
lation of sound, if the whales possessed such a faculty, but think that the size
of its aperture, and the absence of all constricting bands, or apparatus, militate
against that view of its use.

The powerful muscular wall of the sac is unquestionably for the purpose of
“permitting the contraction of the wall on the contents, and as the pouch com-
municates above directly with the glottis, a rapid contraction of the investing
muscle would aid the expiratory act. But there is another purpose to which
this sac may be applied. It may serve the office of a reservoir in which a

* Structure and Economy of Whales.
+ Nieuwe Verhand. van Wetensch. te Amsterdam, 1831.

t Catalogue, pp. 11, 17, 23. § Die Nordischen Wallthiere, p. 103, e. s.
|| Op. cit., p. 236, e. s. ;

240 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNNER WHALE

quantity of oxygenated air may be stored up to be made use of when the animal
remains for some time below the surface, by permitting an interchange, by
diffusion, to take place between the pure air in it and the carbonised air within
the lungs.

It has been customary to consider, that because the Balenoidea have no
vocal cords, therefore they have no voice, and cannot produce sound.* But
although they do not possess a pair of elastic bands, extending horizontally —
across the larynx between the arytenoid and thyroid cartilages, such as we see in
the mammalia generally, yet in the posterior horns of their arytenoid cartilages,
united by the transverse ligament, they possess a pair of structures which can —
be approximated or divaricated, and by the vibration of which, as the air —
passes between them, into or out of the lungs, sounds may very probably be
elicited.t Their vibration would, without doubt, be assisted by their close
relation to the air-filled laryngeal pouch.

The nares consisted of two vertical passages, separated by a cartilage e
septum, which opened superiorly on the dorsum of the head by the external —
apertures or blow-holes, whilst by their deeper orifices they communicated with —
the nasal part of the pharynx. When looked at from below (fig. 39), the
mucous membrane was seen to be pitted with the mouths of numerous gland
follicles, and to cover the surface of an oval fibro-cartilage which formed a
considerable convexity in the outer and anterior wall of the passage, and in —
contact with the outer surface of which was a muscle. When the external
orifice was widely opened, a fold, occasioned by the position of a large postero-
external cartilage, fitted into a corresponding depression on the antero-external -
wall (fig. 40). A muscle, apparently a dilator, lay beneath the skin to the
outer side of the aperture, and was attached to the cartilage at its postero-—
external angle. It is clear that the nostrils can be readily and widely opened, —
and also forcibly and completely closed, during the respiratory movements, so-
as to retain the air within the windpipe and lungs when the animal dives below
the surface of the water. Ng

Genito-urinary Organs.—In fig. 9, the form and relation of the penis —
in the foetus are represented. As all that portion of the organ in front of the
crescentic folds was invested by integument, the penis in this animal seemed in
its flaccid state, not to be altogether retracted within a sheath, but to be in part

* Dr Martyn, in a paper published in the Proc. Roy. Soc., London, 1857, ascribed the supposed —
absence of the voice im the cetacea to the absence of a thyroid gland ; but as I pointed out in a memoir —
published, in 1860, in the Transactions of this Society, a thyroid gland exists both in Phocena and —
Delphinus. %

+ Whilst this memoir is passing through the press, Dr Muriz has published i in the “ Journal of
Anatomy and Physiology,’ November 1870, an interesting paper on Grampus rissoanus, in which he
points out that a laryngeal sac of moderate capacity exists in the toothed whales in the angle of june—
tion between the enlarged epiglottis and the thyroid cartilage. He also describes a pair of folds within”
the larynx of Risso’s grampus, which he regards as representatives of the vocal cords.

STRANDED AT LONGNIDDRY. 241

pendulous from the ventral wall. The organ consisted of a distinct corpus
spongiosum urethre, and of a strong corpus cavernosum. These bodies
extended backwards for eight inches behind the crescentic folds above referred
to. The corpus cavernosum then subdivided at a very obtuse angle to form
the crura penis, which were firmly connected to the perichondrial investment
of the larger and more rounded end of the rudimentary and still cartilaginous
ossa innominata, which represented, therefore, the ischial elements of the
pelvis.*

A strong muscle, which must be regarded as the erector penis, arose from
the ischium, close to the attachment of the crus, and passed forward to: be
inserted into the corpus cavernosum. Large vessels and nerves were also seen
passing to the different subdivisions of the penis. From the posterior border
of each diverging crus, and from the sides of a tendinous raphé, which extended
backwards from the end of the corpus spongiosum, a broad and strong muscle
arose, which passed backwards along the side and under surface of the hinder
end of the rectum, and ended close to the anus. This muscle was apparently
the retractor penis. The corpus spongiosum had unfortunately been torn
across, where the crura diverged, and the rest of the urethra, the bladder,
testicles, &c., were not distinguishable, owing to the soft and injured state of
the parts.

The arrangement of the parts at the entrance to the female passage has
been described on p. 201. The vagina was traced forwards for six feet from the

external orifice. Numerous irregularly arranged, and much subdivided, folds
of the mucous membrane projected from its surface into the canal. The uterus
was not recognised with any certainty, but a bag-like membranous organ, a
part of which was seen to project through a long cut in the wall of the
abdomen, on the day on which the baleen wreath of the foetus was found loose
in the abdominal cavity, was supposed to be a portion of that organ.

The kidneys possessed the lobular construction so characteristic of the form
of those organs in the cetacea.

_ Ishall reserve for another communication the description of the skeleton and
joints, and such observations on the arrangement of the muscles as I was
able to record. I may, however, state that the vertebral formula, both in
the foetus and in the mother, was—Cervical, 7 ; Dorsal, 15; Lumbo-caudal, 41:
in all, 63. The outline of the cranial beak was in conformity with that of the
head, which is so well represented in fig. 10 from the fcetus.
| The following are a few measurements of the skull, taken with a tape-line,
which may be of service in comparing it with the crania of other described
whales :—
* T have described and figured the innominate bones and the sternum in the “ Journal of Ana-
tomy and Physiology,” May 1870.
VOL. XXVI. PART I. oR

242 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

Feet. Inches.

From anterior border of foramen magnum over vertex to tip of beak, . 20 3
From nasal process of superior maxilla to tip of beak, : 3 16 6
From anterior border of foramen magnum to nasal process of superior

maxilla, . : ; ; : 3 9
Breadth across upper ends of nasal processes, 1 8
Breadth of a single nasal process. 0 6

Breadth of dorsum of beak—

3 feet in front of nasal end of superior maxillaries, . 7 0
t 9 Eb 99 6 10
5 2? 7 »? 6 8

Chas, a be 6 64

ioe ” ¥ 6 4}

8 iy 7? ” 6 14
oes; 5 z: 5 9
Uae s 5 4
Auli: :: ; 4 10
125 os 95 4 i)
13 ” ’ ” 3 8

1 Weare hs - 2 «Pie
15 ” ” ” 2 1
Breadth at the tip of the beak, 0 7
Breadth between the orbits, . 9 3
Length of lower jaw along the convexity, 21 2
a 5 in a straight line, ny 5
Depth of ramus at coronoid process, . 2 6
Length of humerus, 2 2
bs radius, 3 9

Comparison with other Finners.—In instituting a comparison between the
Longniddry whale and the other Fin whales which have been described by
different authors, with the view of determining the species to which it should be
referred, there is no need to compare it with either the Balenoptera rostrata, or
the Balcnoptera laticeps. For these animals never, apparently, exceed the length
of 35 feet, and they differ so materially from the Longniddry whale in the number
of vertebree and ribs, that there can be no possibility of confounding it with
either of them. My remarks, therefore, will be restricted to a consideration of
those described specimens of fin whales which have reached the length of 40
feet and upwards. As I have not yet given an account of the skeleton of
this large Finner, I shall almost entirely confine myself, on this occasion, to an
examination of the external characters of these animals.

a. Sir R. SrpBap, in his observations on rare whales cast on the Scottish
coast,* describes two fin whales. One, he says, rostrum acutum habet, et plicas
in ventre; the other maaxidlam inferiorem rotundam, et superiore multo latiorem
habuit. The one with a sharp beak was cast ashore in 1690 near Burntisland,
and measured 46 feet in length. It was in all probability an immature
specimen of the Razor-back. The other with the rounded lower jaw, much
wider than the upper, was stranded on the south side of the Forth, near the
old castle of Abercorn, in 1692. It was a male, 78 feet long, and possessed

* Phalainologia nova. Edinburgh, 1692.

STRANDED AT LONGNIDDRY. 243

various points of resemblance to the Longniddry specimen. From the greater
width of the lower jaw than of the upper, the latter was received within the cir-
cumference of the former. The upper jaw was contracted in front so as to
terminate ina sharp end. The baleen was black, the longest plates having a
vertical diameter of 3 feet, and they were fringed with black hairs. The
bristles near the front of the palate were also black, and the intermediate
substance was similar in character. The flipper was 10 feet in length, and 2
feet in its broadest part. The dorsal fin was 2 feet high, and in it was a
rounded hole made by a leaden ball. Through this hole in its fin the whale had
been recognised by the herring fishermen for nearly twenty years, and was called
by them the Hollie Pyke. The back was black and the belly whitish. The
blubber was 44 inches thick on the sides, and one foot on the head and neck.

Although it is customary for cetologists to regard this broad-jawed whale,
described by SIBBALD, as the Balenoptera musculus,* yet the characters which I
have just related are much more those of the species to which the Longniddry
whale will have to be referred.

b. The best known of the large fin whales is the common Razor-back, the
Baleenoptera musculus, or Physalus antiquorum of Gray, upwards of thirty
specimens of which have come under the notice of, and been more or less perfectly
described by, naturalists. Between the common Razor-back and the Long-
niddry whale there are many characteristic features of difference. In the
former the beak is much more pointed than in the latter, and its width rapidly
contracts from base to apex, instead of the borders forming a gentle convex curve;
the flipper also is absolutely and relatively shorter in proportion to the length
of the animal. In the B. musculus, captured near Gravesend, described by Dr
Murts,t whilst the animal was 60 feet, the length of the pectoral limb along
the anterior curve was only 6 feet 3 inches ; in the specimen 67 feet long, stranded
at Pevensey, described by Professor FLower,{ the flipper was 6 feet 9 inches ;
and in the specimen 61 feet long, beached last year at Langston harbour,§ the
flipper had the length of 5 feet 4 inches. The external or labial baleen plates
are in the common Finner neither so long nor so broad ; their colour is slate-
coloured, mottled, or striped with yellow, or white, or brown, or pale horn
colour, the setze are white, or yellowish-white ; the palatal mucous membrane
is pale, whilst in the Longniddry whale all these structures had a rich deep
black colour. In the Razor-back, whilst the back is black, the belly is white or
yellowish-white, instead of being mottled with silver-grey, or milk-white, tints.
The blubber also is very much thinner in the common Razor-back,—not more

* Escuricut, “Die Nordischen Wallthiere.’” Van BenEpEN and Gervais,“ Ostéographie des Cétacés,”
p. 188. Dr Gray in his Catalogue says, probably it may belong to this species.

+ Proc. Zool. Soc., Feb. 14, 1865.

t Idem., Nov. 28, 1865. § Idem., Dec. 9, 1869.

244 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE

than four tons of oil were extracted from the Gravesend specimen,—so that the
animal possesses very little commercial value, whilst several hundred pounds
have been realised by the sale of the oil from the Longniddry animal. Further, —
it is very doubtful if the Balenoptera musculus exceeds the length of 70 feet ;
the Gravesend and Pevensey specimens, already mentioned, were both adult
males, and yet they did not reach that length. Several specimens which have
been referred to this species are, it is true, stated to have been longer than 70
feet; but of these, some, I believe, belong to another species, whilst it is
doubtful how far the others have been measured with sufficient exactness. More-
over, the vertebree in B. musculus are not so numerous, and do not apparently
exceed sixty-one. ;
It is not necessary to compare the Longniddry whale with the Physalus
Duguidii, described by Mr HEeppLte* and Dr Gray,t as that animal is appa-
rently nothing but a young specimen of the B. musculus.
c. In the year 1827 a fin whale, said to have been upwards of 80 feet long,
was found floating on the North Sea, and towed into the harbour of Ostend, —
from which circumstance it has been customary to term it the Ostend whale.
Unfortunately, no satisfactory account of the dimensions and external charac-
ters of this animal have been recorded, and the descriptions of the skeleton are
in some respects imperfect. Zoologists, therefore, are by no means at one as
to the genus or even species to which this whale ought to be referred. Dr
Gray places it in his genus Sibbaldius, and calls it S. borealis; Escuricut has
termed it the Balwnoptera gigas, or Pterobaliena gigas; whilst VAN BENEDEN ©
and GERVAIS, in their Ostéographie, have regarded it as merely an unusually large
specimen of the B. musculus. Owing to the very imperfect data at my com-
mand, I cannot make any exact comparison between its external form and
that of the Longniddry whale. I may state, however, that the length of the
pectoral fin is said to have been about 10 feet; the distance from the point
of the snout to the dorsal fin 61 feet ; from the point of the snout to the
genital organs 55 feet; that the back was black, and the belly whitish, the
outer surface of the pectoral fin was black, and the baleen sete also black.{
In these respects it more closely approaches the Longniddry whale than it does
the B. musculus. It must be admitted, however, that the measurements, which
have been recorded by those who have described the animal, are not of a very
reliable character, for, whilst VAN BreEDA states its length to be about 84 feet,
Dvusar makes it as much as 105 feet. I shall have again to refer to the Ostend

* Proc. Zool. Soc. 1856. + Catalogue, p. 158.

{ The notices of this animal which I have read, and from which the above statements are drawn
are he M. Van Brepa in Cuvier’s “ Hist. Nat. des Cétacés,” p. 328; by Escuricur in “ Die N: ordischen
Wallthiere,’ p. 176; by Littyepore in the Memoir translated fon the Ray Society, p. 262; by D
Gray in his “ Catalogue of Seals and Whales ;” and by Dusarin his “ Ostéographie de la Baleine,” For
the opportunity of consulting Dusar’s searee pamphlet, I am indebted to my colleague Profess
KELLAND.

STRANDED AT LONGNIDDRY. 245

whale when I describe the skeleton of the Longniddry whale, and to point out
certain other points of correspondence between them. I may on this occasion, °
however, state that the small number of vertebre, 54, described in the former
animal is obviously owing, as DuBAR’s figure shows, to the loss, in the prepared
skeleton, of several vertebre in the caudal series. And there is good reason
for believing that the double headed condition of the first rib which DusBar
figured in this creature, and on the presence of which Dr Gray has to a large
extent based his genus Szbbaldius, is merely an individual peculiarity, and may
occur as a variety in more than one species of whale, just as it occasionally
occurs as a variety in the human subject. .

d. In the month of October 1831, a fin whale was observed floating dead
on the surface of the sea off the mouth of the Firth of Forth, and was brought
ashore near North Berwick, 23 miles from Edinburgh. It was purchased and
anatomised by Dr and Mr Frepericx Knox. The skeleton was carefully pre-
pared and publicly exhibited, and now forms the most noticeable object in the
Natural History Department of the Museum of Science and Art, Edinburgh.
Unfortunately no systematic description of this animal was ever published ; but
from one or other of the publications mentioned below* I have gathered the
following particulars. The animal was a male, and measured 80 feet in length.
The length of the head over the vertex was 21 feet ; the pectoral limb 11 feet
long ; the circumference behind the pectoral limbs 34 feet, and even 52 feet when
_ greatly distended with gas ; the breadth of the tail 20 feet ; the distance from the
anus to the fork of the tail 21 feet.t The whole baleen, with its fringed edge,
was of a clear shining black, and the longest plate measured 2 feet 2 inches in
length, by 15 inches in breadth. Nothing is said as to the colour of the skin

or the thickness of the blubber; but it is stated in the “ Account,” that “the
fluid oil in the abdomen, particularly, was in very considerable quantity, and
often gave a covering to the sea as far as the eye could reach.”

Kwox named the animal the Great Northern Rorqual, or Balena maximus
borealis.

In July 1847, Dr J. E. Gray stated to the Zoological Society of London,t{

* Abstract of a paper on the “Anatomy of the Rorqual (a Whalebone Whale of the largest
magnitude),” by Dr Ropert Kwox (Proc. Roy. Soc. Edin., March 18, 1833). “ Account of the Gigantic
Whale or Rorqual, the Skeleton of which is now exhibiting in the great rooms of the Royal Institution,
Princes Street,” by Freperick Jonn Kwox, surgeon, Edinburgh, 1835. “ Catalogue of Anatomical pre-
parations illustrative of the Whale, particularly the Great Northern Rorqual,” by F. J. Knox, Edinburgh,
1838. Although the name of Mr Freprrick Knox is attached to the catalogue, yet it would appear
that the best part of it was from the pen of Dr Knox (“ Life of Knox,” by Dr Lonspats, p. 168). For
the opportunity of consulting this scarce and valuable catalogue, to which I have referred on various
occasions in the text, I beg to express my acknowledgments to my friend Dr Jonn ALEXANDER SMITH.
The skeleton of this animal is figured in Jarpine’s “ Naturalist’s Library,” vol. vi. Edinburgh, 1837.

+ The Gravesend B. musculus was only 11 feet 1 inch between the points of the tail. The
Pevensey Razor-back, about 13 feet ; the Langston harbour specimen 11 feet. In the Pevensey whale

the distance from the end of the tail to the middle of the anal aperture was 17 feet 9 inches.
+ Proceedings, Part xv. p. 117.

VOL. XXVI. PART I. as

246 PROFESSOR TURNER'S ACCOUNT OF THE GREAT FINNER WHALE

that he had examined, though without being able to take any measurements, on
account of its position, the skeleton of this animal. He considered it to be a
Physalus, very nearly allied to the Physalus antiquorum, though it differed from —
a specimen of that animal taken at Plymouth in some of the characters of its
cervical vertebrae. Since that time it has been customary to describe this great
whale as the Balewnoptera musculus, or Physalus antiquorum.*

A comparison of the measurements, which I have quoted, with those of the
Longniddry whale, given in the early part of this paper, and the very decided —
statement made as to the clear, shining, black baleen and sete, will, I think,
suffice to show that in its general proportions, and the colour of its baleen, the
North Berwick whale resembled closely the Longniddry whale, and differed,

_therefore, in many most material points from the common Razor-back, so that
it can no longer be regarded as of that species. The shape of its cranium,
also, differs most materially from that of the B. musculus. Knox, in his cata-
logue, has given a few measurements of its skeleton, which, if compared with
those of the Longniddry animal, will show that a close resemblance exists be-—
tween these animals in this part also of their frames. The breadth between the
orbits was 10 feet ; the length of the base of the cranium measured in a straight —
line, 19 feet ; the length of the lower jaw, 21 feet 4 inches ; circumference of —
ramus about the middle, 4 feet ; depth of ramus at coronoid process, 2 feet 7—
inches ; depth of body of hyoid, 84 inches ; between the ends of the great
cornua, 2 feet 65 inches ; length of the humerus, 1 foot 11 inches ; of the
radius, 3 feet 10 inches. But it is right also to mention that there are differ-
ences in the skeleton, especially in the form of the sternum and the pelvic
bones, and whilst the North Berwick whale has thirty ribs, it possesses as many
as sixty-five vertebra. The more complete comparison of the skeletons of
these two animals I shall reserve for the second part of this memoir.

e. In 1847 Dr Gray described,t by the name of Physalus Sibbaldii, from
the skeleton of an immature animal 47 feet long, in the museum of the Hull
Literary and Philosophical Society, a new species of Finner, the baleen of which
possessed a uniform deep black colour. In 1864 Professor FLowER { dis-
covered in the collection of the late Professor LiptH DE JEUDE, of Utrecht,
the skeleton of a young finner about 44 feet long, which differed from the —
common Razor-back in possessing a much broader beak. He named it Physalus”
latirostris§, Subsequently, on examining the skeleton in Hull, which Dr Gray —
had observed, he came to the conclusion that the animals were of the same-
species, and he withdrew his specific name in favour of the prior one given by

* Dr Gray, “ Catalogue of Seals and Whales,” p. 144; Van BenepEN and Gervais, “ Ostéographie,”
p. 172 and various other writers on the cetacea.

+ Proc. Zoological Soc., June 8th.

t Idem, Noy. 8, 1864, and June 13, 1865.

§ This skeleton has since been acquired by the British Museum.

STRANDED AT LONGNIDDRY. 247

Gay.R Since then Dr Gray has changed the generic name to Curerius, and
terms the animal C. Sibbaldii.* Those zoologists who do not break up the
great genus Balenoptera ito several smaller sub-genera, prefer to call the
animal Balenoptera Sibbaldii. The Hull and Utrecht skeletons agree in pos-
sessing each 64 vertebre ; but whilst the former has 16 pairs of ribs, the latter
has only 15 pairs. No information existed as to the external characters of
either of the animals from which these skeletons were obtained, so that it was
difficult to identify them with any of the species of whales known to zoolo-
gists, up to that time, only by their external appearances.

In 1867, however, Professor REINHARDT published an important memoir, in
which he gave an account,t from notes furnished him by Mr HA t.as, surgeon
to a whaling ship, of a Finner of which the Danish whalers had captured several
specimens. This whale was known to the Icelanders as the Steypireythr. The
back was blackish grey ; down the sides the colour was lighter ; the belly, behind
the plicze, was uniformly grey, the ridges blackish grey ; the furrows between
them, light grey ; the caudal fin, blackish grey on both sides ; the pectoral fins,
blackish grey, spotted with lighter specks on the outer surface, but milk white
onthe inner. The baleen was uniformly black. The animal was about 80 feet
long, and was said to have a dorsal fin not more than 7 inches high.{ No measure-
ments are given of the caudal or pectoral fins, or, indeed, of the proportions of
the other parts of the body. Mr Hattas also forwarded the skull, hyoid bone,
and atlas of this animal, of which RetnHArpT gives figures. Further, he states
that the animal possessed 64 vertebree and 15 pairs of ribs. In his remarks on
this whale, REINHARDT compares it both with the Balenoptera musculus (Physalus
antiquorum) and B. Sibbaldii, and considers that from its osteological characters
it should be referred to the latter species.

By these observations, it was clearly established that a well-defined species
of Finner exists in the northern seas, which differs from the common Razor-back,
in possessing a greater number of vertebrae, a broader beak to the cranium, a
greyish and not a whitish belly, and a uniform black baleen, instead of one
mottled with various tints. In the distribution of the tints of the skin, in the
uniform black colour of the baleen, and in the length of the animal, the Stey-
pireythr obviously closely corresponds with the Longniddry whale. But what is
even more important for the determination of the species, the cranium, atlas, and
hyoid, as far as one can judge from REINHARDT’s figures, are almost identical in

* Appendix to Catalogue of Seals and Whales, p. 380.

* + Vidensk. Meddelelser fraden Naturhist, Forening iKjobenhayn, 1867. Translated in “ Annals
of Nat. Hist.,” November 1868.

{ Although the end of the dorsal fin had been removed from the adolescent Longniddry whale
before my measurements were taken, yet sufficient had been left to show that this fin had been more
than 12 inches high. Consequently, I do not think that the shortness of the dorsal fin is so definite
a character as REINHARDT supposes.:

248 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE.

form with the corresponding bones in the Longniddry whale. Hence we arrive
at the conclusion, that the Longniddry whale is a specimen of the Balenoptera
Sibbaldi, or Physalus Sibbaldii of GRay.

Two years before the publication of ReEmInHARDT’s memoir, a fin whale, about
54 feet long, came ashore alive at Gothenburg, on the west coast of Sweden.
It was secured by Professor Mam, the superintendent of the Museum in that
city, and was carefully examined by him. He published an elaborate mono-
graph, with numerous photographic illustrations, descriptive of the capture of
the animal, its form, colour, proportions, and dimensions, with a detailed ac-
count of the skeleton, and a number of observations on its visceral anatomy.* The
animal was a male, and had not reached its full growth. Its colour was a deep
slate tint, with somewhat paler tints on the sides, whilst the lower surface was
mottled with patches of milk white, of different sizes and shapes. The flippers
were white on the inner surface, and the lobes of the tail at the under part
whitish. The distance from the anterior part of the base of the flipper to its
free extremity, measured in a straight line, was 7 feet 4 inches, whilst the dis-
tance between the extreme points of the tail was about 11 feet. The baleen
was uniformly of a deep black slate colour, whilst the hairs at the free margins
of the plates were of a brown soot colour. The vertebree were 63 in number,
and there were 15 pairs of ribs. Mat considered it to be a new species, and
named it Balenoptera Caroline.

From a comparison of its osteological characters with those of the B. Sib-
baldii, more especially the resemblance in the breadth of the beak, the form of
the nasal bones, the relative and absolute length of the metacarpals and phalanges,
and the spine of the axis, as well as from the uniform dark colour of the baleen,
Professor FLowER came to the conclusion,t that Maum’s whale ought not to be re-
garded as a distinct species, but was merely another immature specimen of the
Balenoptera (Physalus) Sibbaldii. In this conclusion he has been supported
by Professor REINHARDT, who states{ that, in his opinion, “ Escuricut’s ‘Tun-
nolik,’ the ‘Steypireythr’ of the Icelanders, and, finally, the whale described by
MAM, are only one and the same species, which appears to be one of the most
common in our northern seas, and the systematic name of which must be
Balenoptera Sibbaldi.”

If I am correct in regarding the Longniddry whale as the B. Szbbaldii, then—
Professors FLOWER and REINHARDT being also correct in their supposition—its _
characters should closely correspond, allowance being made for the different sizes
of the two animals, with those of Matm’s whale. In the colour, both of the skin
and the baleen ; in the shape of the tail and pectoral fin ; in the relative pro-

* “Monographie illustrée du Baleinoptére,” Stockholm, 1867. For the opportunity of consulting
this work, three copies only of which are, I believe, in this country, I am indebted to my friend, Mr J.

W. Cuarg, of Cambridge.

+ Proc. Zool. Soc., March 12, 1868. t Memoir, cited above. ¢

STRANDED AT LONGNIDDRY. 249

portions of these parts to the length of the entire body ; in the form of the beak ;
and in the curve of the lower jaw, the resemblances are very striking. The
osteological characters have also much in common; but the consideration of
these I shall not enter into on this occasion.

The comparison I have now made between these different specimens of
Finners, leads me to the conclusion that the following should be referred to
the Balenoptera Sibbaldiit :-—

The North Berwick whale.

The Hull skeleton.

The Utrecht skeleton, now in the British Museum.

The Gothenburg whale.

The Steypireythr.

The Longniddry whale.

And, in all probability, the Ostend whale, and Sibbald’s “ Baleena tripinnis
_ quee maxillam inferiorem rotundam, et superiore multo latiorem habuit.”

EXPLANATION OF THE PLATES.

With the exception of fig. 1 Plate V., of figs. 19, 20, 21, 22, 23, 24, 25, 27, 28, Plate VII., and of
fig. 29 Plate VIIL., the illustrations have been very carefully drawn, under my superintendence, by Mr
J. B. Apercrompis, from nature. Fig. 27 was drawn by Mr Covucurrey, fig. 28 by Mr Foutts, and
figs. 19 to 25 inclusive, and fig. 29, were sketched by myself from microscopic preparations. As far as
possible, the specimens from which the drawings were taken have been preserved in the Anatomical
Museum of the University of Edinburgh. When not otherwise stated, the drawings represent portions
of the adolescent animal.

Prater V.

Figure 1. Side view of the Longniddry Whale. This drawing was constructed from photographs,
from pencil sketches, and from a water-colour sketch by Mr Sam. Bouen. The
lower jaw is represented somewhat out of position so as to give a side view of the baleen
and of the dorsum of the tongue.

Figure 2. The falcate dorsal fin of the foetus.

Figure 3. The horizontal tail of the foetus.

Figure 4. The abdominal plice of the foetus, showing bifurcations of the ridges.

Figure 5. Supero-anterior surface of the left flipper of the foctus. The outlines of the bones of the
antibrachium and of the four digits are represented. The posterior edge of the flipper
was much thinner than the anterior.

Puate VI.

Figure 6. The clitoris, below which is the opening of the urethra, and the folds of mucous membrane,
on the floor of the vestibule. The labia majora have been drawn asunder to expose these

parts.

Figure 7. The orifice of the nipple fossa, displaying the nipple with the pedunculated papillx at its

summit.

Figure 8. The anal orifice, with the rug of the integument converging to it.

Figure 9. The ventral wall of the foetus, displaying the penis with the crescentic folds of skin at its
root, the median perineal raphé, with a rudimentary nipple fossa on each side, and, more
posteriorly, the anal orifice.

VOL. XXVI. PART I. or

250 PROFESSOR TURNER’S ACCOUNT OF THE GREAT FINNER WHALE
Figure 10.
Figure 11.

Figure 12.
Figure 13.

Figure 14,

Figure 15.

Figure 16.

Figure 17.

Figure 18.
Figure 19.

Figure 20.

Figure 21.

Figure 22.

Figure 23.

Figure 24.
Figure 25.

Figure 26.

Figure 27.

Figure 28.

Dorsum of the beak of the foetus. The curved outline of the beak; the dorsal median
ridge ; the form and direction of the blow-holes, which are aie open, and the inter-
mediate groove are all represented.

A portion of the mammary gland to show the rugose character of the mucous lining of the
duct, one-half the size of nature.

One of the large, irregularly quadrilateral, labial, baleen plates, much reduced in size.

A vertical section through the intermediate substance of the baleen, displaying its laminated
appearance. The subsidiary blades are shown, two of which have been cut short. On
the upper or palatal surface, the clefts between the laminz of the plates, into which the
palatal folds of mucous membrane fit, may be seen.

The baleen plates and intermediate substance of the foetus, the size of nature. One of the
plates, with the thin layer of intermediate substance on each side, has been partially sepa-
rated from the others.

Portion of the palatal mucous membrane of the feetus. At the upper end of the figure is
the lip ; lower down the elongated folds for the larger labial baleen plates, and at the lower
end the subconical papillee from which the bristle-like subsidiary plates arise. The tubular —
papille are not represented in the drawing, as they had all been broken off before the —
drawing was made.

Palatal surface of a part of the foetal baleen wreath, with a side view of one of the plates.
The elongated clefts between the laminz of the labial plates and the polygonal pits for the
sub-conical papille are shown; as the tubular papille are still within the blades, their
broken ends may be seen in part occupying the clefts and pits. ;

Puate VII.

Portion of the foetal membranes ; a, the non-villous surface of the chorion ; 8, villous sur-—
face. Between a and b Ae elongated marginal folds of the chorion may be seen.
c, Divided end of one of the arteries oe the cosa

A large triangular fold of the chorion, displaying the reticulated arrangement of the villi on
its surface. ©

Vertical section through a portion of a baleen plate to show the tubes, with the lamellx
and black pigment granules. x 40 diam.

Transverse section through a portion of a baleen plate. The entire antero-posterior diameter
is represented, The tubes are divided transversely. Some are empty, others contain the —
tubular papille, and in some of these the transversely divided ends of the contained blood-
vessels may be seen. Both the tubular and cortical lamellz, with numerous black pig-
ment granules, are represented in the drawing. x 40 diam. .

Epithelial cells from the outer layers of two adjacent tubular systems. At the lower part
of the drawing some interstitial cells are represented. x 200 diam.

Transverse section through one of the setz of a baleen plate. The shaded central portion —
represents the soft papilla, in which a transversely divided blood-vessel is represented.
x 40 diam.

Vertical section through a portion of the intermediate substance ; the clefts extending into
its substance, in which the intermediate papille are lodged, are seen at the upper part of
the section. x 40 diam.

Epithelial cells from the intermediate substance. x 200 diam.

Red blood corpuscles from the blood in the vessels of the baleen plate of the B. rostrata.
x 1200 diam.

Portion of one of the elongated folds (pulp-blades) of the» palatal mucous membrane. The
tubular papille are dependent from the lower edge of the fold. Size of nature.

Vertical transverse section through the pulp-blades, intermediate substance, and imbedded —
parts of the baleen plates of B. rostrata, the blood-vessels of which have been injected ; 2,
the vessels of the intermediate papille; c, the vessels of the cortical papilla; ¢, the elongated —
vessels of the tubular papillz.

Arch of the aorta and great vessels of the foetus. 2, the ductus arteriosus; a, right
coronary artery; 6, brachio-cephalic ; ¢, left carotid; d, left subclavian; e, right carotid ; 7,
right subclavian ; g, right posterior thoracic ; h, right axillary; 7, right internal mammary 5
k, right cervico-facial ; and 7, right internal carotid ; m, left cervico-facial ; and n, left in-
ternal carotid ; 0, left posterior thoracic ; p and ! left axillary and Fiscel mammary
arteries.

Figure 29.

Figure 30.

Figure 31.

Figure 32,

Figure 33.

Figure 34.

Figure 35.

- Figure 36.

STRANDED AT LONGNIDDRY. 251

Prats VIII.

Vertical section through the integument. It shows the elongated papille, the comparatively
thin cuticle containing a quantity of black pigment, and the subcutaneous tissue, with the
small arteries entering the bases of the papilla. x 20 diam. ;

Dorsal surface of the pharynx and commencement of the cesophagus of the foetus; ph, the
pharynx displaying the fibres of the constrictors and the longitudinal raphé. The upper
part of the pharynx has been cut across, and the soft palate v is displayed ; passing under
it is an arrow lying in the bucco-pharyngeal canal. Immediately behind the velum a por-
tion of the epiglottis is visible. :

The interior of the cavity of the pharynx of the foetus opened into by a posterior median
incision ; v, the velum ; e, the epiglottis, the letter is placed on the cushion, which corre-
sponds in position to the bar-like rod of fibro-cartilage ; 7, the lappet-like fold of mucous
membrane which invests the superior horn of the arytenoid cartilage, the outline of which
may be seen in the figure. The upper arrow is in the bucco-pharyngeal canal, the lower
is in the windpipe.

Portion of the intestinal tube. v, the superior mesenteric vem which receives numerous
rootlets from the gut; m, the moniliform tube, giving off numerous small arteries to the
wall of the intestine ; , the sympathetic nerve, also sending branches to the gut ; p, the
peritoneal coat turned down. At the right cut edge of the intestine the valvule conni-
ventes of the mucous coat are shown.

A portion of the beaded mesenteric vessel, displaying the series of dilatations and constric-
tions. At the right side the tube has been opened, and the corrugated folds of the inner
wall may be seen.

One of the dilated portions of the beaded vessel. The lacunary system on its surface is
represented. The darkly shaded, elongated, and globular bodies, J 7, are the small lym-
phatic glands.

Annular and spirally arranged plate of cartilage from a bronchial tube.

Front view of the larynx and hyoid apparatus ; h, the body of the hyoid with the stylo-
hyal and great cornu on each side. Immediately above the hyoid body is the orifice of
the bucco-pharyngeal canal, the arrow lying in which has emerged below through the
cesophagus ; th, the thyro-hyoid muscle; sh, the stylo-hyoid muscle ; ¢, the thyroid
cartilage ; c, the cricoid ; cm, the constrictor muscle of p, the laryngeal pouch. The bifur-
cation of the trachea and the supplementary right bronchus are seen, and the arrow
passed through the left bronchus emerges superiorly, immediately behind the posterior
horn of the left arytenoid cartilage.

Figure 37. Front view of the larynx and trachea ; the laryngeal pouch has been removed and the cartilages

Figure 38.

Figure 39.

Figure 40.

dissected, ¢, the thyroid cartilage ; c, the cricoid with its plate-like processes ; a, the body ;
s, the anterior, and 7, the posterior cornu of the arytenoid cartilage ; ct, the inferior crico-
thyroid membrane. The barb of the arrow passed through the left bronchus, lies im-
mediately behind the posterior horn of the left arytenoid cartilage, and in front of the body
of the cricoid, which is in deep shadow.

View of the interior of the larynx from behind, obtained by cutting through and turning
outwards the body of the cricoid, and the membrane connecting the anterior horns of the
two arytenoid cartilages ; ¢, the epiglottis; c, the cricoid ; J, the lappet of mucous mem-
brane enclosing s, the anterior horn of the arytenoid ; 7, the posterior horn. To the inner
side of the anterior horn is the fold of mucous membrane, which may represent a false vocal
cord.

The posterior nares viewed from below.

The anterior nares or blow-holes viewed from above ; the walls are separated to show the
internal foldings.

Figures 35 to 40 inclusive are from the fcetus.

Pierce ee

R Ni) Farlane Lath? Edint

Plate X.

yy. Soc. Edin™ Vol. XXVI

XII.—On some Points in the Structure of Tubifex. By W. C. M‘Inrosu, M.D.,
Fi Kh. (Plates 1X, X:)

(Read 2nd May 1870.)

At least two species of Tubifex are abundant in Scotland, one of which is,
perhaps, more prevalent in streams, the other in lakes. The former is common
at the margin of the river Tay, when the water is low, in sandy tubes of little
tenacity, and often in tunnels formed amongst the wet sand under stones, just
as many of the marine annelids occur. Its length varies from three-fourths of
an inch to an inch and a-half when stretched, and the segments range from fifty
to seventy, the average number probably being sixty. The body is of various
shades of dull fawn or pale madder-brown, somewhat interrupted by the pale
bands at the junctions of the segments, and streaked by the reddish dorsal
vessel ; or, in summer, marked anteriorly by the whitish mass of the reproductive
organs. It is easily recognised amongst its fellows by its faintly purplish or
lilac hue, as well as by its peculiar mode of progression ; and not a few are
observed with reproducing heads and tails. This form, perhaps, has most claim
to be called Tubifex rivulorum.

The body is tapered towards head and tail, distinctly segmented, and much
shorter and stouter than the succeeding species. Moreover, the length of the
bristles does not equal the diameter of the body. The snout is somewhat
conical, with the puckered mouth at its posterior margin, near the first crenation,
which indicates its separation from the succeeding segment. There are many
motionless microscopic processes on the surface of the snout in well-developed
Specimens, and the same organs occur on the bristle-papille and the caudal
segment. Such have generally been regarded as tactile papille. Though
analogous, they are not quite homologous with similar processes in the Turbel-
Ilaria. The second segment bears a long bristle at each side dorsally, and three
hooks, one of which is small. The third segment has two bristles and three
hooks on each side superiorly ; ventrally four hooks on each side. Generally
the inferior hooks are five or six in number on each side, one being short and
in process of development. The bristles (Plate IX. fig. 1) seem slightly stouter
than those of the longer form, and no serrations are visible on the sides. M.
D'UDEKEM noticed in his form a bristle with a brush-like tip, and has figured

the same,* but though Mr Lanxesrer has seen such in examples of Tubifex

% Hist. Nat. du Tubifex, &c. ; Mém. couronnés et Mém. des say. étrangers, &c. ; Acad. Roy. de
Belgique, tom. xxvi. p. 11, Plate II. fig. 8.

vol. XXVI. PART II. By 10S

254 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

from the Thames, they have never occurred in those found in this neighbour-
hood. The number of pairs of bristles in front ranges from twenty-one to
twenty-three. The hooks are gently curved organs, with a bifid tip, and a dis-
tinct swelling or shoulder about the upper third, from which point they taper
towards the base. Those accompanying the bristles anteriorly (Plate IX. fig. 1)
slightly differ in their curvature from those of the ventral series (Plate IX. fig. 2).
In the other and longer form (with about 150 segments) from the lakes, the
fourth segment has a pair of bristles, and the latter increase in length till the
twelfth segment is reached, after which they gradually diminish and disappear
about the fortieth. There is a small papilla where each bristle-bundle passes
through the skin, and the tips of the hairs are delicately serrated or roughened, ©
the serrations being directed distally. In this form the bristles are larger than
the diameter of the body, and hence it differs from Nazis scotica and the Nais
lacustris of DAtyELL. The hooks commence with the bristles, and besides
those accompanying the latter, form two rows, as usual, inferiorly, which rows —
in front consist of groups of four hooks. Those accompanying the bristles —
(Plate TX. fig. 3) are more closely forked at the tip, and if examined under a
power of 700 diameters show certain processes in the fork (a), a fact first
pointed out to me by Mr LAnKEsTER, whose larger specimens from the Thames
exhibited this and other peculiarities in a marked manner. These are also
less shouldered, less curved, and somewhat more elongated than the inferior
hooks. The latter in each form of 7uhifex continue after the last bristle-
bundle, and thus form four rows posteriorly, the terminal segment only being
bare. M. p’UDEKEMw’s representations of the hooks,* though easily recognisable,
deviate a little from the foregoing, and the same may be said of M. CLAPAREDE’S
figures of the hooks of his Tubifex papillosus.t q

Body-Wall.—M. v’UbDEKEM speaks of the epidermis as being separable from
the chorion by the aid of an alkaline solution, but I have not been able to dis-—
engage it as a distinct layer either by the action of chemicals in the fresh animal,
or in transverse sections of the body-wall. M. CLAPAREDE does not mention
the. superficial layer as a distinct coat, but groups it with the subjacent, under
the name of cuticle.{ The cuticular surface (or layer) is quite homogeneous, |
but the chorion which is incorporated with its inferior surface is distinctly
cellulo-granular, as described by. GRurrHuIsEN in Nais (Chetogaster), and by
Bucuuouz in Enchytreus.. This is most distinctly marked at the snout and
tail, where the layer is thickened. The other layers in Tubifex are a belt of
circular muscular fibres, and a longitudinal muscular coat. M. CLAPAREDE
gives an ideal section of Limnodrilus, a form which differs from Tubifex in the-

* Op. cit. Pl. II. figs. 6 and 7.
+ Beobach. tiber Anat. u. entwicklung. &c., p. 25, Pl. XIII. fig. 15.
{ Recherches Anat. sur les Oligochétes, p. 7.

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 255

absence of the bristles and other points, in which he shows the longitudinal
muscular coat separated into six divisions, viz.,—two dorsal, two lateral, and
two ventral. I have not been able to see this arrangement in the transverse
sections of the minute forms of 7wbifex, and even the separation at the bristle-
sacs is comparatively indistinct. This ambiguity is no doubt due to the small
size of the specimens. In the living animals, however, certain rows of papill
may occasionally be observed, which probably mark the dorsal and ventral lines.
The circular muscular coat is much thinner than the longitudinal. The latter
forms in transverse section numerous well-defined fascicles, and in the fresh
condition these are bounded internally by a membranous layer with many cells.
At the period of reproductive activity the neighbourhood of the eleventh seg-
ment becomes almost opaque from a cellular covering. This appears to be due
to an increased development of the cellulo-granular elements of the chorion.
Adhering by short stalks to the walls of the body posteriorly in both forms,
were numerous parasitic vorticellz (Plate IX. figs. 4 and 5), having an active
crown of cilia, and numerous globules and granules in their interior. In some
examples these were very numerous, often in groups of three or four, but they
rarely occurred on the terminal segment, except in decomposing individuals.
In contraction the base becomes finely corrugated. The free motion of the tail
of the worm in the water places these organisms under very favourable condi-
‘tions for aeration and food. Fungi, also, may frequently be seen growing on the
disorganised anterior segments, while the posterior areinfull activity. Fresh water
annelids, indeed, are prone to have such growths, just as young salmon are
under similar circumstances. The anterior part of the body becomes first
attacked, dissolving into a granular mass swarming with fungi and infusoria.
The segment immediately behind the decayed portion shows its integument
corrugated and thrown into transverse rugee, while the perivisceral corpuscles
and the blood have disappeared. The next septum is strongly contracted, and
in marked contrast to the succeeding segment, within which are many perivisceral
corpuscles of the ordinary appearance, and whose dorsal vessel pulsates
vigorously.

Perivisceral Fluid and Corpuscles.—The perivisceral fluid occupies, as usual,
| the space within the body-wall all round, from the tip of the snout to the tail, sur-
| rounding and bathing the digestive and other structures contained therein. In
| Lubifex rivulorum, the perivisceral corpuscles (Plate IX. fig. 6, a), are very con-
spicuous, and differ characteristically from those of the succeeding form from

the lakes.* They are rounded bodies filled with circular granules of consider-

* Whatever result more extended investigations may give with regard to the specific value of
characters derived from the shape of these bodies, it is right that the name of Dr Tuomas Wittrams
| should be honourably remembered in conjunction therewith. Vide his paper on the “ Blood-proper
and Chylaqueous Fluid of Invertebrate Animals,” Philos. Trans. P. II., 1852.

256 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

able size, and in the living animal undergo various changes in shape by pres-
sure against each other, the body-wall and viscera of the worm. On extrusion
into the surrounding water they become very transparent, and their broken’
contained granules collect together at one point (Plate X. fig. 1). Tincture
of iodine and chromic acid colour them deep yellow, while sulphuric ether
does not materially affect them. Dilute glycerine first corrugates, and then
causes them to burst, giving exit to the contained clear granules (Plate IX.
fig. 7), which measure about sv'ssth of an inch in diameter. Some of the
corpuscles are smaller, and their contained granules less in proportion. Besides
the foregoing, there are many elliptical, curved, and granular corpuscles (Plate
IX. fig. 8) in the perivisceral space. In the elongated form the perivisceral
corpuscles are less conspicuous both as regards number and size. The larger
bodies in this case are rounded cells (Plate IX. fig. 9), filled with much more
minute granules than in the preceding form, and many show a granular nucleus.
The other corpuscules (Plate IX. fig. 10) are elliptical or fusiform, flattened, trans-
parent and non-granular, and often longer than the diameter of the ordinary
granular corpuscle just described. As contrasted with a gland-cell from the
intestinal wall, the perivisceral corpuscle in the former case is widely different,
while in this it has much smaller granules, is pale, and easily distinguished from
the other with its highly refracting yellowish granules. In a form occurring ~
abundantly in certain lakes with the latter, and referable to M. CLAPAREDE’S
genus Limnodrilus, the perivisceral corpuscles are remarkably developed. They —
are larger than usual, very transparent, and frequently show a somewhat
puckered outline within the body of the worm. They also have the peculiarity
of adhering here and there in considerable numbers to the intestinal coating.
Few or none of the ordinary fusiform or other bodies are present. On ex-
truding these corpuscles into the water they swell out, and show a large
granular nucleus, the rest of the cell being quite translucent. Moreover, both
cells and nuclei have a nearly uniform diameter throughout the fluid. On con-
trasting these corpuscles (Plate IX. fig. 11) with the gland-cells from the
intestine (Plate IX. fig. 12), a very marked difference is observable. The for-
mer are quite transparent—with the exception of the nucleus, becoming slightly
cranular only after remaining many hours in the water. They have nuclei
of definite size and structure, which retain their shape and appearance after the
bursting of the cell-wall. Acetic acid and ether only show the structure just —
described more clearly. On the other hand, I have watched the gland-cells
from the intestine under water and various reagents side by side with the others,
and for a considerable time; but I do not feel able to say that nuclei or
other contents than the refracting granules have been detected. The gland-cell
may be set in motion, rolling over and over, so as to expose it all round, yet
not a trace of a nucleus is seen. In all the forms there is thus a considerable

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 257

histological difference between the two sets of cells; and both differ very
much from the cells on the inner surface of the intestinal wall (Plate IX.
fig. 13.)

I have been somewhat minute in observing this point, because it has
generally been stated that the perivisceral corpuscles have their origin from, or
are closely connected with, the gland-cells which cover the intestine and dorsal
vessel. M. p’UDEKEm is stated by Mr Lanxester, in his recent paper on
Cheetogaster,* to have connected the two in his memoir on Tubifex, but such is
not my impression. It is true the author describes two kinds of “ glandules ”
covering the intestine and the dorsal vessel, viz., nucleated brownish “ glandules,”
and colourless ‘“ glandules” having oily contents, and says they represent the
liver of the higher animals,—secreting a liquid for assisting digestion. In speak-
ing of the perivisceral fluid, moreover, he omits all notice of the origin or rela-
tionship of the corpuscles ; and adds that the number and large size of these
“olobules lymphatiques” constitute one of the differences between the young
Tubifex, on its extrusion from the egg, and the adult. He hints at no connec-
tion between the two structures. Mr LANKESTER, also, in the same paper does
not fully express the published opinions of M. CiLapar&pe on this point. He
says, “There is a very distinct relation between the abundance of the perivisceral
granules and cells, and the abundance of the brownish granules which surround
in masses the dorsal vessel and its ramifications on the stomach or intestine.
CLAPAREDE, in his‘ Recherches sur les Oligochétes,’ has spoken of the brown-yellow
‘hepatic’ tissue of the intestine in Lumbricus being continued to and surround-
ing the dorsal vessel, and has hinted (but only obscurely) at some connection
between the perivisceral cells and the supposed hepatic tissue.”+ Now, in the

first place, in the memoir alluded to, M. CLApAREDE did not specially refer to

Lumbricus so much as to those genera included under his family of ‘‘ Oligochetes
Limicoles” (Tubifex, Limnodrilus, Clitellio, Lumbriculus, Stylodrilus, Enchytreus,
&c.), whose structure formed the text of his work. Chctogaster, of course,
would come under the same head. M. CLAPAREDE states that the pigment-
cells of the intestine have generally been considered as hepatic, and points out

that the said cells have as much connection with the dorsal vessel as with the

intestine ; that in Lumbriculus variegatus, for example, the cellular coating,
which ceases to cover the intestine at the sixth segment, continues on the
dorsal vessel to the fourth, and, moreover, the coating follows certain branches
of the dorsal vessel. Further, he adds, that the intimate connection between
the supposed hepatic structure and the vascular system is extremely evident in
the true Lumbrici, and concludes with the following :—“ Il est donc trés-im-

* Trans. Linn. Soe. vol. xxvi. p. 637.
{ Mr Lanxesrer has altered his views here——Vide his paper in the Quart. Jour. of Micros. Sc.
vol. v. N.S. p. 109.

VOL. XXVI. PART. II. 3X

258 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

probable que ces cellules versent de la bile dans la cavité de Vintestin. II est
beaucoup plus vraisemblable qu’elles déversent leur contenu dans la cavité
périviscérale.” He reiterates this opinion in his recent beautiful and accurate
memoir on the Histology of the Earthworm.* Dr GRvUITHUISEN, indeed,
clearly anticipated most of the subsequent observers in regard to the connection
between this glandular coating and the perivisceral fluid, which he termed the
chyle. In describing the glands which envelope the intestine of his Mais
(Cheetogaster) diaphana, he observes, “ Diese Driis’chen bilden das, was bei
héhern Thieren die Chylusdriisen sind, und ergiessen den Chylus unmittelbar
in den Raum zwischen der musculdsen Haut und dem Darmeanale.”+ This
author, moreover, notes the peculiarity that in a single “ Mutternaide” of
Cheetogaster diastropha, he found in December that the chyle-corpuscles were
larger than usual, and seems to think that there is a connection between chyli-
fication (referring to the perivisceral fluid) and generation. Dr THomas
WILLIAMS,{ again, was strongly of opinion that the long coils of the blood-
vessels anteriorly in his Nais jiliformis (probably Tubi/ex) and the perivisceral
branches elsewhere in the body of this worm, were specially intended for
absorbing from the perivisceral (his “‘chyl-aqueous”) fluid elements by which
the blood-proper is formed and replenished. It will therefore be seen that the
supposition thrown out by my friend Mr LAnxesTEr that “the yellow glandular
tissue” surrounding the alimentary canal “may have but little to do with the
secretion of digestive juices, or, at any rate, may have an additional and most
important connection with the production of the corpuscles of the perivisceral
fluid, and may serve to place that fluid in organic relation with the liquid of the
closed vascular system of the intestine, and the contents of the digestive tract,”
is by no means new. It appears, indeed, to be the result arrived at by Dr
Fritz Ratze from an examination of the literature of the Oligochétes previous
to the appearance of the foregoing ; the author, moreover, assigning the peri-
visceral fluid the function of a communicating medium between the digestive and
circulatory systems. Further, I have not seen anything to support the idea of
Mr LanxesTer that the abundance or scarcity of the “granules” in the perivis-
ceral fluid depends on the condition of the glandular coating of the intestine
and dorsal vessel. The glandular investment presents the same appearance
whether the corpuscles be few or many, and the agglomerations of granules
shown by him most readily take place ‘in such a highly coagulable fluid. For
my part, I have no objection to offer to any of the theories advanced on this
subject, so far as they rest on actual observation and not on mere conjecture.
In Tubifex and its allies the perivisceral fluid is an eminently coagulable and vital

* Zeitsch. f. wiss. Zool. Bd. xix. (1869), p. 614.
t+ Ueber die Nais diaphana, &c., Nova Acta Acad. Leop. Carol. Tom. xiv. Pt. i. p. 411.
{ Report Brit. Assoc. 1851, p. 182; and Philos. Trans. Part ii. 1852, p. 625.

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 259

fluid, and, as has long been supposed, doubtless performs important functions
in the animal economy; as, indeed, the observations of Frey and Leuckart, Dr
QUATREFAGES, and Dr WILLIAMs, show it does in other invertebrate animals.
I think, however, that too little attention has been bestowed upon the in-
herent properties of the fluid itself. Perhaps the remarkable corpuscles
contained therein are the products of such inherent properties, and not
necessarily derived from its surroundings. If the glands covering the intestine
discharge their contents into the perivisceral fluid, as most authors believe, such
a discharge probably only furnishes materials for the evolution of the special
properties of the liquid. It is well to bear in mind, also, that in the clearly
defined group of the Nemerteans, a fluid identical in appearance, coagulable
nature, and in the presence of definite corpuscles, occurs within.a special
muscular sheath on the dorsum of the intestine. This chamber has smooth
walls, and contains, besides, the proboscis, which, as it were, is invaginated
within it. The glandular elements, which exist in vast numbers in the walls of
the digestive tract, cannot thus communicate directly with the fluid.

In the perivisceral space of one example was a curious parasitic larva
(Plate X. fig. 2) which moved backwards and forwards in its chamber. It
lengthened its body into the shape shown at a, then contracted itself as in 6,
forming a club with a large rounded head, and finally assumed the appearance
figured at c; after which it again elongated itself and repeated the same con-
tractions. Its structure was minutely granular, with a streak at the anterior
end. It appears to be the same form as that described subsequently from the
lobule of the testis (p. 265).

Digestive System.—Granular Gilands.—Anteriorly there are some finely
granular glands at the sides of the cesophagus ; and by-and-by numerous larger

glands cover the entire external surface of the alimentary canal, and envelope
_ the dorsal blood-vessel. These are somewhat pedicled structures, consisting of
a fine external membrane containing numerous distinct granules of an orange
or pale brownish hue (Plate X. figs. 3and 4). When these bodies are extruded
into the water, the contained granules show very evident molecular movements,
and in a short time escape by the bursting of the cell-wall, their movements con-
tinuing in the surrounding fluid. The yellowish granules also occasionally
group themselves together, and larger granules are formed here and there, ap-
parently by the union of several. All the granules refract the light very strongly.
The gland-cells are rendered more translucent by acetic acid, which, however,
does not affect the granules. On the addition of sulphuric ether the residue
clearly shows that their composition is of a fatty nature. Dr Bucuuo.z* is of
opinion that the pigment-granules of these cells in Lumbriculus variegatus is a

L * Beitrage zur Anat. der Gattung Enchytreus, &c. Schriften der Physik. Skonom. Gesellschaft
in Konigsberg, 1862, p. 108.

260 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

modification of chlorophyll. This author, moreover, shows a distinct nucleus
in all his figures of the gland-cells of the same worm, and M. CLAPAREDE
describes a nucleus in those of Lumbricus. So long as the cell is filled with the
granules, it would be a very difficult thing to make out a nucleus, and the large
number of nucleated cells from other parts which get mixed up with these in
the field of the microscope necessitates some caution in observing. As pre-
viously stated, I have not succeeded in seeing a nucleus while the granules
were within their investment, nor on watching their extrusion has such been
observed. The same result was arrived at after a careful scrutiny of highly
favourable longitudinal and transverse sections of the alimentary canal of
Tubifex, and after manipulation of the fresh specimens with carmine.

Amongst the sandy mud and Diatomacez in the intestinal canal, are nume-
rous examples of an Opalina (Plate IX. fig. 14, a, 6, c, d). Some are about 5th
of an inch long, and had the various shapes shown in the figures. The species is
probably identical with that found in other minute fresh-water annelids. A few
had a swollen anterior end covered with fine striz like a Pecten or Lima, while
the posterior or caudal portion was filiform. The usual clear granules and-
vesicles were present. An elongated granular structure like a canal was occa-
sionally visible in the central line, but this could not be traced throughout
the entire length of the animal. When freed by laceration of the worm, they
rush through the surrounding water very actively by aid of their cilia, for it is
to be remembered they are but in their native medium, since the intestinal
canal is ciliated, and often gives passage to currents of water. Although a
little glycerine is added to the water, their cilia continue in active motion, and
the contained globules are very distinct.

In a few specimens minute parasitic Nematode worms (Plate IX. fig. 15)
were coiled at the sides of the intestine near its termination. There is a streak
at the snout, and some faint central markings along the body. They appear to
be undeveloped or partially developed Anguillulide, numerous examples of
which are frequently found in Lwmbricus.

Circulatory System.—The following observations on the circulatory system
are in the main confirmatory of the investigations of M. CLAPAREDE, who added
considerably to the descriptions of M. D’UpEKEm. In Tubifex rivulorum the
course of the blood-vessels is very regular, and I have met with very few
abnormal arrangements. There is a large contractile dorsal vessel which is
thrown in the usual state of the animal into many zig-zags or figure of eight
crossings, from the fibrous contractions at the septa. A ventral vessel less in
calibre and paler in colour has a similar direction, but lies on the opposite side
of the alimentary canal. The dorsal trunk, in conjunction with the alimentary
canal, is covered by the gland-cells previously described, while the ventral is
free. The course of the dorsal vessel is as follows :—

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 261

At the posterior end of the worm it is joined by the ventral (in a similar
manner to that shown in Plate X. fig. 5) and proceeds forwards as a deep red
trunk, the depth of its colour being, perhaps, due more to the larger calibre of
the vessel as contrasted with the ventral, than to its thicker coats. In this
region, moreover, it pulsates with a swift and clear stroke from behind forwards
—the action being noticeable in the penultimate segment. The ventral trunk,
on the other hand, so far as I could observe, remained of the same calibre at
this part, except when affected by the wave of perivisceral fluid. In the stasis
following the introduction of chloroform, the dorsal vessel becomes moniliform
posteriorly, being constricted, apparently, by the spasm of the septal fibres ;
while the perivisceral corpuscles rush with great vehemence through the
narrowed apertures. From the colour of the central region of the last segment,
it would seem that, before joining, the ventral and dorsal vessels form a slight
plexus, and, from the vigorous motions usually occurring in this part, there
could be no better region for the aération of the blood.

In each segment two great branches pass off from the dorsal and ventral
vessels respectively. Towards the posterior border a large trunk (the perivis-
ceral) springs on each side from the dorsal, and, proceeding outwards towards -
the body-wall, divides into numerous capillary branches, which again unite to
_ form a trunk, nearly as large as the original, that on each side enters the ventral
vessel. The extensive coils formed by the perivisceral branch of the dorsal
provide ample freedom of motion, an arrangement so necessary during the con-
tortions of the worm. ‘The coils are especially distinct towards the posterior
part of the body. About the middle of each segment, again, the ventral vessel
on each side gives off a branch, which passes upwards round the intestine ; but
whether it terminates by anastomosing with its fellow of the opposite side, or
_by joining the dorsal, could not be determined. Certainly no branch of any
size joined the dorsal in this region.

In some views there are, besides the perivisceral branch of the ventral, one
‘or two vessels towards the anterior part of the segment, which course outwards
from the ventral, and anastomose on the body-wall with branches of the peri-
| visceral. Such a branch or branches are not strictly “intestinal,” for they like-
Wise send twigs to the body-wall. I must alsoadd that in one specimen the
ventral main trunk was observed to bend outwards in a simple curve, without
being fixed in the centre by any vessel or fibrous tissue. The “intestinal”
branch thus does not always attach the main trunk closely to the alimentary
canal. After the addition of a little aconite (which causes spasm of the fibres
at the septa, so as to render the worm moniliform) the dorsal and ventral
vessels become much contracted posteriorly, while the periviscerals remain
well filled, indeed, so as to constitute a broad red bar across the segment.
Anteriorly some of the intestinal branches are similarly distended.

VOL. XXVI. PART II. 3 Y

262 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

In the eighth segment the perivisceral branch is considerably enlarged, and,
curving outwards and downwards, enters the ventral as a large trunk, only
slightly less than at its commencement. The dorsal half of the vessel pulsates;
the ventral does not. It would thus seem that the vessel, probably where fixed
to the body-wall, ceased to pulsate. This arrangement constitutes the so-called
“hearts” of these annelids, and in this species both contracted simultaneously.
The perivisceral branches of the seventh and ninth segments acted similarly,
though in a less conspicuous manner. The perivisceral vessel of the latter
(ninth) segment is often noticed to give off large branches at the body-wall.

In specimens whose generative organs are much developed, the periviscerals
of the ninth and tenth segments are of considerable size, but neither approach
the periviscerals of the eleventh, which are enormously dilated, indeed, nearly
as large as the main trunk itself. This enlarged perivisceral sends branches
over the succeeding segments, sometimes as far backwards as the twenty-third
in ripe animals (Plate X. fig. 6, e, e). The periviscerals of the twelfth segment
under the same circumstances are also dilated, and those of the thirteenth
more so, three large branches being directed forwards. The arrangement of
these trunks would seem to countenance my view of the circulation in the
ordinary condition, viz., that the periviscerals as a rule do not proceed as con-
tinuously cylindrical trunks into the ventral, but that they communicate by
their branches on the body-wall.

Continuing forwards, the main trunks (dorsal and ventral, Plate IX. fig. 17)
have the same arrangements in the fifth and sixth segments as previously de-
scribed, the only noteworthy change being an occasional abnormality in the
origin of the intestinals—one coming off before the other, and thus affording a
better view of their distribution. It is to be observed, also, that the ventral
trunk has in this region faint contractions, which render the vessel pale ; it then
fills again. ‘The periviscerals of the fourth segment are somewhat smaller than
usual. At a point corresponding to the posterior border of the third segment,
the dorsal gives off two large trunks, doubtless the homologues of the perivis-
cerals, though they generally slant obliquely forwards and outwards rather
than transversely, and divide into many small branches towards the margin of
the body. The dorsal then pursues its course straight forwards, again gives
off two branches behind the mouth—the branches subdividing and inosculating
with others in front and behind, and finally terminates by forking in the snout
in front of the mouth. The two divisions thus formed split, after bending
backwards, into numerous twigs, which unite with the capillaries that go to
form the feeders of the main ventral vessel. The latter originates, by the
junction of the two great branches constituted by the feeders just mentioned,
towards the posterior border of the third segment. In many positions under
pressure, the whole anterior part of the animal is one vast series of vascular

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 263

reticulations. The latter quite differ from the long simple coils described and
figured by Dr WituiAMs* in Nais jiliformis, but doubtless he was misled by
their complexity.

In some views, where congestion had been produced by the addition of
chloroform, atropine, or muriate of morphia,t the intestine was observed to
be covered by an extensive series of minute blood-vessels, longitudinal and
circular. This arrangement was due to the presence of two or three vessels in
each segment winding round the canal, and sending off lateral branches to meet
others from the adjoining trunks, as shown on the supero-lateral surface of
the intestine in Plate X. fig. 7, the ventral vessel in this case not being seen.
A series of nearly parallel anastomosing branches course from the secondary
trunk in a longitudinal direction. A lateral view of the seventh segment after
the addition of chloroform (Plate X. fig. 8) exhibits a much coarser reticulation,
in which the main trunks arise from the ventral. In such experiments, of
course, the trunks do not remain of their normal calibre, but are irregularly con-
tracted. These statements with regard to the vascular ramifications on the
surface of the intestine are fully borne out in the transverse and longitudinal
sections of the worm, the former exhibiting a complete mesh-work of blood-
vessels surrounding the alimentary canal in certain positions. The same has
been noticed by M. Perrier{t in Dero obtusa, one of the Nais-group, and he
aptly compares the arrangement to a very elegant trellis with rectangular open-
ings. According to Mr LAnxKester they would appear to be more easily
observed in Chetogaster. Under the action of chloroform, also, many fine
cutaneous branches were seen in Zubifex forming a network and a series
of parallel longitudinal vessels. These ramifications remained comparatively
still during the motions of the worm, and were probably fixed branches of the
perivisceral. Some of the twigs (which extended over most of the body-wall)
coursed towards the septa, and inosculated with the same set of vessels in the
adjoining segments. There is thus a series of vascular communications between
Segment and segment independently of the main trunk. It would also appear
that some of the branches, which proceed from the dorsal towards the ventral,
do not join the latter trunk, but inosculate with twigs from the opposite side of
the body.

The circulatory system in the elongated form (Plate X. fig. 9,) much
resembles that of the foregoing, though it is proportionally more developed.
The swollen periviscerals or “hearts” occur in the eighth segment, and
pulsate vigorously and alternately from eight to twelve times per minute. Very
slight contraction of the tissues anteriorly causes the mobile vessels to assume

* Report Brit. Assoc. p. 182, Pl. III. fig. 8.
} The addition of a small quantity of this drug in solution was speedily fatal to ciliary action.
¢ Ann. Nat. Hist. 4th Se. vol. 6, p. 264 (Extr. Comptes Rendus).

264 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

so many curves and spirals that it is impossible to unravel them, (wde Plate
IX. fig. 16), until a more favourable condition of extension ensues. It is a fact
of interest, that while the vascular distribution is more apparent in this species,
the perivisceral corpuscles are less developed.

Generative Organs.—The first swollen segmental organ in the shorter form
(Tubifex rivulorum), as M. CLAPAREDE observes, occurs in the eighth segment.
In one specimen the dilated portion of the organ was at the anterior border,
and on the same side the coils of the duct had entered the seventh; while on
the other side neither coils nor dilated portions were visible in either seventh
or eighth segments, but both were present on that side in the ninth. Consider-
able room, therefore, exists for misunderstandings. The segmental organs
(Plate IX, fig. 18, from behind the middle region,) vary a little in shape, and
some in developed specimens are tinted brownish by transmitted light. The
shorter tube (a) is attached to the septum in front, opening through the mem-
brane by a slightly dilated and ciliated opening. The long coil, again, opens
externally, also by a very slightly enlarged extremity, at a point a little posterior
to the former, but in the same segment.

The sexual pore lies a short way behind the ventral bristles of the eleventh
segment, and has the form of a conical papilla (Plate X. fig. 10), which is per-
forated at the summit. Occasionally spermatozoa are observed to issue from
the tip. In this instance the copulating organ is slightly protruded.

The integuments, as already noted, from the tenth to the fifteenth segment
become at the reproductive season very opaque, and hence the difficulty in
making an accurate description is much increased. The tenth and eleventh
segments especially swell out, and become opaque white at the period of
perfection.

Male Organs.—In those with undeveloped (or only slightly developed)
generative organs the testicles are found at the anterior border of the tenth
segment, and the segmental organ in the eleventh is small, showing that it only
becomes enlarged with the other structures subsequently. Under the same
circumstances coils of the ciliated duct of the segmental organ are found at the
posterior border of the twelfth segment. At a further stage of development
the testicles form large opaque-whitish masses, which are at first granular
(Plate X. fig. 19). What appears to be a further stage is shown in fig. 20,
Plate IX., the cells being filled with a vast number of awl-shaped bodies,
measuring 73!55 of an inch long, rounded at one end, and having a slender style
at the other. There are also numerous minute ovoid granules in the cell, their
long diameter ranging from 5,95 tO gogo Of an inch. The bodies represented
in fig. 6, b, c, d, Plate IX., are probably also stages in the development of these
structures.

In their fully developed state the spermatozoa resemble wavy or zig-zag

Ba. -

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 265

lines (Plate IX. fig. 21), sometimes with attached globules or loops resembling
heads, but more frequently without them. They do not swim actively about
on escaping through a wound, but spread themselves insensibly over the field
of the microscope. They often, as M. p’UDEKEm shows, surround a sperm-
cell so completely as to resemble a seed with its downy pappus.

The first testicle, in those with developed organs, occupies to a greater
or less degree one side of the ninth segment; and occasionally it is little
developed while the second stretches to the sixteenth segment. I have also
seen the first testicle slip entirely out of the ninth segment, and lie towards
the posterior part of the tenth. It is attached to the septum between the
ninth and tenth segments, in the angle next the intestine (in ordinary views,
¢, fig. 6, Plate X.) In a large specimen there was the unusual appearance of a
glandular organ resembling a testis with sperm-cells in the fifth segment, and
the seventh and eighth had each two of a similar nature. The ninth had the
vas deferens with its trumpet-shaped aperture fixed in the septum between it
and the eighth. The developed organ (testis), moreover, stretched from the
bulged septum last-mentioned to the fifteenth segment. The ordinary condition,
however, is that the first testis occurs in the ninth segment, the receptacles in
the tenth, the ovaries in the eleventh, and the second testis in the twelfth
segment.

The eleventh segment also holds the large coils of the vas deferens (Plate
X. fig. 6, 6), which, moreover, often slip into the twelfth. The trumpet-shaped
aperture is connected with the septum between the tenth and eleventh segments.
It (vas deferens) is clearly a development of an ordinary segmental organ, as
indeed most authors state.

In the sixteenth segment of one example there was a large parasite (Plate X.
fig. 11) in the lobule of the testis, and extending throughout the entire length
of the division. Its interior was filled with cellulo-granular matter, and in
contraction its sides were distinctly crenated, while the body was crossed by
transverse ruge, like a larval cestode. There is a short median furrow passing
from a notch in its anterior or smaller end. It is not ciliated.

Female Organs.—In the early condition the ovaries are observed at the
anterior border of the eleventh segment, attached on each side to the septum,
close to the dorsal vessel and intestine (Plate X. fig. 12, a, 6). In this state
they are composed of granular cells. The developed organs (sometimes of an
orange colour) stretch to the fourteenth, fifteenth, and subsequent segments,
and when compressed give exit to a vast mass of fatty granules.

The seminal receptacles (Plate X. fig. 6, @) are amongst the most distinctly
marked organs in the developed animal, and are at once distinguished by the
lively contractions which ensue when they are filled with spermatozoa. They
occupy nearly the whole of the tenth segment, and the apertures have the form

VOL, XXVI. PART II. 3 Z

266 DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX.

of trumpet-shaped organs, one on each side anteriorly. When empty the sacs
have a somewhat coarse granular appearance, and do not show the rolling
contractions.

In a specimen with largely developed ovaries, a curious ovoid organ (Plate
X. fig. 13) occurred at the anterior part of the eleventh segment, and another
at the anterior part of the twelfth. Externally there was a dense capsule, and
internally a minutely granular mass altogether different in appearance from the
structure of the ovaries or their contents. They measured about 5th to 735th
of an inch in diameter, and were of so unyielding a nature that they soon
escaped by rupture through the body-wall of the worm.

EXPLANATION OF THE PLATES.

Puate 1X.

Figure 1. Bristle and dorsal hook of Tubifex rivulorum, from the anterior segments. x 350 diam.

Figure 2. Ventral hooks of the same species. x 350 diam.

Figure 3. Anterior hooks (accompanyingthe bristles) of the elongated form from the lakes; a, pro-
cesses in the fork. x 350 diam.

Figure 4. Body-wall (a) of T. rivulorum, with parasitic Vorticelle. x 210 diam.

Figure 5. One of the parasitic Vorticelle in a partially expanded state. x 350 diam.

Figure 6. Perivisceral corpuscles of T. rivulorum ; a, cells in the ordinary condition ; 6, a cell from
the perivisceral chamber after the addition of chloroform; ¢ and d, the contents of the
latter more highly magnified. In all probability, however, this (b) is only a sperm-cell in
course of development. x 350 diam.

Figure 7. The same after the action of dilute glycerine, with extruded clear granules. x 350

diam.

Figure 8. Variously shaped corpuscles from the perivisceral fluid. x 350 diam.

Figure 9. Perivisceral corpuscles of the elongated Tubifex from the lakes. x 280 diam.

Figure 10. Elliptical and other corpuscles from the same fluid. x 350 diam.

Figure 11. Perivisceral corpuscles of a species referable to CLAPAREDE’S genus Limnodrilus. x 350
diam.

Figure 12. Gland-cells from the intestinal wall of the same species. x 350 diam.

Figure 13. Ciliated epithelial cell from the interior of the digestive tract of the same animal. x 350
diam.

Figure 14. a, b, c, d. Various forms of the Opalina parasitic in the alimentary chamber; 0 represents
an outline of the anterior end of a large specimen, the cilia being omitted. x 210 diam.

Figure 15. Parasitic Nematode worm from the same region. x 400 diam.

Figure 16. Anterior region of the elongated Tubifex from the lakes, showing the dense coiling of the
blood-vessels anteriorly in the semi-contracted condition; a, dorsal blood-vessel; d,
ventral.

Figure 17. Anterior segments of Tubifex rivulerwm in a somewhat contracted and flattened condition,
exhibiting the arrangement of the ,vascular system; a, fissure at the mouth; d, dorsal
blood-vessel ; v, ventral. The figures indicate the segments.

Figure 18. Segmental organ of the same species from behind the middle region of the body ; a, the
septum.

Figure 19. Sperm-cell in the granular stage. x 350 diam.

Figure 20. Awl-shaped bodies and granules from a sperm-cell in course of development. x £05
diam.

Figure 21. Spermatozoa of 7. rivulorum. x 350 diam.

Figure
Figure
Figure

Figure
Figure

Figure

Figure 10
Figure 11
Figure 12

Figure 13

DR M‘INTOSH ON SOME POINTS IN THE STRUCTURE OF TUBIFEX. 267

Puate X.

_ Perivisceral corpuscles from Tubifec rivulorum after extrusion into the water. x 210

diam.

. a, b, c, Various forms assumed by the parasitic larva from the perivisceral chamber of

Tubifex. x 210 diam.

. Granular glands from the wall of the digestive cavity in situ. x 350 diam.
. Isolated gland-cells similarly magnified.
. Posterior end of the Tubifex from the lakes, showing the junction of the dorsal and ventral

blood-vessels ; a, cuticle ; 6, chorion and muscular coats ; c, ending of dorsal blood-vessel ;
c’, last perivisceral ; d, perivisceral and other corpuscles. x about 50 diam.

. Portion of Tubifex rivulorum at the reproductive season ; a, seminal receptacle in the tenth

segment ; 5, coils of ciliated duct (vas deferens) ; c, testicle ; d, atrium (?) with a double
outline under pressure ; e, ¢, elongated vascular branches from the eleventh and other
segments for the supply of the developed generative products. x 55 diam.

7. Vascular ramifications on the supero-lateral surface of the intestine ; a, dorsal blood-vessel ; 0,

septum. x 210 diam.

8. A lateral view of the seventh segment after the addition of chloroform ; d, dorsal vessel ; v,

ventral ; p, perivisceral. x 210 diam.

9. Anterior portion of the elongated form from the lakes, showing the circulating system; a

dorsal blood-vessel ; 6, ventral; c, enlarged pulsating cavities or “ hearts” of the peri-
viscerals in action ; d, forking of ventral vessel.

. Tip of the copulating organ protruding through the sexual pore; a, hooks. x 210 diam.
. Larval parasite from the lobule of the testis. x 210 diam.

The eleventh segment, showing the development of the ovaries; a, cellulo-granular con-
tents ; 6, investing membrane ; ¢, perivisceral corpuscles ; d, granular glands of intestine.

' x 210 diam.

Curious ovoid structure from the anterior part of the eleventh segment of a specimen with
largely developed ovaries. x 210 diam.

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( 269 )

XIIL—On the Place and Power of Accent in Language. By Professor BLACKIE.

(Read 6th March 1871.)

On the subject of accent and quantity as elements of human speech, there has
been such an immense amount of confusion, arising from vague phraseology,
that in renewing the discussion nothing seems more necessary than to start
with a careful and accurate definition of terms; and that a definition not taken
from books, and the dumb bearers of a dead tradition, but from the living facts
of nature, and the permanent qualities belonging to articulated breath. Now,
if we observe accurately the natural and necessary affections of words in human
discourse, considered merely as a succession of compact little wholes of arti-
culated breath, without regard to their signification, we shall find that all the
affections of which they are capable amount to four. Either (1), the mass
of articulated breath which we calla word, is sent forth in a comparatively
small volume, as in the case of a common gun, or it is sent forth in large
volume, as in the case of a Lancaster gun ; this is a mere affair of bulk, in virtue
of which alone it is manifest that any word rolled forth from the lungs of a
Stentor must be a different thing from the same mass of sound emitted from
a less capacious bellows. In common language this difference is marked by
the words /oud and ow. A broader wave of air impelled against the acoustic
machinery of the ear will always make a more powerful impression independent
of any other consideration. But (2), an equal or a stronger impression may
be made on the ear by a less volume of sound, if it be sent forth with such
an amount of concentrated energy and force as to compensate for its deficiency
in mass. A more sharp and intense clap of thunder, for instance, may in this
way affect the ear more powerfully than a greater peal less vigorously sent
forth and more widely spread. The affection of sound brought into action here
is what in language we generally call stress or emphasis; and it depends altogether
on the intensity of the projectile force, and gives to speech the qualification of
more or less forcible. But (3), this force may often be, and very naturally is,
accompanied with another affection of sound altogether distinct, viz., the
sound may be deep and grave, or high and sharp, corresponding to what in
music we call bass and treble notes. The analogy between music and articulate
speech is here so striking, that it has passed into common use ; as when we talk
of a person speaking in a high or a low key, in a monotone, or in a deep low
sepulchral tone, and so forth. And in reference to single words, we are

VOL. XXVI. PART II. 4A

270 PROFESSOR BLACKIE ON THE

accustomed to say, that the acute accent stands on syllables pronounced in an
elevated tone, and the grave on those pronounced with a low tone. The only
difference between the musical scale and the scale of articulate speech in this
view, is that the latter, besides being much narrower in its compass, rises or
sinks, not by mathematically calculated intervals, but by a mere upward or
downward slide, not divided by any definite intervals. The true connection of
these slides with the general doctrine of accent has been well set forth by Mr
Watker, the author of the Pronouncing Dictionary, in a separate treatise.*
(4). The fourth affection of articulated sound is that which is familiarly known
to scholars and schoolboys under the name of quantity, and signifies simply the
greater or less duration of time during which the sound continues to impress
the ear. For it is manifest that any sound may be produced either by a sudden
stroke, or jerk, or by a traction prolonged to any extent. In grammar a short
vowel corresponds to a quaver or semiquaver in music, and a long vowel to a
crotchet or minim, according to the ratio of the movement.

Now it should seem to be pretty plain at the outset, to all persons whose ears
have been exercised in a very slight degree to discern the differences of articulate
sounds, that what is called accent in grammar has to do only with the second
and third of the above four elements, and not at all with the first or fourth ; in
other words, that the accent of a word is totally distinct both from the volume
of voice with which the word is enunciated, and the length of time during which
the speaker dwells on the syllable. Nevertheless, such is the confusion which
learned writers have introduced into this subject, that it is necessary at the very
outset to enter a caveat against a very prevalent notion that the placing of the
acute accent on a syllable, naturally or necessarily implies a prolongation of
the sound of the accented vowel; or, in other words, that to accent a syllable
withoutmaking it long is impossible. In music no performer ever dreams that
the rhythmical beat on the first, we shall say, of three quavers—that is jig time—
necessarily turns the quaver into a crotchet. A musician making such an
assertion would simply be deemed drunk or mad ; nor does it make the slightest
difference in the quantity of the note receiving the musical accent, whether in
respect of elevation of tone it stands high or low in the scale. It is understood
by every girl who fingers the piano, that the elevation of the note, the duration
of the note, and the rhythmical emphasis upon the note, are three essentially
different things which never interfere with one another. But the moment we
transfer this case to the analogous domain of spoken accent,—the certus quidam
dicendi cantus, as CICERO called it,—we find ourselves involved in a region of con-
fusion and contradiction with regard to the simplest matters, than which few
things can be imagined more humiliating to human reason. For however diver-

* A Key to the Classical Pronunciation of Greek and Latin Proper Names; with Observations
on the Greek and Latin Accent and Quantity. By Joun Watker. London, 1827.

PLACE AND POWER OF ACCENT IN LANGUAGE. Di

gent the printed opinions of the learned may sound, that the relative facts are
exactly the same in the case of spoken speech, as of song or played notes, is
beyond question. A single example will make this evident. The first syllable
of po’-tent, for instance, according to a well-known rule in the English language
is long ; but the first syllable of the Latin word from which the English comes
is short, pot’-ens, while the accent is on the same syllable in-both languages.
Now, it surely will not be alleged, in obedience to the dictates of any sane ear,
that in pronouncing the Latin word I am obliged to call it pd’-tens, after the
English fashion, on account of the tyrannic force of the acute accent. It seems,
nevertheless, that British schoolmasters and professors have acted under the
notion that some compulsion of this kind exists ; for as a rule they say b0’-nus,
and not bd6n’-us, though they know very well that the first syllable of this word is
not long, as in the English word pd’-tion, but short, as in m67’-al. Such confusion
of ideas on a very simple matter is a phenomenon so strange, that some reason
may justly be demanded for its existence ; and on reflection I find two reasons
principally that seem to account for it. The first is the confounding of a really
long quantity with that predominance of a sound to the ear which is a necessary
element of all accentuation. Thus, when I take the word ¢ép’-id, and form the
abstract substantive from it—iepid’-ity, by changing the place of the accent from
the first syllable of the adjective to the second, I certainly have given a pro-
minence to the short 2 which it did not possess before, and a prominence, no
doubt, which though it consists principally in force, emphasis, or stress, may
also carry along with it a certain dilatation of the tenuous vowel, so that it is
really longer in the substantive, being accented, than when it was slurred over
without emphasis in the adjective. But though this is quite true, it is altogether
false to say that the vowel has been made long according to the comparative
value of prosodial quantity ; for, if the second syllable of tepi’d-ity be compared,
not with the last syllable of the adjective zepid, but with the same syllable of
the substantive, as mispronounced by some slow, deliberate Scot—tepi-dity,
tepé e-dity—we shall see that the vowel #, for all rhythmical purposes, still remains
short. The other cause which presents itself to explain the confusion of
English ears on this subject, is the doctrine of what the Greek and Roman
grammarians call length by position. According to this doctrine, a vowel
before two consonants is long. What this means we may clearly conceive by
the example of such words as géld, ghost, in English, or Pabst or Obst in German ;
but though the vowels in these words are unquestionably long in both
languages, they are so only exceptionally, the rule both in English and German
being that a vowel before two consonants is short. Of this rule the word
short itself may be taken as an excellent example; which, if it occurred in a
Greek chorus, by the law of position, would be sung shdrt, with the o prolonged,
like 0 in shore. Now, with this classical analogy in their ears, or rather in their

22, PROFESSOR BLACKIE ON THE

head (for it is by no means certain that all those authors who have written on
this subject did use their ears), when I pronounce such a word as prim’-rése or
él’-bow, it is not at all uncommon for English scholars to say, and obstinately to
insist, that the accent on the first syllable of these words is necessarily accom-
panied by a prolongation of the vowel. But this is a judgment of the question,
not by the living fact of the sound, but by the doctrine of an old book about
the sound. And as to what the old book says, we in fact do not know whether
length by position meant a habitual prolongation of the vowel sound in common
discourse, as in our words gold, told, sold, ghost, most, or only a poetical license ;
that is to say, whether the genitive plural of avjp, of which the penult is short,
was really pronounced awndré’ne or dndrone in prose. I for one am strongly
inclined to think that the latter is the true fact of the case; for, if it had been
otherwise, would it not have been a more correct phraseology in the grammarian
to say, that a vowel before two consonants is naturally long? But when they
tell us that a vowel which is naturally short becomes long when two consonants
follow, this looks rather like an artificial exception than a natural rule. And I
am inclined to think that such an exceptional rule was introduced from sheer
necessity, like the long o in certain comparatives, such as cofdrepos, because,
without such a license, really long syllables in sufficient abundance would not
have been found in the language for the necessities of the early dactylico-
spondaic poetry. As to any inherent natural necessity in the rule, such an idea
cannot be entertained for a moment; for the vowel is then most easily pro-
longed—as in the English words pé’-tent, no’-tion, na’-tion, pd’-tent, where it is kept
separate in spelling from the influence of the succeeding consonant or con-
sonants, which, as in po7’-tion, rather act by cutting the breath short, and pre-
venting the prolongation of the vowel. The influence of the consonant in
shortening the vowel will be apparent in comparing the words ném’-cnal and
Leéb’-anon with nd’-tional and ld’-bial; nor does the addition of a second consonant
in any perceptible way alter the case. If the first syllable in prim is manifestly
short, it is certainly not made long by the addition of the long syllable rose in
the noun prim'rése—a word which, in the relative values of its final and penult
syllables, corresponds exactly to a whole host of Greek words which usher in a
long final by a short accented penult, as in I]\dérwv, the name of the great
philosopher of Idealism, in Anglicising which, we, besides attenuating the
vowel, elongate the short penult, according to the practice of our own language.

It will now be distinctly understood, as a starting-point to the present
inquiry, that by accent I mean merely a certain predominance, emphasis, or stress
given to one syllable of a word above another, in virtue of a certain greater
imtensity of force in the articulated breath ; this increased intensity being natu-
rally in many cases, but not necessarily in all cases, accompanied by an elevation
in the key of the voice. My observations do not include either rhetorical accent,

PLACE AND POWER OF ACCENT IN LANGUAGE. 273

which affects whole sentences and clauses, or national accent, which, in addition
to rhetorical accent, often includes some favourite sound, note, or vocal man-
nerism characteristic of different peoples.

The general question to which we shall now attempt a scientific answer is
the following—What are the great leading principles on which accent, as a
phenomenon of articulate speech, depends? Are there any such principles, or is
it a matter of mere arbitrary association, fashion, and habit? and in the com-
parison of different languages what is the standard of value in respect of their
accentual character? Does esthetical science contain any general rules which
might enable us to measure the value of accents, as we do the value of sounds in
language, when, for instance, we say that Italian is a more harmonious language
than Gaelic, and Greek a more euphonious language than Latin? In answering
this question, I would remark, in the first place, that there is no such thing as
a language altogether without accent; only a machine could produce a con-
tinuous series of sounds in undistinguished monotonous repetitions like the tim,
tim, tim of a drum; a rational being using words for a rational purpose to
manifest his thoughts and feelings, necessarily accents both words and sentences
in some way or other. When, therefore, we find it stated in ApAmM SmIrTH’s
Essay on Language, and other English writers, that the French have no accent
in their words, this is either a gross mistake, or it must be understood to mean
that the French do not give such a decided and marked preponderance to one
syllable of the word as the English do ; which is very true, as any man may see in
comparing the English velocity with the French velocité. But this is merely a
difference in the quantity and quality of accent, not a contrast betwixt accent
and no accent. The second postulate of all rational discussion on this subject
is, that the significant utterance of articulate breath, like every other mani-
festation of reason-moulded sense, is a part of esthetical science, and subject
to the same necessary laws which determine the excellence of a picture, a poem,
or a piece of music. No doubt in the enunciation of words, as in all the fine
arts, fashion may often prevail to such an extent, as in some cases to usurp the
place of reason and propriety; but the prevalence of false taste in any depart-
ment of art does not effect the certainty of the eternal principles by which it is
regulated, any more than the prevalence of murders or lies amongst any people
can take away from the essential superiority of love to hatred, and of truth to
falsehood in all societies of reasonable beings. We are, therefore, justly entitled
to look for a standard of excellence in the matter of orthoepy, no less certain
than the standard of truth in morals or mathematics ; as, indeed, all things in
the world being either directly or indirectly the necessary eftluence of the
Divine reason, must, in their first roots and foundations, be equally rational and
equally necessary. Now, in looking for the necessary conditions on which the

VOL. XXVI. PART II. 4B

274 PROFESSOR BLACKIE ON THE

comparative excellence of accentual systems may depend, we find that they may
be reduced to the four following heads :—

1. Significance. 3. Variety.
2. Euphony. 4. Convenience.

And first, that SIGNIFICANCE must be a main point in all accentual systems,
is manifest from the very nature of accent. For why should a man give pre-
dominance to one syllable in a word more than to another, unless that he means
to call special attention to the significance of that syllable? Nay, it may often
be essential to the effect intended to be produced by the word, that its most sig-
nificant syllable should be emphasised—as when Lord Derby lately said that the
adoption of the Prussian system of making every citizen a soldier, would not
be a progression but a 7rétrogression. No doubt, in order to express such an
accentual contrast as this, the English language departs from its usual fashion
of accenting these words; but this only proves that the English method of
accentuation in this case is a mere fashion, founded on no natural law, and
which accordingly must yield to the higher law of emphatic significance, when .
nature, like murder, will out. And here we may observe that the English, as a
merely derivative and mixed language, is by no means a favourable one for ex-
hibiting the natural and normal laws of a rational accentuation. Neither, so
far as I know, is there any language whose orthoepy presents so many anomalies,
and where changes entirely reasonless and arbitrary, require only the stamp of
aristocratic or academic whim to give them currency. With regard, however,
to the natural preponderance of the contrasting element in compound words,
the Saxon part of our language affords obvious examples of its recognition, as,
when we say, out’-side and in’-side, back'-wards and jor’-wards, up'-hill and down’-
hill, male and fémale. So in the names of the Highland clans, as MacBain,
MacDonald, MacG'rigor, &c., the emphasis does not lie on the common element,
the Mac, but on the distinctive element to which the other is attached ; and in this
view our Saxon pronunciation of MacIntosh and MacIntyre, affords two very
good examples of words where custom and fashion have inverted the natural
and significant place of the accent. Im the Greek language, this most natural
of all accentual laws, operates in all such compounds, as dékapzos, drais, ctvo8os,
mapooos, with which we may contrast the English /rwitless, childless, where the
accent is on the root, and not where it ought naturally to be on the contrasting
element of the compound. In the same category with this I am inclined to
place the accent on the augment in Greek, as in erua, rérv~par; for it is the
augment here manifestly that contains the element of past time which is dis-
tinctive of the tense, bemg equivalent in effect—whatever its original meaning
may have been—to J vip strike, as opposed to J am striking. The same desire
to call attention to the distinctive element may have determined the Greeks to

PLACE AND POWER OF ACCENT IN LANGUAGE. ano

accent the penult of all diminutives, contrary to their usual practice in words,
with a short final syllable, as in wadiov, wadioxos, K.7.X.

Under this head I am sorry to record my dissent from a German writer of
acknowledged excellence on this subject—Dr Karu Goértiine.* This learned
writer lays down the maxim in the first place, that im the Greek language the
accent falls on the syllable containing the principal idea of the word; and,
accordingly, he says that in \€yw and other verbs not pure it falls on the penult,
because this syllable is the root, and the root, as containing the principal idea of
the word, is naturally emphasized. Now, looking back to the first framers of a
language, I cannot see in this case any reason why the root syllable should have
received the accent rather than the termination, which, for the sake of distinc-
tion and contrast, is added to the root. If we say dkapzos, because we wish to
call attention to the negative particle, why should we not say eyo calling
attention to the personal pronoun ; as, in fact, we do say in English, quoth J’,
quoth H&% And in the same way with regard to nouns, as the terminations of
the cases were originally expressions of relation, attached to the noun for the
sake of emphasis and contrast, I do not see why the schoolboy fashion of declining
dominis-t-o’-im—should not have been the original one. And so in the case
of the German brauerti and the Scotch brewerthe as contrasted with the English
bréwery ; for though no doubt it may be said, that as the root brew contains the

principal idea, the accent should naturally be there, and this is what GorTTLING
says, yet it may with more right be said, that what is intended to be emphasized
here is not mere brewing, but a place for brewing, and that the syllable denoting
the place receives the accent as appropriately as the terminations ypuov, eiov,
and #v, when used for the same purpose in Greek. Only so much truth,
therefore, can I perceive to lie in GoETTLING’s principle, as to admit that, so soon
as the original signification of terminations is lost, and people commence to
supply their place by prepositions, pronouns, and other separate words, whose
significance is felt—then, and not till then, can the accent on the root syllable be
regarded as natural and normal in language. Thus, when the German says

__ Habe, laying the stress on the first syllable of the first person singular present

indicative of the verb to have, it is natural and normal, because the termina-
tion ¢ has no significance to him, and could receive an accent only from
a senseless fashion, not from a natural propriety. On the other hand, in
A’‘bgabe, Hingdbe, Zigabe, and similar compounds, the accent is properly placed
on the contrasting element of the compound, of which the significance is strongly
felt.

The next element we have to take into consideration in measuring the value
of different accentual systems is EupHony. The simple mention of this word
will suffice to show how very one-sided a notion it was in Gorrr.ine, that the

* Elements of Greek Accentuation, from the German. London: Wuiraker. 1831.

276 PROFESSOR BLACKIE ON THE

accent, as a general principle, should always be on the root syllable, as being
the most significant. If man were only a logical animal, this might be all very
well as an @ priori ideal of a perfect accentual system ; but he is also, if not
always at starting, certainly when fairly developed, an esthetical animal, who
may be allowed on occasions to sacrifice the significance of ideas to the luxury
of sounds. And if this is true of man generally, it is certainly so a@ fortiori of
the Greeks, whose whole culture grew out of music, and remained in the closest
connection with it to the very end of their classical period. Supposing,
therefore, that with this most musical and artistic of all peoples a regard to the
mere luxury of sound had, in certain cases, determined the position of the accent,
let us ask in what way this determination would naturally manifest itself? The
answer is obvious. In richly terminational languages such as the Greek, where
the terminations are not insignificant little short vowels or syllables as in the
German Gabe, Buche, Briider, &c., but deep, full-rolling, prolonged vowel-
syllables such as wy, ows, ao, awv, and oo, there might exist a very natural
tendency to place the accent on these syllables,—not, of course, because there is
any necessary connection, aS some persons say, between accenting a syllable
and lengthening it, but because when a syllable by the presence of a long vowel
actually is long, the placing of the accent on it, is the most certain way both to
bring out the full length of the vowel, and to ensure the permanence of the full
musical value of the syllable, so long as the language lasts. For whatever
other syllables of a word may from carelessness, or haste, or reasonless fashion,
be cheated of their natural quantity, the accented syllable will always most
stoutly maintain its rights, even if it be a short syllable, much more if it be
along. To illustrate this by a familiar example ; in the famous Homeric line
(Il. i. 49), in which the twang of Apollo’s bow is described :—

“Sewn S€ khayyn yever apyvpéo.o B.010,”

it is manifest both that the euphony of the line lies mainly in the two termi-
nations in o.o, though these syllables are certainly not the significant ones in the
verse ; and further, that this verse is much more beautiful when recited with the
rhythmical accent on both the full-sounding penults, than when, according to
the prose accentuation, it emphasizes only the ot of the last word. The coinci-
dence of the termination with the accent therefore is favourable to music ; and
it is favourable also as a bar to the injury which time is always ready to inflict
on final unaccented syllables. Now, with this principle to guide us, we shall have
no difficulty in seeing the cause of one peculiar excellence which the ancient
Roman critics recognised in the Greek, as contrasted with their own tongue, in
respect of the accentual system. For, as the Romans in no word placed the
accent on the last syllable, it followed that they could enjoy the rich auricular
luxury of a grand terminational unison of accent and quantity, only in the case

PLACE AND POWER OF ACCENT IN LANGUAGE. HT a

of words whose terminations are dissyllabic. Thus, they dealt largely in final
trochees—trochees both by accent and quantity, in such words as sermd‘nis,
pennarum, domino’rum, legis, proba’ vit, voluptatem, and so forth, but could not
say domino’s, or Macend’s, or any word accented in the same way as in English
our enginéer, voluntéer, evdde, capsize, theori’se. On the other hand, the Greek
terminational accent is pretty equally divided between trochaic terminations,
such as oto, diovan, tupGeica, pvOos, caua, waddov, and oxytone endings, such
as ayalav, haBav, tupbeis, pureis. Of the prevalence of the oxytone accent in
Greek, especially in large groups of adjectives and substantives, not to mention
the whole army of prepositions, and certain familiar parts of verbs, any one may
convince himself by taking a sentence at random from a Greek book ; and the
effect of this on the music of the sentence will be evident to the dullest ear.
Sometimes a whole sentence runs on with a succession of accented terminational
syllables, a peculiarity which, without any rhetorical intention, arises naturally
from the number of oxytone substantives and adjectives, and the additional
fact that all substantives of the first declension, whatever the accent of their
termination may be, receive a long rolling accent on the last syllable of the
genitive plural, while all monosyllables of the third declension, by a law common
both to Greek and Sanscrit, transfer the accent from the radical syllable to
the termination in the genitive and dative cases of both numbers. Take a
passage from Plato’s Republic as an example :—

“OU re Onpevtal wavres, ot TE pysnTat, TodAoL pev Ob TEPL TA OXHpaTA TE Kal
Xpopara, modol S€ of TEpl povotKyV, TonTal TE Kai TOVTwY VaNpETaL, Parwdol,
UroKpiTal, xopevTat, épyohdBou, cKevav TE TavTOdaTav Synmuoupyol, TAY TE GANoV Kal
TOV TEPL TOV yuvatKEtoy KdopoV, Kal 67 Kal SiaKdvey TreLdvav Senoduela. 7 od SoKEt
Seqoew TaWaywyar, TiTAGY, TpoPaV, KOMPwTPLOY, KOUPEwV, Kal ad dioToL@Y TE Kal
payeipwr ; eri € Kal cvBwrav tpoadenodpucBa.”*

Greek, therefore, is superior to Latin in this respect, just as an instrument
with a larger is superior to one with a smaller compass of notes. And taking
Italian, under this point of view, into the comparison, we observe that the few
oxytone accents which that beautiful language possesses all arise out of Latin
words, with an accented penult, whose last syllable has fallen away ; thus,
podesté from potestate, amé from amavit, and so forth. The same is the case
with the French, as in velocité, varieié, valeir ; and most of our English oxy-
tones, whether Latin or Greek, are merely curtailed forms of a final trochaic
accent, as evdde from evddo, volunteér from volontiére, proceed from procédo,
theorise from Oewpitw. And it is this systematic curtailment by the way, caused
by the dropping of the final unaccented vowel both in Latin and Saxon words,
which has produced that lamentable deficiency in trochaic endings which makes
our rhythmical language so much narrower in compass than that of Greek,

* Rep. ii. 373, B.
VOL. XXVI. PART II. 4c

278 PROFESSOR BLACKIE ON THE

Latin, German, and Italian. Only for short lyrical efforts can we manage the
rhymed trochaic ending with graceful effect ; all attempts to go beyond this
natural limit have ended either in a manifest artificial strain, or an admixture
of the comic element which is fatal to the effect of serious composition.*

If this rich and various disposition of the accent on terminational syllables
is thus manifestly a plain element of euphony, that accentuation, on the other
hand, will be justly esteemed cacophonous which, by drawing the accent back
to the beginning of the word, that is to the third, fourth, or even fifth syllable
from the end, has a tendency to cheat the ultimate or penultimate syllable of
its full musical value ; we say a tendency, because it is only in this tendency that
the evil lies ; for, if by careful elocution the tendency is corrected, the blot may
be turned into a beauty on a principle to be mentioned under the next head.

The remark here made is a very serious consideration for us English, as our
predominant accent is decidedly antepenultimate, and the fashion seems to be
increasing of throwing back the accent from the penult to the antepenult, and
from the antepenult sometimes to the fourth syllable from the end. Thus we
used to say contem'plate and illistrate, whereas we say now con’template and
Wustrate, dispitable has become disputable, and contemplative, of course, must
become con’templative. The tendency of this practice to deprive our syllabifica-
tion of its natural melody is obvious enough. In such words, for instance, as
signify and purify, the tendency to rob the final y of its natural long quantity
is strong, while in co’lumbine, bro'gardine, from the fuller quality of the final
syllable it is less. But if the danger be great in the case of the final syllable of
such words, it is greater in the case of the penult, that is, the syllable imme-
diately following the accented antepenult ; for, im the case of the final syllable, a
secondary accent may come in to save the prominence of the vowel, while the long
unaccented penult lies under the double disadvantage of a sinking inflexion
and a feeble stress, after the combined force, it may be, of an elevated accent
and a long quantity. From this cause it is that in vulgar speaking the second
syllable of the verb dicate is so liable to be shortened and turned into édicate ;
and so strong is this tendency, that many English scholars will tell you that to
pronounce the Greek word av@pwzos, with the accent on the first syllable and
the second syllable long, is impossible ; and it is no doubt true that it is not so
easy as saying avOpo7os, which the modern Greeks generally do; but as to the
alleged impossibility, we have only to look to such words as lan‘dholder, codl-
heaver, corn’dedler, to see that it exists only in the unpractised orthoepic organs
of the objectors. Of all languages that I know, the Gaelic is that whose euphony
has suffered most from the habit of throwing the accent back to the beginning
of the word. Of this there cannot be a more striking instance than words com-

* This is one among half-a-dozen reasons for the general want of success in our English hexa-
metrical experiments.

PLACE AND POWER OF ACCENT IN LANGUAGE. 279

pounded with the element mdr, signifying great, which may be divided into two
classes, those in which the termination mor, recognised in its full significance, is
accented, and those in which it falls under the category of the German /ich and
our y—in Giliicklich and lucky—being used for flexional purposes without a distinct
appreciation of its meaning, and therefore naturally unaccented. Of the one class
of words, Liosméor and Ben More, i.e., large garden and great mount, may serve as
familiar examples ; of the other, setmhor, fat, pronounced seltur, and grasmhor,
gracious, pronounced grdsvur, are excellent illustrations. For in these two last
words we see that the adjective mor, in losing its separate significance, loses both
its quantity and its natural accent ; and the compound word becomes a paltry
pyrrhic ~~, instead of a respectable iambus, ~ -, or a majestic spondee, — —.

Under this head it only remains to mention the extraordinary theory of
Bopp with regard to the place of the accent both in Sanscrit and Greek.
That illustrious philologer, in a work entitled “System of Comparative
Accentuation, or concise Exhibition of the Points of Agreement between Greek
and Sanscrit in the Doctrine of Accent, Berlin, 1854,” lays it down as a ruling
principle, that the most perfect kind of accentuation generally, and that which
prevailed originally in the Sanscrit language, was that im which the acute
intonation is placed as nearly as possible to the beginning of a word, however long.
Into the historical proofs of any such system of accentuation ever having existed,
of course only a profound student of the Vedas could enter. I am authorised,
however, by Professor Max MULLER and Professor AUFRECHT to say, that the
theory of Bopp is universally recognised as baseless; and this is just what
might have been expected. The mere assertion of such a principle to a man
whose ears have been trained to a rich and various orthoepy is monstrous. If the
accentuation of the first syllable, as in the well-known case of the Greek voca-
tives of the third declension, [ldrep, “Awo\ov, and such like, may well be
explained by the eager energy with which the call was made ; it does not there-
fore follow either that eager energy is the only thing to be looked at in a good
orthoepy, or that such oxytone words as d@yaly and Oeds may not be so enun-
ciated as to carry an intense expression of energy to the ear of the hearer. Let
this notion of Bopp, therefore, stand as only another instance of the great
blunders to which great wits are subject, and which, as large experience teaches,
are the natural consolation of the dunces.

That VARIETY is a necessary element of all esthetic presentation of the highest
order, needs no special proof. Variety is both an indication of wealth and a
preventive of monotony ; and as such is no less a natural source of delight to
the recipient of esthetic pleasure than of just boast to the producer.

Alles in der Welt lisst sich ertragen,
Nur nicht eine Rethe von schénen Tagen

says Goethe,

280 PROFESSOR BLACKIE ON THE

and what the Weimarian sage here says of beautiful days, is equally true of
beautiful verses or of beautiful words. Hence arises the sure canon—

That language is superior in point of accentual effect which gives no partial
predominance to any one accentual place, but gives the rising inflexion free play
over all the syllables of a word, so far as the range ts consistent with a full
vocalisation. Now, when we compare the Greek and Latin language by this
rule, we find a decided and universally admitted superiority in favour of the
Greek ; for this language admits of the acute on any one of the three last
syllables, while Latin allows it to fall only on the penultimate and the antepen-
ultimate. English, on the other hand, im this view, asserts one point of decided
superiority over both the classical languages; for words so accented as
lamentable and héritable, on the fourth syllable from the end, are not at all
uncommon with us, while the Greeks and Romans, who had no such accents,
fell into the very natural error of thinking that they were contrary to nature.
But, though with help of this peculiarity we are able to marshal a much larger
army of what the ancients called proceleusmatic feet in words than either Greeks
or Romans, we have gained this small advantage at a great risk in point of
general weight and majesty ; and we may be thankful to the graceful pedantry
of our classical scholars, who, in retaining the penultimate accent of many
Latin words, have done something to balance our habit of flinging the principal
accent far back and skipping over the remaining part of the word. The next
canon deducible from the test of variety is, that of any two compared languages
that is the more rich and beautiful in respect of accent, in which the acute accent
is placed not on the long syllable but on the short, so that, while the accent gets
fair play in one syllable, the quantity stands out in another, and thus a richer
and more various melody is distributed over every part of the word. For this
reason such words as célumbine, rénegade, are more beautiful than glo’riotis and
victo'riotis, enginéer and volunté’er, because in these last words, whether oxytone
or proparoxytone, all the wealth of sound is spent upon one syllable, while the
others remain comparatively weak and ineffective. On the same principle the
Greek avOpwos is richer than the same word accented in the Latin way,
avOperros, and *Apiorodavns is more beautiful than Aristophanes, if, as the English
habit has generally been, the final es of the word is pronounced short.

On the fourth principle, by which the comparative excellence of accents
may be determined, I place very little value. No doubt, as languages, like
buildings, are intended for use, convenience as well as theoretic excellence must
be consulted; but as utilitarian considerations have changed many an archi-
tect’s noble plan for a great building into a grand incongruity, so considerations
of mere convenience have spoiled many a fine language. For convenience,
really, ina great majority of cases, means haste and carelessness, or sloth and lazi-
ness, and in all such cases proves eventually a hostile and destructive force acting

PLACE AND POWER OF ACCENT IN LANGUAGE. 281

against all excellency of organism in articulate speech. We shall only say
generally, therefore, that it is always an imperfection in language when words
are so accented as to produce a lumbering unwieldy heaviness in the march of
syllables ; and we may say also that accents ought, if possible, to be so placed
as to admit of suffixes or prefixes being added without changing the intonation
of the word. In this view, contem'plative is a more convenient accentuation
than con’/templative, because it admits of a substantive contem'plativeness, and an
adverb contem’platively, being formed from it, without the necessity of either
advancing the accent or allowing it to remain on the fifth syllable from the end
of the new word, where its influence on the following syllables must naturally
be feeble in proportion to their remoteness from the point of vocal energy.

Of the effect of fashion and whim and caprice, in determining the accent of
certain words, and even of whole classes of words, contrary to every principle
whether of significance, euphony, or convenience, I say nothing, because such
arbitrary freaks belong not to the domain of scientific knowledge, and are
merely noticeable as casual aberrations or monstrosities.

Such are the grand principles of the general doctrine of accents, so far as I
have been able to discover them. It will be observed that they are based
on a wide induction, and apply to Latin and Greek as well as to Gaelic or
Italian. It is, however, a poit which has been long maintained in the
learned world, that the Greek accents have something altogether peculiar, and
not peculiar only, but peculiarly mysterious about them, which prevents
them from being used along with examples from any modern language as illus-
trations of general propositions about accent. It is against this notion—a notion
peculiarly English, and prevalent in high quarters—that I must proceed now
to make a distinct and deliberate protest ; for, till it be removed, it will be impos-
sible to say a single sensible word on the doctrine of accents, from which the most
interesting language in the world shall not be withdrawn asan example. I pro-
ceed, therefore, to show, both from the nature of the case and from the most
authoritative evidence, that there is not the slightest ground for the imagination
that accent in the classical languages meant anything substantially different from
what it means in English, or Italian, or modern Greek ; and, as a natural sequel
to this, I will trace the long course of scholarly opinion on the subject, from the
doctrine of Erasmus to that of Professor Munro, Mr Getpart, and other English
scholars ; and conclude by showing practically, what I have proved in the actual
work of teaching, how all the strange contradictions of this singular controversy
can be reconciled, and all the imaginary difficulties be made to disappear.

As a foundation for all argument on this subject, we may assume—what no
well-instructed scholar in the present state of learning will question—that the
-accentual marks now seen in every Greek book were first invented by AristTo-
PHANES of Byzantium, about 250 B.c., for the very same purpose that the marks

VOL. XXVI. PART II. 4D

282 PROFESSOR BLACKIE ON THE

of emphasis stand in our pronouncing dictionaries, viz., to ensure a correct
orthoepy in the reading and recitation of the language. The assertion once
boldly flung forth by the early opponents of Greek accents, that they were pro-
perly marks of musical intonation, having nothing to do with spoken eloquence,
can now be hazarded by no philologer. Whatever the accents meant, they
were intended to direct the reading of prose; had they been anything else
indeed, it is impossible to understand how they ever found their way into
the familiar notation of prose. But for the sake of those who may not be
familiar with the evidence on which this poimt rests, we shall here set down
the testimonies of two eminent grammarians: first, Dionysius THRrAx, who
lived at Rome about 80 B.c., and whose réyvn ypappaticy, quoted by SEXTUS
Empiricus (Adv. Math., i. 12), has been recently printed in the second volume
of Bexxer’s Anecdota (p. 629). This grave authority tells us that the art ot
erammar, as it was then practised, consisted of six parts—

1. dvdyvoots evtpiBys Kata Tpoc@diayv—assiduous reading, according to accen-
tuation.

2. Explanation of the meaning, according to the significance of the tropes
used by the writers.

3. Explanation of the historical facts and of the glosses or peculiar words.

4, Etymology.

5. Consideration of linguistical analogies.

6. A critical appreciation of the work expounded, in its beauties and defects.

Now, there can be no doubt here as to what zpoowdia means ; for, though
the plural of this word sometimes is used in a wider sense, as we tall of the
Hebrew points, so as to include aspirations, pauses, quantities, and every affec-
tion of which spoken and written words are capable, when used in the singular
as a special technical term, it signifies accent, and nothing else. The second
grammarian whom I quote is THEopostus, who lived in the time of the Emperor
CONSTANTINE, and whose treatise on grammar was published by GoETTLING in
the year 1822. This author, in the chapter (p. 58) entitled més ypy dvayty-
vooKew, says that good reading consists in three things—

1. wrdxpiois, dramatic expression, arising out of a sympathetic conception
of character.

2. mpoowdia—or reading kata Tovs axpiBels tovovs—according to the exact
accents—zpoo@dia yap 6 tovos—for accent and tone, are the same.

3. diacrody, attention to pauses and punctuation.

Now, if any person further inquires whether the ancients did not read their
prose according to quantity also, I answer that of this there can be no doubt ;
but that the prominence in correct reading is naturally given to accent, because
quantity is the specialty of poetry, and unless where we talk specially of poetry,
by the word reading we are understood to mean prose. But that correct read-

PLACE AND POWER OF ACCENT IN LANGUAGE. 283

ing of prose included quantity also, is evident from what the same grammarian
says a sentence or two below, viz., that under tpoc@dia, in a wider sense we
understand both accent and quantity, and in this wider sense correct prosodial
reading arises ex Tov mapapuddtTew Tos Tévouvs Kal Tovs xpdovous, from observing
the tones and the times, and all the other affections of articulate speech. Now,
as there was an uninterrupted succession of grammatical teachers, from the age
of the Alexandrian Ptolemies to the time of the Roman Emperors, and from the
establishment of the Eastern Empire by ConsTanTINE to the taking of Constan-
tinople by the Turks, no historical fact can be more certain than this, that the
living accentuation with which Greek was spoken in the great seats of learning
and culture in the third century before Christ, and by which a just ortheopy in
reading was determined, has been handed down to us in an unbroken chain of
the most authoritative testimony. If this is not true, there is nothing now
credited in the wide sphere of linguistic tradition that rests on a surer basis.

If, then, the ancient Greeks both spoke and read by the rule of those
accents which we now see on our printed books, what are we to understand by
that accent? Now, here the field of definition is happily well narrowed. That
Greek accent did not mean quantity, every page of tradition on the subject
distinctly testifies; that it did not mean mere volume of mass of articulate
sound is equally certain ; and no man, ancient or modern, ever dreamt that it
did. There remain, therefore, under which it may fall to be subsumed, only
the other two affections of articulate speech with which we started, viz., eleva-
tion of tone and intensity of utterance. Greek accent must be either the one
or the other of these, or both together. That it means the first, viz., elevation
of tone, is plain from the mere terms és and Bapis, sharp and heavy, or high
and dow, by which the two familiar accents are designated. It is also distinctly
stated by both Greek and Roman grammarians that accent implies change of
tone in the voice, by alternate elevation and depression. The phraseology,
indeed, of this matter was borrowed by the grammarians from the musicians,
and had reference to the high and low notes in the musical scale, these minute
speculators having justly observed that, as the voice in music rises or falls by a
series of measured intervals, so in articulate speech it rises and falls by a suc-
cession of slides, what our great orthoepic teacher calls the rising and falling
inflexions. Either, therefore—the acute accent of the Greeks, which is the
accent properly so called—means the rising inflexion of the voice on particular
syllables of a word, or it means this, p/ws a stress or emphasis on a certain
syllable of a word, produced by the greater force, or stretch, or tension of
the voice on that particular syllable. Now that it does not mean elevation of
the voice merely, but also, and at the same time, that greater stretch or tension
of the voice which produces the emphatic syllable of a word, will, I think, be
evident from the following six considerations :—

284 PROFESSOR BLACKIE ON THE

1. From the natural difficulty of elevating the voice, and not at the same
time giving an increased emphasis to the elevated vowel; or, may I not say,
rather the natural impossibility—for, though it is. certainly possible to give a
great emphasis to a syllable, and keep the voice at a low key, that is to say,
though stress does not necessarily imply elevation—it certainly does not seem
very natural or very easy to raise the pitch of the voice without accompanying
that high pitch with a certain emphasis. JI may, for instance, pronounce the
Greek word dvarohy, with a stress on the last syllable, and yet with the
whole pronounced in monotone ; but, if I raise my voice on that syllable, it will
be difficult for?me to withhold from the syllable the stress which naturally
accompanies the act of elevation.

2. But that Greek accent implies stress as well as elevation is manifest
from the natural and obvious meaning of the terms used by the grammarians
in describing the phenomena of accent. For what does tao.s mean but stretch
or tension? and is it not quite plain that as contrary as light is to darkness, so
contrary is ériracis to aveois,—t.é., intension to remission, strain to slackness of
sound—the constant phraseology of the grammarians with regard to this matter.
The word xpotopma, also signifying beat or strike, which is sometimes used, of
the acute accent,* sufficiently indicates its analogy to the emphatic note in a
musical bar, which certainly does not signify elevation or depression.

3. The analogy of the ictus metricus in rhythmical composition, suggested by
the word kpotopa, supplies another argument to prove that the Greek and
Roman accent meant stress as well as elevation. For there are some places in
the poets where we can observe that a word naturally short is made long for no
other reason that can be seen than that the spoken accent on the syllable
favoured the poetical license, just in the same way that the rhythmical accent
sometimes does. Mere elevation has no effect on quantity; but stress or
emphasis can easily be so manipulated by the voice as to pass over into a long
syllable, or, to use the language of the grammarians, ésiraois may become
extacis, intension may spread itself out into extension.

4. That the acute accent meant stress is plain from the inherited intonation
of the modern Greeks; for accent is one of the most obstinate affections that
belong to spoken speech ; and no man can hear such words as kado 7awdt, 2xomd,
and Ilapyacoé in the mouth of the living Greeks without feeling that the dead
mark on our books has here received its living interpretation ; and, if any per-
son objects that the modern Greek not only acutes the last syllables of these
words, but makes their quantity long, this is all in favour of my argument ;
for the length arose and could arise naturally only from an exaggeration of
that tension of voice which was the necessary accompaniment of the accent.

* THEODOSIUS, GoErrTLine, p. 61 ; KpovaTtexotépa yuyvouevn 1) Nets GEvveTaL, Schol. Dionys.
Thrax. Bekker, ii. p. 690.

PLACE AND POWER OF ACCENT IN LANGUAGE. 285

With regard to the modern Greek dialect generally, I would observe that
though the place of the accent has been changed in a few classes of words, in
the great majority of cases it has been retained ; and that in the case of cur-
tailed words, as pds for éuds, riow for dice, dpi for didprov, mardi for zasdior,
dev for ovdev, &c., it is the stress upon the medial accented syllable which has
secured its permanence after the initial or final unaccented syllable had
dropped off.

But the most incontestable proof that accent means emphasis lies in the
doctrine of Encuitics ; for in Greek as in English there are certain little words,
such as the pronouns or the negative no, which in common cases are purposely
kept unemphatic, and pronounced so rapidly as to appear to lean upon (éyrdivo),
or be taken up by the previous or followmg word; but the moment that the
necessity of speech demands these words to become prominent, they receive
the accent, and become emphatic. Thus we say, “ give me the book,” like datemi
in Italian, as one word, but “ give mE the book,” that is, give it to me, not to
you. Now, there could not be a stronger fact than this to prove that Greek
accent meant emphasis; for this use of the acute accent to emphasize in
particular cases otherwise unemphatic words is quite common, as, for example,
in the case of the negative particle wa Ata odk éywye, contrasted with ovrws héyes
H ov, do you say so, or do you nol?

6. Lastly, the analogy of the modern Italian compared with the ancient
Roman, plainly shows us both the obstinacy of accent as a fact in the life of
language, and what accent really meant in ancient Rome and Greece, as in
modern Rome. For nothing is more certain than that, though its special laws
were different in the two learned languages, accent, as an accident of articulate
speech, did not mean one thing in Greece and another thing in Rome; but
the Greek and Latin accent were in their nature and operation identical; so

| that what is predicated of the essence of the one must be considered as predicated

of the essence of the other. If, therefore, the modern Italian accent, in its position
and power so evidently identical with the old Latin, possesses the element of
stress as a prominent feature, it is a legitimate conclusion that the Greek accent
did so too. Altogether, it may be remarked as a very extraordinary fact, and in-
dicative of the operation of some strange deluding prejudice, that, while the most
formidable artillery of erudite arguments have been brought to bear against pro-
nouncing Greek with Greek accents, no learned Latinist has yet written a book
to prove that Latin ought not to be pronounced with Latin accents. When
reading Latin we put the stress on the accented syllable exactly where Cicero,
and QUINCTILIAN, and PriscrAN say it was placed ; but the moment a Hellenist
gives the natural predominance to the accents which he finds marked on his
books, he is immediately told that accent does not mean stress, but means some-
thing that no man can understand or make use of. Whence this inconsistency ?
VOL. XXVI. PART II. 4k

286 PROFESSOR BLACKIE ON THE

Having thus proved, by what may surely seem sufficiently strong arguments,
that accents mean nothing in Greek, which they do not equally mean in Eng-
lish, or Latin, or Italian, there remains only to take a bird’s-eye view of the
somewhat remarkable literature of this subject, from the revival of letters
down to the present hour. Such a review will at once be the best justification
of the principles above set forth, and will place vividly before the reader the
partial and inadequate points of view from which the opposing doctrines have
taken their rise.

Now, in tracing the stream of confusion which this matter exhibits to its
fountain head, it is most natural that we should, in the first place, turn to
Erasmus, both because he was the most prominent scholar of European reputa-
tion in the eventful age to which he belonged, and because it is quite certain that
before his time no learned man ever dreamt or could have dreamt of disown-
ing the pronunciation of the Greek language, which Europe had received as a
common legacy from the Constantinopolitan Greeks. The early scholars, indeed,
were occupied with matters of far more serious import than the exact accen-
tuation and quantification of syllables. They read the Greek books for the
information they contained: Heropotus for history, Straso for geography,
TuucypDIpEs for political wisdom, Puato for philosophy, ARISTOTLE for science.
So long as this appetite for the stores of Hellenic thought and knowledge was
the one thing needful, no man had either leisure or desire to put curious ques-
tions to himself with regard to the auricular luxury of a just orthoepy.
But the time must come when this matter also would be examined : Homer
and SopHocLes could not be read in their mother tongue by men who used
their ears as well as their eyes, without provoking questions as to the best
method of bringing out the full music of that most musical of human languages
which it was the happy fortune of these great poets to employ. If Greek was
the language of the gods, there seemed a manifest impiety in allowing it to be
enunciated by a confused, degraded, and irrational elocution. And, if such
questions were to be raised, ERAsMus was precisely the man, who, from his fine
genius, cultivated taste, and broad human sympathies, was best fitted to raise
them. Accordingly, in the famous dialogue, “ De recta Latini, Groecique
sermons pronuntiatione,” published at Basle in the year 1528, the whole subject is
brought under review ; and the text of his discourse is in the broadest terms, that
“nunc tota fere pronuntiatio depravata est tam apud G'recos, quam apud Latinos ;”
and this is proved in a very exhaustive style in an argument extending to above
two hundred pages. The powers of the different letters are critically discussed,
and the relation of accent and quantity illustrated both by learned rules and
by living examples. With regard to the vowel sounds, which is the first point
handled, he had an easy task to prove that the slender sound the characteris-
tic of the Byzantine Greeks could not have been the original sound of so

PLACE AND POWER OF ACCENT IN LANGUAGE. 287

many distinct vowels and diphthongs. Signs of different vowels were certainly
not made originally to confound, but to distinguish. The confusion in this case
is always of a later birth. What Erasmus, however, failed in here, and what,
from want of materials, he could not but fail in, was to show at what period
this confusion commenced ; for, as the most polished nations in modern times
display in their speech abnormal tendencies and depravations of all kinds,
which are consecrated by usage and fashion, so there is no reason why the
itacism of the theologians of Byzantium should not have been practised by the
philosophers of Alexandria, and even, to a certain extent, by the orators of the
Periclean and Demosthenic age. However, this was not curiously looked into;
and the result was that, by this assault of Erasmus, the faith of scholars in the
orthoepic traditions of the Byzantine elders was shaken in all the most learned
countries of Europe, and every nation set up vocalizing Greek according to
what seemed good in its own eyes. Hence the motley babblement of Greek
which now prevails. The old foundations were removed before the ground was
opened, or the materials ready, to make new ones. And thus it has happened
that an orthoepic reform, well intended, and in so far conducted on rational
principles, has issued in an extremely irrational and altogether unsatisfactory
result. So much for Greek vocalisation. With regard to that other matter
with which we are specially concerned here, we do not find, what we might per-
haps have expected to find, that the great modern innovation of disowning Greek
accents in reading Greek, receives the slightest countenance from Erasmus.
On the contrary, part of the bad pronunciation which it was his object to reform
was precisely the ignorance or loose observance of the proper accents in Greek
and Latin, according to the characteristic laws of each language. He saw also
everywhere amongst careless, tasteless, or ignorant speakers, that confusion of
things so distinct as accent and quantity, which from the same causes prevails
so largely at the present day. Scholars still tell you that accent and quantity
annihilate each other, and cannot both be observed, meaning only, in fact, that
for their particular ill-tutored and perverted auricular organs, it has become
difficult, and is perhaps impossible. It certainly is impossible for a sharp, hard
Aberdonian to speak with the rich silvery mellowness of a high-bred English
lady ; but the difficulty lies in bad habit, not in Scottish nature. On the super-
induced habitude which erudite ears have so often displayed in not being able
to distinguish accent from quantity, there is a passage in the Erasmian tractate,
which we shall be excused for inserting at length :—

“ Sunt quidam adeo crassi, ut non distinguant accentum a quantitate, quum sit
longe diversa ratio. Aliud est enim acutum, aliud diu tinnire: sicut aliud in-
tendi, aliud extendi: quanquam mhil vetat eandem syllabam et acutum habere
tonum, et productum tempus, velut in vidi, et legi preeteritis. At eruditos novi, qui,
quum pronunciarent illud avéxov kai amréxov, mediam syllabam, quoniam tonum habet

288 PROFESSOR BLACKIE ON THE

acutum, quantum possent producerent, quum sit natura brevis, vel brevissima
potius. Et ferée qui Greca legunt, accentus observatione confundunt spatium more,
sie enunciantes pevédaos, quasi penultima sit brevis, et wevédnmos quasi duce postre-
me sint breves, quemadmodum in Oeddwpos trapaxdyros, eidwda, aliisque innumeris.
Nec ita multis contingit sonare Greca, ut accentuum simul et morarum rationem
observent, vel in carmine. Loquor autem non jam de vulgo, sed de eruditissimis
quoque. Minus est erroris in Latinis, sed tamen ilic quoque tonus acutus ac in-
flexus obscurat ceeterarum sonum, ut in videbimus, congruit accentus cum quanti-
tate, at in legebaimus, sola penultima videtur esse producta, quum secunda sit ceque
longa: in amavérimus sola antepenultima, quum ea sit brevis, secunda producta.
LE. Omnino sic obtinuit usus, quem dediscere difficillimum est. UR. Atqui qui
degustarunt musicam, nullo negotio distinguunt inter longam, brevem, et inter acu-
tam et gravem. Nihil enim est aliud pronunciatio, quam modulatio queedam
vocum numerosa. Est enim et in oratione soluta pedum ratio, licet non perinde
certis astricta legibus ut in carmine: que si confundatur, non magis erit oratio
quam cantio in gua graves cum acutis, longe cum brevibus temere confunduntur.
Unde quidam priscorum grammaticorum non inscite dixerunt, accentum esse ani-
mam dictionis. Et tamen hodie talis est etiam eruditorum pronunciatio, qualis
esset illa ridicula cantio. Scis opinor canere cithara. LE. Utcunque. UR.
Nonne frequenter imam chordam pulsans producis sonos, et summam tangens
brevibus insonas aut contra? LE. Frequenter, quanquam hoc discrimen eviden-
tius est in flatili musica. UR. Unde igitur nos sumus usque adeo dpovoo., ut
omnes acutas syllabas sonemus productiore mora, graves omnes corripiamus? Vel

ab asinis licebat hoc discrimen discere, qui rudentes corripiunt acutam vocem,

imam producunt. LE. Idem propemodum facit cuculus.”

The only other interesting point, with regard to the present matter, which
requires to be mentioned here, is that Erasmus distinctly teaches that verses,
both in Greek and Latin, are to be read with an accurate observance both of accent
and quantity. The difficulty and alleged impossibility of doing this, so much
spoken of by modern scholars, he supposes to arise only from the gross neglect of
the art of elegant reading in modern education. How far he is right in apply-
ing the spoken accent thus sweepingly to the rhythmical recitation of poetry,
we shall have occasion to consider afterwards.

But what to the fine genius and well-trained ear of ERAsmus presented no
difficulty, to the gross majority who take everything without discrimination in
broad masses was so formidable, that they do not even seem to have had the
courage to look the difficulty in the face, but quietly settled down into a habit
of confounding accent and quantity, and making all accented syllables long.
This is distinctly mentioned by the next champion in the field, ADoLPH von
MEETKERCHE (vulgarly Mexircu), a Flemish nobleman, born at Bruges in the
very year when Erasmus’ book was published, and well known in high circles

PLACE AND POWER OF ACCENT IN LANGUAGE. 289

in England, from his having lived and died at London as an attaché of the
Belgian ambassador at the court of ExizasetH. He was, besides an able diplo-
matist, an accomplished scholar, and in the year 1576 published a Discourse “ de
vera et recta pronuntiatione lingue Grece,”* which seems to have given the first
impulse to the paradoxical movement which caused the Greek accentuation, so
laboriously preserved by the Alexandrian grammarians, to be thrown overboard
in the general practice of scholars, and the vulgar Latin accentuation substi-
tuted in its place. The principal part of this work is occupied with the ques-
tion which then loomed most large, whether the Byzantine vocalisation should
be retained, or a reformed one introduced, as suggested by Erasmus; but, ina
short appendix, the doctrine of accents is stated succinctly, and, what is more
important, the author’s practice with regard to their observance. In the first
place, he tells us the important fact that, im his day, Greek was so read by
many, confounding accent and quantity, as altogether to destroy the perception
of any poetical rhythm. “ Manifestus est corwm error qui tonos cum temporibus
confundunt, ita ut quecunque acuenda vel flectenda est syllaba, eam producant :
quecunque deprimenda vel equabiliter pronuncianda, eam corripant. Ea quo jit
ut in Greecd oratione vel nullum vel potius corruptum numerum intelligas, dum
multe breves producuntur, et contra plurime longe corripuntur ; ut pene prestt-
terit Greeca vel Latina non legere quam ita foedé depravare” (p. 175). . And no
wonder ; if, as he says, the accent was allowed such a power that, in the second
line of the Iliad, é@yxev was read as a dactyle, and the two final syllables of
ovromevnv as a Spondee. And then he tells us of a general practice of school-
masters, which by the way prevails in England almost universally to the present
hour. ‘Solent enim peedagogi vulgo ita suos erudire ut in omnibus dissyllabis
penultimam producant.” Just as in Eton and Harrow the boys had, till very
recently, if indeed they are not still, taught or carelessly allowed to say, bonus, and
not bonus. He then goes on to show how this practical assumption that a penul-
timate accent must necessarily lengthen the vowel has no foundation in the real
nature of accent and quantity, of which the one expresses the quality of the
sound, the other the dimensions. And then anticipating an objection often
made im modern times, he goes on to say, “ Neque tamen nego brevi syllabe
temporis aliquid accedere, quando acuto signo signatur, quantum scilicet necesse
est in acuendd syllaba consumi ; ged, ut minus sit brevis quam antea, minime tamen
consequitur habendam esse pro longa, sicut ab iis habetur qui MALUS arborem @ MALO
adjectivo non distinguunt” (p. 178). This is exactly what Erasmus had said ; and
one should think it would be sufficiently patent to all ears, except those of stupid
schoolmasters, careless schoolboys, and bookish scholars, whose learning is all in
their eyes, and notin their ears. But things easy in speculative thought become
in the hasty practice of life, sometimes tolerably difficult ; and, in fact, a just

* Reprinted in “ Havercamr’s Sylloge.” 1736. Vol. i. p. 9.
VOL. XXVI. PART II. ; 4*F

290 PROFESSOR BLACKIE ON THE

and true pronunciation, even in the case of the mother tongue, is not attaimable
without a certain amount of trouble. MEETKERCHE accordingly finds that
his argument for accents, however just, is liable to be met with the objection
which nullified so many of Soton’s well-conceived legislative reforms. The
laws were no doubt very good, but they were too good for the people. The
best for them was not the best absolutely, but the best which they could endure.
“ At enim,” he continues, “ dices, ista (t.e., the right pronunciation both of quan-
tity and accent) esse perdificilia, et fortassis etiam advvara, tis quidem qui diverse
pronuntiationt assueverunt. Id ego vero fateor, et in me ipso non invitus agnosco.
Sed nihil vetat rectam viam aliis ostendere, etiam ut illam ingredi non possis.
Certé veritas mihi dissimulanda non fuit, ut paullatim meliora probare et sequi
condiscamus. Ergo, ut libere dicam quod sentio, vel tonos prorsus sublatos esse
velim, tantis per dum depravata ila pronunciatio tonorum pro temporibus emen--
detur (quum presentim veteres constet istos apices in scribendo non usurpasse) vel
nullam eorum rationem habert.”* Which simply means that he is in favour of
suspending the operation of Greek accents till such time as schoolmasters—
proverbially not a very teachable race—shall have learned to distinguish 6s, a
bone, from 6s, a mouth, and that cdn'o is a possible combination of articulate
sounds, as much as caw’no or caéno.

The next important work which falls to be noticed indicates plainly by its
title—“ De Poematum cantu et viribus Rhythm ;” Oxon. 1673—from what
quarter the attacks of a section of the learned world were now to be directed
against the traditional sway of Greek accents. The author of this tract was
the celebrated Isaac Vosstus, “‘ unquestionably,” to use the words of MArKLAND,
“a very learned man, but whose whimsicalness and love of paradox scarce
leaves room for him to be considered a reasonable one.” t Vossrus, like MEgEt-
KERCHE, had got his ear possessed with a genuine living appreciation of the
beauty of measures and rhythm in poetry, which justly resented the barbarism
of those scholars who read ancient verse by accents, just as if it was so
much German or English verse. In expressing his indignation strongly against
these systematic murderers of the regal majesty of Latin, and the luxu-
riant swell of Greek verse, Voss did well; but, when he went farther, and
not content with the interim act of suspension passed by MEETKERCHE,
stood up in violent revolt against the whole gccredited system of accentua-
tion in the Greek language, and cast it, to save the ship, like a Jonah
overboard, he committed a great mistake, and kicked vehemently against
the pricks, where he could only wound his own legs. He declared roundly that
the whole system of Greek accents, as we now have them, was a modern
invention, or, at least, a corruption, or a monstrous compound of both;

* Havercamr’s Sylloge. Ludg. Bat., 1836. Vol. i. p. 179.
+ Letter to Fosrrr in the Essay on Accent and Quantity. 3d edit. London, 1820. P. 207.

-PLACE AND POWER OF ACCENT IN LANGUAGE. 291

that accents were originally musical marks, and had nothing to do with
the pronunciation of the language ; that the best proof of this was the un-
rhythmical jar which they produced, when actually applied to the recitation
of verse, whether Greek or Latin ; and that therefore the only course left to the
scholar of taste was to disregard them altogether, and use only such accent as
was manifestly dictated by the march of the metre. While, however, this
ingenious scholar found it comparatively easy work to pronounce a dictatorial
sentence of eternal exclusion against Greek accents, of which few had any real
knowledge, he found himself obstinately met by an obvious objection from the
familiar practice of the Latin tongue, which, while it distinctly disowns (except
in a very few exceptive cases) all oxytone accentuation, nevertheless, in verse,
constantly uses an emphasis, which falls with marked effect on the last syllable
of one or more words in the verse. In answering this objection, Voss fell upon
an aspect of the case, which, if he had applied it to Greek poetry, might have
saved him from the trouble of beating vainly against the strong bulwarks of
Alexandrian and Roman and Byzantine tradition in the matter; for he distinctly
says that singing is one thing and reading another, and that the Romans may
have followed a different law of accentuation with regard to each. “‘ Quare non
quidem multum refragabor, st quis in recitatione Latinorum poematum ultimas
syllabas unquam productas Juisse negaverit: sed vero in CANTU id ipsum fiert
potuisse si quis contendat, idem etiam merito afirmet et Latinos canere nescivisse.”*

Close upon the traces of Vossrus comes a German, HENRY CHRISTIAN HENNIN,
whose work entitled ““EdAyuopds dp8@d0s, Traject ad Rhenum, 1684,” with a
great flourish of trumpets on its title-page, proclaims itself to prove “ Graecam
linguam secundum accentus, ut vulgo ab omnibus hucusque fiert consuerit, pronun-
ciandam non esse.” The inspiration of this book—for it is full of fervour and
emphasis, and a sort of lofty protestation—manifestly is the same as that of
Voss’ treatise ; a certain school of scholars with whom the writer had been
familiar, or it may be all the scholars of his time and place had got into a habit
of sacrificing the rhythmical recitation of Greek poetry to the traditional accen-
tuation of Greek prose, a usurpation, no doubt, of a most gross kind, and which
it was obvious to think could best be got rid of by not only dethroning the
usurper and telling him to keep to his proper place, but by killing him outright,
and casting him down among the dead men with a triple volley of curse and
execration. It wasa procedure akin to that in political history, when democracy
dethrones despotism, and acts ten times more despotically than the tyrant whom
it overthrew. In conducting his indictment against the accents, the author com-
mits in the outset the very transparent blunder of confounding the marks of the
accents in printed books, with the living accents in the mouth of the people who
spoke the Greek language. These marks, whether present or absent in books,

* Deviribus rhythmi, p. 44. N.B.—By productas in this passage he evidently means accented.

292 PROFESSOR BLACKIE ON THE

do not in the slightest degree affect the question; they do not exist in English
books, and yet English words have a well-known accent in the voice of the
English people, and as made visible artificially to the eye inthe pronouncing
dictionaries of WALKER and other orthoepists. The next great error made by
HENNIN lies in the theory—for it is a mere baseless theory—that the accents were
invented by ARISTOPHANES of Byzantium, for some purpose quite different for that
which they now subserve. This is simply to leap over the testimonies of the
most learned Greek grammarians from the time of the Alexandrian scholars to
the taking of Constantinople by the Turks. And in order to make such a
hypothesis possible and even plausible, he draws a flaming picture of the
barbarism which corrupted the Greek language at a fever pace from the
Roman to the Turkish conquest. All this, however, is purely imaginary, as any
person who has looked even superficially into Byzantine literature must con-
fess. Whatever changes in the course of time naturally might take place in
the spoken language of the Greeks, the last element that would be touched by
the change was the accentuation; and that not only from its own natural
obstinacy, but from the very fact that the proper place of the accent visible in
most written books presented a stereotyped norm, that checked all arbitrary
deflexion in the start. Any other arguments that make a parade in HENNIN’s
book are based on the fact of which we hear so much in these days, that certain
persons could not pronounce avOpw7os without saying avOporos, and certain
other persons imagined that it was impossible to do so. After overleaping
heroically the bristling fence of historic testimony on the matter, the author
proceeds to lay down four rules of accentuation, which, both in the Greek and
Latin languages are, “ sine alld exceptione weterne veritatis.” These rules are
as follows :—

(1.) “ Omnis vox monosyllaba modulationem habet in sud vocali ut Bas, vovs,
mons, Pons.”

(II.) Omnis vox dissyllaba modulationem habet in syllabd priori, ut hoyo.,
adou, povn.”

Ill. “Omnis vox polysyllaba penultimam longam modulatur ut dvOparos
turtapev, Greecorum, jucinda, Romanorum.”

IV. “ Omnis vox polysyllaba, penultima brevi, modulatur antepenultimam ut
déminas, adoyov.”

This is certainly one of the most cool pieces of insolent one-sided dogmatism
that the history of learning presents, the whole affair being simply an assertion
that the particular method of accentuation in the Latin language, which the
author had inherited from secular and ecclesiastical Rome, should be stilted up
into an eternal norm of accentuation for all languages, while the most plain and

obvious facts, both in ancient Greek and modern English, which contradict the

theory are held as non-existent, and excluded from the calculation; an instructive

dl de eis

PLACE AND POWER OF ACCENT IN LANGUAGE. 293

example of the truth of GorTHE’s remark, that truth is often disagreeable to us,
because it limits the despotic sweep of our one idea, while error is grateful for
this, above all other reasons, because it prostrates fact and thought and
history before the triumphant march of our infallible conceit.

It was not to be supposed that the sweeping dictatorial dogmatism of this
book of HeEnnin, backed as it substantially was by the high authority of Voss,
would pass without comment from the learned of the Continent ; and accordingly
we find that in the year 1686 it received a long and able reply from Joun
RupoiprxH WETSTEIN, professor of Greek in the university of Basle. WETSTEIN’s
book, by an overwhelming array of historical testimony, enforced by sound
argument, demonstrates the utter untenableness of the proposition of his
adversary, unwarrantable equally in the wholesale swamping of the Greek by
the Latin accent, and in the elevation of this latter into a rational norm of
accentuation, by which the excellence of all articulate speech is to be measured.
With regard to the main difficulty which had staggered MEETKERCHE, the Basle
professor quietly reminds his antagonist, in the words of QUINCTILIAN, that the
recitation of verse is in many respects different from the speaking of prose,
“imprimis lectio virilis et cum suavitate quadam gravis, et non quidem prose
similis, quia carmen est.”

The infection of this notable dispute now comes to England, and the first
oracle to whom we feel inclined to propound the question for solution is, of
course, the great Benttey, This massive and masculine scholar, in the short
treatise on metres prefixed to his edition of “TERENCE,” has the following
passage :—“ Tam vero id Latins comicis, qui fabulas suas populo placere
cuperent magnopere cavendum erat ne contra linguee genium ictus seu accentus in
quoque versu syllabas verborum ultimas occuparent. Id in omni metro, quoad
hicut, observabatur ; ut in his

‘ Ay’ma virumque cano, Trdjae qui primus ab Oris,
Italiam fito préfugus, Lavinia vénit

Litora; multum ille et térris jactatus et alto

Vi superum, saévae mémorem Junonis ob iram.’

Qui perite et modulatae hos versus leget sic eos, ut hic accentus notantur, pro-
nuntiabit, non ut pueri in scholis, ad singulorum pedum initia ;

Italiam faté profugus, Lavinaque venit, sed ad rhythmum totius versus.”

Now, it in no wise concerns us to discuss the value of the remark here
made as to the practice of the Latin poets ; that is a delicate matter, we believe,
not so easily settled as the stout Cantab seems to have imagined. The only
significance of the passage for our present inquiry is, that the writer believed that
in some way or other the structure of Latin verse was regulated by a regard to
the spoken accent, and not simply by the law of quantity and the metrical beat.

VOL. XXVI. PART II. 46

294 PROFESSOR BLACKIE ON THE

What truth there may be in this notion will appear in the sequel ; meanwhile
it is quite plain that it leaves the matter in a state of considerable uncertainty,
an uncertainty which is not at all diminished by the unquestionably rash
assertion in the letter to MILL, that Greek accents were an invention of later
times, which could only mislead the accurate scholar.* An obiter dictum of
this kind, even from a BENTLEY, on a confessedly difficult question, cannot be
regarded as having any real weight. It may, however, along with other causes,
have contributed to produce that strange aversion to Greek accentuation so
characteristic of English scholarship.

We now advance by a long stride into the middle of the great battle of
accent and quantity that was fought in this country about the middle of the
last century. The protagonist of this warfare is the Rev. Henry GALLy, a
Kentish Doctor of Divinity, and chaplain to His Majesty King Grorce IJ.

His dissertation against Greek accents was first published in the year 1754, —

seventy years after the famous works of HENNINIUS and WETSTEIN ; and quite
recently on the back of two treatises on the same subject, which had appeared
in Rome.t Dr Gatty wrote, quite aware of the achievements of his predecessors,
but convinced that their attempts to untie the Gordian knot were unsatisfactory,
and that his own method was altogether new and original; and so it is, no
doubt, in some things, but novel only in the daringness of its assertions and the
glaringness of its absurdity. Its absurdity consists mainly in the writer’s
belief that he can overturn the whole principles and practice of the Greek
accentuation, by simply saying that it is irrational and absurd, as if some
famous philosopher, some thousand years after this, when the English orthoepy
may have become a field for learned debate, were to say that Mac’Intosh and
MacIntyre could not have been pronounced with the accent on the first syllable,
because it is irrational to place the accent on the common element of the Mac,
instead of on the distinguishing element, the clan; which rational method of pro-
nunciation, as above remarked, exists not only in all the other Macs, but im all
the Saxon names ending in son, as An’derson, Péterson, not Anderson’, Peterson.
A writer belonging to a people whose pronunciation is in all points so various,
so arbitrary, and so dependent on fashionable caprice as the English, might
surely have spared himself the inconsistency of such an argument. In the
other parts of this learned divine’s book we find merely a repetition of what had
been said by MEETKERCHE, HENNINIUS, Vosstus, and others. Accents, we are told,

were entirely musical, and had nothing to do with the intonation of colloquial

speech : then it is broadly asserted that accent necessarily constitutes quantity,

* “ Note: accentuum quorum omnis hodierna ratio prepostera est atque perversa.” Works by
Dyce, vol. ii. p. 362,

t (1) Sarpedonii dissertatio de vera Atticorum pronunciatione. Romae, 1750. (2) Velaste disser-
tatio de literarum Greecarum pronunciatione. Romae, 1751.

i ate sl

tie 2 See

PLACE AND POWER OF ACCENT IN LANGUAGE. 295

and therefore must be wrong; and that, whatever the advocates of accents
might preach in theory, in practice they never did, because they never could
observe the accents without destroying the quantity. This practical difficulty
is, in fact, the gist of his whole treatise, as is manifest from the very notable
words with which he concludes :—“ If, therefore, we would observe uniformity,
and keep to what we can safely rely on, we must not admit of any use of
accents in the pronunciation of the ancient Greek language but what is con-
sistent with quantity ; and if we have lost the nicer part of the ancient pronun-
ciation, we have the more reason to adhere to the essential part which still
subsisteth.” And this way of putting the case, viewed as an argumentum ad
hominem addressed to the great mass of the English scholars and teachers, is no
doubt perfectly just ; for these gentlemen had got into a monstrous and irrational
habit of writing Latin and Greek verses with much labour and wonderful
dexterity, by help of their understanding only, against the verdict of their
ears, and treated both accent and quantity as an affair of dead rules, not of
living vital action.*

But English scholarship—whatever might be the absurdities of professional
pedagogy—was not destined to surrender one of the strongholds of venerable
philological tradition at the trumpet-blast of such a windy dogmatist as Dr
GatLy. In the year 1767, a reply to his pretentious heresy was sent forth from
Eton, by Foster, in which, so far as the learning of the subject is concerned,
he showed himself as superior to GALLY as WETSTEIN was to HENNINIUS.
He proved, beyond all possibility of denial, that accent had always been a
recognised element in Greek orthoepy, and was in no sense the barbarous
creation of a decadent age and a degraded taste. He stated also most distinctly
that, while elevation of tone was the most characteristic element in Greek
accent, it also necessarily included the element of stress—which Dr GALLy also
saw clearly—but that this stress or emphasis was in no case to be confounded
with the length or duration of syllables. Hence, indeed, the great superiority
of his argument to that of the Kentish D.D.; for he not only maintained that
accent was not to be confounded with quantity, but that, from the very nature
of the case, the intense energy of the acute accent might, in many cases, have a
tendency to shorten rather than to prolong the emission of breath by which it
was enunciated.t With regard to the main difficulty, however—the practice of
the theory, which, as we have seen, was the stumblingblock of Dr GALty—he
does not seem to advance the matter far. Hear his words :—

“Nor let it be said, if we should retain these sounds, we can never apply

* On this notable inconsistency of those champions of quantity who denounce accent, Mr Fosrrr
is justly severe ; ch. x., on accent-quantity.

+ On this point he produces a remarkable passage from Sumas, in voce o€v, vol. ii. p. 1136.
BERNHARDY.

296 PROFESSOR BLACKIE ON THE

them to their proper use in practice. Who can affirm that with certainty? An
English voice was capable of doing this in the time of Henry VIII., and why
not now? Sir JoHN CHEKE declares it not only practicable, but that it was
actually practised, and that he knew many persons who could express these
sounds consistently with accent and quantity perfectly well. I know one
person who, after a few trials, is now able to do the same.” By this one
person, the reader will naturally suppose that he means himself, though it is a
pity he did not say so in a manner that could not admit of ambiguity. But who-
ever the individual might be who in the year of grace 1761 had solved this
easy vocal problem, curiously imagined to be so difficult, schoolmasters who

sinned against this high ideal of classical recitation might well reply, that to_

attempt to indoctrinate the ears of schoolboys with such delicate distinctions
would prove as hopeless as to bring out the beautiful harmony of one of
HANDEL’s operas from a hurdy-gurdy. On another point also, FosTEr’s Essay,
though victorious against GALLy, did perhaps more harm than good to the
question of orthoepic reform in the great schools. He does not always suffi-
ciently distinguish between the emphasis, or stress, or intensity of utterance,
which he rightly considers to belong essentially to accent, and the prolongation
of sound with which that intensity may sometimes be accompanied. Hence he
speaks of the effect of the accent in English being habitually to lengthen the
syllable ; whereas, if we attend to our ears, words like vapid and rdp’id, are just
as common in our language as po’tent and pa’tent, and no person feels himself
under any tendency or compulsion to assimilate the pronunciation of the first
two words to that of the other pair.

Three years after the appearance of Mr FostTeEr’s Essay, the ‘‘ Accentus
Redivivus” of Primattr appeared, the title of which seems sufficiently to indicate
that in England at least MEETKERCHE, and Voss, and GaAtty, had ‘practically
won the day, and that accents had retired from the schools, and even from the
typographic theatre in Oxford ; for in the year 1759 an edition of ARISTOTLE’S
Rhetoric, without accentual marks, had appeared under the imprimatur of
THomas RanpoipyH, Vice-Chancellor of the University. How many more
Greek books, in the same nude fashion, may have issued from the same quarter

about the same time, I do not know ; but there was certainly just cause for the _

champions of accents to take the alarm ; and so Mr Primatt marched forth, an
accentual cataphract, bristling all over with Alexandrian and Byzantine erudi-
tion, through which it was impossible to pierce him. In his learned work, he
first shakes himself free from the notion flung out by Vossrus, and the extreme
men of the rhythmical party, that accents, however they might have been
observed afterwards, were originally a musical, and not an orthoepic notation.
He then shows, by a long historical deduction, that the reading of Greek prose

always was accentual, and that nothing can be more illegitimate than to —

PLACE AND POWER OF ACCENT IN LANGUAGE. 297

transfer to prose the laws of quantitative rhythm, which belong to poetry. But
in this second proposition unfortunately, he is only half right, and entangles
himself and the whole subject in a network of the most hopeless confusion ;
for, in defining accent, besides asserting with Fosrerr, that there is an over-
bearing tendency in English to lengthen all accented syllables, and an invariable
rule in Latin to accentuate long penults, he lays it down in the strongest terms
that the acute accent necessarily lengthens the syllable on which it falls,
and that, in fact, when properly read, every accented syllable in Greek prose is
long. Nay, more, so confused are his ideas on the whole terminology of the
subject which he treats, that he actually tells us “we can hardly read a verse
in Vireit or Homer in which the rhythm does not more than once break in upon
the quantity” (p. 157), a sentence which, according to the usage of all who
write intelligibly on such subjects, is pure nonsense, or true only of such
accented verse as we have in English and other modern languages. This ex-
traordinary confusion of two things by the ancient grammarians, kept so
distinct as accent and quantity, rendered his whole discourse nugatory. To
accept accent according to this theory was to make a formal transference of
quantity from one syllable to another, and to acquire a habit of reading prose,
which, in the point. of quantity, would require to be reversed the moment a
scholar threw down Piato, and took up SopHocies. In a country where the
most elegant scholars, under the guidance of such a Titan as BENTLEY, had
already begun to look with a curious preference on everything connected with
metrical composition, such a startling doctrine could not be expected to make
converts. }
After these violent but practically ineffective efforts, the great strife about
accents in England stopped for thirty years, when in the year 1796 another re-
markable combatant entered the lists in the person of SamueL Horsey, one of
the most notable of the smgular army of erudite polemical bishops of which
the Anglican Church has been so fertile.* Into the weakness and utter un-
tenableness of the received method of reading Greek in this country the Bishop
‘casts a piercing eye, and with an outspoken emphasis calls black black, and
white white in the matter, after a fashion to which it might have been expected
that in a country where the Church has so much to say in the school, some
Serious attention might have been given. “A practice,” he says, “is adopted
in this country of reading Greek verse with the Latin accent, and this is most
absurdly called reading by quantity; and having adopted this strange practice of
reading one language by the rules of another, it is not unnatural that we should
wish to prove the practice right” (pp. 26, 27). This is indeed hitting the nail on
the head; but the strange practice, like many strange things in England, still

* On the Prosodies of the Greek and Latin Languages. Lond. 1796. The author's name was
not given on the title page.

VOL. XXVI. PART II. 4H

298 PROFESSOR BLACKIE ON THE

continues, and we still make ourselves ridiculous by awkward endeavours to
prove that what is altogether unnatural and monstrous is justifiable and even
beautiful. How is this? Not only, I believe, because the patient was self-
willed and obstinate, but because the physician who pronounced a most scien-
tific diagnosis of the disease had not the sagacity to discover the proper cure.
He suggested a cure more flattering to his own ingenuity than true to the
state of the case, or beneficial to the patient. He was as original as Dr
GALLY, in a more subtle indeed, but not in a more practical way. GALLY’s
originality, as we have seen, consisted simply in calling everything on the doc-
trine of Greek accents irrational and absurd which was contrary to his
orthoepic habits or fancies, and nonsuiting it, without more ado, as a defaulter 7
foro rationis. Hors.ey, with that respect for historical fact and erudite testimony
which became a bishop and a theologian, admitted the doctrine of accent in its
full weight, as an element of which no sane reasoner on the matter of Hellenic
orthoepy could get rid; but, in order to explain its operation as part of the
harmony of Greek verse, he invented a theory altogether novel and altogether
arbitrary, which nobody had ever proposed before, and which nobody, we may
feel pretty certain, will ever propose again. This theory consists simply in
acknowledging the Greek accents, as we find them in the books, as the law for
the pronunciation of the separate words, but refusing to allow them their
natural force under certain rhythmical conditions. Thus, he says, that at the
end of a hexameter verse such a word as eOyxe must be pronounced €Oyxe,
because the last syllable of a hexameter verse being long, the accent, according
to a well-known canon of Greek orthoepy, must fall on the penult! Now, the
objection to this theory is threefold—(1.) It is not true that the last syllable of
hexameter verse, as €0yxe, is long; it is short, and the time is filled up by the
pause which belongs to the end of the line, like a rest in music ; (2.) The theory
proceeds on a supposed connection between prose accent and rhythmical
emphasis, which is fundamentally false ; and (3.) The whole theory is a figment
spun out of the brain of the writer, without a shadow of authority from ancient
srammarians and metricians. This being so, the natural consequence fol-
lowed ;—the book explained nothing, and changed nothing. If everybody
could not answer it, nobody cared to understand it.

Immediately upon the back of the learned Bishop’s treatise, in 1797, appeared
a little book entitled “Metron Ariston; or, a new Pleasure Recommended,”
with a ruffed and bearded effigy of MEETKERCcHE fronting the title-page, and a
motto which sufficiently indicates the temper and direction of the writer—

“ Tollite barbarum
Morem perpetuum, dulcia barbare
Laedentem metra, que Venus
Quinta parte sui nectaris imbutt.”

—_ ee -

PLACE AND POWER OF ACCENT IN LANGUAGE. 299

This book was not written by a scholar, but by a man of taste and
vivacity, and a gay self-reliance which stands him in good stead against a
whole host of scholastic cuirassiers. In poimt of tendency and contents, this
book is nothmg more than a repetition of MEETKERCHE and Voss, and
those writers who have maintained the right of rhythmical as opposed to the
accentual recitation of Greek and Latin verse; but the striking fact which
the title of the book suggests is, that the masters and teachers of the great
English schools, who certainly could not be accused of paying any partial
attention to accent, were the very persons who had so thoroughly ignored the
practice of rhythm in their teaching, that it was a discovery to the author of
the book to find that there was such a thing as rhythmical reading of classic
verse ; and this discovery, with a prompt philanthropy, he hastens to com-
municate to the ingenuous youth of the nation under the inviting name of “a
new pleasure.” This entirely agrees with the complaint which we have just
heard the right reverend Bishop make with regard to the absurdity of reading
Greek poetry with Latin accents and calling it reading by quantity. No wonder
that clever schoolboys on occasions should begin to dream that the learned and
reverend doctors, by whom their ears had been indoctrinated in the unpleasant
mysteries of long and short syllables, at bottom knew less about the matter
than they might have known themselves with the help of a little unsophisticated
juvenile instinct. And accordingly the writer of ‘ Metron Ariston ” tells us
that “he always indeed had an idea that our very anomalous and irrational
way of reading Greek and Latin poetry was founded on error ; yet, from indo-
lence, he had conformed, though reluctantly, to the general practice, because it
was not his business to examine the error and seek its remedy.” But what he
did not seek for, he goes on to tell us, like WorcEsTER’s rebellion, came in his
way, and he found it; and the good Hermes, on whom he stumbled to direct
him in his rhythmical wanderings one day, was a learned Italian ecclesiastic,
while they were walking together in the Campo Vaccino at Rome one morning,
and talking of Horacez, and quoting the well-known line—

“ Tham forte vid sacra sicut meus est mos.”

The full musical weight with which the learned Italian recited this verse struck
the Englishman with a pleasant surprise; whereupon the priest, divining the
cause of his satisfaction, began to expound to him the correct theory of classical
recitation according to MEETKERCHE, “the great ambassador of a little state.”
Against this true doctrine, without which verse had no meaning, and lost more
than half of its suavity, the English scholars and schoolmasters were in the
systematic habit of sinning, by pronouncing éwus, for instance, a horse, as if it
Were aequus, equitable—by shortening the final syllables of all words, and pro-
nouncing dém’inds as if it were déminds and sacrd, the ablative singular, like

300 PROFESSOR BLACKIE ON THE

sacrdé the nominative plural ; and by turning anapests into dactyles, dactyles into
tribrachs, spondees into trochees, iambi into pyrrhics—in fact, doing everything
that could be done systematically to turn order into disorder in this region, and
“by this most abominably absurd custom, destroying at once both sound and
sense, and seeming to sin from a love of the very ugliness of sinning.” These
are hard words, but not, in fact, one whit more strong than those which we
have quoted from the English Bishop; nor is it possible, indeed, to conceive
anything at once more unscientific, more tasteless, and more unpractical than
the way in which prosody and rhythm have been handled in the great English
classical schools up to the present hour. On this point, certainly, the author of
“Metron Ariston,” a single light horseman, could triumphantly ride up and
attack without fear a whole army of big blundering and self-contradictory
hoplites. As to accents, however, about them he wisely said nothing; but
allowed them quietly to lie in the state of suspended animation to which they
had been condemned by his patron-god MrErxercue. If these mute, mysterious,
little oblique and curved lines were ever to revive into speaking significance
at the touch of some philological wizard, the author of “Metron Ariston”
certainly did not possess the secret for their disenchantment ; nor, indeed, if he
had possessed it, would he have cared to use it; for the accents, whatever
virtue they might possess, could add but little to the luxury of the new
rhythmical pleasure which he had discovered.

But what were the great German scholars doing all this while,—the HEynEs,
the WotFs, and the HERMANNS, the founders of that stable and splendid edifice of
philological learning which has placed Germany in the van of erudite and
thoughtful research during the whole of the present century? In the preface
to the second edition of his Odyssey, WoLr remarks that in the matter of the
accents, “the editors of the previous centuries had shown a great laxness of
procedure, a fault which had commenced with so illustrious a name as HENRY
STEPHANUS, who in this respect had declined from the accuracy of his prede-
cessors, CHALCONDYLUS and Atpus.” And after a few remarks on points of

detail, follows a remarkable witness to the practical disuse into which accents —

had fallen in Germany just as in England towards the end of the last century.
“Tn fact, no person now-a-days—and for many centuries back—ever hears
a Greek accent; and only a few, indeed, seem to believe that the doctrine of

the grammarians on this subject is a thing that belongs to a complete course of

2936

teaching. This passage is decided as to the general disuse of accents among
the Germans in Wotr’s time; but the phrase sect vielen Jahrhunderten is certainly
too strong ; for the works of MEETKERCHE, VossIus, and HENNINIUs, are sufficient
to prove the living predominance of the Byzantine tradition in respect to

* These extracts are taken from an historical review of the opinions of scholars about accents in

Wacner’s “ Accent Lehre.” Helmstadt, 1807.

:
'

—

i
x

PLACE AND POWER OF ACCENT IN LANGUAGE. 301

accents in the scholastic practice of their time. An equally emphatic declaration
in favour of accents is made by HERMANN in his famous work ‘‘ De emendandd
ratione Grammatices Greece ;*” but whether these two illustrious scholars
contented themselves with publishing an authoritative manifesto on the neces-

sity of maintaining accents as an inherited doctrine of genuine Hellenic ortho-

doxy, or took any steps to put their views into that practical shape which alone

could give them significance to articulate-speaking mortals, I have not been
able to learn. Certain it is, however, that the stagnant waters of the schools—
in Germany much more apt than in England to deduce practice from principle—
began to be moved in this matter; and, according to information which I
have from continental scholars of high reputation, the accents are now pro-
nounced in a great number of the best German gymnasia. I myself, some
forty years ago, heard Professor BorcxH, in Berlin, reading the Iambic verse of
the tragedians with a distinct and well-marked observance both of accent and
quantity. The matter appears to have been left pretty much to the arbitration
of the scholastic world ; and we may feel perfectly convinced that the natural
conservatism of teachers would have resisted all change in this matter, unless it
had been incontestably proved that the change carried with it the double
advantage of scientific truth and practical convenience. Whilst the matter was
thus not only fairly ventilated, but to a large extent embodied in the scholastic
practice of Germany, in England not a single step seems to have been taken
either to the recognition of the principle or the settlement of the practice of
Greek accents. The well-known declaration of Porson, no doubt, in a note to
the Medea,t gave the imperial amprimatur to certain traditional marks as a fact
on paper, and of course put a stop for ever to the inchoate practice of printing
Greek books without such marks; but it was a fact which seemed to remain as
mysterious as a row of hieroglyphics on an obelisk before the great decipherment

of CHampot.ion. In fact, to use Scripture language, notwithstanding the authori-

tative dictum of the great Cantab, the doctrine has remained in England up to the
present hour a meaningless thing, ‘‘ having a name to live while it is dead.” In
Scotland, indeed, a country too much accustomed slavishly to follow English
authority in classical matters, twenty years ago I published a short protest
against the gross inconsistency and grave practical grievance of inculcating
rules about a host of mysterious marks which gave neither ideas to the intellect
nor direction to the ear ;{ it had become clear to me as sunlight, not only from

‘Meditation on the nature of the case, but from an accurate study of the ancient

* Ch. xiii. De accentu.

t “Si quis igitur vestrum ad accuratam Grecarum litterarum scientiam aspirat, is probabilem
sibi accentuum rationem quam maturrime comparet in propositoque perstet, scurrarum dicacitate et stul-
torum derisione immotus.”

Sabo! The Pronunciation of Greek; Accent and Quantity; a Philological Inquiry. Edinburgh,

VOL. XXVI. PART II. 41

302 PROFESSOR BLACKIE ON THE

grammarians, that Greek accents contained the two elements of elevation and
stress of voice, and are, in fact, practically identical with the accents in English,
Italian, German, and other modern languages. And this truth I have carried
out in practice for twenty years with increasing profit and satisfaction. In
England, however, as was to have been expected, no attention was paid to a
Greek argument coming from the north side of the Tweed ; and, accordingly, in
the next work, that of CHANDLER,* which issued from the Oxford press, we find
the whole subject flung back into a grim limbo of despair, and involved in a
mantle of impenetrable darkness. In the preface to his work, this author goes so
far as to assert that neither Porson nor any other scholar, ‘‘ while sanctioning
the practice of accentuating Greek by their example, has condescended to
justify it by sound and conclusive reasons. Porson specially, it is hinted in
terms more vigorous than polite, “ gave assertion for proof in the matter,
actuated partly by his contempt for WAKEFIELD, who happened to entertain a
different opinion from his own.” Then he goes on to proclaim the utter hope-

lessness of being able to arrive at any certainty with regard to the meaning of

accents ; it is not even certain that they did not “ indicate the length or short-
ness of syllables ;” he denounces “the absurdity of those who perpetuate in
writing a something to which they never attend in reading, and who persist in
ornamenting their Greek with three small scratches, the very meaning of which
is doubtful and perhaps unknown,” and laments in the most pathetic terms his
own evil destiny in having had anything to do with the tangled disorder of
“these troublesome appendages.” .

“ There’s something wrong i accents—cursed spite
That ever I was born to set it right !”

In fact, it appears not a little extraordinary that a writer who uses such
strong language, should not have followed out consistently the practice of his
predecessor Hennintvs, and flung the whole cargo of Byzantine lumber over-
board ; for what task can be imagined more irksome and more fruitless than to
spend long months of painful inquiry, with fret of brain and vexation of vision,
upon every mappik and dagesh of a gospel in which the writer does not believe 4
Almost contemporaneously with this remarkable book of Mr CHANDLER, ap-
peared an interesting paper on accent and quantity by Professor Munro of
Cambridge.t The occasion of this discourse was a Latin inscription in accentual
hexameters from Cirta in Numidia, and supposed by the professor to belong to
the third century of our era. In commenting on these verses, of course, the
writer was led to explain both what accent meant, and how it came to pass

* A Practical Introduction to Greek Accentuation. By H. W. Cuanpurr, M.A. Oxford, 1862.
t On a Metrical Latin Inscription, copied by Mr Buaxxstey, at Cirta— Transactions of the
Cambridge Philosophical Society,” vol. x. part 2. 1861.

prey

PLACE AND POWER OF ACCENT IN LANGUAGE. 308

that accentual verse, at so very early a date, came to usurp the place of quanti-
tative, which only we now acknowledge as classical. In making this explana-
tion, Professor Munro lays down the following propositions :—

(1.) That the acute accent of the ancients was a mere elevation of the voice,
without any stress on the accented syllable.

(2.) That in the composition of Greek and Latin verse, the metre was
determined by quantity alone, and that accent had no influence on it direct or
indirect.

(3.) That, nevertheless, the quantity of syllables was a matter which swine-
herds in the days of Homer, and ploughmen in those of Puautus, had imbibed
with their mother’s milk, and could discriminate with the nicest precision.

(4.) That by some strange and, to us, unaccountable process, the nature of
the Greek and Roman accent was.suddenly changed in such fashion that, from
being a mere raising or sharpening of the tone, “‘it became a stress,” “a mere
stress,” ‘‘ a stiff and monotonous stress,” a stress which is always accompanied
with “the lengthening of the quantity,’ having nothing in common with the
genuine classical accent except the name ; and that by this strange and inexpli-
cable plunge, the accentual poetry of the medieval hymns, and the whole of our
modern metrical system, so early as the third century had started into recog-
nised existence.

So much for the theory of the matter. With regard to the strange and un-
scientific practice of the English great schools and colleges, the following
passage is notable :—

“Tt appears from what has been said, that we English, in reading Latin,
place the accent generally, but by no means always, on the proper syllable.
But then, we have entirely changed its nature, making it a mere stress, instead
of a simple raising of the tone, without any lengthening of the quantity. And
Praciius and his cotemporaries already did the same. From them, and their still
more degraded descendants, the Italians, and other western nations, we inherit
this debased accent, which had usurped and overthrown the rights of quantity.
In the second line of the A‘neid we read Jtaliam fato profugus with the accent
on the right syllable ; but on the same principle we ought to say—and Pract-
Lius, indeed, and the Romans for centuries after him, did say—Lavindque, with
the accent on the second a. We flatter ourselves that we thus preserve the
quantity, but that is a mere delusion. It we feel by a mere mental process.
Whether we pronounce prdfugus or profigus, quantity is equally violated. In
the same way we read Greek with this debased Latin accent, and fancy that
we preserve the quantity while sacrificmg the accent. The modern Greeks
read old Greek with the ancient Greek accent, debased in the same way into a
mere stress. We think them, they think us, in the wrong; and in different
ways we are both equally in the wrong. Myvw deide Oéa in an English or

304 PROFESSOR BLACKIE ON THE

Italian, and pjvw dee Oecd in a modern Greek mouth, are equally remote from
the accent and quantity given to the words by Homer or DEMOSTHENES.”

It will be observed that this passage touches exactly on the same absurdity
which, sixty years earlier, had roused the sprightly mdignation of the author of
“Metron Ariston,” and the grave episcopal censure of Dr Horstey.

In the “Cambridge Journal of Philology,” vol. i., for 1868, appeared an
article on the English pronunciation of Greek, by W. G. Ciark, then public
orator, Cambridge. Mr Cuarx is a scholar particularly well entitled to
speak on this subject, both from his general accomplishments, which are far
from being confined to the ordinary routine of an English classical scholar, and
specially from his having travelled in Greece, and taken note of the actual
accents of the language, as at present spoken by the people. In theory, Mr
CLARK entirely agrees with Professor Munro, that the ancient Greek accent
consisted merely in the elevation of the tone, while the accent of the modern
Greek includes “a stress precisely like our own, which is given by prolonging
the sound, as well as by raising the note.” When it falls upon a syllable it
lengthens the vowel except before a double consonant. Thus dAdyos is pro-
nounced AWyos, dvos wvos, and so forth. With regard to scholastic practice, Mr
CiaRK is of opinion that, while our English Greek vocalisation is altogether
anomalous and indefensible, and must be abandoned, the present system of
reading Greek with Latin accents should not be touched, because the modern
system of accentuation is widely different from the ancient, and its adoption
could only tend “to confuse such ideas as we at present possess of the rhythm
of ancient Greek verse.” And again, “It is impossible in practice to recur to
the ancient system of accentuation, supposing that we have ascertained it in
theory. Here and there a person may be found with such an exquisite ear,
and such plastic organs of speech, as to be able to reproduce the ancient dis-
tinction between the length and tone of syllables accented and unaccented, and
many not so gifted may fancy that they reproduce it when they do nothing of
the kind. For the mass of boys and men, pupils as well as teachers, the dis-
tinction is practically impossible.” So Mr Ciark leaves us, so far as action is
concerned, in a plight little better than that in which we were left by CHANDLER,
—not enveloped, indeed, in impermeable mystery, but clogged with impracticable
fetters, and groaning under a yoke of grammatical tradition which neither we
nor our fathers were able to bear.

A strange and a grateful contrast to the general current of English scholar-
ship on this subject is presented by Mr Getpart, of Balliol College, Oxford, in
his interesting and ingenious book, entitled “The Modern Greek Language in
its Relation to Ancient Greek; Oxford, 1870.” In the third chapter of this

work, the author states views with regard to accent and quantity which lift

him completely out of what has always appeared to me the sort of enchanted

PLACE AND POWER OF ACCENT IN LANGUAGE. 305

circle of confusion and delusion in which English scholars are involved the
moment they approach this subject. Mr GeELparr is a decided advocate for
accents, both in theory and practice, and he says roundly that “our prejudice
against accents is for the most part insular, and deepened, to boot, by the pecu-
liarities of our own insular pronunciation.” He blows to the wind in a single
sentence the vulgar error of English scholars, so often noticed in these pages,
that accent has the necessary effect of lengthening the syllable on which it
falls, the accented syllable in English being, in fact, as often short as long, as in
gét'-ting, pick'-ing, while a long syllable is often unaccented, as findncial, fertile,
a priori, in which last the first syllable is nearly always pronounced long, in
spite of the fact that it is short in Latin. It is accordingly a complete delusion
to imagine “that the Latin accent is either an indispensable or an infallible
device for marking the right quantity of Greek syllables.” With regard to
accent, he makes the just remark that the raising of the note, and the increase
of the stress generally go together. He farther denies altogether—and on this
point he is a witness of great authority—that the modern Greeks always, or
even in a majority of cases, lengthen the syllable on which the accent falls ;
and in regard to the relation of accent and quantity, he shows that neither is
modern poetry always governed by the mere spoken accent, nor is ancient poetry
altogether regardless of it, but that the real regulator, both of ancient and of
modern poetry, though in very different ways, is RuytHM, which is determined
by the musical beat. How far the spoken accent was heard, as it were, through
the rhythmical movement, depended principally upon whether the verse was
sung or recited. In pure singing there might be heard only a faint glimmer of
the spoken accent; in prose it was the prominent element, and directed the
flow of the period ; while between these two extremes there might be several
intermediate styles of utterance in which the spoken accent was more or less
prominent, according to the greater or less approach of the style of recitation
to colloquial prose.

It will not be difficult, after this long and strange historical survey, to sum
up the conclusions to which, by the consideration of the various facts and argu-
ments, we are inevitably led. We find ourselves, in fact, after more than three
centuries of confusion, one-sidedness, and hallucination, arrived at a point of
view where no fact or principle, necessary to a just conclusion, is concealed,
and all apparent contradictions find a happy conciliation. In particular, the
whole history of the controversy displays the fact that in one form or another
quantity is the bugbear, and that from Voss and MEETKERCHE, to Munro,
CHANDLER, and CLARK, a sacred regard for the rights of metre is the apology
for the monstrous invasion of the province of Greek by Roman accents. But
those who have attended to the course of our argument and historic survey will

VOL. XXVI. PART II. 4k

306 PROFESSOR BLACKIE ON THE

easily perceive that the interference of Greek accents with the laws of Greek
metre is a pure hallucination ; inasmuch as—

1. It has been amply proved that in the case of individual words the pre-
dominance given to one syllable by the stretch, stress, or emphasis of the voice
with which the acute accent is naturally accompanied, has no necessary tendency
to lengthen the syllable on which it is laid. Through the whole argument of
those who oppose Greek accents a confusion runs between two things, which
in this matter must be kept carefully apart—a confusion between a short sylla-
ble unaccented compared with the same syllable accented, and a short accented
syllable with along syllable accented. When the three terms jyépa, jpépa,
and ypy’pa are compared, the middle syllable of the middle term, while it is more
prominent, and may be in some degree longer than the same syllable of the
first term, is decidedly short when compared with the same syllable of the
third term. If, therefore, any short syllable, whether in Greek or English, on
which the accent falls, is in danger of being pronounced long, it arises not
from the nature of the case, but from the ignorance, carelessness, or stupidity
of the teacher ; and, in fact, a great part of the strange confusion which has so
long prevailed on this subject may not unreasonably be traced to the want of the
directing presence of a living rhetorical and musical culture in our great English
schools and colleges.

2. The second great element of confusion which has been introduced into
this matter is the gratuitous and altogether unauthorised assumption, that
because our metrical composition follows the laws of spoken accent, therefore
in Greek and Latin the same law was necessarily observed. In the writings of
HYPHAESTION and of those who lay down the canons of classical verse, there is
not a single word said about the spoken accents ; and the sure inference is, that
in metrical composition they were, as Professor Munro justly remarks, systema-
tically ignored, or, if attended to at all, only in a subordinate, exceptional, inci-
dental, and even accidental way. Nothing, therefore, could be more mistaken
than the attempt of Horsey to give a new theory of Homeric scansion, founded
on the doctrine of the spoken accents. On what principle, then, it will be
asked, was the.Greek poetry written? Can it be supposed that a nation of
refined taste and high culture could be delighted with the barbarism of pronounc-
ing words, one way in prose, and another way in verse? We answer, there is
nothing at all strange in this supposition ; and that, whether it appear strange
or not, it was certainly the fact. To understand this, instead of transferring
the laws of our modern poetry wholesale to the poetry of the Greeks, let us
rather transfer ourselves from an age of books, reviews, newspapers. and read-
ing-rooms into an age where there was no such thing as books or reading at
all, where prose composition was altogether unknown, and where every com-
position, not purely ephemeral, was made to be sung, and had its existence

eo © ~e

PLACE AND POWER OF ACCENT IN LANGUAGE. 307

only in the element of music. Now, we need not at the present day set fortha
formal proof that Homer and the pre-Homeric teachers of Greece were not
dvayoora but dowoi, and that all hexameter verse, the current form of the
oldest Greek metrical compositions, was originally sung, and not recited.
Under these conditions, it naturally conformed to the laws of musical compo-
sition ; and what these laws were, especially in relation to spoken accent, it is not
difficult to realise. What music principally demands from poetry is a mass of
rich and full vocalisation, to correspond with the measured flow of the notes ; for
the vowels are the musical element in human speech, and especially the deep
broad vowels pronounced long, and not rapidly rattled over. This element,
therefore, was naturally preserved in the first place: that is to say, Hellenic
poetry was founded on quantity. But what of accent? The rhythmical march
of speech adapted to music, as every one knows, is secured by the element of
equality expressed in the succession of equal spaces of sound, marked by recur-
rent emphasized pulsations; these pulsations constitute what is called the
musical accent, or beating of time, as it is vulgarly called. Now, it certainly
might have been desirable to make this rhythmical accent of the music cor-
respond in every case with the spoken accent of the words ; but this was not
done, for the very simple reason that the choice of poetical language would
have been too much fettered by the constant double demand on the poet of
conformity in every case, both with the spoken quantity and the spoken accent.
Nor should this appear at all strange. As it is, we see how often HomER—as
in a&varos and other words—is obliged to put an artificial length upon tribrachic
feet in order to get them admitted into the dactylic march of his verse ; and how
impossible it would have been to compose a long poem under the strict law of
both quantitative and accentual conformity, we may see from the fact, that, in
our own poetry, we have contented ourselves with fettering one of the elements
_ and leaving the other free ; that is to say, that, while we never, or very rarely,
allow our spoken accent to clash with the rhythmical beat, we constantly take
the liberty in our sung psalms and songs of drawing out short syllables to any
length, and skipping over long ones with any amount of metrical celerity.
Here, therefore, the Gordian knot is untied: the Greek poetry made to be
sung is governed by quantity, the musical element of language; the modern
poetry made to be read is governed by accent, the colloquial accent. What
Nature, or rather the necessities of Art, have kept asunder, let no man bring
together. Let no man imagine that colloquial accents, whether Greek or
Roman, can possibly come into collision with the laws of a poetry so essentially
musical in its character as the Greek.

3. But the ancients, it will be said, though their poetry was all musical in
its birth, and a verse had no meaning except as sung, certainly did recite their
poetry at an early period. Of course; and in this case it is obvious, that a

308 PROFESSOR BLACKIE ON THE

poetry constructed as part of the musical art was to a certain extent put out
of Nature the moment it was translated into the region of spoken verse. In
this case a collision between the musical beat and the accented syllables was
unavoidable, and some sort of compromise would naturally be the result. This
compromise, however, would on the whole be decidedly to the advantage of the
musical rhythm, as opposed to the colloquial accent. For metre, as we have
seen, was metre only in virtue of the regularly recurrent musical beat ; and to
abolish this was to destroy metre, and to turn verse into prose, as, in fact, we
often hear English schoolboys do, when reading Horace, and as the modern
Greeks do when they read Homer accentually. But that the ancients could
not have done this is manifest both from the prominence of music in their
national culture and from the effect of the rhythmical stroke in lengthening the
shortest vowels, even in the verse of VireiL, which certainly was not sung.
The poet who wrote

Liminaqué’ laurusque Dei,

must have had his ear tuned to the march of a verse which gave that marked
preponderance to the first syllable of a foot, which is musically given to the first
note of a bar, and which allowed the license of lengthening a short vowel in such
a position after the example of Homer, specially before a word beginning with
a liquid. MErETKERCHE and Voss were therefore right in reading classical verse
mainly by this rhythmical beat, and practically disregarding the spoken accent.
It does not follow, however, that though the rhythmical accent remained
dominant even in spoken verse, it therefore exercised an exclusive sway. In
many cases, of course, there would be no clash, and this, indeed, regularly
happened in the two last feet of a Latin hexameter. But im other cases, where
a clash did occur, the occasional bringing forward of the spoken accent might
serve to break the monotony of a merely musical rhythm, and cause it to
approach nearer to the march of dignified prose eloquence. Thus, the first line
of VirGiL may either be accented

Arma viumque cand’ Trojeé’ qui primus ab oris,

or
Arma viumque can'o Tro'jae que primus ab oris ;

and in both cases the true quantities are preserved ; but in the second method
the spoken accent is allowed to control two words to the prejudice of the
musical beat, by whose regular recurrence the hexameter verse was originally
framed. In this way it was quite easy to recite Latin hexameters or Greek
iambics in such a manner that, while the rhythmical beat mainly ruled, and no
short syllable was ever heard where the music had a long note, the spoken
accent te which the ear had been habituated im conversation did neverthe-
less generally shine through, and in special cases assert itself with that natural

PLACE AND POWER OF ACCENT IN LANGUAGE. 309

emphasis which subordinates rhythm only to aid expression, and to prevent
monotony.

4. It will now be evident how entirely Professor Munro was mistaken when
he expressed surprise at the fact, that, while the rudest boor in the days of
PLautTus was familiar with the exact laws of quantitative metre, even well-
educated gentlemen of the middle class before the time of CoNSTANTINE were
apparently unable to write anything but accentual metre, constructed on the
same principle as the Byzantine orixo. wodurixoi. The rudest boor, no doubt,
could distinguish a long syllable from a short, and could discriminate the penul-
timate vowel in péter and md@ter in a way that seems impossible to the gross
ears of some of our English teachers. Our own peasants will distinguish godt
from goat, or god from goad, exactly in the same way ; but it will require more
than a rhetorical flourish from Cicero to prove that the peasants of Italy, or
even Attica, at any time were perfectly master of the complete doctrine of
quantity as taught in the musical schools. For it must always be borne in
mind that the practice of these schools was to a certain extent artificial ; it
was founded on certain concessions which the currency of common life had
made to the necessities of art; and the common people, whose ears were
trained mainly by the spoken accent, could .not be expected either to
compose verses in neglect of that accent, or to sympathise fully with its
neglect in the case of verses composed by cultivated poets, except in so far
as their own education had kept them in living connection with those schools
of music from which the cultivated poetry had emanated. Now, in the best
ages of Greece this living connection naturally existed ; and the effect of custom
and association would be such, that no other verses but those composed on the
original quantitative principle would be recognised as legitimate even by the
vulgar ear. But the moment that a great national decay commenced, and
schools of popular culture-were neglected, from that moment the common
people, left to themselves, if ever they tried poetical composition, could do so
only in obedience to the instinct which governs all poetry not intimately associated
with the musical art. Poetry now became a species of measured conversation to
which laws were given by the spoken accent, and where the fixed musical
recurrence of long and short syllables was systematically ignored. In this
change there is nothing strange or mysterious; on-the contrary, it was the
natural, and, we may say, necessary consequence of passing from a musical to a
colloquial epoch in literature; and as a fleet-footed man, when he leaves the
ice and takes off his skates, passes to a kind of locomotion governed by different
conditions and subject to different laws, so a people, shaken loose from all
musical tradition and left to form a poetry for itself, will infallibly fall upon a
form of verse in which the musical value of vowels will be sacrificed to the
familiar control of accentually preponderant syllables.

5. One word remains on the question of scholastic practice, which has

VOL. XXVI. PART II. 41

310 PROFESSOR BLACKIE ON THE

been such a bugbear to our teachers. Now, with regard to this problem, it is
one of those to which, as GELDART says, the old adage applies, solvitur ambulando.
What appears impossible in theory, is often easy in practice. If you wish to
learn how to use your legs, just rise up and walk. If you imagine that there is
any difficulty in saying Zoxpa’rns without saying Ywxpa’rys, or bdn’us without
saying bonus, just put yourself under a master of elocution for five minutes, and
you will shortly be drilled out of your difficulty. But why should the ears of
teachers be haunted by such a hallucination as that by placing the Roman
accent on the penult of all dissyllabic words, they are furnished with some sure
spell against the violation of quantity? Is it not quite evident, rather, that the
short quantity of the first syllable of Buds, a bow, is much more easily preserved
by the natural oxytone accent than by the Latin accent Bi’os on the penult ?
And if the quantity of the long penult in the verb d.arpiéw is more effectively
brought out by the accent on that syllable than if it had been on the last, is it
not manifest that the same syllable, being short in the substantive duarpity, is
more certainly pronounced short—according to the. argument of the Latinising
Hellenists themselves—with the native oxytone accent than with the imported
Latin one? Take, again, the word kayapa, a vault, where all the vowels are
doubtful, and where, of course, the quantity of each syllable can be recognised
only by utterance. According to the current method, the accent, laid on the first
syllable of this word, should inform me, that the syllable is long by virtue of the
stress, and it does inform me also, if I am to believe my ears, that the other two
syllables are short. But three parts of the information thus given are false ; for
the accent is not on the first syllable, and the quantity of the first syllable is short,
and that of the last long. On the other hand, if I pronounce the same word
according to the principles laid down in this paper, I learn not only where the
accent is, but that the two first syllables are short, and the last long. The fact of
the matter is, that, while the Greek accent, rightly placed, informs the ear rightly
both as to the accent and the quantity of the syllables of which a word is com-
posed, the Latin accent inverts and perverts both, and teaches, with regard to
accent and quantity, only what must be unlearned. The opponents of accents,
who absurdly call their Latinising method the quantitative pronunciation of
Greek, ought to bear in mind that, in practical teaching, next to pronouncing
the long syllables long and the short short, the best way to teach quantity is to
pronounce the accent, which either stands upon the long syllable and favours
its prolongation, or stands in such a definite relation to that syllable that the
quantity of the unaccented syllable is known from the place of the accented.
But the great practical difficulty to which teachers allude is, perhaps, rather
rhythmical than prosodiacal. The pronunciation of the Latin accent, says Mr

CLARK, is the only way we have of teaching our pupils to appreciate the

measure of classical verse. Abolish the Latin accentuation of Greek prose,
and you turn the organ of Homer into a hurdy-gurdy. Now, with regard to

—-

PLACE AND POWER OF ACCENT IN LANGUAGE. 311

this matter, I would observe, in the first place, that if the young gentlemen
who usually come to our universities were to lose all the rhythmical apprecia-
tion of Greek verse that really lives in their ears, and not merely in their
understanding, they would lose little that is worth keeping. For what are the
facts of the case? The observation of the Latin accent facilitates the rhythmical
reading of the two last feet of a hexameter verse; this is an accident of the
Latin language, that is all. But not even in the reading of Latin does the
reading, according to the Latin prose accents, prevent the constant occurrence
of a clash between the spoken accent and the rhythmical beat. Inthe Ovidian
pentameter such a clash must always occur twice, and in the two most marked
places of the verse, And, if the absence of the oxytone accent causes this
opposition in Latin, is it not strange that we should banish this same accent
from its natural place on a Greek word, in order, as we say, to avoid, but
actually in a great number of cases to produce, a collision between the rhyth-
mical beat and that accent? Take, for instance, this second line from “the
Wasps” of ARISTOPHANES—

“ Dvd\akny katahvew vuKreowynv SudacKomer,”
poy BEL,

and it is plain that in the only two places where a clash does occur between
the spoken accent and the rhythmical beat, according to the Latinised accent,
that clash disappears the moment the words are read according to their natural
Greek accentuation. And so,not only in Iambic verse, but in every verse whatever,
the introduction of the Latin accent must jar with the rhythmical flow of the
line wherever the rhythmical stroke falls, as it constantly does, on the last
syllable of a word. This practical objection therefore vanishes in smoke. That
eross-eared and ill-trained persons may be enabled to receive the harmonies of
the two last feet in a Homeric line, with a little less trouble, or with no trouble
at all, no wise educator can deem a sufficient reason for invading the whole
inherited intonation of the finest language in the world, with sounds which,
however proper on the banks of their native Tiber, on the banks of the Ilissus
must be felt to be a gross barbarism. The rhythmical objection from the prac-
tical side is, in fact, only an ingenious apology to cover carelessness, to prop
prejudice, and to mask with an attitude of apparent utility a pedagogic pro-
cedure, alike unscientific in principle and self-contradictory in practice.

Finally, if those who delight themselves in exaggerating imaginary difficul-
ties have any honest desire to see how they disappear in the actual business of
teaching, let them come to me; for I am a practical man, and speak from the
experience of half a lifetime. I teach Greek on the principle that the ear is the
natural and legitimate organ which must be addressed in the first place. I
pronounce every word according to its just accent and quantity, allowing its
own natural emphasis to sway the proper syllable of the Greek word, just as
the Latin accent emphasizes the proper syllable of the Latin word, taking

312 PROFESSOR BLACKIE ON THE

special care at the same time that in no case shall the emphasis of the accent
be drawn into a prolongation of a short vowel. In the matter of quantity, I
allow length by position to be pronounced short, according to the English habit,
partly because I do not feel sure that this length was anything but a metrical
license unknown to prose, partly because I should not think it advisable to
encumber the English lighthorseman with a greater weight of heavy Spondaic
armour than he can conveniently carry. On the elevation of tone which natu-
rally accompanies the stress, and indeed always seems to have done so at the
end of a clause, I do not curiously insist, the accent being sufficiently
marked without it. As little do I endeavour to distinguish between a long
accented syllable, as in pjvn, and a circumflex, as in waAdov, though I have not
the slightest difficulty myself in bringing out the combination of rising and falling
inflexion on the same syllable which the circumflex properly denotes. Thus, in
the reading of prose, which should be continued assiduously for six months or
a year before poetry is meddled with: I then take up Homer, and forthwith
intimate to my students that, as the whole doctrine of Greek metres was a part
of the science of music, it necessarily followed the laws of that science, and
can be understood only by an entire subordination or sinking of the spoken accent
in the first place, and a recitation according to the regularly recurrent beats of
the rhythm. This, which teachers imagine to be so difficult, is one of the
easiest things in the world. Most human beings have ears, and can beat time.
Even serpents, and elephants, and dancing bears can do this. And in order
that the rhythm may be thoroughly worked into the ear, I have no objec-
tion even to what may be called a little sing-song at starting ; but the pupil, of
course, as he advances, must be trained to counteract the monotony of mere
rhythm by that variety which a proper attention to expression and punctuation
produces. In this way, the whole perplexing and tedious doctrine of accent
and quantity is learned from beginning to end by the ear ; the pain of prosody
becomes a pleasure ; accent and quantity learn to observe their proper bounds,
each, happy in his recognised domain, forgetting all thought of making a hostile
invasion into the territory of the other. The only difficulty in the matter arises
from the necessity of teaching a number of thoughtless and idle young men to
unlearn all that lumber of false quantities and false accents which has either been
systematically built up, or carelessly allowed to accumulate in the schools ; but
this is a difficulty which it is in the power of schoolmasters, and of schoolmas-
ters alone, radically to remove. And I feel convinced that, so soon as a radical
reform in this matter shall be seriously undertaken by teachers, not only will
the inculcation of classical Greek be much facilitated, but the organs of utter-
ance being rendered more flexible and more amenable to training, will accom-
modate themselves to the characteristic peculiarities of German, French, and
other living orthoepies, with an aptitude the want of which is now so frequently
lamented.

(313 )

XIV.—On the Average Quantity of Rain in Carlisle and the Neighbourhood.
By Tuomas Barnes, M.D., F.R.S.E.

(Read 17th April 1870.)

In the year 1827, I communicated to the Royal Society of Edinburgh some
meteorological journals, kept at Carlisle by the late Mr Pirr, extending over a
period of twenty-four years, viz., from 1801 to 1824 inclusive. An abstract of
these journals, with explanatory remarks and tabular results, were drawn up
by me, and read before the Society, and were afterwards published in their
Transactions. I now beg to offer some remarks to the Society on journals
kept by Dr CaRrLyLg, in the city of Carlisle, from 1757 to 1783 inclusive, by the
Rev. Jos. Gotpine at Aikbank, near Wigton, Cumberland, fourteen miles west
of Carlisle, from 1792 to 1810 inclusive, and by myself at Bunkers Hill, two
and a half miles west of Carlisle, which is situate 184 feet above the sea level,
according to the late Ordnance Survey, from 1852 to 1870 inclusive. I shall
confine my remarks to the quantity of rain that fell during the several periods
of our journals. The accompanying tables show the quantity of each month and
year included in these periods. I regret much that I am not able to givea
description of the instruments used by Dr CartyLe and Mr Goxpine ; but as
they both were gentlemen of considerable ability and of liberal education, and
devoted much time and attention to meteorology, there is no reason to doubt
either the quality of their instruments or the correctness of their observations.
Dr CARLYLE’S rain-gauge was placed in his garden, near the head of Abbey
Street, and is about the same height as the ground on which the Cathedral
stands, eighty-two feet above the level of the sea. My own rain-gauge consists
of a copper funnel, twelve inches in diameter at the top, and is inserted into a
strong tinned iron vessel, placed in a box on my garden wall, the height of the
funnel being six feet above ground. It is examined from time to time, and
particularly after a fall of rain. The water is measured by means of a glass
tube of half an inch diameter, with an attached scale of inches and tenths. By
this means, the rain that falls on a circular area of twelve inches diameter is
collected on an area of half an inch diameter, so that inches and tenths in the
tube correspond to 5+, and 57g, of an inch of rain on the surface of the gauge.
To prevent waste by evaporation, the communication between the funnel and
the receiver is very narrow; and to prevent the rain that falls within the gauge
from splashing over, the upper edge or rim of the funnel is turned upwards
from the inclined direction of the under part, so as to stand vertically, and the

VOL. XXVI. PART II. 4M

314 DR BARNES ON THE AVERAGE QUANTITY OF

top of the gauge is parallel to the horizon. I have abstracted from the journals
the quantity of rain that fell at Carlisle, Aikbank, and Bunkers Hill, during
the periods they were kept, and have drawn up in a tabular form the quantity
of rain for each month and year of these periods. I have also taken the aver-
ages of the observations, and have found some remarkable coincidences in the
results. By these tables we observe the wet and dry months of every year for
long series of years; we also observe the wet and dry years, and the wet and
dry seasons of every year.

A TABLE EXHIBITING THE QUANTITY OF RAIN OF EACH MONTH AND YEAR, FOR TWENTY-SEVEN YEARS, TAKEN FRON !
THE METEOROLOGICAL JOURNAL OF THE LATE GEORGE CARLYLE, M.D., KEPT AT ABBEY STREET, CARLISLE, RO! v

1757 To 1783.

Years. Jan. Feb. | March. | April. May. | June. July. Aug. Sept. Oct. Nov. Dec. |Q

1757 44.) 1°097) 2°117) 2°23 | 2-206} 1°014) 2°005| 3:102| 544] 1°457| 2°703| 1111
1758 °832| 2°9 1°319| 1°59 | 1152) -759) 5°66 | 1774) 2°254) 2°46 | 2°196) 3354
1759 1°348| . 284) 27099) 1°479| -668) 3°773) 1°76 | 2°368) 3°016| 4°137| 1494)" 525
1760 1°722| 2°519| 445] °687| 1°:106| 2°795| °‘577| 4:°166| 3°442] 3-433] 3°903| 3:848
1761 °357| 2°808| 1°534| -96 | 1°925| 2°322| 2°617) 1976) 5079} 1°432| 3°698) 1:735
1762 2°327| 1-487] 1386) 2°157| -905) 648) 2°271) °96. | 4°393/-1°348| 2296) “36

1763 291} 17608] 1:094] 1°884) 1°909}| 3°018| 3°668] 3°261) 2°412] 2°16 | 1°844)| 5-204
1764 4°18] | 2°538| 1°497| -718| 1568) -772| 2°764| 2°097| 1°897| 2°565| 2-525) 1°04

1765 2°079| 596] 3°343) 2°368] °408| 1°575| ‘386) 2°195| 1°903| 2:°147| 1°572) -814
1766 173] 1:257| °259) 1:°371| 2°927) 3°316| 2°241) 1°794) 2°948| 2-566) 1-541] 1:079
1767 1°647 | 2°426| 1°586| °211] 3°41 *559| 3°941] 2°03 | 3°065| 2°954] 4°084| ‘624
1768 893] 6°504) °654|) 1°73 | 1°114] 3°475| 4°49 | 1:43 | 3°236] 2°578) 3°099| 2°598
1769 1°016| 1°557| °902) 1:°447| ‘886}) 1°753|] 1°488] 3°427| 5°138] 1°37 | 1:242) 1577
1770 L111) 1505), 1:521 | 1°32 |.1-277) 4:009| 1°969| -°831)| 3:8 1:299'| 2d7. oleae

eral 1°58 476) °632| °805| 1894] 694] 3°027|] 3°619| 1°728| 4°374| 2°887| 2-266
1772 1365} 1-398) 1°8 “772 | 1°239| 2°679'| 3°035| 3°256} 3°517).3:09. | 4°991%) 16376
1773 2°927:) 1:24 | L077 | 1-993) 189%) 835) 17460), 1-9 5°62 | 5°24 | 2°378}| 1-666
1774 2°01 | 2°222) °565) 1°481|] 1°859| 1°757| 2°212) 1°953) 2°006| -737) -947) 1°595
1775 3°136| 2°958} 2°099| °902|] 1°154}| °645) 2°857] 3°903] 3°489] 4°104/] 2°58 | 1°305
1776 “620i 2 oval) deta) sg 857] 1°93 | 3°645| 3°237| 3°252| 1531} 1°336| 1-601
1777 (2 OO es el oA aoa ie led 3°308| 2°606| 2°771| 962] 4°392| 2°3 “416
1778 1405; 691] 1°861} “42 | 2°688] 2°154|] 4°177| 2°179| 1°504| 3°519| 2°328)| 3°36

1779 °258| °626| °324] 1607] 2°49 | 1°376| 4°058| 1°01 | 5°829)| 4°5 1°651} 3°643
1780 D "988 | 2°303:|.1°799)| 2°137 |.1:347 | 2043). °833) 3°561) 35°161) 1591 |) 722
1781 824) 2°081| 551} 1:024) 1°075) 1°417] 1-891] 3°16 833] ‘63 | 4403} 1°517
1782 3°531| 678) 2°041) -767] 2°104| 1°362| 1°674] 4:229] 3°392| 3608] -840] 1:271
1783 2°076) W074. A458) 5-17.) 1°93) 1981) 20s | 36763592 | 2-743 | 1684 eee

praemee 379 |47-281 |35°332 134:075 |43°886 |51:273 |70°632 |67°137 |82°412 |73°535 |64°883 |48°893 | 6

Sane 1458] 1-751] 1:309| 1-262] 1°625]| 1-899] 2°616| 2-486| 3-052] 2-723] 2-403] 1-812

58718

24°396

RAIN IN CARLISLE AND THE NEIGHBOURHOOD. 315

| TABLE EXHIBITING THE QUANTITY OF RAIN OF EACH MonTH For NINETEEN YEARS, AND THE ANNUAL QUANTITY
OF EACH YEAR, TAKEN FROM THE REY. JOSEPH GOLDING’S METEOROLOGICAL JOURNAL, KEPT AT AIKBANK, NEAR
| Wicron, CUMBERLAND, From 1792 To 1810.

11792 1:5599| 1°41 | 2°348] 3°578| 3:067| 1:979| 3°757| 4°968| 5°843| 2°623) 1°781| 4:495| 37-448
11793 1:388 | 4°1 2°405| °901) 1°620] 2°095| 1°594| 5°328) 1°617| 2°426) 2:°034| 2°988|] 28:496
11794 2°594| 4:255| 1-426] 3°357| 1:°948} +995] 2:°564] 2°594| 3:207| 4:°813] 2°675| 2°862| 33°290

682 | 3°505| 3°256| 2°503) 1°342] 3°377| 1°836| 4:117| °674}| 6°216| 6°830] 6:022] 40°360
6°505| 2°502| 928] 1°559| 3515} 3°31 | 5633) 1192] 3°027) 3-558] 1:°988] 1627) 35°344
3°776| °655}| 1°437| 1646] 3°757| 1°921| 4°146| 4:°336| 5°668| 3°239|) 3°534| 4°458| 38°573
3294) 2-715) 1-401) 1°671| 1:15 | 17193) 4°604| 3°053| 3°349| 3:775) 2°847| 1°731] 30°783
2°781| 2°379| 1:276| 2-717) 2°357| -558) 3°657| 8476] 5°228) 4:°777| 3°916| -242) 38-364
3°267| ‘880| 2°368| 3°499) 3°559| 647) 1°641] 1°362|] 4°659| 5-243] 4°813}] 1:929| 33°867
3117) 3°953| 4°898) 1:19 | 1:°257| 318) 5°544| -857) 4°459| 5°321| 2°394) 3-915) 37-223
2°564| 3°429| 1°611| 3-441} +394] 2°627| 6883] 3°368] 2°597) 5:199| 501] 3:260| 35-874
1:398| 3°007] 1°734| 1°849| 4°612] 3°41 694) 3°794| 3°17 | 2°158| 2°715| 3°192| 31°733
6°101| 1°695| 2°543| 1°811] 2°189] 2°164) 1°758] 4°324] 2°013] 5°699| 2°028| 1-463] 33-788
2°934| 3°404| 2°769| 946] 1°75 | 2°938| 3°628] 2°31 | 3°011| °235| ‘547| 4°869| 29°341
4°88 | 2°669|) ‘73 | 1:253| 1°744] 1:956| 3°97 | 6°896] 4°87 | 1°639]| 5-424) 5:18 41211
1:385 | 4°77 967 | 2°129| 3°069| 1°628] 4°246] 3°179| 6°415| 4:16 | 3°893| 2°743|] 38584
3°839|-1°595| -267| 1662} 2°942| 1:773)| 3°269| 3°816] 2°103| 5-162] 3°478| 1-886) 31-792
3°963| 2°967| 636) ‘887| 4°547) 3:°194| 2°194| 7°386| 4:036| 562] 1°525| 5-492] 37-389
1°886 | 1°348| 5°123| -719| -642| 1°386) 5:°174| 3°046| 1:254) 3°033| 2:976| 3-771] 30°358

37°318 |45°461 |37°469 |66°792 |74-402 |67°200 |69°838 |55-899 |62°125 | 663°818

——q—| qq _ | | | EE ee

4 305 | 2°697| 2°006) 1-964) 2°393| 1°972] 3°515| 3:°916) 3°537| 3°676| 2°942| 3:27 34°938
: |

It is worthy of remark that Dr Miter states in his “ Synopsis of the Fall
of Rain, &c., in the English Lake and Mountain District in the year 1853,”
| that—‘‘ Among several abnormal and opposite atmospheric conditions presented
_ by the years 1852 and 1853, the departure from the average in the rain fall is
the most obvious and remarkable. While the former was the wettest, the
_ latter was the driest year since the experiments were begun in 1844. In 1852
the depth of water precipitated at Seathwaite was equivalent to 156°74 inches,
and, in 1853, to 113°69 inches—a difference of 43 inches, corresponding to the
_ average annual fall at Whitehaven in the last ten years.” A similar departure
from the average rain-fall took place at Bunkers Hill—the greatest fall during
the period of my journal being in 1852, and the least in 1853, as appears by the
accompanying Table—31‘825 inches in the former year, and 19°613 inches in
the latter.

e
A TABLE EXHIBITING THE QUANTITY OF RAIN OF EACH MONTH FOR NINETEEN YEARS, AND THE ANNUAL QUANTIT

Barnes, M.D., rrom 1852 To 1870.

316 DR BARNES ON THE AVERAGE QUANTITY OF

p
NA
56°04

Years. Jan. Feb. | March. | April. May. June. | July. Aug. Sept. Oct. Nov. Dec.
1852 | 3571] 1360) -625| 1:189| 2°363| 4:88 | 2:114| 3-446] 1°947| 2694] 2:192| 5-444
1853 | 3033] -732| -621| 1:303| -881| 2°151| 2°814] 1:904| 2°07 | 2351] 1243] -51
1854 | 1:665| -788| -81 | -072| 3:43 | 3-447] 1501] 3-277 | 1°817| 2°326| 18895 | 2-178
1855 097 | -746| 1:449| 1-442] 1-486 | 2°718] 2°819] 3-098; 1197] 4°322| 1375] -598
1856 | 2:072| 1°843| -062| 1:152| 2°829| 3:906| 1-277] 4140] 2°175| 2°706 | 1:137 | 3°618
1857 | 1:657| -753| 2:47 | 1:18 | 1:138] 2:413] 2°347] 1-934] 2°895 | 2-263] 2°09 | 2:003
1858 | 1:027| :458] 1:75 | 1:055| 2965 | 2:157| 3-402] 3-276 | 3°718| 3:072| 1158] 1:75
1859 | 2:004| 1-222] 2:59 | 1:984] -05 | 1°531| 3:111] 2-407] 4:185| 1°378] 3611] 2:361
1860 | 3°381| -854|] 2:588| 1:187| 1-807] 3-114] 1:4 | 3684] 1:06 | 45 | 1:25 | 2:187
1861 | 1:093| 1-42 | 2-944] -564]| 1-223] 1:916] 4:02 | 3-407] 4623] 1°815| 6-704 | 1:935
1862 | 2°593| -781| 1:756| 2218] 3-066] 2°854| 4:159| 3:469| 2°609| 4145] 1:698| 2:312
1863 | 3572} 1:371| -468|] 2:°593| 2:376| 2°61 | -625| 2:583| 4-711] 4:003| 3177] 2:274
1864 | 20 | 1781] 2966] 1156] 1:868| 2°388| -541] 1-72 | 4:607| 3-407] 1°921 | 1:935
1865 | 1:374] 1656] 1:093| -796| 4:311| -783| 1:097| 3:671| -89 | 5:0 | 2°631| 1-412
1866 | 3:772| 2184) 1593] -684| 1-064] 1:937] 2°967| 4:0 | 3°965| 1-281 | 3592] 3-457
1867 | 2:281] 1:718] 1-407] 2°92 | 2:49 | 1:16 | 3:-416| 1°665 | 231 | 1857] °5771 1:27
1868 | 2393] 2:01 | 3355| 2:5 | 1:993| -986| -281| 3°125| 2:116| 2187] 1646] 4-408
1869 | 2:225| 3062) -468|] 1°871| 2:02 | 1:17 | -743| -871| 4:25 | 1°92 | 3-057) 2-239
1870 | 2°673| 1:7 ‘468 | -998| 1°354| 1:629| -859| 2:333| 1-436] 3868] 2:21 | -972
Totals for) 15.483 26-439 |29-483 [26-864 [38° ‘75 |39°493 [54-01 (59-581 (55-095 [43-164 |42- 93
ee 483 |26°439 |29°483 |26°864 |38°714 |43°75 54 52 55-095 |43-164 |42°863 | 49493)
ieee 9-236] 1:391| 1:552| 1-414] 2°037| 2°303| 2°078| 2°843| 2°767| 2-9 | 2:272| 2-256

The Tables from Dr Cartyte’s and Mr Go.pine’s Journals, I made nearly
forty years ago, but they were never published. Mr GoLpIne set a great value
upon his Journals, and, for their safe keeping, gave them into the custody of
the Rev. RicHarp Marruews of Wigton Hall, with a request that he would
place them in his library. Mr Marruews died many years ago ; his library was
sold after his death; and Mr Go.tpine’s Journals disappeared. They have

probably been torn up as waste paper. Dr Cartyze’s family are all dead, and

what has become of his Journals I know not.

On comparing the averages of our observations with those of Mr Pirt, which

I have also added, I find in three out of four, viz—in Dr Cartyiz’s, Mr
GoLpinc’s, and Mr Pirv’s, April to be the driest month of the year. According
to my own, February was the driest, and April stands next on the list. July,
August, September, and October were wet months, according to all the Journals.

The following are the averages or mean quantities of rain for the several
months of the year, during the different periods. They are arranged in the
progressive order of the increasing quantity of rain in each month, according to

RAIN IN CARLISLE AND THE NEIGHBOURHOOD. 317

the several Journals, beginning with the driest month, and proceeding to the
wet months :—

Dr Cartyte’s Journal. Mr Gotpine’s Journal. Mr Pirt’s Journal. Dr Barnes’ Journal.
27 Years Mean. 19 Years Mean. 24 Years Mean. 19 Years Mean.

September, . 3°052 | August, . % 3916 |oulyss = . 3317 | October, eS)

From the averages it appears that about twice as much rain falls in each of
the latter months of the Table, as in the month of April; and about one-third
less rain falls in the first six months of the year, than in the last six months.

This distribution of rain answers wise and important purposes in the economy
of nature. Were the reverse the case, 7.e. did more rain fall in spring than in
autumn or winter, very disastrous consequences would ensue. The great benefit
of dry weather in spring to agriculture is obvious ; the value of an ounce of March
dust is proverbial. The great fall of rain in the latter months of the year may,
I think, in some measure be accounted for by the cold increasing as the sun
recedes from us in autumn, and the vapours, which had been raised by the heat
of summer, are then condensed and precipitated in the form of rain.

Mr Gotpine, whom I had the pleasure of numbering among my friends, on
seeing the comparative Table I had drawn up of the mean quantities of rain for
_ the several months of the year, made the following remarks :—

“This mode of exhibiting the subject is both curious and useful; and not-
withstanding the great attention which I formerly paid to the phenomena of
the weather, I confess that till now I never knew that April was the driest
month of the year. April showers are so frequently mentioned as to give a
general idea that it is rather a wet month than otherwise ; but it may be remarked
that the showers in April are seldom stormy, or attended with great falls of
rain, as some of the summer months are after the solstice is turned. This dryness
of April is most probably occasioned by the less development of the electric
fluid at that particular season of the year; for when by means of the summer
heats the air begins to be more strongly electrified, then it is that the showers
become heavy, and often send down immense quantities of rain in a very short
space of time. This it is which makes July and August generally the wettest
months of the year; and happy it is for us poor mortals, that such is the

VOL. XXVI. PART IL. 4N

318 DR BARNES ON THE AVERAGE QUANTITY OF

arrangement of nature ; for if there were not very heavy falls of rain during the
excessive heats of summer, the ground would be exhausted of moisture, and
vegetation entirely at a stand.”

There is a remarkable difference between the Journals of the late Dr
CARLYLE and Mr Pirt, in regard to the mean annual quantity of rain. Both
Journals were kept at Carlisle, and both of the gentlemen, I have reason to
believe, were careful and accurate observers. According to Dr CARLYLE’
Journal, the average annual quantity of rain is 24°396 inches, and according to
Mr Pirt’s it is 30°571 inches. How is this to be accounted for? Has the
climate of this country undergone some change? It is evident from an inspection
of the Journals, or of the Tables formed from them, that the quantity of rain is
different in different years, and that sometimes there are a few wet years and
sometimes a few dry years in succession. Is this the case with long periods of
time ? So that Dr CartyLe’s Journal may have been kept when there was a
dry series of years, and Mr Pirt’s when there was a wet series? Perhaps the
difference may be explained by their rain-gauges being placed in different
situations. Dr CARLYLE’sS gauge was placed at the head of Abbey Street, 82
feet above the sea-level, higher than Mr Pirt’s, which was kept in Shaddongate,
40 feet above the sea-level. It has been frequently remarked, that when one
rain-gauge is placed on the top of a high tower, and another at the bottom, more
rain falls into the lower gauge than into the higher one. But there is another
reason which may be assigned for the difference. The situation of Dr CARLYLE’s
rain-gauge was in the vicinity of his dwelling-house, which would occasionally
prevent some rain falling into the gauge. It was placed on a wall on the 8. W.
side of his house. This I am inclined to consider the principal cause of the
difference of the two Journals. Still the difference of altitude between Abbey
Street and Shaddongate might have considerable effect, and may in some
measure account for the different results. These causes, however, would have
very little influence on the comparative monthly averages of the fall of rain.

The Cumberland Infirmary stands on elevated ground on the S. side of the
river Eden, 30 feet above the bed of the river, and about one mile N.W. of my
garden at Carlisle, which is nearly of the same height as the site of the Cathedral.
For several years I kept a rain-gauge at each place, constructed on the plan
recommended in BrewstTEr’s Cyclopedia, and I always found a greater fall of
rain at the former than at the latter place. The following is the quantity
registered at each place in the years 1837 and 1838, and shows the difference
of rain-fall at these two places in these years :— |

RAIN IN CARLISLE AND THE NEIGHBOURHOOD. 319

Jan. Feb, | March.| April. | May. | June. | July. | Aug. | Sept. Oct. Nov. | Dec.
1837
mmary, . | 2:278| 1-866] -91 | -535/1-0 | 2:38 | 5-537] 4:53 | 2-28 | 319 | 317 | 3-66
den int} 1-59 | 3:16 | 1-02 | 1-2 | 1-13 | 2:23 | 437 | 1-67 | -98 | 99 | -82 | 1-78
astle St.,
1838
mmary, . | 1-215] -59 | 2-436] 1-51 | 1-49 | 4-69 | 3-195] 3-45 | 9-28 | 2-65 | 2-47 | -78

den 0}! 1.95 | -o9 | -976] 1-384] 1:69 | 3:172/ 1:85 | 1-08 | 1-035/ 1-38 | 1-4 | 1-15
astle St.,

Annual
quantity of

each year.

30°972
22°85

26-756
16257

With regard to the difference that exists between the annual mean quantity
of rain of Mr Gotp1ne’s and Mr Pirv’s Journals, Mr Goipine’s being 4°4 inches
more than Mr Pirt’s, I shall give you the explanation of the former gentleman

in his own words, contained in a letter written by him many years ago :—

“T find that the annual quantity of rain, according to my diary, is somewhat
greater than that shewn by Mr Pirt’s; and this might naturally be expected
from the difference of situation, for my observations were chiefly made at
Aikbank, which borders on the hilly part of the country,—and it is well known
that in a hilly, and more especially in a mountainous district, there is much
more rain than in a level one. Besides, Carlisle has a further cause of exemp-

- tion from rain :—It lies nearly in the direction of the Solway Firth, and when
storms come from off the Irish Sea, as they frequently do, the vapour on entering
the Firth is attracted either by the Scotch or the English mountains, which
will occasion more rain to fall on each side of the Firth, than in the direction

of the Firth itself.”

Plate XI.

Trans Roy. Soc. Edin? Vol. XXV1.

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Plate XV!.

Trans. Roy. Soc. Edin* Vol. XXVI

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( 321°)

XV.—On the Physiology of Wings, being an Analysis of the Movements by
which Flight 1s produced in the Insect, Bat, and Bird. By James BELL
Petticrew, M.D., F.R.S., Pathologist to the Royal Infirmary of Edinburgh,
and Curator of the Museum of the Royal College of Surgeons of Edinburgh.
Communicated by Professor TuRNER. (Plates XI. to XVI.)

(Received 2d August 1870. Read 16th January 1871.)

INTRODUCTORY REMARKS.
(For Table of Contents see end of Memoir.)

In order to determine with exactitude the movements made by the wings in
flight, and the part which the air plays in modifying them, I was induced several
years ago to collect a large number of facts, and to undertake an extensive
series of experiments with natural and artificial wings. My observations and
experiments, I may remark, were not wholly confined to flight. On the con-
trary, I traced the analogy between flying, swimming, and walking ; a circum-
stance which compelled me to pay particular attention to the size, shape, and
movements, not only of wings, but also of the travelling surfaces of quadru-
peds, amphibia, and fishes. By adopting this method, I obtained suggestions
which have proved of the utmost importance to me in my attempts at elucidat-
ing the very intricate problem of flight.

As there are, strictly speaking, only three highways.in nature (the land, the
water, and the air), so there are three principal varieties of locomotion. There
are, however, a limited number of mixed forms, the animal in such cases being
furnished with travelling surfaces, modified in such a manner as to enable it to
progress upon, or in, two essentially different media. The mixed movements are
alike interesting and instructive, as they prove that movements apparently very
dissimilar are in reality only links of a great chain of motion, which drags its
weary length over the land, through the water, and extends skyward. That,
therefore, is not wanting which connects the motions peculiar to walking
animals with those peculiar to swimming and flying animals. Thus the seal
furnishes the link between the land and water, and the galeopithecus between
the land and air; while the flying fish supplies the link between the water
and the air.

On making a careful examination of the structure and movements of the

VOL. XXVI. PART II. 40

322 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

ereat pectoral fins or pseudo-wings of the flying fish, I felt persuaded that a
close analogy existed between the flippers, fins, and tails of sea mammals and
fishes on the one hand, and the wings of insects, bats, and birds on the other ;
in fact, that theoretically and practically these organs one and all formed
flexible helices or screws, which, in virtue of their rapid reciprocating action,
operated upon the water and air after the manner of double inclined planes.

Guided by these indications, I especially directed my attention to the
twisting flail-like movements of the wings of insects ; of the flippers and tails
of sea mammals, and of the fins and tails of fishes. These I found all acted
upon the air and water by curved surfaces, the curved surfaces reversing,
reciprocating, and engendering a wave pressure, which could be continued
indefinitely at the will of the animal.

In order to prove that sea-mammals and fishes swim, and insects, bats, and
birds fly, by the aid of curved figure of 8 surfaces, which exert an intermittent
wave pressure, I constructed artificial fins, flippers, and wings, which curved
and tapered in every direction, and which were flexible and elastic, particularly
towards the tips and posterior margins. These fins, flippers, and wings were
slightly twisted upon themselves, and when applied to the water and air by
a sculling or figure of 8 motion, curiously enough not only reproduced the
curved surfaces referred to, but all the other movements peculiar to the fins
and tail of the fish when swimming, and to the wings of the insect, bat, and
bird when flying.

HISTORY OF THE FIGURE OF 8 OR WAVE THEORY OF FLYING.

The Wing a Twisted Lever or Helix.—t announced this view in a lecture
delivered at the Royal Institution of Great Britain in the early part of 1867.
An abstract of the lecture appeared in the Proceedings of the Institution under
date the 22d of March 1867.* At pages 99, 100, and 101 of the abstract in
question, the spiral conformation of the wing in the insect and bird is adverted
to at length, and there described as a twisted lever or helix, which owes its
peculiar elevating and propelling power in a great measure to its shape. Par-
ticular emphasis is also placed upon the partial rotation of the wing on its long
axis during extension and flexion, and to its screwing and unscrewing action
during the down and up strokes, this being a “ sine gua non” in flight. In the
pages alluded to, the subjoined passages occur :—“ The wings of insects and
birds are, as a rule, more or less triangular in shape, the base of the triangle
being directed towards the body, the sides anteriorly and posteriorly. They are
also conical on section from within outwards and from before backwards ; this
shape converting the pinion into a delicately graduated instrument, balanced
with the utmost nicety to satisfy the requirements of the muscular system on

* On the Various Modes of Flight in relation to Aéronautics.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 323

the one hand, and the resistance and resiliency of the air on the other.
The neure or nervures in the insect’s wing are arranged at the axis or root of
the pinion, after the manner of a fan or spiral stair; the anterior one occupy-
ing a higher position than that farther back, and so of the others. As this
arrangement extends also to the margins, the wings are more or less twisted
upon themselves, and present a certain degree of convexity on their superior
or upper surface, and a corresponding concavity on their inferior or under
surface; their free edges supplying those fine curves which act with such
efficacy upon the air, in obtaining the maximum of resistance and the minimum
of displacement ; or what is the same thing, the maximum of support with the
minimum of slip. . . . . All wings obtain their leverage by presenting
oblique surfaces to the air, the degree of obliquity gradually increasing in a
direction from behind forwards and downwards during extension, when the
sudden or effective stroke is being given, and gradually decreasing in an oppo-
site direction during flexion, or when the wing is being more slowly recovered
preparatory to making a second stroke. The effective stroke in insects, and
this holds true also of birds, is therefore delivered downwards and forwards,
_and not as the majority of writers believe, vertically, or even slightly backwards.
To confer on the wing the multiplicity of movement which it re-
quires, it is supplied at its root with a double hinge or compound joint, which
enables it to move not only in an upward, downward, forward, and backward
direction, but also at various intermediate degrees of obliquity. . . . The wing
of the bird, like that of the insect, is concavo-convex, and more or less twisted
upon itself. The twisting is in a great measure owing to the manner in which the
bones of the wing are twisted upon themselves, and the spiral nature of their
articular surfaces, the long axes of the joints always intersecting each other
at nearly right angles. Asa result of this disposition of the articular surfaces,
the wing may be shot out or extended, and retracted or flexed in nearly
the same plane, the bones of the wing rotating in the direction of their
length during either movement. This secondary action, or the revolving of the
component bones upon their own axes, is of the greatest importance in the
movements of the wing, as it communicates to the hand and forearm, and con-
sequently to the primary and secondary feathers which they bear, the precise
angles necessary for flight. It, in fact, insures that the wing, and the curtain
or fringe of the wing, which the primary and secondary feathers form, shall be
screwed into and down upon the wind in extension, and unscrewed or with-
drawn from the wind during flexion. The wing of the bird may therefore be
compared to a huge gimlet or auger, the axis of the gimlet representing the
bones of the wing; the flanges or spiral thread of the gimlet the primary and
secondary feathers.”
The lecture referred to formed part of a memoir which was communi-

324 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

cated by Professor Hux.ey to the Linnean Society, and read before that body
on the 6th and 20th of June 1867. It is published 2” extenso in the 26th
volume of the Transactions of the Society, with upwards of eighty illustrations.*

The principal object of the memoir is to establish an analogy between the
walking surfaces of quadrupeds, the swimming surfaces of fishes, and the flying
surfaces of insects, bats, and birds. These are all describedt and figured{ as
twisted levers or screws in an anatomical sense (pages 361 and 362, figures 37,
38, 39, and 40), and as flexible reversing screws in a functional or physiological
sense (pages 336 and 362, figures 2, 41, 42, and 43).§ As a consequence,
the quadruped and biped|| are represented as walking,‘ and the seal and

* On the Mechanical Appliances by which Flight is attained in the Animal Kingdom, &e.

+ Op. cit., from page 199 to page 267 inclusive.

t Op. cit., Plate XV. figs. 49, 51, 57, 68, 69, 70. Likewise Diagram 18 A d’e’/’, a’l’, page 253.

§ Op. cit., Plate XV. figs. 58, 59, 61, 73, 74, and 75.

|| Op. cit., Plate XV. fig. 78.

q I think it proper to state that various anatomists have carefully examined the form of the articular
surfaces of the joints in the limbs, more especially in man. ‘The researches of the brothers WEBER and
Professor Mryer of Zurich are so well known, that it may suffice simply to refer to them. I would also
direct attention to the writings of Lancnr, Henke, Meissner, and the late Professor Goopsir. LANGER,
Henke, and MEIssNver succeeded in demonstrating the “ screw configuration” of the articular surfaces of the
elbow, ankle, and calcaneo-astragaloid joints, and Goopsir showed that the articular surfaces of the knee- -
joint consist of ‘‘a double conical screw combination.” The last-named observer also expressed his belief,
“ that articular combinations, with opposite windings on opposite sides of the body, similar to those in
the knee-joint, exist in the ankle and tarsal, and in the elbow and carpal joints ; and that the hip and
shoulder joints consist of single-threaded couples, but also with opposite windings on opposite sides of
the body.” The following are the views of Lanczr as interpreted by Goopstr :—(Proc. Roy. Soe. Edin.,
Jan. 18, 1858, and Anatomical Memoirs, vol. ii. p. 231.) ‘ Lancer, acting on the happy idea of pro-
longing the screw by uniting, in one direction, a number of plaster casts of the same articular surface,
succeeded in forming continued screws from the upper articular surface of the astragalus in the horse,
panther, and human subject. Lancer concludes that the ‘ go line’ (a line obtained from the scratch of
a steel point fixed on one of the articular surfaces, and which marks the opposite surface when the joint
is moved) of the ankle-joint in all the mammalia is a portion of a helix, and that therefore the astraga-
loid surface is a segment of a cylindrical or conical male screw, while the tibio-fibular surface is a
segment of the corresponding female screw. The right ankle-joint is a left-handed screw combination ;
the left ankle-joint a right-handed. When therefore the foot is conceived to be fixed, the leg, in passing
from a position of extension to flexion, moves laterally outwards along the axis of rotation, and the sine
of the angle of inclination of the thread—that is, in proportion to the extent of flexion and the rapidity
of the screw.” Goopsir, in attempting by Lancrr’s method to develop those articular screw-models, found
that when two casts were united, an apparently satisfactory helix was produced ; but in adding to the
series, the spire diminished, and the helix closed upon itself; so that it appeared that not only the
angle of inclination of the thread, but also the radius of rotation, diminished. He was, therefore, of
opinion, that the tibio-astragaloid articular surfaces could not be regarded as segments of a cylindrical
series, and thought it extremely probable that, abstracting the terminal facets, the acting areas on each
surface consist each of a segment of a conical screw—the convex portions of these two screws being on
the astragaloid, the concave on the tibial articular surface ; the one screw coming into action in flexion,
the other in extension. Goopsir’s experiments on the knee and ankle-joints, conducted with extreme
care, by the aid of fresh specimens, casts, and models, led him to conclude that both joints were ‘ spiral
in their nature’—that in fact they were ‘screwed structures,’ and that the movements of the knee-joint
are combined gliding and rolling movements of conical screwed surfaces upon one another. The follow-
ing are his own words :—“ The general character of the curves observed, and the corresponding move-
ments and structure of the joint (knee-joint) leave little doubt in my mind that the flexion and extension,
combined gliding and rolling movements of the knee, are performed between two conical double-threaded —
screw-combinations, an anterior and a posterior—the anterior being a left-handed screw, and the posterior

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 325

fish* as swimming in figure of 8, or looped curves. The wings of the insect,
bat, and bird, are also described and figured as executing figure of 8 movements
when the animals are hovering before an object, or when their bodies are
artificially fixed (page 336, figure 2) ;+ the figure of 8, as I explained, being
opened out or unravelled when the animals are flying at a high horizontal
speed to form a looped and then a waved track (pages 341, 342, 344, and 345,
figures 10, 13, 14, and 15).t

The following brief passages from my memoir in the Transactions of the
Linnean Society§ will, I hope, serve to elucidate the peculiar figure of 8 move-
ments made by the wings in flight :—

The Wing Twists and Untwists during its action —“ That the wing twists upon
itself structurally, not only in the insect, but also in the bat and bird, any one
may readily satisfy himself by a careful examination,|| and that it twists upon
itself during its action I have had the most convincing and repeated proofs.4
The twisting in question is most marked in the posterior or thin margin of the
wing, the anterior and thicker margin performing more the part of an axis. As
a result of this arrangement, the anterior or thick margin cuts into the air
quietly, and as it were by stealth, the posterior one producing on all occasions
a violent commotion, especially perceptible if a flame be exposed behind the
insect. Indeed, it is matter for surprise that the spiral conformation of the
pinion, and its spiral mode of action, should have eluded observation so long ;
and I shall be pardoned for dilating upon the subject when I state my convic-
tion that it forms the fundamental and distinguishing feature in flight, and must
be taken into account by all those who seek to solve this most involved and
interesting problem by artificial means.” The importance of the twisted confi-
guration or screw-like form of the wing cannot be over-estimated. That this
shape is intimately associated with flight is apparent from the fact that the
rowing feathers of the wing of the bird are every one of them distinctly spiral
in their nature ; in fact, one entire rowing feather is equivalent—morphologi-
cally and physiologically—to one entire insect wing. In the wing of the martin,
where the bones of the pinion are short and in some respects rudimentary, the
primary and secondary feathers are greatly developed, and banked up in such a

a right-handed screw in the right knee-joint ; the anterior a right-handed, and the posterior a left-handed
screw in the left knee-joint. The movements which take place round these two combinations are
alternate, those round the anterior completing extension and commencing flexion, those round the
posterior completing flexion and commencing extension of the joint.”

* Op. cit., Diag. 2, page 204; Plate XV. fig. 76.

t+ Op. cit., page 233, Diag. 5; Plate XV. fig. 61.

t Op. cit., page 233, Diag. 6; Plate XV. fig. 59.

§ Op. cit., pages 231, 232, 233, and 234.

|| Op. cit., Plate XV. figs. 68, 69, and 70.

{ Op. cit., Plate XV. figs. 58, 61, 73, and 74.

VOL. XXVI. PART. II. 4p

326 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

manner that the wing as a whole presents the same curves as those displayed
by the insect’s wing, or by the wing of the eagle where the bones, muscles, and
feathers have attained a maximum development. The conformation of the
wing is such that it presents a waved appearance in every direction—the waves
running longitudinally, transversely, and obliquely. The greater portion of the
pinion may consequently be removed without essentially altering either its form
or its functions. This is proved by making sections in various directions, and
by finding that in some instances as much as two-thirds of the wing may be
lopped off without materially impairing the power of flight. Thus, in the summer
of 1866,* I removed the posterior two-thirds from either wing of a blow-fly,
and still the insect flew, and flew well. The only difference I could perceive
amounted to this, that the fly, while it could elevate itself perfectly, flew in
circles, and had less of a forward motion than before the mutilation. It had
in fact lost propelling or driving power, the elevating or buoying power remain-
ing the same. I took another blow-fly and removed the tip or outer-third of
either wing, and found that the driving-power was the same as before the muti-
lation, while the elevating or buoying power was slightly diminished. These
experiments prove that the posterior or thin elastic margin of the wing is more
especially engaged in propelling, the tip in elevating.t “The spiral nature of
the pinion is most readily recognised when the wing is seen from behind and
from beneath,{ and when it is foreshortened.§ It is also well marked in some
of the long-winged oceanic birds when viewed from before, || and cannot escape
detection under any circumstances, if sought for,—the wing being essentially
composed of a congeries of curves, remarkable alike for their apparent sim-
plicity and the subtlety of their detail.”

The Wing during its action Reverses its Planes, and describes a Figure of 8
“ The twisting or rotating of the wing on its long axis is parti-
cularly observable during extension and flexion in the bat and bird, and like-
wise in the insect, especially the beetles, cockroaches, and others which fold
their wings during repose. In these in extreme flexion the anterior or thick
margin of the wing is directed downwards, and the posterior or thin one up-
wards. In the act of extension, however, the margins, in virtue of the wing
rotating upon its long axis, reverse their positions, the anterior or thick mar-
gins describing a spiral course from below upwards, the posterior or thin
margin describing a similar but opposite course from above downwards.
These conditions, I need scarcely observe, are reversed during flexion. The
movements of the margins during flexion and extension may be represented

* Op. cit., pages 219, 220, 221, 229.

For further experiments in oie direction, see footnote to pages 361 and 362.
Op. cit., Plate XV. figs. 68, 69, 70, 73, and 74.

Op. cit., Plate XV. Bes 61 and 62.

| Op. cit., page 253 ; Dinars 18 A, aU’, der’

Pre

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 327

with a considerable degree of accuracy by a figure of 8 laid horizontally.* .. .

It may likewise happen, though more rarely, that the anterior or thick margin
of the pinion may be directed upwards and backwards during the return or up
stroke. I infer this from having observed that the anterior margin of the wing
of the wasp (when the insect is fixed and the wings are being driven briskly)
is not unfrequently directed upwards and forwards at the beginning of the
down stroke, and upwards and backwards at the commencement of the up or
return stroke. A figure of 8, compressed laterally and placed obliquely with
its long axis running from left to right of the spectator, represents the move-
ment in question. The down and up strokes, as will be seen from this
account, cross each other, the wing smiting the air during its descent from
above, as in the bird and bat, and during its ascent from below, as in the flying
fish and boys’ kite. The pinion thus acts as a helix or screw in a more or less
horizontal direction from behind forwards, and from before backwards ; but it
has a third function—it likewise acts as a screw in a nearly vertical direction
from below upwards. .... If the wing (of the larger domestic fly) be viewed
during its vibrations from above, it will be found that. the blur or impression
produced on the eye by its action is more or less concave. This is due to the
fact that the wing is spiral in its nature,t and because during its action it twists
upon itself in such a manner as to describe a double curve,{—the one curve
being directed upwards, the other downwards. The double curve referred to is
particularly evident in the flight of birds from the greater size of their wings.§
The wing, both when at rest and in motion, may not inaptly be compared to the
blade of an ordinary screw propellor as employed in navigation.|| Thus the
general outline of the wing corresponds closely with the outline of the propellor,
and the track described by the wing in space is twisted upon itself propellor
fashion. The great velocity with which the wing is driven converts the
impression or blur‘i into what is equivalent to a solid for the time being, in
the same way that the spokes of a wheel in violent motion, as is well under-
stood, completely occupy the space contained within the rim or circumference
of the wheel. .... From these remarks it will appear that not only the
margins, but also the direction of the planes of the wing, are more or less
completely reversed at each complete flexion and extension ; and it is this
reversing, or screwing and unscrewing, which enables the wing to lay hold of
the air with such avidity during extension, and to disentangle itself with such
facility during flexion,—to present, in fact, a more or less concave, oblique, and

* Op. cit., page 233, Diagram 5. Compare this diagram with figs. 59 and 61 of Plate XV.
t Op. cit., Plate XV. fig. 68.

{ Op. cit., Plate XV. figs. 58 and 59a a’. Compare with aq’ of fig. 52.

§ Op. cit., Plate XV. figs. 73 and 75 bae.

|| Op. cit., Plate XV. fig. 52 aa’.

{ Op. cit., Plate XV. figs. 58 and 59.

328 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

strongly resisting surface the one instant, and a comparatively narrow, non-
resisting cutting edge the next. The figure of 8 action of the wing explains
how an insect or bird may fix itself in the air, the backward and forward
reciprocating action of the pinion affording support, but no propulsion. In
these instances, the backward and forward strokes are made to counterbalance
each other.”

The Wing, when advancing with the body, Describes a Waved Track.—

“ Although the figure of 8 represents with considerable fidelity the twisting of
the wing upon its axis during extension and flexion, when the insect is playing
its wings before an object, or still better, when it is artificially fixed, it is other-
wise when the down-stroke is added, and the insect is fairly on the wing, and
progressing rapidly. In this case the wing, in virtue of its being oa for-
wards by the body in motion, describes an undulating or spiral course. *
The down and up strokes are compound movements,—the termination of the
down-stroke embracing the beginning of the up-stroke, the termination of the
up-stroke, on the other hand, including the beginning of the down-stroke.
This is necessary in order that the down and up strokes may glide into each
other in such a manner as to prevent jerking and unnecessary retardation,—
the angle made by the under surface of the wing with the horizon during the
first part of the down-stroke being increased to support and propel the insect,
and decreased during the second part to prepare it, for making the up-stroke,
and to diminish the friction oe by the sage itself, while it does not inter-
fere with its sustaining power.”

The Margins of the Wing gs noise into Opposite Curves during Extension
and Flexion.—< The anterior or thick margin of the wing and the posterior or
thin margin present different degrees of curvature, so that under certain con-
ditions the two margins cross each other, and form a true helix (page 361,
fig. 37).t The anterior margin (7, s) presents two well-marked curves, a corre-
sponding number being found on the posterior margin (¢,«). These curves may,
for the sake of clearness, be divided into axillary curves and distal curves, the
former occurring towards the root of the wing, the latter towards its extremity.
The curves (axillary and distal) found on the anterior margin of the wing are
always the reverse of those met with on the posterior margin, 7.¢., if the con-
vexity of the anterior axillary curve be directed downwards (r),{ that of the
posterior axillary curve (¢) is directed upwards,§ and so of the anterior and
posterior distal curves (s, w). The two curves, axillary and distal, occurring on
the anterior margin of the wing, are likewise antagonistic, the convexity of the
axillary curve (7) being always directed downwards,|| when the convexity of the

* Op. cit., page 233, Diagram. 6. + Op. cit.. Plate XV. figs. 70, 73, and 74.
ft (Op. cit., Plate XV), fies Tose: § Op. cit., Plate XY. fig. 73, a,c.
|| Op. cit., Plate XV. fig. 73, c.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 329

distal one (s) is directed upwards,* and vice versa. The same holds true of the
axillary and distal curves occurring on the posterior margin of the wing
(é. w).t .... The anterior axillary and distal curves completely reverse them-
selves during the acts of extension and flexion, and so of the posterior axil-
lary and distal curves. This reversal of the curves is seen to most advantage
in the posterior margin of the wing, formed in the bird, by the primary, second-
ary, and tertiary feathers.{ When the wing is partially flexed the convexity
of the distal curve (occurring on the posterior margin of the wing) is directed
downwards (page 362, figure 41 a, 6),§ that of the axillary curve upwards (a, c).|
When the wing is rather more than half extended the curves are obliterated,
the posterior margin of the wing becoming straight (page 362, figure 42 3, c).4
It is at this stage of extension that the axillary and distal curves reverse.
When the wing is fully extended the Coupee of the axillary curve is directed
downwards (page 362, figure 43 a, c),** that of the distal one upwards (a, 0),tt
which is just the opposite of what happens in flexion. This antagonism in the
axillary and distal curves observed in the posterior margin of the wing of the bird
is referrible to changes induced in the anterior margin of the pinion, as the
subjoined paragraph will show.”

The Tip of the Bird’s Wing describes an He “The movements of the
wrist are always the reverse of those occurring at the elbow joint. Thus,
during extension, the elbow and bones of the forearm are elevated, and describe
one side of an ellipse ; while the wrist and bones of the hand are depressed, and
describe the side of another and opposite ellipse.{{ These movements are reversed
during flexion, {§ so that when the elbow is raised and carried backwards, the
wrist is lowered and carried forwards, and vice versa.” ||| .

The Wing capable of Change of Form in all its Parts.—*‘ From this descrip-
tion it follows that when the different portions of the anterior margin are ele-

ep, cit,, Plate XV, fig. 73, f. i Op» Cit. Plate XVerie, (3,60, D, ¢.
meOp. cit., Plate XV. figs. 73, 74, 75. § Op. cit., Plate XV. fig. 73, b.
|| Op. cit., Plate XV. fig. 73, a, c. I Op. cit., Plate XV. fig. 74, b, ¢.

a Op. cit., Plate XV. fig. 75, c. ++ Op. cit., Plate XV. fig. 75, a, 0.

tt Op. cit., p. 249, Diagram 14. §§ Op. cit., p. 249, Diagram 15.

\\|| Similar movements occur in the body and tail of the fish in the act of swimming. “The double
curve or spiral into which the fish throws itself when swimming may be conveniently divided into an
upper or cephalic curve,* and a lower or caudal one.t When the concavity of the caudal curve is biting
or laying hold of the water, and when the concave surface of the tail is being forced during extension
with great violence in the direction of the axis of motion,} where the concave surface is suddenly converted
into a convex one, the concavity of the cephalic curve, 7.¢., the concave surface of the upper half of the
fish, is being urged, with less vigour, in the direction of the same line from the opposite side of it. As the
caudal and cephalic curves are obliterated when the line in question is reached, there is, consequently, a
period (momentary it must be), between the effective and non-effective strokes, in which the body of the
fish is comparatively straight, and, consequently, in a position to advance almost without impediment.’ §

* Op. cit., Diag. 2, d, p. 204. + Op. cit., Diag. 2, c, p. 204. + Op. cit., Diag. 2, a, t, p. 204
§ Op. cit.. p. 205.

VOL. XXVI. PART TI. 4Q

330 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

vated, corresponding portions of the posterior margin are depressed, the dif-
ferent parts of the wing moving in opposite directions, and playing, as it were,
at cross purposes for a common good—the object being to rotate or screw the
wing down upon the wind at a gradually increasing angle during extension,
and to rotate it Im an opposite direction and withdraw it at a gradually
decreasing angle during flexion. It also happens that the axillary and
distal curves co-ordinate each other and bite alternately, the distal curve
posteriorly seizing the air in extreme extension with its concave surface
(while the axillary curve relieves itself by presenting its convex surface),
the axillary curve, on the other hand, biting during flexion with its con-
cave surface (while the distal one relieves itself by presenting its convex
one). The wing may, therefore, be regarded as exercising a fourfold func-
tion, the pinion in the bird bemg made to move from within outwards, and
from above downwards during extension, in the effective or down stroke; and
from without inwards, and from below upwards, during flexion in the up or
return stroke.”

The Wing during its Vibration produces a Cross Pulsation.—“ This oscillation
of the wing on two separate axes—the one running parallel with the body of the
bird, the other at right angles to it—is well worthy of attention, as showing that
the wing attacks the air on which it operates in every direction, and at almost the
same moment, viz., from within outwards, and from above downwards, during
the down or effective stroke ; and from without inwards, and from below upwards,
during the up or return stroke. Asa corollary to the foregoing, the wing may
be said to agitate the air in two principal directions, viz., from within outwards,
or the reverse, and from behind forwards, or the reverse, the agitation in question
producing two powerful pulsations—a longitudinal and a lateral; the longitu-
dinal running in the direction of the /ength of the wing, the lateral in the
direction of its breadth. As, however, the curves of the wing glide into each
other when the wing is in motion, so the one pulsation merges into the other by
a series of intermediate and lesser pulsations. ;

The longitudinal and lateral pulsations occasioned by the wing in action
may be fitly represented by wave-tracks running at right angles to each other,
the longitudinal wave track being the more distinct.”

Analogy between the Wing in Motion and the Sounding of Sonorous Bodies.—
“Tt is a remarkable circumstance that the undulation or wave made by the wing
when the insect and bird are fixed or hovering before an object, and when they
are progressing, corresponds in a marked manner with the track described by
the stationary and progressive waves in fluids,* and likewise with the waves of
sound.t This coincidence would seem to argue an intimate relation between

* Handbook of Natural Phil. (vol. on Electricity, Magnetism, and Acoustics), by Dr LarpNER
(Lond. 1863), pp. 366-7. + Op. cit., pp. 378, 379, 380.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 331

the instrument and the medium on which it is destined to operate—the wing
acting in those very curves into which the atmosphere is naturally thrown in the
transmission of sound, in order, as appears to me, to secure the maximum of
progression with the minimum of slip. Can it be that the animate and inani-
mate world reciprocate, and that animal bodies are made to impress the inani-
mate in precisely the same manner as the inanimate impress each other? This
much seems certain :—The wind communicates to the water similar impulses to
those communicated to it by the fish in swimming ; and the wing in its vibrations
impinges upon the air as an ordinary sound would. The extremities of quad-
rupeds, moreover, describe spiral tracks on the land when walking and run-
ning; so that one great law would seem to determine the course of the insect
in the air, the fish in the water, and the quadruped on the land.”

Various other passages might be adduced in elucidation and support of the
curve, wave, or figure of 8 theory of flying, as originally propounded by me, but
a sufficient number have, I trust, been cited to prove that the theory owes its
origin and development to no hasty generalisation from a few scattered and
imperfectly known facts, but that it rests upon a broad basis, such, in reality, as
nature herself supplies.

In order that the reader may form his own conclusions on this point, I pro-
pose to lay before him in the course of my subsequent remarks the observations
and experiments on which the theory was originally founded. The present
memoir is illustrated by upwards of ninety original diagrams and drawings, the
intricacy of the subject being such as to necessitate a free use of the pencil.
The drawings have been made by myself from the life. I have gone into the
origin and development of the figure of 8 theory of flymg somewhat in
detail; first, because the passages selected have an obvious bearing on the
subject of the present communication ; and second, because nearly two
years after I had made my views known, Professor E. J. Margy (Col-
lege of France, Paris), published a series of lectures and papers in the
“Revue des Cours Scientifiques de la France et de L’Etranger,’* and in
the “Comptes Rendus hebdomadaires des Séances de L’Académie des
Sciences,” + in which the figure of 8 theory of wing movements is put
forth as a new discovery. Professor Marry made no allusion to my
researches, which was the more remarkable, as an abstract of my lecture,
already referred to (p. 322), as published in the Proceedings of the Royal
Institution of Great Britain in March 1867, was translated into French, and

* Les mouvements de l’aile chéz les insectes, p. 171, 13th Février 1869. Mécanisme du vol chez
les insectes—comment se fait la propulsion, p. 252, 20th Mars 1869. Du vol des oiseaux, p. 578,
14 Aout 1869. Du vol des oiseaux (suite), p. 601, 21 Aout 1869. Du vol des oiseaux (suite), p.
646, 11 Septembre 1869. Du vol des oiseaux (fin), p. 700, 2 October 1869.

{ Determination expérimentale du mouvement des ailes des insectes pendant le vol. Par
M. E. J. Manny. Tome LXVII. p. 1341, Tome LXVIIL p. 667.

332 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

appeared on the 21st of September of that year in the same Journal* in which
Professor MARreEy’s lectures were originally published. Having had my attention
directed to this circumstance, I addressed a letter to the French Academy on
the 28th of March 1870, which appeared in the “Comptes Rendus ” (p. 875) on
the 18th of April 1870. In it I claim to have been the first to describe and
illustrate the following points, viz :—

That quadrupeds walk, and fishes swim, and insects, bats, and birds fly by
figure of 8 movements.

That the flipper of the sea bear, the swimming wing of the penguin, and
the wing of the insect, bat, and bird, are screws structurally, and
resemble the blade of an ordinary screw propellor.

That those organs are screws functionally, from their twisting and un-
twisting, and from their rotating in the direction of their length,
when they are made to oscillate.

That they have a reciprocating action, and reverse their planes more or less
completely at every stroke.

That the wing describes a jigure of 8 track in space when the flymg animal
is artificially fixed.

That the wing, when the flying animal is progressing at a high speed in a
horizontal direction, describes a looped and then a waved track, from
the fact that the figure of 8 is gradually opened out or unravelled as the
animal advances.

That the wing acts after the manner of a kite.

Previous to replying to the foregoing, Professor MAREY wrote me, to inquire
how he could respond to my “juste reclamation,” without entering into a dis-
cussion which would needlessly complicate the question. I thereupon asked
him to admit in a letter addressed to the French Academy my claim to have
described and illustrated before him the figure of 8 movements made by the
wings of insects, bats, and birds, when those animals are artificially fixed, and
of the spiral and undulatory wave tracks made by the wings of said insects,
bats, and birds, when the animals are flying at a high horizontal speed. This
he has done, as the subjoined extract from his letter, printed in the ‘“‘ Comptes
Rendus” for May 16, 1870 (p. 1093), will show :—“J’ai constaté qu’ effective-
ment M. Pettigrew a vu avant moi, et représenté dans son Mémoire, la forme
en 8 du parcours, de l’aile de linsecte: que la méthode optique a laquelle j’avais
recours est & peu pres identique ala sienne . . . . je m’ empresse de
satisfaire a cette demande légitime, et je laisse entiérement la priorité sur moi,
a M. Pettigrew relativement A la question ainsi restreinte.”

Mode of Investigation pursued by the Author.—I obtained my results by

* Revue des Cours Scientifiques de la France et de I’Etranger.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 333

transfixing the abdomen of insects with a fine needle, and watching the wings
vibrate against a dark background ; by causing dragon-flies, butterflies, blow-
flies, wasps, bees, beetles, &c., to fly in a large bell jar, one side of which was
turned to the light, the other side being rendered opaque by dark pigment ;
by throwing young pigeons and birds from the hand into the air for the first
time ; by repeated observation of the flight of tame and wild birds ; by stiffen-
ing, by tying up, and by removing portions of the wings of insects and birds ;
by an analysis of the movements of the travelling surfaces of quadrupeds,
amphibia, and fishes; by the application of artificial fins, flippers, tails, and
wings to the water and air; and by repeated dissections of all the parts directly
and indirectly connected with flight.

Professor Marey obtained his results by gilding the extremities and mar-
gins of the wings of the insect with minute portions of gold leaf; by the
application of the different parts (tip and anterior margin) of the wing of the
insect to a smoked cylinder rotating at a given speed, the wing being made to
record its own movements ; by the captive and free flight of birds, which carried
on and between their wings an apparatus which, by the aid of electricity, regis-
tered the movements of the wings on a smoked surface, travelling at a known
speed in a horizontal direction ; and by the employment of an artificial wing,
constructed on the plan recommended by Bore.11, CHABRIER, STRAUS-DURCK-
HEIM, GIRARD, and others.

Professor Margy describes and figures a captive insect (the wasp) with its
wings forming figure of 8 loops,* and a free insect, with its wings describing a
waved track,+ precisely similar to what I described and figured in a variety of
ways in my memoir.{ He also shows that the tip of the wing of the bird,
because of its alternately darting out and in during extension and flexion,
describes an ellipse. This, curiously enough, is another of the many points in
which I have anticipated this author, and one which I took special pains to
establish,§ having in my memoir devoted no less than ten figures || to its illus-
tration. Professor MArgy’s views may therefore be regarded as confirmatory
of my own, as the following brief passage, selected from one of his papers, will
show. He writes :—‘ But if the frequency of the movements of the wing vary,
the form does not. It is invariably the same—vt is always a double loop—a

* Revue des Cours Scientifiques de la France et de I’Etranger, 13 Février 1869, page 175, figure
5. Professor Margy represents the wing of the wasp as fanning the air in a vertical direction. In
reality, the wing of the wasp and of most insects is made to vibrate very obliquely, and in a more or
less horizontal direction.

t Revue des Cours Scientifiques et de la France et de l’Etranger, 13 Février 1869, pages 173,
174, and 176.
? { Trans. Linn. Society, Vol. XXVI, page 233, Diagrams 5 and 6; page 249, Diagrams 14, 15,
and 16; Plate XV. figures 59 and 61. Vide introduction to present memoir.

§ Op. cit., pages 247, 248, 249, and 250.

|| Op. cit., pages 248 and 249, Diagrams 7, 8, 9, 10, 11, 12, 13, 14, 15, and 16.

VOL. XXVI. PART II. 4R

304 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

Jigure of 8. Whether this figure be more or less apparent, whether its branches
be more or less equal, matters little ; it exists, and an attentive examination will
not fail to reveal it.”*

Professor Marery’s experiments, I may add, have been repeated and verified
in England by Mr Senecat. This investigator also represents and describes the
double loop and figure of 8 movements.t These two sets of experiments con-
ducted independently, and after a considerable interval, by M. Marry and Mr
SENECAL respectively, will, I hope, suffice to establish the absolute correctness
of the “ Figure of 8 or Wave theory of Flight.”

NATURAL FLIGHT.{

Method of Testing the Accuracy of the Figure of 8 Theory of Wing Mote-
ments.—The correctness of the figure of 8 theory of flymg may be readily estab-
lished by a careful study of the rapidly vibrating wing of the wasp or common
blow-fly.

If the body of the former be held, and the wing made to vibrate in front of
a dark screen, it will be found that not only the tip but also the margins of the
wing describe a figure of 8 track in space.

It will further be observed that the planes of the wing are as a rule
reversed during the down and up strokes ; nay, more, that the angles of inclina-
tion made by the surfaces of the wing with the horizon vary at every stage of
the wing’s progress, this variation in the angles being accompanied by a varia-
tion in the curves occurring on the anterior and posterior margins, as already
explained. As a consequence, the wing is moving in all its parts at the same
time—a somewhat remarkable occurrence, and calculated, it appears to me, to
excite the curiosity, if it does not rivet the attention of physiologists. The wing
of the insect is, with few exceptions, more flattened than that of the bat and
bird, a circumstance which enables it, when it is made to vibrate in a more or
less horizontal direction, and when its planes are reversed at the end of each
stroke, to apply its under or ventral surface to the air when it is urged
from behind forwards, and its upper or dorsal one when urged from before
backwards (figures 3 and 4, page 338). It sometimes happens that the
posterior margin of the wing is rotated in an upward direction at the end
of the forward stroke, and in this case it is the under surface of the wing which
is effective during the backward stroke (vide g hij k J of figure 19, page 351).

* Méchanisme du vol des insectes—comment se fait la propulsion. Revue des Cours Scientifiques
de la France et de lEtranger, 20th March 1869. :

+ Fifth Annual Report of the Aéronautical Society of Great Britain for 1870, pages 42-47.
Figures 1, 2; Diagrams 1-4.

{ Artificial flight is described at page 402.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 305

When the wing acts in this manner, it is the under or ventral surface which is
effective both during the forward and backward strokes. The wing, during
the back stroke occasions very little friction, from its being placed in a more
or less horizontal position—this position being favourable to its affording a
maximum of support. The upper and under surfaces of the wing are applied
to the air alternately, more particularly when the insect is fixed, or when it is
hovering in one spot. When it is flying at a high horizontal speed, and when
the wing is made to oscillate in a slightly vertical direction, as in the butterfly
(figures 29, 30, 31, 32, 33, and 34, page 360) and dragon-fly (figures 35, 36, 37,
and 38, page 361), it is the under or concave surface of the pinion which does
the principal part of the work, this attacking the air both during the down or
forward stroke and the up or backward stroke, like a boy’s kite, as explained
at pages 349 and 350, figures 16 and 17. The direction of the stroke varies
slightly according to circumstances, but it will be quite proper to assume that
the wing of the insect is made to vibrate in a more or less horizontal direction,
and that of the bird and bat ina more or less vertical direction. By a slight
alteration in the position of the body, or by a rotation of the wing in the
direction of its length, the vertical direction of the stroke is converted into
a horizontal direction, and vice versa. The facility with which the direction of
the stroke is changed is greatest in insects ; it is not uncommon to see them
elevate themselves by a figure of 8 horizontal screwing motion, and then, sud-
denly changing the horizontal screwing into a more vertical one, to dart rapidly
forward in a curved line. The horizontal screwing movement is represented at
figures 2, 3, 4, 5, 6, 7, and 10, pages 336, 338, 340, and 341; and the vertical
screwing at figures 12 and 13, page 342. The horizontal action of the insect’s
Wing is described at pages from 336 to 341 inclusive, and the vertical action at
pages from 347 to 355 inclusive. The vertical action of the bat and bird’s wing
is described at page 342, and at pages from 366 to 397 inclusive. Whether the
Wing is made to vibrate vertically or horizontally, it, practically speaking, in
progressive flight, strikes downwards and forwards during the down stroke, and
upwards and forwards during the up stroke, as fully explained at pages 344
and 345.
Compound Rotation of the Wing.—The wing during its vibration rotates
upon two separate centres, the tip rotating around the root of the wing as an
axis (short axis of wing), the posterior margin rotating around the anterior margin
(long axis of wing). This compound rotation goes on throughout the entire
down and up strokes, and is intimately associated with the power which the
wing enjoys of alternately seizing and evading the air.
The Wing inclined Forwards at the End of the Down Stroke and Backwards
at the End of the Up Stroke.—-1 had my attention first strongly directed to the
screwing figure of 8 action of the wing by closely observing the twisting figure

306 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

of 8 movements made by the pectoral fins and tails of fishes, and from finding that
in the beetle, blow-fly, and wasp (anterior wings), the posterior margin and body
of the wing were inclined forwards (fig. 1 a) with reference to the head of the
insect, at the end of the down stroke, and backwards (fig. 1 6) at the end of the
up stroke.

Fig. 1.

The Wing Rotates upon its Long Axis.—This at once suggested a rotation of the
wing upon its long axis along its anterior margin, or, what is practically the
same thing, a folding and plaiting of the posterior or thin yielding margin of
the wing around the anterior semi-rigid and comparatively unyieldmg margin
—a certain amount of rotation, or what is equivalent thereto, being necessary
to reverse and change the planes of the wing at each stroke.

The Wing Twists and Untwists during its action.—I further observed that the
planes of the wing were not only changed at the end of each stroke, but that the
wing at this juncture was twisted upon itself, the outer portion of the posterior
margin of the wing at the end of the down or forward stroke being inclined
Jorwards (g of fig. 2), while the inner portion was inclined backwards (r of fig. 2) ;
whereas at the termination of the up or backward stroke, the outer portion of
the posterior margin was inclined backwards (a of fig. 2), while the inner
portion was inclined forwards (s of fig. 2).

The Image produced on the Eye by the Wing in Motion is Concavo-Convea,
and Twisted.—I likewise discovered that the blur or impression produced on the
eye by the rapidly oscillating wing was twisted upon itself (fig. 1cdh, eg f), and
more or less concave above (c d ¢ fig. 1), and convex below (fg h fig. 1), a circum-
stance which, while it strongly corroborated the opinion that the wing rotated
upon its long axis during its vibration indicated that the twisting and reversal

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 307

of the planes of the wing occurred more especially at the end of the down and
up strokes. I inferred this from observing that the angle made by the wing
with the horizon is greater towards the termination than towards the middle of
the strokes. This could readily be ascertained by looking at the blur produced
by the oscillating wing edgewise, and this view revealed what is perhaps the
most important feature in wing movements, viz., that the tip of the wing during
its vibrations describes a scooped out (cde fig. 1) figure of 8 track as repre-
sented at 1,2, 3, 4, 5, 6,7, 8, 9,10, 11, 12, 13, and 14 of fig. 2.

The Direction of the Stroke of the Wing in the Insect—what Effective and what
Non-effective—the Kite-like Action of the Wing.—This view also showed that the
wing of the insect is made to vibrate in a more or less horizontal direction (figs.
3 and 5, page 338, Plate XI. fig. 4), in which respect it differs somewhat from
the wing of the bat and bird, these being worked more or less vertically (Plate
XI. figs. 5 and 6, and Plate XIV. figs. 18 and 19). The oblique action of the
pinion is necessary to avoid the resistance of the air during the up stroke, the
wing of the insect being in one piece, and having in many cases no adequate
apparatus for diminishing its area during its ascent. One great advantage
gained by the wing of the insect reversing its planes at the end of each stroke
consists in the great length of the effective stroke—the wing flying backwards
and forwards like a true kite, and tacking upon the air so suddenly as to
occupy very little either of time or space.* The period occupied by the wing
in reversing does not apparently amount to more than one-eighth of the time
taken up by one entire stroke, so that something like seven-eighths of the
area mapped out by the rapidly vibrating wing represents buoying area—the
remaining eighth slip. This, put in other words, simply means that in one
passage of the wing from behind forwards (down stroke) the pinion is effective
in seven-eighths of its course and non-effective in one-eighth, the same remark
being applicable to the passage of the wing from before backwards (up stroke).

The Wing Atiacks the Air at various Angles.—It is Just possible that even less
than one-eighth is devoted to slip, from the fact that the wing when it is being
reversed is slowed and applied to the air at an increased angle—a surface
which makes a large angle with the horizon, giving, when forced against the
air at a low speed, as much support as a similar surface whose inclination is
less, but whose speed is higher. As the wing attacks the air during the down
and up strokes at various angles, those angles being greatest when the wing
travels slowest, and least when the wing travels most rapidly, it follows that the
wing adapts itself to the resistance opposed to its passage by the air, and always
extracts the maximum of support from it, The wing, in this respect, differs

* The movements of the wing somewhat resemble those of a sailing ship. The wing and ship
both tack upon the wind, and both change their tack or reverse abruptly. The changing of the tack
1s moreover always accompanied by a slowing or diminution of the speed.

VOL. XXVI. PART II. 4AS

398 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

widely from the screw propellors at present in use—the blades of these propel-
lors always striking at a given angle and in the same direction. The advantage
in favour of the wing as compared with the screw as employed in navigation
is very great, and not at present understood.* The area mapped out by the
wing during the effective stroke and while reversing; the various angles made
by the surfaces of the wing with the horizon in its passage to and fro; the
rotating and twisting of the posterior or thin margin of the wing round the
anterior or thick margin ; and the figure of 8 track made bythe tip of the wing
during its action, as seen in the wasp, are shown at figs. 3, 4, 5, and 6.

Analysis of the Movements of the Wing of the Wasp, Reversal of the Planes
of the Wing, Reciprocating Action, &c.—In the wasp the wing commences the
down or forward stroke at a of figures 3 and 5; and it will be observed that
the angle which it makes with the horizon (x of fig. 5) is something like 45°.
At 6 (figures 3 and 5) the angle is slightly diminished, partly because of a rota-
tion of the wing along its anterior margin (long axis of wing), partly from
increased speed, and partly from the posterior margin of the wing yielding to a
greater or less extent.

At ¢ the angle is still more diminished from the same causes.

At d the wing is slowed slightly, preparatory to reversing, and the angle
made with the horizon (2) increased.

At e the angle, for the same reason, is still more increased; while at / the
wing is at right angles with the horizon. It is, in fact, in the act of reversing. —

For specific differences between the screws formed by the wings and the propellors employed in
navigation, see memoir by the author, Trans. Linn. Society, vol. xxvi. pages 228, 229, 230, and 231.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 3909

At g the wing is reversed, and the up or back stroke commenced.

The angle made at gis, consequently, the same as that made at a (45°), with
this difference, that the anterior margin and outer portion of the wing, instead
of being directed forwards, with reference to the head of the insect, are now
directed backwards.

During the up or backward stroke all the phenomena are reversed, as shown
atghijk/ of figures 4 and 6 ; the only difference being that the angles made by
the wing with the horizon are somewhat less than during the down or forward
stroke—acircumstance which facilitates the forward travelof the body, while it en-
ables the wing during the back stroke still to afford a considerable amount of sup-
port. This arrangement permits the wing to travel backwards when the body
is travelling forwards; the diminution of the angles made by the wing in the
back stroke giving very much the same result as if the wing were striking
in the direction of the travel of the body. The slight upward inclination of
the wing during the back stroke permits the body to fall downwards and for-
wards to a slight extent at this peculiar juncture, the fall of the body, as will be
more fully explained hereafter, contributing to the elevation of the wing.

If figure 5, representing the down or forward stroke, be placed upon figure
6, representing the up or backward stroke, it will be seen that the wing crosses
its own track more or less completely at every stage of the down and up strokes.
As, moreover, the wing draws a current after it, and is pursued in its passage
from above downwards by a stream of air which it meets in its passage from
below upwards, it follows that the pinion, during the down or forward stroke,
creates a current on which it operates during the up or backward stroke, and
vice versa; hence the reciprocating action of the wing.

The wing reciprocates most perfectly, and the figure of 8 is most dis-
tinct when the insect is fixed artificially, or when it is hovering of its own accord
in a given spot, as is well shown at a bc defghijkimnop of fig. 8,
p. 340, where the wing is represented as screwing steadily downwards.

Points wherein the Wing differs from the Scull of the Boatman.—The down-
ward screwing movement of the wing somewhat resembles the action of an oar
in sculling, as represented at ab, cd, wx s, of fig. 7, the
cross movement occasioned by the rotation of the
Wing on its long axis as it darts. to and fro being
shown at mn, op, qr. There is, however, this
marked difference. Itis the wyper surface of the oar
which is effective in sculling, whereas it is the wnder
surface of the wing which is effective in flying.* This 3
is accounted for bythe fact that the oar simply propels Fic. 7.
—the boat being buoyant, the wing propelling and

* A precisely similar difference is found to exist between the aérial or flying wing and the subaquatic
VOL. XXVI. PART II. aradl

340 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

likewise elevating. There is this further difference. The margins of the blades of
the oar are of the same thickness, the axis of rotation running midway between
the two ; the anterior margin of the wing, on the contrary, is much thicker than
the posterior one, the axis of rotation corresponding to the former. The oar, as
far as the margins of its blade are concerned, is as it were concentric, the wing
eccentric. As the downward screwing movement of the wing, in virtue of the
action and reaction of the wing and air upon each other, is at once converted
into an upward screwing movement, as shown at dU ed ¢é fof Wilf Kk Um wv
o’ p’ of fig. 9, it follows that the body of the insect is rapidly but steadily elevated
in an almost vertical wave-line. The impulse is communicated to the wing at
points corresponding to the heavy portions of the line in figure 8, and the
corresponding upward recoil is indicated at similar points in figure 9.

Fig. 8. Fig. 9.

How the Figure of 8 is Unravelled, and becomes a Waved-Track.—When the
insect flies in a horizontal direction, and the speed attained increases with the
duration of flight, the wing reciprocates less and less perfectly, because the figure
of 8 sweeps described by it are converted into a looped and then a waved track,
asrepresentedatabcdefghijkimnopqrst of figure 10 (p. 341); the cor-
responding looped and waved track due to recoil being shown at similar letters
of figure 11 (p. 341). When the horizontal speed attained by the insect is high,

or diving wing. In the gannet, cormorant, merganser, grebe, &c., which fly under the water, it is the upper
or dorsal surface of the pinion which gives the effective stroke, whereas in aérial flight it is the under or
ventral surface. This is proved by the fact that in the penguin and great auk, which are incapable of flying
out of the water, and confine their efforts to diving or swimming under it, the wing is actually twisted
round, so that the dorsal surface of the pinion occupies the position normally occupied by the ventral surfaces
in all other birds. This is necessitated by the fact that a diving bird, seeing it is of lighter specific
gravity than the water, must always fly downwards ; in other words, it must counteract buoyancy
as the flying bird counteracts gravity—buoyancy forcing the diving bird to the surface of the water in
the same way that gravity drags the flying bird to the surface of the earth. Levity and weight are
therefore separate forces, and act under diametrically opposite conditions, levity being quite as useful to
the diving bird as weight to the flying one. The wings of diving birds are applied to the water
precisely in the same manner as the flippers of the seal, sea bear, walrus, turtle, porpoise, whale, manatee,
&c, All these animals are lighter than the water, and, as a consequence, their travelling surfaces
to be effective must act from below as in the case of the scull. It is the reverse in the air, the

travelling surfaces acting invariably from above. For further development of this view see footnote to
page 371.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 341

the wing is successively and rapidly brought into contact with innumerable
columns of undisturbed air. It consequently is a matter of indifference whether
the wing is carried at a high speed against undisturbed air, or whether it operates
upon air travelling at a high speed (as, e.g., the artificial currents pro-
duced by the rapidly reciprocating action of the wing). The result is the same
in both cases, inasmuch as a certain quantity of air is worked up under the
Wing, and the necessary degree of support and progression extracted from it.
It is, therefore, quite correct to state, that as the horizontal speed of the body
increases the reciprocating action of the wing decreases, and vice versa. In
fact, the reciprocating and non-reciprocating function of the wing in such cases
is purely a matter of speed. If the travel of the wing is greater than the hori-
zontal travel of the body, then the figure of 8 and the reciprocating power of
the wing will be more or less perfectly developed, according to circumstances.
If, however, the horizontal travel of the body is greater than that of the wing,
then it follows that no figure of 8 will be described by the wing, that the
wing will not reciprocate to any marked extent, and that the organ will describe

a= _'—s=_g
ue L

Fig. 10,

ss a

Fig. 11.

a waved track, the curves of which will become less and less abrupt, z.¢., longer
and longer in proportion to the speed attained. The downward looped track
represented at fig. 10, is at once converted into an upward looped track, as shown
at figure 11, in virtue of the action and reaction of the wing and air upon each

342 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

other, the body of the insect being carried along a waved line obliquely upwards
and forwards (q 7's ¢, fig. 11, p. 341). The waved track made by the wing is gene-
rated by the figure of 8 loops being gradually opened out, these becoming less and
less distinct as the insect advances, as is more especially shown at nopqrst of
both figures (10 and 11, p. 341). The impulse is communicated to the wing at
acegikmoqs of fig. 10, and the upward recoil at corresponding letters of fig. 11.

The waved track formed by the ascent and descent of the wing of the bat
and bird is originated in a similar manner, but in this case the figure of 8
loops are disposed more vertically, because of the more vertical direction of
the stroke, as shown at efghijk/ of figure 12. (Vide also Plate XI. figures
5 and 6). In this figure (12) the oar, as seen at a6, xs, and ed, represents the

Fig. 12.

wing of the bat and bird at the beginning, middle, and termination of the
down stroke—the little oar, mn op qv7, indicating the cross action of the wing.
The large oar is more especially engaged in elevating, the little one in pro-
pelling. The manner in which the figure of 8 loops made by the wing of the
bat and bird during its ascent and descent are opened out or unravelled by the

horizontal travel of the body is shown at abcdefghijkimnop of figure 13;
the completed waved track being seen at stu vw of the same figure.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 343

When the Wing Ascends the Body Descends, and vice versa.—As the body
of the insect, bat, and bird falls forwards ina curve when the wing ascends, and
is elevated in a curve when the wing descends, it follows that the trunk of the
animal is urged along a waved line, as represented at 1, 2, 3,4, 5 of figure 14, p. 344,
the waved line acegi of the same figure giving the track made by the wing.
I have distinctly seen the alternate rise and fall of the body and wing when
watching the flight of the gull from the stern of a steam-boat.

The direction of the stroke in the insect (figs. 3, 5, and 8, pp. 338, 340), as
I have already explained, is much more horizontal than in the bat or bird (figs.
12 and 13, p. 342). In either case, however, the down stroke must be delivered
in a more or less forward direction. This is necessary for support and pro-
pulsion. <A horizontal to and fro movement will elevate, and an up and down
vertical movement propel, but an oblique forward motion is requisite for pro-
gressive upward flight.*

The Wing during its Vibrations moves on the Surface of an Imaginary
Sphere.—All wings are convex above and concave below. ‘This shape is neces-
sary to enable the wing to evade the air during the up stroke, and to seize it
during the down one. The concave surface is presented during the up stroke,
and the concave one during the down stroke—the resistance experienced by a
concave surface when compared with a convex one being something like two to
one. The resistance is further increased by the wing being made to descend
with greater rapidity than it ascends. In whatever direction the wing turns
during the up stroke its movements are calculated to evade the air, and in
whatever direction it turns during the down stroke they are calculated to
seize it. This arises alike from the shape of the wing and the manner in which
itis applied to the air. Thus, in the insect in progressive flight the wing during
the up stroke describes a curve which is directed upwards and forwards. In the
bat and bird, where the wing is drawn towards the body during the up stroke,
the wing describes a second curve, this curve being directed upwards and inwards
with reference to the body. The under or concave surface of the wing may,
therefore, be said to be moving on the surface of an imaginary sphere during
the up stroke—an arrangement which enables it to avoid the superincumbent
air by its upper or convex surface, while it affords a certain amount of support
and ascensional power by its under or concave surface, this latter acting partly
as a kite and partly as a parachute. The wing may, in fact, be said to climb
during the up stroke ; and this climbing is so adroitly performed that two objects
are served by it—the superimposed air being avoided, and the body bearing
the wing being supported. In the climbing movement the anterior margin of
the wing cleaves a passage from behind upwards and forwards for the body

: Mi aE the Mechanism of Flight, by the Author, Trans. Linn. Society, vol. xxvi. pages 214, 255,
and 256,

VOL. XXVI. PART II. £0

344 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

and posterior margin, the root in like manner cleaving a passage from without
inwards and upwards for the body and tip. It is in this way that the wing
presents a sharp cutting edge during the up stroke, a remark which applies even
to the rowing feathers (quill feathers) of the wing of the bird. The ascent of
the wing, as will be subsequently explained, is favoured by the reaction of the air
on its under surface, and by the downward and forward fall of the body. If
the wing was not concavo-convex in form, and made to oscillate on the surface
of an imaginary sphere, it would be impossible for it alternately to avoid and
seize the air while it is rismg and falling. When the wing descends or
makes the down stroke, as it is termed, it also rotates on the surface of the
imaginary sphere in question. In this case, however, it is the concave or under
surface of the wing which is active, and the rolling takes place in such a manner
(it is outwards, downwards, and forwards) as actually greatly to increase the sup-
port afforded—the air, which was dispersed and avoided during the up stroke,
being now collected together and seized with avidity. It would be difficult to
conceive a more simple or effective arrangement.

The Natural Wing, when Elevated and Depressed, must move Forwards.—It
is a condition of natural wings, and of artificial wings constructed on the prin-
ciple of living wings, that when forcibly elevated or depressed, even in a strictly
vertical direction, they inevitably dart forward. This is well shown in figure 14.

If, for example, the wing is suddenly depressed in @ vertical direction, as
represented at a b, it at once darts downwards and forwards in a curve to ¢, thus
converting the vertical down stroke into a down oblique forward stroke. Wf,
again, the wing be suddenly elevated in a strictly vertical direction, as at ¢ d, the
wing as certainly darts upwards and forwards in a curve to e, thus converting
the vertical up stroke into an upward oblique forward stroke. The same
thing happens when the wing is depressed from e to /, and elevated from g to
h. In both cases the wing describes a waved track, as shown at ¢ g, g 7, which
clearly shows that the wing strikes downwards and forwards during the down
stroke, and upwards and forwards during the up stroke. The wing, in fact, is
always advancing, its under surface attacking the air like a boy’s kite. If, on
the other hand, the wing be forcibly depressed, as indicated by the heavy waved
line a c, and left to itself, it will as surely rise again, and describe a waved
track, as shown at ce. This it does, in virtue of its flexibility and elasticity,

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 345

aided by the recoil obtained from the air. In other words, it is not necessary
to elevate the wing forcibly in the direction ¢ d to obtain the upward and for-
ward movement ¢ ¢. One single impulse communicated at a, causes the wing
to travel to ¢, and a second impulse communicated at e, causes it to travel to 7.
It follows from this that a series of vigorous down impulses would, ¢/f a certain
interval was allowed to elapse between them, beget a corresponding series of up
impulses, in accordance with the law of action and reaction, the wing and the
air under these circumstances being alternately active and passive. I say if a
certain interval was allowed to elapse between every two down strokes, but
this is practically impossible, as the wing is driven with such velocity that
there is positively no time to waste in waiting for the purely mechanical
ascent of the wing. That the ascent of the pinion is not, and ought not to be,
entirely due to the reaction of the air, is proved by the fact that in flying
creatures (certainly in the bat and bird) there are distinct elevator muscles and
elastic ligaments, delegated to the performance of this function. The reaction
of the air is therefore only one of the forces employed in elevating the wing ;
the others, as I shall show presently, are vital and vito mechanical in their nature.
The falling downwards and forwards of the body when the wings are ascending
also contribute to this result.

The Wing acts as a true Kite both during the Down and Up Strokes.—lIf, as
I have endeavoured to explain, the wing, even when elevated and depressed in
a strictly vertical direction, inevitably and invariably darts forward (figure 14, p.
344), it follows as a consequence that the wing, as already partly explained, flies
forwards as a true kite, both during the down and up strokes, as shown at
cdefghijkim of fig. 15, and that its under concave or biting surface, in
virtue of the forward travel communicated to it by the body in motion, is closely
applied to the air, both during its ascent and descent, a fact hitherto overlooked,
but one of considerable importance, as showing how the wing furnishes a per-
sistent buoyancy, alike when it rises and falls.

In figure 15 the greater impulse communicated during the down stroke is
indicated by the double dotted lines. The angle made by the wing of the bat and
bird with the horizon (a } of figure 15) is constantly varying, as in the insect wing,
as a comparison of ¢ with d, d with e, e with 7, and fwith g of figure 15 will
show, these letters having reference to supposed transverse sections of the

346 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

wing. Figure 15 also shows that the convex or non-biting surface of the wing
is always directed upwards, so as to avoid unnecessary resistance on the part
of the air to the wing during its ascent, whereas the concave or biting surface is
always directed downwards, so as to enable the wing to contend successfully
with gravity.

On comparing ¢d e/g of figure 15, p. 345, with a bcd of figures 3 and 5, p. 338,
it will be seen that the principle involved in the flight of the wing of the insect,
bat, and bird is essentially the same. The wing is, in short, in every instance, a
true kite, and flies forward in accordance with natural laws.

Where the Kite formed by the Wing differs from the Boy’s Kite.—The natural
kite formed by the wing differs from the artificial kite only in this, that the former
is capable of being moved in all its parts, and is more or less flexible and elastic,
thelatter bemgcomparativelyrigid. The flexibility and elasticity of the kite formed
by the natural wing is rendered necessary by the fact that the wing is articulated
or hinged at its root ; its different parts travelling at various degrees of speed in
proportion as they are removed from the axis of rotation. Thus the tip of the
wing travels through a much greater space in a given time than a portion nearer
the root. If the wing was not flexible and elastic, it would be impossible to
reverse it at the end of the up and down strokes, so as to produce a continuous
vibration. The wing is also practically hinged along its anterior margin, so that
the posterior margin of the wing travels through a much greater space in a
given time than a portion nearer the anterior margin. The compound rotation
of the wing is greatly facilitated by the flexible and elastic properties of the
pinion. It causes the pinion to twist upon its long axis during its vibration, as
already fully explained (see g, 7 and a, s of fig. 2, p. 336). The twisting
referred to is partly a vital and partly a mechanical act ; that is, it is occasioned
in part by the action of the muscles, and in part by the greater momentum
acquired by the tip and posterior margin of.the wing, as compared with the
root and anterior margin ; the speed acquired by the tip and posterior margin
causing them to reverse always subsequently to the root and anterior margin,
which has the effect of throwing the anterior and posterior margins of the
wing into figure of 8 curves. It is in this way that the posterior margin of the
outer portion of the wing is made to incline forwards at the end of the down
stroke (fig. 2 g, p. 386), when the anterior margin is inclined backwards, and
that the posterior margin of the outer portion of the wing is made to incline
backwards at the end of the up stroke (fig. 2 a, p. 336), when a corresponding
portion of the anterior margin is inclined forwards.

The Angles formed by the Wing in Action.—Not the least interesting feature
of the compound rotation of the wing, of the varying degrees of speed attained
by its different parts, and of the twisting or plaiting of the posterior margin
around the anterior, is the great variety of kite-like surfaces developed upon

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 247

its dorsal and ventral aspects. Thus the tip of the wing forms a kite
which is inclined upwards, forwards, and outwards, while the root forms a
kite which is inclined upwards, forwards, and inwards. The angles made by
the tip and outer portions of the wing with the horizon are less than those
made by the body, and those made by the body less than those made by
the root and inner portions. The inclined surfaces peculiar to any portion of
the wing become more inclined as the speed peculiar to said portion decreases,
and vice versa. The wing is consequently mechanically perfect, the angles made
by its several parts with the horizon being accurately adjusted to the speed
attained by its different portions during its travel to and. fro. From this
it follows that the air set in motion by one part of the wing is seized upon
and utilised by another, the inner and anterior portions of the wing supplying,
as it were, currents for the outer and posterior portions. This results from the
wing always forcing the air outwards and backwards. These statements admit
of direct proof, and I have frequently satisfied myself of their exactitude by
experiments made with natural and artificial wings.

In the bat and bird the twisting of the wing upon its long axis is more of a
vital and less of a mechanical act than in the insect, the muscles which regulate
the vibration of the pinion in the former (bat and bird), extending quite to the
tip of the wing.

The Body and Wings move in Opposite Curves.—I have stated that the wing
_ advances in a waved line, as shown at aceg i of figure 14, p. 344; and the same
remark holds true, within certain limits, of the body as indicated at 1, 2, 3, 4,
and 5 of the same figure. Thus, when the wing descends in the curved line
a ¢c, it elevates the body in a corresponding but minor curved line, as shown at
1, 2; when, on the other hand, the wing ascends in the curved line ¢ e, the body
descends in a corresponding but smaller curved line (2, 3), and so on ad infinitum.
The undulations made by the body are so triflmg when compared with those
made by the wing that they are apt to be overlooked. They are, however,
deserving of attention, as they exercise an important influence on the undula-
tions made by the wing, the body and wing swinging forward alternately, the
one rising when the other is falling, and vice versa. Flight may be regarded
as the resultant of three forces:—the muscular and elastic force, residing in
the wing, which causes the pinion to act asa true kite, both during the down
and up strokes; the weight of the body, which becomes a force the instant
the trunk is lifted from the ground, from its tendency to fall downwards and
forwards; and the recoil obtained from the air by the rapid action of the wing.
These three forces may be said to be active and passive by turns.

Analysis of the Down and Up Strokes in the Insect—the Terms Extension and
Flexion defined—As considerable confusion exists in the minds of most inves-
tigators as to the precise changes induced in the wing during the down and up

VOL, XXVI. PART It. 4x

348 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

strokes respectively, and in especial as to the manner in which the wing is
elevated, so as to avoid the resistance of the air and yet afford support, I have
felt it ncumbent upon me carefully to analyse the movements as observed in pro-
gressive flight. In insects the wings are variously arranged during the period
of repose. In some they are elevated above the body, as in the butterfly ; in
others, they are disposed on the same level with the body, and rest upon the
dorsal surface of the abdomen, as in the common fly ; in a third, the wings are
arranged partly on the sides and partly upon the dorsal aspect of the body, the
anterior or thick margin of the wing being in such cases directed downwards,
as in the cicada. -Thisis also the position occupied by the wings of the bat and
bird, the pinions, when not employed in flying, being folded upon themselves to
economise space. In some insects, as the ephemera or mayfly, the beetles,
locusts, &¢., the wings are also folded upon themselves during the intervals of
rest. The power which some wings possess of alternately folding, flexing, or
crushing their component parts together, and of extending and widely separating
them, has introduced the terms extension and flexion: extension, strictly speak-
ing, signifying the opening out or spreading of the pinion, and the carrying of
it away from the body in the direction of the head of the animal; flexion sig-
nifying the folding of the pinion, and the drawing of it towards the body in a
direction from before backwards. The terms extension and flexion, when
applied to insect wings, which are in one piece, and which consequently do not
admit of being alternately opened and closed to any great extent, are only
partly correct,—extension in the insect, signifymg the carrying of the
wing away from the body in a plane nearly on the same level with it in the
direction of the head ; flexion the drawing back or recovering of the wing until
it regains its original position.

The terms extension and flexion have, unfortunately, got mixed up with the
expressions the down and up strokes, from the fact that the wings of bats, birds,
and some insects are always extended towards the termination of the up strokes,
and flexed towards the termination of the down ones. This confusion is the
more natural as all wings when extended rotate upon their long axes in such a
manner that their posterior margins are screwed downwards and forwards.

In all wings, whatever their position during the intervals of rest, and whether
in one piece or in many, this feature is to be observed in flight. The wings are
slewed downwards and forwards, 7.¢., they are carried more or less in the direc-
tion of the head during their descent, and reversed or carried in an opposite
direction during their ascent. In stating that the wings are carried away from
the head during the back stroke, I wish it to be understood that they do not
therefore necessarily travel backwards in space when the insect is flying for-
wards. On the contrary, the wings, as a rule, move forward in curves, both
during the down and up strokes. The fact is, that the wings at their roots are

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 349

hinged and geared to the body so loosely that the body is free to oscillate in a
forward or backward direction, or in an up, down, or oblique direction. As a
consequence of this freedom of movement, and as a consequence likewise of the
speed at which the insect is travelling, the wings during the back stroke are for
the most part actually travellmg forwards. This is accounted for by the fact
that the body falls downwards and forwards in a curve during the up or return
stroke of the wings, and because the horizontal speed attained by the body is
as a rule so much greater than that attained by the wings, that the latter are
never allowed time to travel backward, the lesser movement being as it were
swallowed up by the greater. For a similar reason the passenger of a steam-
ship may travel rapidly in the direction of the stern of the vessel, and yet be
carried forward in space,—the ship sailing much quicker than he can walk,
While the wing is descending, it is rotating upon its root as a centre (short axis).
It is also, and this is a most important point, rotating upon its anterior margin
(long axis), in such a manner as to cause the several parts of the wing to
assume various angles of inclination with the horizon.
Figures 16 and 17 will supply the necessary illustration.

g EE LLM LLL LL 4
% PVQ vr

~.
'~

Sus
st
SS

%
“” Ss; 4
4- LLZLL 2
Tea
Fig. 17.

If, for example, we take the common blow-fly when reposing we will find
that the plane of the wing (fig. 16 a’) is arranged in the same plane with the
body, and that both are in a line with the horizon (zx’).* When, however, the

* Tt happens occasionally in insects that the posterior margin of the wing is on a higher level than

the anterior one towards the termination of the up stroke as shown at a (dotted line) of fig. 16. In
such cases the posterior margin is suddenly rotated in a downward and forward direction at the

300 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

wing is made to descend, it gradually, in virtue of its simultaneously rotating
upon its long and short axes, makes a certain angle with the horizon as repre-
sented at b. The angle is increased at the termination of the down stroke as
shown at ¢, so that the wing, particularly its posterior margin, during its descent
(A), is screwed or crushed down upon the air with its concavity or biting
surface directed forwards and towards the earth. The same phenomena are -
indicated at a bc of fig. 17, p. 349, but in this figure the wing is represented as
travelling more decidedly forwards during its descent, and this is characteristic
of the down stroke of the insect’s wing—the stroke in the insect being delivered in
a very oblique and more or less horizontal direction, as shown at Plate XI. fig. 4.
The forward travel of the wing during its descent has the effect of diminishing the
angles made by the under surface of the wing with the horizon. Compare 0 ¢ d
of fig. 17 with the same letters of fig. 16. At fig. 15, page 345, the angles fora
similar reason are still further diminished, and this latter figure gives a very
accurate idea of the kite-like action of the wing both during its descent and
ascent. The downward screwing of the posterior margin of the wing during
the down stroke is well seen in the dragon-fly at page 361, fig. 38. (In this
figure the arrows 7 s give the range of the wing.) At the beginning of the down
stroke (dragon-fly) the upper or dorsal surface of the wing (¢d//) is inclined
downwards and backwards, the under or ventral surface downwards and for-
wards. In other words, the anterior margin (¢ d) of the pinion is directed -
slightly upwards and forwards, the posterior margin (/) slightly downwards
and backwards. As the wing descends, which it does in a downward and
forward direction, the posterior margin (/) is screwed downwards and for-
wards until it assumes the position indicated by 7; the anterior margin (@ d)
inclining more and more upwards and backwards, as shown at gh. This rota-
tion of the posterior margin (7) round the anterior margin (g h) has the
effect of causing the different portions of the under surface of the wing to
assume various angles of inclination with the horizon, the wing attacking
the air like a boy’s kite. The angles are greatest towards the root of the wing
and least towards the tip. They accommodate themselves to the speed at which
the different portions of the wing travel—a small angle with a high speed giving
the same amount of buoying power as a larger angle with a diminished speed.
The screwing of the under surface of the wing (particularly the posterior margin)
in a downward direction during the down stroke is necessary to insure a sufficient
upward recoil, the wing being made to swing downwards and forwards pendulum
fashion, for the purpose of elevating the body, which it does by acting upon the air
as a long lever, and after the manner of a kite. During the down stroke the wing
beginning of the down stroke—the downward and forward rotation securing additional elevating

power for the wing. The posterior margin of the wing in bats and birds, unless they are flying down-
wards, never rises above the anterior one, either during the up or down stroke.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. BDA

is active—the air passive. In other words, the wing is depressed by a purely
vital act. This is proved by taking a living or dead blow-fly, and forcibly
depressing its wing in the direction of the head by the aid of a slender rod.
This act causes the wing to make various angles of inclination with the horizon,
as shown at abcde/g of fig. 18; but the instant the rod is removed the wing
obliterates the angles in question, and flies in an upward and backward direction
to its original position as indicated at g hij & lm of fig. 19.

= Z Z. L ZS
Se ee é ee ee —
5 a peer ee ee
Fig. 19.

This shows very satisfactorily that while a voluntary effort is required to
depress the wing, it is in some measure elevated, and the various inclined
surfaces which it makes with the horizon changed by the aid of an elastic
ligament or spring common to all wings. The down stroke is readily explained,
and its results upon the body obvious. The real difficulty begins with the up or
return stroke. Ifthe wing was simply to travel in an upward and backward direc-
tion from ¢ to a’ of fig. 16, page 349, itis evident that it would experience much
resistance from the superimposed air, and undo or negative the advantages secured
by the descent of the wing. What really happens is this. The wing does not travel
upwards and backwards in the direction ¢ b a’ of fig. 16 (the body be it remembered
is advancing), but upwards and forwards in the direction cdefg. This is brought
about in the following manner. The wing is at right angles to the horizon (72’) at ¢.
It is therefore caught by the air because of the more or less horizontal travel of
the body at 2, the elastic ligaments and other structures rotating the posterior
or thin margin of the pinion in an upward direction, as shown at g hi of figure
19, page 351, and de/g of figure 16, page 349. The wing by this partly vital
and partly mechanical arrangement is rotated off the wind in such a manner as
to keep its dorsal or non-biting surface directed upwards, while its concave or
biting surface is directed downwards. The wing, in short, has its planes so
arranged, and its angles so adjusted to the speed at which it is travelling, that
it darts up a gradient like a true kite, as shown at cd e/g of figures 16 and 17,
page 349. The wing consequently elevates and propels during its ascent as well
as during its descent. It is, in fact, a kite during both the down and up strokes.
The ascent of the wing is greatly assisted by the forward travel of the body.
Tt is further assisted by the downward and forward fall of the body. This
view will be readily understood by supposing, what is.really the case, that
the wing is more or less fixed by the air in space at 2 of figure 16, page 349,
and that the body, the instant the wing is fixed, falls downwards and forwards

VOL. XXVI. PART II. 4¥

302 DK PETTIGREW ON THE PHYSIOLOGY OF WINGS.

in a curve, which, of course, is equivalent to placing the wing above, and, so
to speak, behind the insect—in other words, to elevating the wing prepara-
tory to a second down stroke, as seen at g of figures 16 and 17, page 349.*

The Body ascends when the Wing descends, and vice versa.—The manner in
which the body falls downwards and forwards in
progressive flight is illustrated at figs. 20, 21, and
22.

At fig. 20 the body is represented at @ and ¢,
the wing at 6 and d, x supplying the fulcrum or
pivot on which the body and wing oscillate.

If the body (a) is elevated in the direction
c, the wing (0) of necessity descends in the direc-
tion h. If, on the other hand, the body (ce)
descends in the direction /, the wing (d) ascends
in the direction g. The ascent or descent of

Fig. 20. the wing is always very much greater than that
of the body, from the fact of the pinion acting as
a long lever. The remarks just made are true more especially of the body

* When a bird rises from the ground it runs for a short distance, or throws its body into the air
by a sudden leap, the wings being simultaneously elevated. When the body is fairly off the ground,
the wings are made to descend with great vigour, and by their action to continue the upward impulse
secured by the preliminary run or leap. The body then falls in a curve downwards and forwards, the
wings, partly by the fall of the body, partly by the reaction of the air, on their under surface, and
partly by the contraction of the elevator muscles and elastic ligaments being placed above, and to
some extent behind the bird—in other words, elevated. The second down stroke is now given, and
the wings again elevated as explained, and so on “ ad infinitum,” the body fallmg when the wings are
being elevated, and vice versa, as shown at fig. 14, p. 344. When a long-winged oceanic bird rises
from the sea, it uses the tips of its wings as levers for forcing the body up, the points of the pinions
suffering no injury from being brought violently in contact with the water. A bird cannot be said to
be flying until the trunk is swinging forward in space and taking part in the movement. The hawk,
when fixed in the air over its quarry, is simply supporting itself. ‘To fly, in the proper acceptation of
the term, implies to support and propel. This constitutes the difference between a bird and a balloon.
The bird can elevate and carry itself forward, the balloon can simply elevate itself, and must rise and
fallin a straight line in the absence of currents. When the gannet throws itself from a cliff the inertia
of the trunk at once comes into play, and relieves the bird from those herculean exertions required to
raise it from the water when it is once fairly settled thereon. A swallow dropping from the eaves of
a house, or a bat from a tower, afford illustrations of the same principle. Many imsects launch them-
selves into space prior to flight. Some, however, do not. Thus the blow-fly can rise from a level sur-
face when its legs are removed. This is accounted for by the greater amplitude and more horizontal —
play of the insect’s wing as compared with that of the bat and bird, and likewise by the remarkable
reciprocating power which it possesses when the body of the insect is not moving forwards. (Vide
figs. 3, 4, 5, and 6, page 338). When a beetle attempts to fly from the hand it extends its front
legs and flexes the back ones, and tilts its head and thorax upwards so as exactly to resemble a
horse in the act of rising from the ground. This preliminary over, whirr go its wings with immense
velocity, and in an almost horizontal direction, the body being inclined more or less vertically. The
insect rises very slowly, and often requires to make several attempts before it succeeds in launching
itself into the air. I could never detect any pressure communicated to the hand when the insect was —
leaving it, from which I infer that it does not leap into the air. The bees, I am disposed to believe,
also rise without anything in the form of a leap or spring. I have often watched them leaving the

petals of flowers, and they always appeared to me to elevate themselves by the steady play of their

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 353

and wing when oscillating on either side of the fixed point 2, this furnishing
the fulcrum on which the body and the wing alternately act. The pecu-
liarity, however, of the wing consists in the fact that it is a flexible lever and

kK g
bX

, NX
JN

Fig. 22.

acts upon yielding fulcra (the air), the body participating in, and to a certain
extent perpetuating the movements originally produced by the pinion. The
part which the body performs in flight is illustrated at fig. 21. Ata the body
is depressed, the wing being elevated and ready to make the down stroke at 0.
The wing descends in the direction ¢ d, but the moment it begins to descend
the body moves upwards and forwards (see arrows) in a curved line toe. As
the wing is attached to the body it is made gradually to assume the position 7.
The body is now elevated and the wing depressed, the under surface of the
latter being so adjusted that it strikes upwards and forwards as a kite would.
The body now falls downwards and forwards in a curved line to g, and in doing
this it elevates or assists in elevating the wing to 7. The pinion is a second
time depressed in the direction & /, which has the effect of forcing the body
along a waved track and in an upward direction until it reaches the point
m. ‘The ascent of the body necessitates the descent of the wing as at m. The
body and wing, as will be seen from this figure, are alternately above and beneath
a given line 22. The same points are shown at fig. 22, the only difference
being that the sweep of the wing is greater and the undulation made by the
body less abrupt, as seen in vigorous flight. At @ the body is depressed, and
the wing (2) elevated high above the body. The pinion (b) descends in the
direction ¢ d, and forces the body in an upward curve to e. The body (e) is
now elevated and the wing (/) depressed. The body (¢) falls downwards and
wings, which was the more necessary, as the surface from which they rose was in many cases a yield-

ing surface. The falling forward of the body during flight was indicated in my Memoir “On the
Mechanism of Flight,” Trans. Linn. Society, vol. xxvi. p. 226.

354 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

Jorwards ina curve to g, the pinion (/) by this act being made to describe the
segment of a circle h 77, its under concave surface being applied to the air like
a kite all the time. (It is thus that the wing elevates and sustains during the
up stroke.) The wing (7) is made to descend in the direction & /, and forces
the body (g) along an upward curve until it arrives at m, its subsequent fall
elevating the wing (7) in the direction 0 p. Here again, the body and wing
play alternately on either side of a given line 2 2’,

A. careful study of figs. 20, 21, and 22 (pages 352, 353) shows the great im-
portance of the twisted configuration and curves peculiar to the natural wing.
If the wing was not curved in every direction it could not be rolled on and off
the wind during the down and up strokes, as seen more particularly at fig. 22.
This, however, is a vital point in progressive flight. The wing (0) is rolled on to
the wind in the direction ¢ d, its under concave or biting surface being crushed
hard down with the effect of elevating the body to e. The body falls to g,
and the wing (/) is rolled off the wind in the direction / 2, and elevated partly
by the action of the elevator muscles and elastic ligaments, and partly by the
reaction of the air, operating on its under or concave biting surface, until it
assumes the position 7, The wing is therefore to a certain extent resting
during the up stroke. The concavo-convex form of the wing is admirably
adapted for the purposes of flight. In fact, the power which the wing possesses
of always keeping its concave or under surface directed downwards and more
or less forwards enables it to seize the air at every stage of both the up and
down strokes so as to supply a persistent buoyancy. The action of the natural
wing is accompanied by remarkably little slip—the elasticity of the organ, the
resiliency of the air, and the contraction and relaxation of the elastic ligaments
and muscles all co-operating and reciprocating in such a manner that the
descent of the wing elevates the body, the descent of the body aided by the
reaction of the air and the contraction of the elastic ligaments and muscles
elevating the wing. The wing during the up stroke arches above the body after the
manner of a parachute, and in turn prevents the body from falling. The
sympathy which exists between the parts of a flying animal and the air on
which it depends for support and progress is consequently of the most intimate
character.

The up stroke (B of figures 16 and 17, page 349), as will be seen from the fore-
going account, is a compound movement due in some measure to recoil or resist-
ance on the part of the air—to the contraction of the muscles, elastic ligaments,
and other vital structures, to the elasticity of the wing, and to the falling of the
body in a downward and forward direction. The wing may be regarded as rotating
during the down stroke upon 1 of figure 16, page 349, which may be taken to
represent the long and short axes of the wing, and during the up stroke upon
2, which may be taken to represent the yielding fulcrum furnished by the air.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 305

The same points are illustrated at 1 and 2 of figure 17, page 349, allowance
being made in this case for the greater horizontal travel of the body during
the down (A C) and up (BD) strokes, the increased horizontal travel of the
body, as already pointed out, having the effect of diminishing the angles made
by the under surface of the wing with the horizon during its vibrations.

The Wing acts upon Yielding Fulcra.—The chief peculiarity of the wing, as
has been stated, consists in the fact that it is a twisted flexible lever specially
constructed to act upon yielding fulcra (the air). The points of contact of the
wing with the air are represented atabedefghij kl respectively of figures 16
and 17, page 349, and the imaginary points of rotation of the wing upon its long
and short axes at 1, 2, 3, and 4 of the same figures. The assumed points of
rotation advance from 1 to 3, and from 2 to 4 (vide arrows marked r and
s, fig. 17). The actual points of rotation correspond to the little loopsabcdef
ghijkTlof same figure ; the descents of the wing to A and C, and the ascents
to Band D. When the wing is in the position represented at g of figures 16
and 17, page 349, it is ready to begin a second down stroke, that is, it is
screwed in a downward and forward direction. At z the second down stroke
(C) is completed ; at 7 the second up stroke is begun, the posterior margin of
the wing being gradually rotated in an upward direction to prepare it for making
the return or up stroke (D), as shown at 74/m. A third down stroke (E, fig.
16) is commenced at m and completed at o.

Weight contributes to Horizontal Flight.—That the weight of the body plays
an important part in the production of flight may be proved by a very simple
experiment. If two quill feathers are fixed into an ordinary cork, as repre-
sented at fig. 23, p. 356, and the apparatus is allowed to drop from a height,
the cork does not fall vertically downwards, but downwards and forwards in a
curve, and for the following reasons. The feathers a 6 are twisted flexible
inclined planes, which arch in an upward direction. They are, in fact, true wings
in the sense that an insect wing in one piece is a true wing. When dragged
downwards by the cork (c), which would, if left to itself, fall vertically, they
have what is virtually a down stroke communicated to them. Under these
circumstances they inevitably dart forward ; a struggle ensuing between the cork
tending to fall vertically and the feathers tending to travel in a horizontal
direction. As a consequence, the apparatus describes the curve d e/g before
reaching the earth, #7. This is due to the action and reaction of the feathers
and air upon each other, and to the influence which gravity exerts upon the
cork. The forward travel of the cork and feathers, as compared with the space
through which they fall, is very great. Thus, in some instances, they advanced
as much asa yard and a half in a descent of three yards.

_ When artificial wings constructed on the principle of natural ones (v7de fig. 24,
p. 357), with stiff roots (¢, a), tapering semi-rigid anterior margins (a 0, ¢ d), and
VOL. XXVI. PART IT. 47%

356 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

thin yielding posterior margins (¢/, g /), are allowed to drop from a height (7),
they describe double curves in falling, as shown at mnol, ij kl, the roots of
the wings (c¢,@) reaching the ground first, a circumstance which proves the
greater buoying power of the tips of the wings. I might refer to many other
experiments made in this direction, but sufficient have been adduced to show
that weight, when acting upon wings, or, what is the same thing, upon elastic
twisted inclined planes, must be regarded as an independent moving power, But

oN
[le

-
-
- f-
aa
re

Lies VR obec ih Zz
Fig. 23.

for this circumstance flight would be at once the most awkward and laborious form
of locomotion, whereas in reality it is incomparably the easiest and most graceful.*
The power which rapidly vibrating wings have of sustaining a body which tends
to fall vertically downwards, is much greater than one would naturally imagine,
from the fact that the body, which is always beginning to fall, is never per-
mitted actually to do so. Thus, when it has fallen sufficiently far to assist in
elevating the wings, it is at once elevated by the vigorous descent of those
organs. The body consequently never acquires the downward momentum
which it would do if permitted to fall through a considerable space uninter-
ruptedly. It is easy to restrain even a heavy body when beginning to fall,
while it is next to impossible to check its progress when it is once fairly
launched into space and travelling rapidly in a downward direction (see foot-
note to page 371).

* The importance to be attached to weight in flight is variously explained in my Memoir on the
subject, Trans. Linn. Society, vol. xxvi. pages 218, 219, 246, 260, and 261,

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 307

Mechanical Theory of the Action of the Insect’s Wing as stated by CHABRIER.—
In one instance only, according to CHaprigr,* are the muscles of flight in
insects inserted directly into the root of the wing. This solitary example is

\
K y
R /
, /
\ /
‘ /
\ Hany bs
\ He KG

Seco

Fig 24.

the dragon-fly. CuHasrier regards the action of the insect’s wing as purely
mechanical. His argument may be stated in a few words. He observes,
that whereas the muscles which propel the wings of the insect are, with one
exception (the dragon-fly), confined to the interior of the thorax, that there-
fore they exert no direct influence upon the wings. He further gives it as
his opinion, that the wings are actuated by the muscles only during the down
stroke, and that the up stroke is entirely due to the reaction of the air—in
fact, that if the wings only be depressed rythmically, the air will do the
remainder of the work. Unfortunately for this theory there is no time to
wait for the reaction of the air, the wings being driven with such velocity as
necessitates their being partly elevated either by elastic ligaments or elevator
muscles, in addition to the reaction of the air (vide page 345). CHABRIER, as will
be seen, delegates to the air the task of reversing the planes of the wing, and
of conferring upon it those peculiar curves which, overlooked by him, I have

* Mémoires du Muséum d’Histoire Naturelle. Tome septiéme. Paris, 1821. Essaisur le vol des
Insectes, par I, Cuaprier, p. 297. Plates x. xi, and xii.

358 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

endeavoured to show are indispensable in flight. In short, he confides to the air
the delicate task of arranging the details of flight—those details constituting in
reality the most difficult part of the problem.

Oljections to the Mechanical Theory of Wing Movements.—There are many |

facts which militate against CHABRIER’S mechanical theory of the movements of
the insect’s wing. I find, for example, that if the wing of the wasp, fly, humble bee,
or butterfly be depressed by a delicate rod, its posterior margin is made to curve
downwards, and to make various angles with the horizon (fig. 18, abede/fg,
page 351) ; the wing, the instant the rod is removed, being flexed and elevated
by the action of elastic ligaments which obliterate the angles formed during the
depression (fig. 19, ghigkim, page 351). This implies the existence of a
muscular system for depressing the wing, and a fibro-elastic system for elevating
it, similar to what is found in the bat and bird, to be described presently. It
also proves that the wing is jointed to the body in such a manner that it cannot
either descend or ascend without changing the direction of its planes—the air
taking no part in the change of plane referred to.

I find, secondly, that insects have the power of vibrating either wing by
itself in any part of a radius not exceeding a half circle, and that the wing may
be played above the body or on a level with or beneath it, as circumstances
demand. These facts argue a much more intimate relation between the muscular
system and the wings than CHABRIER is inclined to admit.

Thirdly, The wing in most insects is composed of two distinct portions at its
root (figure 25, a b, p. 359), those portions being endowed with independent move-
ments, which enable the insect to incline the anterior or thick margin (a ¢ fe) of
the wing in one direction, and the posterior or thin margin in another—to twist,
in fact, the wing upon its long axis. This twisting of the wing upon its long axis

exerts upon the organ precisely the same influence which the extending and ~

flexing of the pinion does upon the wing of the bird and bat (figures 39, 40, 41, 42,
and 43, p. 362). tin short developes double figure of 8 curves along the anterior
and posterior margins, and converts the wing into a screw capable of change of
Sorm.

Fourthly, In the humble bee and other insects supplied with two pairs of —

wings geared to each other by hooklets, the posterior or thin margin of the first
wing glides along the anterior or thick margin of the alula or second wing,
which latter, acting as a long lever, has the power of adjusting the posterior
or thin margin of the first wing.

Fifthly, In the wasp the first wing can be distinctly folded upon itself in the
direction of its length, the alula or second wing folding upon the first wing previ-
ously folded,so that the area of the two wings is reduced to about one-third of what

it was before the folding took place. When the wing is so folded it is very compact, —
and presents a well-defined cutting edge, which points in a backward direction. —

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 3509

I am induced to believe that the wing is folded after this fashion im certain
cases during the back or return stroke, although the action of the pinion is so
rapid that I have hitherto failed to make it out. The folding of the first wing
upon itself in the wasp occurs in the line gs of fig. 25; the folding of the first
wing upon itself and of the second upon the first, being seen at fig. 26 (hd) ;
and the two wings, when folded and ready to make the return stroke, at fig.
27 (ds). The course pursued by the folded wings during the back stroke is
indicated at ghijkim of fig. 19, page 351, Figure 28 represents the wing of

Sie A ;
A c ad
ha aa a 1 eivic
% ‘ = ———_—— ~
t &
oe are
Fig. 27. Fig. 28.

the crane-fly, which has, I believe, a similar action, the thin posterior margin,
Fgh, being folded during the back or return stroke, and opened out during
the forward stroke.

Sixthly, Many insects, such as the ephemera, beetles, locusts, &c., have
assuredly the power of more or less completely crushing their wings together,
and of alternately increasing and diminishing the wing area during the down
and up strokes. The wings of most insects, moreover, are during the up stroke
thrown into rugee, which are flattened or altogether disappear during the down
stroke. They further have the power of arching their wings during the up
stroke, and of opening them out so as to increase their area during the down
one. The butterfly affords an admirable example.

The Down and Up Stroke of the Wing of the Butterfly; Increase and
Diminution of the Wing Area; Development of Figure of 8 Curves on the Margins
of the Wing.—In the butterfly, as I have sufficiently satisfied myself, the first
wing is made to pass above or over the second wing towards the termination of
the down stroke, the convexity of both wings increasing meanwhile. This
reduction in the wing area is necessary to destroy the momentum acquired by
the wings during their descent, and to prepare them for making the up or return
stroke. In the butterfly the wings strike downwards and forwards, and have
amore vertical play than in almost any other insect. The wings are elevated

VOL. XXVI. PART II. DA

360 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

in the overlapped arched condition, and towards the end of the up stroke they
are gradually separated to increase the area and prepare them for making the
down stroke in a manner precisely analogous to what happens in bats and birds.
They are then made to descend in their flattened condition, the first wing passing
over the second towards the termination of the down stroke as just stated.
Nor is this all. While the wings are being depressed and made to overlap
more or less completely, and while they are being elevated and spread out,
double and opposite curves are being developed along their anterior, posterior,
and outer margins. ‘This isa somewhat remarkable circumstance, as the butter-
fly is perhaps the most awkward flying creature that exists. It seems to prove
that the presence of double or figure of 8 curves, is indispensable to flight. These
points are illustrated at figs. 29, 30, 31, 32, 33, and 34. At a, of fig. 29, the

Fig. 32. Fig. 33. Fig. 34.

concavity of the first wing is directed downwards, the concavity of the second
wing being directed slightly upwards as at 6. The two curves taken together
give a double or wave curve. In this figure the two wings are separated or
spread out and ready to give the down stroke. At fig. 30 the two wings are
separated to the utmost, and in the act of making the down stroke. Here the
concavity of both wings is directed downwards as at a, a very small portion of
the second wing only curving upwards (0). At fig. 31 the down stroke is com-
pleted, the first wing overlapping the second, and both being deeply concave on
their under surfaces, as shown at a. They are now in a condition to make the
up stroke, which is the reverse of the down one, and need not be described.
The curves produced along the anterior and posterior margins of the wings of
the butterfly during the up and down strokes are seen at figs. 32, 33, and 34.
At fig. 32, the curves formed along the anterior (c d) and posterior (¢/) margms
of the first wing at the beginning of the down stroke, are represented. At fig.
33 the wing is represented, as seen at the middle of the down stroke, and the

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 361

curves referred to are nearly obliterated (vide 7's, tu). At fig. 34 the wing is
shown at the end of the down stroke, and the curves are reversed, as a com-
parison of cd, ¢f of fig. 32 with gh, 77 of fig. 34 will satisfactorily prove.
In the dragon-fly similar figure of 8 curves are developed along the anterior
and posterior margins of the wings at the beginning, middle, and termination of
the down stroke, as an examination of figs. 35, 36, 37, and 38 will show. If

Fig. 37. Fig. 38.

the letters ed, ¢/ of fig. 35 (dragon-fly) be compared with corresponding letters
of fig. 32 (butterfly) ; the letters 7s, ¢u of fig. 37 (dragon-fly) with similar letters
of fig. 33 (butterfly), and the letters gh, 77 of fig. 36 (dragon-fly) with the same
letters of fig. 34 (butterfly), it will at once be perceived that the curves which
these letters represent are identical in both cases. At fig. 38 the wings are
represented as seen at the beginning and end of the down stroke, the arrows 7, s
giving the range or play of the wings. ‘The letters df of this figure (anterior
Wing at beginning of down stroke) correspond with d/ of fig. 35; the letters gh
ij (anterior wing at end of down stroke) corresponding with similar letters in fig.
36. Fig. 38 shows how the posterior margin of the wing (/) is screwed downwards
and forwards (7) during the down stroke (compare with a, 0, ¢ of figs. 16 and
17, page 349, and read remarks on the dragon-fly’s wing at pages 335 and 350).*
* The wing area in insects is usually greatly in excess of what is absolutely required for flight, as
the following experiments made with the common white and brown butterfly and dragon-fly will show :
1. Removed posterior halves of first pair of wings of white butterfly. Flight perfect.
2. Removed posterior halves of first and second pairs of wings. Flight not strong but still per-

fect. If additional portions of the posterior wings were removed, the insect could still fly, but with
great effort, and came to the ground at no great distance.

3. When the tips (outer sixth) of the first and second pairs of wings were cut away, flight was in
no wise impaired. When more was detached the insect could not fly.

362 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

Curves in all respects analogous to those occurring in the wing of the butterfly
and dragon-fly are observed in the wing of the bat and bird, as a reference to

Fig. 43.

figs. 39, 40, 41, 42, and 43 will satisfy. They are also found in the rowing
feathers of the wing of the bird, as shown at fig. 50, page 379.

4, Removed the posterior wings of the brown butterfly. Flight unimpaired. '

5. Removed in addition a small portion (one-sixth) from the tips of the anterior wings. Flight
still perfect, as the insect flew upwards of ten yards. ; ; ;

6. Removed in addition a portion (one-eighth) of the posterior margins of anterior wings. The
insect flew imperfectly, and came to the ground about a yard from the point where it commenced its
flight.

¥ 7. In the dragon-fly either the first or second pair of wings may be removed without destroying
the power of flight. The insect generally flies most steadily when the posterior pair of wings are
detached, as it can balance better ; but in either case flight is perfect and in no degree laboured,

8. Removed one-third from the posterior margin of the first and second pairs of wings. Flight
in no wise impaired. ‘ ‘

If more than a third of each wing be cut away from the posterior or thin margin, the insect can
still fly, but with effort. : ‘ ,

Experiment 8 shows that the posterior or thin flexible margin of the wing may be dispensed with
in flight. It is more especially engaged in propelling.

9. The extremities or tips of the first and second pair of wings may be detached to the extent of
one third, without diminishing the power of flight.

If the mutilation be carried further, flight is laboured, and in some cases destroyed.

10. When the front edges of the first and second pair of wings are notched, or when they are
removed, flight is completely destroyed.

This shows that a certain degree of stiffness is required for the front edges of the wings, the front
edges indirectly supporting the back edges It is, moreover, on the front edge of the wing that the
pressure falls in flight, and by this edge the major portion of the wing is attached to the body. The
principal movements of the wing are in addition communicated to this edge. : .

Note.—Some of my readers will probably infer from the foregoing experiments, that the figure

of 8 curves formed along the anterior and posterior margins of the pinion are not necessary to flight,

since the tip and posterior margin of the wing may be removed without destroying it. To such I
reply, that the wing is flexible, elastic, and composed of a congeries of curved surfaces, and that so long as
a portion of it remains, it forms, or tends to form, figure of 8 curves in every direction. ;

* Figures 39, 40, 41, 42, and 43 slow the double curves which occur on the anterior (bac) and posterior (def)
margins of the wing of the bat and bird.

‘

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 363

Consideration of the Forces which Propel the Wings of Insects.—Proceeding
now to a consideration of the forces which propel and regulate the wings of
insects, I find that in the thorax of the insect the muscles are arranged in two
principal sets in the form of a cross—.e., there is a powerful vertical set which
runs from above downwards, and a powerful antero-posterior set which runs
from before backwards. There are likewise a few slender muscles which proceed
in a more or less oblique direction. The antero-posterior and verti7al sets of
muscles are quite distinct, as are likewise the oblique muscles. Portions, how-
ever, of the vertical and oblique muscles terminate at the root of the wing in
jelly-looking points which greatly resemble rudimentary tendons, so that I am
inclined to believe that the vertical and oblique muscles exercise a direct
influence on the movements of the wing. The contraction of the antero-pos-
terior set of muscles (indirectly assisted by the oblique ones) elevates the
dorsum of the thorax by causing its anterior extremity to approach its posterior
extremity, and by causing the thorax to bulge out or expand laterally. This
change in the thorax necessitates the descent of the wing. The contraction of
the vertical set (aided by the oblique ones) has a precisely opposite effect, and
necessitates itsascent. While the wing is ascending and descending the oblique
muscles cause it to rotate on its long axis, the bipartite division of the wing
at its root, the spiral configuration of the joint, and the arrangement of the
elastic and other structures which connect the pinion with the body, together
with the resistance it experiences from the air, conferring on it the various
angles which characterise the down and up strokes. The wing may therefore
be said to be depressed by the contraction of the antero-posterior set of
muscles, aided by the oblique muscles, and elevated by the contraction of
the vertical and oblique muscles, aided by the elastic ligaments, and the reac-
tion of the air. If we adopt this view we have a perfect physiological expla-
nation of the phenomenon, as we have a complete circle or cycle of motion,
the antero-posterior set of muscles contracting when the vertical set of muscles
are relaxing, and vice versa,an arrangement which gives an equal period of
activity and repose to both sets. This, I may add, is in conformity with all
other muscular arrangements, where we have what are usually denominated
extensors and fiexors, but which, as I have shown elsewhere,* are simply the two
halves of a circle of muscle and of motion, an arrangement for securing diametri-
cally opposite results in limbs and the condition of activity and rest in muscles.

CHABRIER’s account, which I subjoin, virtually supports this hypothesis :—

“Tt is generally through the intervention of the proper motions of the
dorsum, which are very considerable during flight, that the wings or the elytra
are moved equally and simultaneously. Thus, when it is elevated, it carries

* On the Mechanical Appliances by which Flight is attained in the Animal Kingdom, Trans.
Linn. Society, vol. xxvi. pages 200, 201, and 262.

VOU, XXVI, PART II. »B

364 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

with it the internal side of the base of the wings with which it is articulated,
from which ensues the depression of the external side of the wing ; and when
it approaches the sternal portion of the trunk, the contrary takes place. During
the depression of the wings the dorsum is curved from before backwards, or in
such a manner that its anterior extremity is brought nearer to its posterior, that
its middle is elevated, and its lateral portions removed further from each other.
The reverse takes place in the elevation of the wings; the anterior extremity
of the dorsum being removed to a greater distance from the posterior, its
middle being depressed, and its sides brought nearer to each other. Thus its
bending in one direction produces a diminution of its curve in the direction
normally opposed to it; and by the alternations of this motion, assisted by
other means, the body is alternately compressed and dilated, and the wings
are raised and depressed by turns.”

Objections to Mechanical Theory of Insect WingMovements specified—The
objections to CHABRIER’s mechanical theory of the action of insects’ wings
may be briefly stated :—

First, The movements of the wings of insects are not necessarily absolutely
synchronous. On the contrary, insects have the power of moving their wings
independently.

Second, Insects can twist or plait theirwings at the root—the butterfly having
the power of causing the one wing to overlap the other when required.

Third, Insects can increase the convexity of their wings during the up stroke
and decrease it during the down stroke.

Fourth, They can in some cases fold and diminish the area of the wing
during the up stroke and increase it during the down one.

Fifth, In the dragon-flies we can without difficulty trace the muscles termi-
nating in the roots of the wings—a presumptive proof that in other insects there
is a direct connexion between the muscles of the thorax and the wings they
are destined to move.

Sixth, All insects have the power of elevating their wings when dressing
them, so that the reaction of the air is not necessary to the up stroke, although
it certainly contributes to it in flight. They can, moreover, during the intervals
of rest, develope figure of 8 curves along the anterior and posterior margims
of the pinion independently of the air.

Seventh, There are muscles in the dragon-fly, and I believe in other insects
also, delegated to elevate as well as depress the wing.

Eighth, There are elastic ligaments which recover or flex and partly elevate
the wing when the organ is depressed artificially and not engaged in flight. In

* “General Observations on the Anatomy of the Thorax in Insects, and on its Functions during”
Flight.” By E. T. Beynezrt, F.LS., &c. (Extracted chiefly from the “‘ Essai sur le vol des Insectes,” Pas
J, Catenms Mém. du Muséum @’ heme Naturelle. Zool. Journal, vol. i. art. xlvi. 1825.)

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 365

such cases the air can exert no influence whatever, as the wing is depressed
gently, expressly to avoid recoil.

We have therefore the conditions of flight developed to nearly as great an
extent in the insects as in the bats and birds. That distinct elevator and de-
pressor muscles exist in the bat and bird, and that these act in conjunction with
elastic ligaments there can be no doubt whatever, see pages 395, 396, and 397.

Wings Mobile and Flexible as well as Elastic—Elasticity, Fleaibility, and
Mobility not to be confounded—Mobility and Flexibility necessary to Flight.—
Much importance has been attached by ancient and modern authors to the
elastic properties of the wing, and not a few recent investigators are of opinion
that flight is mainly due to the yielding of the wing to the impact of the air on
its under surface during the down stroke. That, however, the mere elasticity
of the pinion, if regarded apart from its mobility and flexibility, avails little
may be proved in a variety of ways. By mobility I mean that power which
the wing enjoys of moving at its root m an upward, downward, forward,
backward, or oblique direction, and likewise the remarkable property which
it possesses of rotating or twisting in the direction of its length. I also include
under the term mobility the additional power possessed by bats and birds of
opening and closing, 7.¢., of flexing and extending the wings during the up and
down strokes, as well as the power enjoyed by the bat of moving its fingers,
and by the bird of moving its individual primary, secondary, and tertiary
feathers at their roots. By the flexibility of the wing, I mean that power
which the wing possesses of throwing itself into a great variety of curves during
its action—these curves being formed, reversed, or obliterated at the will of the
flying animal. It is necessary to distinguish between mobility, flexibility, and
mere elasticity, because any rotation of the wing along its anterior or thick
margin is at once followed by an elevation or depression of its posterior or thin
margin, which elevation or depression is almost invariably and wrongly attri-
buted to elasticity. That the wing is elastic throughout, and that its posterior
or thin margin yields slightly (to prevent shock) when it attacks the air there
can be no doubt. The yielding, however, is very slight, and it is always accom-
panied by a certain degree of rotation or torsion. If it were otherwise—if
the posterior margin of the wing yielded to any marked extent in an upward
direction when the wing descended, it is evident that the air on which the wing
depended for support would escape from under it, and flight as a consequence be
rendered abortive. It is the air more than the wing which yields or gives way
in flight, and the yielding that occurs in the wings, is to be traced for the most
part, to a rotation of the wing along its anterior margin—to movements occur-
ring in the muscles and ligaments, and in the bones and feathers when present,
particularly at the root of the feathers. These remarks are true of living wings.
It is not, however, to be inferred from what is here stated that natural wings

366 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

may not be successfully imitated, both in their structure and movements, by
mechanical appliances in which elasticity plays a very prominent part. On the
contrary, Iam prepared to show further on, that flight may be regarded as a
purely mechanical problem, and that it admits of a mechanical solution. I am,
however, desirous of showing in the first place what movements are vital,
what vito-mechanical, and what mechanical 7m natural flight. This done, we
will then be in a position to enter upon a consideration of artificial flight. That
elasticity of itself will not produce flight may be inferred from the following
experiments. If, for instance, we lash light unyielding reeds to the anterior
margins of a pigeon’s wings so as to prevent flexion at the elbow-joints, we
instantly destroy flight. In this experiment the e/asticity of the wings, and
particularly of the rowing feathers, is in no wise impaired; in reality the
mobility and flexibility of the wings only are interfered with. A still more
conclusive proof is to be found in the fact that in insects the most elastic
portions of the wings can be altogether removed without destroying the power
of flight. Thus I have cut away as much as two-thirds from the posterior
margin of either wing of the blow-fly, and yet the insect flew with remarkable
buoyancy. I have also removed portions of the tips of the wings with impunity.
I made similar experiments with the dragon-fly, butterfly (pages 361 and 362),
and sparrow, and with nearly uniform results.

Analysis of the Down and Up Strokes in the Wing of the Bird and Bat.—
What was said of the movements of the wing of the insect holds equally true
of those of the bat and bird, if allowance be made for the more vertical direc-
tion of the down and up strokes, and for the fact that the wings of the bat and
bird are in several pieces and jointed.* The joints, like the muscles, extend in
the direction of the length of the wing ; thus, in addition to the shoulder-joint,
we have the elbow, wrist, and finger joints. The insect, bat, and bird have the
shoulder jot in common, and this joint is so constructed that the wing is
free to move in an upward, downward, forward, backward, and oblique direc-
tion. It also admits of a certain amount of rotation or torsion in the direction
of the length of the wing. The joint is in fact universal in its nature. Another
feature possessed in common by insects, bats, and birds, is the elastic igaments
which recover and partly elevate the wing during the up stroke. Those liga-
ments in the bat and bird are not confined to the root of the wing, but extend
along its margins even to its tip.

The presence of those ligaments shows that the wing is not elevated exclu-
sively by the reaction of the air. There are, moreover, distinct elevator muscles
in the wing of the bat and bird. The presence of voluntary muscles, and of
elastic and other ligaments, afford important indications in the construction and

* The beetles have also their wings jointed.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 367

application of artificial wings, and I find that by employing a ball and socket
joint, and a cross system of elastic bands at the root of the wing, I can imitate
the movements of the natural wing with remarkable precision. By adopting
the springs referred to—by making the wing elastic in all its parts, even along
its anterior or thick margin (natural wings are elastic in this situation), and by
applying a power which varies in intensity, I can communicate to an artificial
wing a vibratory motion, completely devoid of pauses or dead points. The
working of the wing in question is accompanied with very little slip. Indeed,
the slip is so little that the wing may be said to supply a persistent buoying
and propelling power. When the wing is made to vibrate briskly in a more or
less vertical direction, it leaps forward in a series of curves, the down stroke run-
ning into the up one and vice versa, to form a continuous upward wave track.
The power applied is greatest at the beginning of the down stroke. It is
decreased at the end of the down stroke, slightly increased at the beginning of
the up stroke, and again decreased towards the termination of that act. Those
changes in the intensity of the driving power are necessary to allow the air
time to react on the under surface of the wing, and to bring the elastic pro-
perties of the springs and of the wing into play. The springs should be arranged
at right angles and obliquely, that is, there should be a superior, inferior, ante-
rior, and posterior set running at right angles to each other, and between these
as many oblique springs as are deemed necessary. The springs ought to vary
as regards their length and their strength. Thus, the superior springs, which
assist in elevating the wing, ought to be longer and stronger than the inferior
ones ; and the posterior springs, which restrain the wing from leaping forwards
during its vibrations, should be longer and stronger than the anterior ones, the
wing having no tendency to travel backwards. A detailed account of the structure
and movements of artificial wings will be found at the end of the present memoir.
In the bat and bird the wing is extended or pushed away from the body prior
to the down stroke, and folded or drawn towards the body prior to the up stroke.
The unfolding or extending of the wing prior to the down stroke, as seen in
the gull, is shown at Plate XI. figures 3, 2, 1,5; Plate XIV. figure 18.
When the wing is being extended or opened out it is also being elevated,
as shown at 1, 2,3 of Plate XI. figure 5, and Plate XIV. figure 18. When
the wing is flexed, as at ¢ p of figure 3, Plate XI., the under surface of the wing
(s g) is nearly on a level with the horizon (6 d). When, however, the wing
is partially extended, as at Plate XI. figure 2, the angle which its under surface
makes with the horizon is considerable, c b d representing the angle, and b d
the horizon. When the wing is fully extended, and ready to give the down
stroke, the angle which the under surface of the wing makes with the horizon
is still more increased, as shown at Plate XI. figure 1, c 6 d indicating the
angle, and 6 d the horizon. The angle made by the under surface of the root
VOL. XXVI. PART II. 5 C

368 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

of the wing with the horizon considerably exceeds that made by the tip, and
is much greater than a casual observer would be inclined to admit. It
is obscured by the curving downwards and forwards of the anterior mar-
gin of the wing towards the root, as seen at a@ of figure 7, Plate XII. In
this figure the apparent angle made by the root of the wing with the horizon
(ef) is a b d, the real angle being ¢ 6 d. The wing of the bird rotates in
opposite directions during extension and flexion. The various angles of
inclination made by the wing of the gannet in extension and flexion is well
shown at Plate XIII. figures 16 and 17.

In figure 17 (flexion) the posterior margin of the wing (s q p 0) is ona
level with the body of the bird ; whereas in figure 16 (extension) the posterior
margin (7 p 0) is directed downwards and forwards, as indicated by the arrows.
The same thing is seen in the pea-wit, at Plate XII. figure 8. In this figure
the wing to the right of the observer is flexed, and in the act of making the
up stroke, the anterior margin of the pinion being slightly directed down-
wards (vide arrow). The wing to the left of the observer is, on the contrary,
extended, and in the act of making the down stroke, the anterior margin of
the pinion being directed upwards (cide arrow).

The rotation of the posterior margin around the anterior as an axis during ex-
tension, is occasioned by the points of insertion of the pectoralis major and other
muscles, by the attachments and directions of the elastic and other ligaments,
and by the spiral nature of the articular surfaces of the bones of the wing—
the mere act of extension on all occasions involving the rotation in question.

The Wing of the Bird Descends as a Long Lever.—Let us imagine the wing
fully extended and elevated, and making a certain angle with the horizon, as
indicated at ¢ bd of figure 1, Plate XI, at 3 of figure 5, Plate XL. and at 3’ of
figure 18, Plate XIV. The wing is now prepared to make the down stroke, and
descends in a spiral swoop, successively assuming the position 4 in figure 19,
Plate XIV., and 4in figure 6, Plate XI. It acts with extreme energy as a long
lever (vide c d of figure 6, Plate XI.), the purchase which it has on the body
being much greater than is usually anticipated.

During its descent the angle which the wing makes with the horizon is
increased, as shown at abc of figures 16 and 17 (page 349), the horizon in these
figures being indicated by the straight line 2 2’.

In the bird, therefore, as in the insect, the posterior or thin flexible margin
of the wing is screwed down upon the air while the wing is descending.

The Rotation of the Posterior Margin of the Wing in a Downward Direction
increases the Elevating, but diminishes the Propelling Power of the Wing.—The
additional hold which the bird can cause its wing to take of the air by resorting
to a greater or less degree of rotation, is truly surprising. If the wing is
depressed minus the rotation, it darts forward, but takes no very decided catch

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 369

of the air. As a consequence, the recoil is feeble. If, however, the rotation is
added, the wing seizes the air with such avidity as in all cases to produce a very
violent reaction. The tendency of the wing to dart forward is diminished by the
rotation, but the actual elevating power of the pinion is greatly augmented.
This point can be readily ascertained by depressing and screwing, in the manner
described, the wing of the swan or of any other large bird, previously dried, in
the extended position. In preparing the wing for the experiment care should
be taken not to destroy the curves peculiar to the natural extended wing. I
mention this fact because, of many swans’ wings prepared by me for this purpose,
I found one had been inadvertently flattened, and gave quite an indifferent result.

The Importance to be attached to the Concavo-Convex Form of the Wing in
Birds.—The downward screwing of the concave or under surface of the wing,
which is so efficacious in securing a powerful hold of the air during the down
stroke, is followed during the up stroke by an upward screwing of the convex or
upper surface, which is not less effective in evading the air. In fact, when the
wing ascends it is drawn towards the body, and deeply arched, so that it is
literally made to roll upwards, its convex or dorsal surface being directed
upwards throughout the entire up stroke. It is thus the wing evades the super-
incumbent air during the return stroke. This account will be readily under-
stood by a reference to figures 13, 14, and 15, Plate XIII.

At figure 15, Plate XIII., the wing is represented as seen in the middle of the
down stroke. It is widely spread out, and finely arched. At figure 14, Plate
XIIL., the wing is shown as observed towards the end of the down stroke—the
wing being partly flexed or drawn towards the body, and the arch rendered more
abrupt, particularly towards the root of the pinion. . At figure 13, Plate XIII,
the wing is seen quite at the termination of the down stroke. It is fully flexed,
and drawn still closer to the body. It is, moreover, more deeply arched than in
either of the other figures. It has, in fact, assumed the shape which offers
least resistance in an upward direction, and is prepared to make the up stroke.

The Under or Concave Biting Surface of the Wing of the Bird effective both
during the Down and Up Strokes.—If, mstead of believing that the wing is
elevated, we believe what, as I have already stated is actually the case, viz., that
the body of the bird falls downwards and forwards, we at once transfer the
resistance from the dorsal or convex non-biting surface of the wing to the ven-
tral concave or biting surface—the body being supported while the wings are
being elevated by a beautifully arched natural parachute formed by the wings.
The elevation of the wings is, in short, in a great measure a consequence of
the falling of the body. It is in this way that the air comes to assist in elevating
the wings. The air, in short, caught under the wings is instrumental in elevat-
ing and extending them in proportion as the body falls (vide figures 13, 14,
and 15, Plate XIII.) The small size of the elevator muscles of the wing of the

070 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

bird and bat, as compared with the very powerful depressor muscles, is thus
accounted for. The elevation of the wing, as will be inferred, is to a certain
extent a mechanical act, and is due to the reaction of the air, the contraction
of the elastic ligaments, and the downward and forward fall of the body. It
is, however, not altogether mechanical, the wing, as I shall show subsequently,
being perfectly under control both during the down and up strokes.

Lax Condition of the Shoulder Joint in Birds, &c.—The great laxity of the
shoulder joint readily admits of the body falling downwards and forwards during
the up stroke. This joint, as has been already stated, admits of movement in
every direction, so that the body of the bird is like a compass set upon gimbals,
2.é., 1t swings and oscillates, and is equally balanced, whatever the position of
the wings. The movements of the shoulder joint in the bird, bat, and insect,
are restrained within certain limits by a system of check ligaments and pro-
minences; but in each case the range of motion is very great, the wing being
permitted to swing forwards, backwards, upwards, downwards, or at any degree
of obliquity. It is also permitted to rotate along its anterior margin, or to
twist in the direction of its length to the extent of nearly a quarter of a turn.
This great freedom of movement at the shoulder joint enables the insect, bat,
and bird, to rotate and balance upon two centres—the one running in the
direction of the length of the body, the other at right angles, or in the direc-
tion of the length of the wings.

The Wings Elevated Indirectly by the Operation of Gravity.—I have explained
that during the up stroke the body falls, and the wings are elevated. Let us now,
for the sake of argument, advocate an opposite view. Let us take for granted
that the body is fixed in space, and that the wings are elevated by a purely
vital act. From this it follows that the wings during their ascent will of necessity
experience much resistance from the superimposed air, the rounded form of the
upper or dorsal surfaces of the pinions diminishing, but not removing the evil.

The resistance experienced by the wings during their ascent is obviated in the
simplest manner possible, the movement, as has been explained, being dex-
terously transferred from the wings to the trunk in such a manner that the under
or concave surfaces of the wings are made to act in lieu of the upper or convex
surfaces. The body, in a word, is dragged downwards by the inexorable power of
gravity ; but the descent of the bodyinvolves the ascent of thewings. The bodyand
wings, therefore, reciprocate, the body being elevated by the descent of the wings
in conjunction with other means, while the wings are elevated to a great extent
by the descent of the body, as shown at figures 20, 21, and 22, pages 352 and 353.*
The wings are also partly elevated by the reaction elicited from the air—the
contraction of the elevator muscles and elastic ligaments and the forward travel —

* The alternate ascent and-descent of the wings and body during the down and up strokes are
well seen in the butterfly and in all animals whose wings are large for their bodies.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. ofl
of the body. The space through which the body descends when the wings
ascend is very trifling, from the fact that the body is situated at the roots of
the wings—a very slight movement at the roots of the pinions necessitating an
extensive movement at the tips. This explains the very small waved track made
by the body in progressive flight as compared with that made by the wings.
(Contrast 1, 2, 3, 4, 5 of figure 14 page 344, with ac egi of the same figure. )

The Wings of the Bird form a Natural Parachute from which the Body
Depends both during the Down and Up Strokes.—The falling downwards of the
body, and the gradual expansion and elevation of the wings during the up
stroke, is seen at Plate XIII. figures 13, 14, and 15. At figure 13 the wings
and the body are in the position peculiar to them at the end of the down stroke,
1.¢., the body is elevated and the wings depressed. The up stroke is com-
menced, and the body falls, while the wings are somewhat expanded and
elevated, as at Plate XIII. figure 14. The body falls still more, and the wings
are further elevated and expanded, as seen at Plate XIII. figure 15. The
wings are now on a level with the body of the bird, and mark how beautifully
the latter is buoyed up. The body is attached to, and suspended from, a wide-
spread finely arched parachute. The body goes on falling, and the wings rising,
till the body is depressed and the wings elevated, as seen at 2, 2’ and 3, 3’ of
figure 18, Plate XIV. This terminates the up stroke, and it will be observed
that the position of the body is just the reverse of what it was at the beginning
of the up stroke. At the beginning of the up stroke, the body was highest and
the wings lowest (vide figure 13, Plate XIII.) At the end of the up stroke,
the body is lowest and the wings highest (vide 3, 3’ of figure 18, Plate XIV.)
That the body is supported and carried forward during the up stroke of the wings
is proved beyond doubt by the experiment described at pages 355, 356, and
illustrated by figure 23. Ifthe quill feathers «, 4, of figure 23 (p. 356) be compared
with the two wings 3, 3’ of figure 18, Plate XIV., and the cork ¢ of figure 23
with the body of the bird in figure 18, Plate XIV., it will be found that the con-
ditions are the same in both, and that both are to a great extent sustained and
carried forward in space, the one by the overarching feathers and the other by
the overarching wings.* Perhaps the simplest illustration that can be given of

* Weight necessary to Flying Animals as at present constructed—Weight and Levity relatively
tonsidered with regard to Aérial and Subaquatie Flight (Diving).—Captain W. F. Hurron, in a recent
pamphlet (On the Sailing Flight of the Albatros, Phil. Mag., August 1869), contends, that whereas a
bird lighter than the water can fly in it, so, in like manner, a bird lighter than the air could fly in this
medium, and that therefore weight is not necessary to aérial flight. Captain Hurron, however, forgets
that a bird destined to fly above the water is provided with travelling surfaces so fashioned and so
applied (they strike from above downwards and forwards), that if it was lighter than the air, they would
carry it off into space without the possibility of a return ; in other words, the action of the wings would
carry the bird obliquely upwards, and render it quite incapable of flying either in a horizontal or down-
ward direction. In the same way a bird destined to fly wnder the water (auk and penguin), if it was

not lighter than the water, such is the configuration and mode of applying its travelling surfaces
(they strike from below upwards and backwards), they would carry it in the direction of the bottom

VOL. XXVI. PART Il. 5D

372 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

the mutual action and reaction of the body and wings during the up stroke is
that furnished by a partly opened umbrella, whose handle has been intention-
ally weighted. If the umbrella thus prepared be dropped from a height, the

without any chance of return to the surface. In aérial flight, weight is the power which nature has
placed at the disposal of the bird for regulating its altitude and horizontal flight, a cessation of the play
of its wings, aided by the inertia of its trunk, enabling the bird to approach the earth. In subaquatiec
flight, levity is a power furnished for a similar but opposite purpose ; this, combined with the partial
slowing or stopping of the wings and feet, enabling the diving bird to regain the surface at any
moment. Levity and weight are auxiliary forces, but they are necessary forces when the habits of the
animals, and the form and mode of applying their travelling surfaces are taken into account. If the
aérial flying bird was lighter than the air, its wings would requite to be twisted round to resemble the
diving wings of the penguin and auk. If, on the other hand, the diving bird (penguin or auk) was
heavier than water, its wings would require to resemble aérial wings, and they would require to strike
in an opposite direction to that in which they strike normally. From this it follows that weight is
necessary to the bird (as at present constructed) destined to navigate the air, and Jevity to that destined
to navigate the water. If a bird was made very large and very light, it is obvious that the diving
force at its disposal would be inadequate to submerge it. If, again, it was made very small and very heavy,
it is equally plain that it could not fly. Nature, however, has struck the just balance ; she has made
the diving bird, which flies under the water, relatively much heavier than the bird which flies in the
air, and has curtailed the travelling surfaces of the former, while she has increased those of the latter.
For the same reason, she has furnished the diving bird with a certain degree of buoyancy, and the
flying bird with a certain amount of weight—levity tending to bring the one to the surface of the
water, weight the other to the surface of the earth, which is the normal position of rest for both. The
action of the subaquatic or diving wing of the king penguin is well seen in the annexed woodeut
(Fig. 44).

Fig. 44.

At A, the penguin is in the act of diving, and it will be observed that the anterior or thick margin of
the wing is directed downwards and forwards, while the posterior margin is directed upwards and back-
wards. This has the effect of directing the under or ventral concave surface of the wing upwards and
backwards, the effective stroke being delivered in this direction. The efficacy of the wing in counter-
acting levity is thus obvious. At B, the penguin is in the act of regaining the surface of the water,
and in this case the wing is maintained in one position, or made to strike downwards and forwards like —
the aérial wing, the margins and under surface of the pinion being reversed for this purpose. ‘The
object now is not to depress but to elevate the body. Those movements are facilitated by the alter-

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 373

falling of the weighted handle will have the effect, in conjunction with the
resistance which the under concave surface of the umbrella experiences from
the air, of opening it up, precisely as the wings are opened up and elevated at
figures 13, 14, and 15, Plate XIII. And if it so happens that the steels of the
umbrella are feeble, and the weight attached to the handle sufficiently great,
the umbrella will be more or less everted, as shown at 2, 2’ and 3, 3’ of figure
18, Plate XIV. If the frame of the umbrella was endowed with vitality, and
had the ‘power of quickly regaining its original form, it would elevate the
weighted handle, and so attain its original position. A repetition of those
changes, if the proper degree and kind of power were added, would result in
flight, particularly if one side of the umbrella was rendered more rigid than the
other, as this would have the effect of conferring an eccentric action upon it.
The parachute principle here advocated is corroborated to a certain extent by the
flight of the beetles. In these, in some cases, the e/yira or wing cases are deeply
concavo-convex. The membranes or true wings strike in a downward, forward,
and more or less horizontal direction, and in so doing they force the air forward
under the ventral or concave surfaces of the elytra or false wings, which are thus
converted into parachutes or tiny sustaining balloons. That the elytra perform
a very important function in flight is proved by the fact that when they are
removed the insect cannot fly. I had ocular demonstration of this at Somerton,
Wexford, in the summer of 1868. When I amputated the elytra close to the
roots, the insects could not rise, although they made frequent attempts to do
so. The elytra or false wings and the membranous or true wings form, when
extended, deeply concave or umbrella shaped surfaces, the peculiarity in such
instances being that the umbrellas formed by the true wings move and are
active ; whereas those formed by the elytra are fixed or immobile, and conse-
quently passive.

The Wing of the Bird elevated as a Short Lever.—tIn birds with short rounded
Wings, and in others with longer wings, in forced flight the wing is usually
elevated as a short lever, as shown at 6 of figures 6 and 19, Plates XI. and
XIV., and 1 of figures 5 and 18, Plates XI. and XIV. ; it being extended or
spread out quite towards the end of the up stroke, as represented at.1, 2, 3 of
figures 5 and 18, Plates XI. and XIV. In birds with long pointed wings, when
flying leisurely, the wing is not unfrequently expanded at the middle of the up

nate play of the feet. What strikes one in the present woodcut is the comparatively small size of the
diving or swimming wing, which resembles the flipper of the turtle, seal, sea bear, and walrus. At
Plate XIII. figure 15, the aérial wing, as seen in the gull, is represented, and the large size of the
flying pinion, as compared with the diving subaquatic one, is at once apparent. Here the anterior
margin (x s t v w) of the wing is directed upwards and forwards, the posterior one (0 p g) downwards
and backwards. This causes the under or ventral concave surface of the pinion to look downwards and
forwards, the direction in which the effective or down stroke is delivered. The aérial wing, like the
subaquatic wing, is twisted upon itself. It strikes downwards and forwards, because this is the direc-
tion in which a body in motion would naturally fall.

374 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

stroke, as seen at 4 of figure 19, Plate XIV., and at figure 15, Plate XIIL ;
and it is no doubt this circumstance which has induced hasty generalisers to
deny that the wing is flexed during the up stroke. This is a pardonable mis-
take, as the wing in such cases may be actually extended for two-thirds of the
up stroke. When the wing is fully flexed and elevated as a short lever, its
rowing feathers are separated and opened up, and the bird draws largely upon
its vital resources. When, on the other hand, the wing is elevated as a long
lever, and is wielded in one piece, after the manner of the insect wing, the bird
takes advantage, to a great extent, of the numerous mechanical adaptations ©
with which nature has endowed it. The flight of the albatros furnishes the
best example. The opening up of the feathers during the up stroke facilitates
the ascent of the pinion, and permits a more rapid action. The separation of
the feathers is, however, not necessary to successful flight, the bat flying
remarkably well by the aid of a continuous membrane which, as is well known,
is destitute of feathers.

The Wing Vibrates Unequally on either Side of a given Line.—The wing, during
its vibration, descends further below the body than it rises above it. This is
necessary for elevating purposes. In like manner the posterior margin of the wing
(whatever the position of the organ) descends further below a given line than
it ascends above it. This is requisite for elevating and propelling purposes, the
under surface of the wing being always presented at a certain upward angle to
the horizon, and acting as a true kite. This view is fully explained at p. 345.
If the wing oscillated equally above and beneath the body, and if the posterior
margin of the wing vibrated equally above and below the line formed by the
anterior margin, much of its elevating and propelling power would be sacrificed.
The tail of the fish oscillates on either side of a given line, but it is otherwise
with the wing of a flymg animal. The fish is of nearly the same specific
gravity as the water, so that the tail, as a rule, only propels. The flying
animal, on the other hand, is very much heavier than the air, so that the wing
requires both to propel and elevate. The wing to be effective as an elevating
organ must consequently be vibrated rather below than above the centre of
gravity ; at all events, the intensity of the vibration should occur rather below
that point. In making this statement, it is necessary to bear in mind that the
centre of gravity is ever varying, the body rising and falling im a series of curves
as the wings ascend and descend.

To elevate and propel, the posterior margin of the wing must rotate round
the anterior one, the posterior margin being, as a rule, always on a lower level
than the anterior one (vide pages 414, 415, and 416). By the oblique and more ~
vigorous play of the wings wnder rather than above the body, each wing expends
its entire energy in pushing the body upwards and forwards. Fig. 12, page 342,
will illustrate my meaning. Let the oar x, s, represent the wing. If the wing

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 375

be made to play equally above (a b) and below (cd) the body, the tendency is to
drive the body in an undulating line, away from a, in the direction s 2. As,
however, the opposite wing tends to push the body im a precisely contrary
direction, the forces exercised by the two wings neutralise each other in the mesial
line of the bird, the force which ultimately prevails being that of gravity. To
destroy the power of gravity, and to elevate and propel the bird, it is necessary
that the wings descend further than they ascend, and that the posterior mar-
gins of the wings be constantly kept on a lower level than the anterior ones.
It is also necessary that the wings be convex on their upper surfaces, and con-
cave on their under ones, and that the concave or biting surfaces be brought
more violently in contact with the air during the down stroke, than the con-
vex ones during the up stroke. The greater range of the wing below than
above the body, and of the posterior margin below than above a given line, may
be readily made out by watching the flight of the larger birds. It is also well
seen in the upward flight of the lark. The range of the wing of the gull in
ordinary flight is shown at Plate XIV. fig. 19. When the wing is elevated high
above the body, as represented at 3 of figures 5 and 18, Plates XI. and XIV.,
it is generally in the effort of rising, or in picking up garbage from the surface
of the sea, or in suspending or letting the body down gradually prior to alight-
ing. In such cases the wings expend their greatest force when a little above or
on a level with the body, as is well exemplified in the hovering of the kestrel.

Compound Rotation of the Wing of the Bird.—To work the tip and posterior
margin of the wing independently and yet simultaneously, two axes are neces-
sary, one axis (the short axis) corresponding to the root of the wing ; the second
(the long axis) to the anterior margin. This renders the wing eccentric in its
nature. The primary or rowing feathers are also eccentric, the shaft of each
feather being placed nearer the anterior than the posterior margin, an arrange-
ment which enables the feathers to open up and separate during the up stroke,
and approximate and close during the down one. ‘The axes of rotation in the
wing of the bird are given at figure 19, Plate XIV., a@ representing the short
axis around which the tip of the wing rotates with a radius ¢ bf; ¢, the long axis,
around which the posterior margin of the wing revolves with a radius g d h.

_ These points are more fully illustrated at figure 45, p. 376, where a b repre-
sents the short axis (root of wing), with a radius ¢/; ¢ d, the long axis (anterior
| Margin of wing), with a radius g p.

The Wing of the Bird cranked slightly Forwards—the Compound Rotation
ofthe Rowing Feathers.—It will be observed from figure 45 (p. 376), that the wing
is cranked somewhat forwards (compare position of axis a } with that of axis
ed), avery slight movement of rotation along the anterior margin (cd) being
accompanied by a considerable rotation of the posterior margin (hijh/). This
figure also shows that the individual primary, secondary, and tertiary feathers

VOL. XXVI. PART II. DE

376 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

of the bird’s wing have each what is equivalent to a long and a short axis. Thus
the primary and secondary feathers marked 477, k/are capable of rotating on
their long axes (7 s) and upon their short axes (m 7). The feathers rotate upon

Fig. 45.

their long axes in a direction from below upwards during the down stroke, to
make the wing impervious to air; and from above downwards during the up
stroke, to enable the air to pass through it. The primary, secondary, and
tertiary feathers have thus a distinctly valvular action.* They rotate upon their
short axes (mm) during the descent and ascent of the wing, the tips of the feathers
rising slightly during the descent of the pinion and falling during its ascent.

The Primary, Secondary, and Tertiary Feathers are Geared to each other,
and Actin Concert.—To admit of the primary, secondary, and tertiary feathers
rotating upon their long axes (7 s), a very elaborate combination of fibrous and
elastic structures, with a certain admixture of muscular substance, is necessary.
The arrangement, as witnessed in the crested crane, is given at Plate XVI.
figures 24, 25, 26, 27, and 28.

The roots of the primary, secondary, and tertiary feathers are imbedded
behind the muscular mass (f, 7, 2), fig. 24, Plate XVI. The insertions of the
roots of the feathers are shown in figure 28, Plate XVI. Each root is enveloped
by a continuous elastic ligament (0 p q of fig. 24), this ligament being provided
with fibrous bands, which run in the direction of the length of the wing (7 s, ¢ u,
vw of figs. 25 and 27, Plate XVI.) and obliquely (g 2, gh). Two oblique bands (g
and h) run between every two feathers (~), and are joined to the longitudinal ones
(rs tu vw), and to the feathers in such a manner that the whole are geared
together, an arrangement combining great freedom of movement with great
strength. The longitudinal bands run along the roots of all the feathers, and

* The valve action, as explained, is called more or less into play according to circumstances.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. BYE

are three in number, the outermost band breaking up at the root of each
feather, and giving off two processes (a d, b e, c f of figure 25, Plate XVI.), the
one of which coils round the root of the feather in a spiral manner from right
to left ; the other coiling in an opposite direction, or from left to right (m, » of
figure 26, Plate XVI.) The root of each feather is consequently enveloped by a
fibrous investment, capable of rotating it in opposite directions. The fibrous
bands referred to are arranged with much precision, and as they are geared to
each other at stated intervals, they cause the feathers (right wing) to rotate
at nearly the same instant from right to left, and from below upwards, during
extension ; and from left to right, and from above downwards, during flexion.
The arrangement of the fibrous bands is much the same on the dorsal and
ventral aspects of the wing (compare figs. 24 and 28). It varies slightly in dif-
ferent species of birds, but the function of the bands is the same in all.

The tips of the primary, secondary, and tertiary feathers are prevented from
rising too high during the descent of the wing by the oblique overlapping of the
feathers forming the primary, secondary, and tertiary coverts (m, n, 0 of figure
28, Plate XVI.), those feathers acting as buffers and limiting the action.

The Up or Return Stroke of the Wing of the Bird—Diminution of Area of
Wing— Valvular Action, &e.—Towards the termination of the down stroke, the
wing is suddenly flexed and drawn towards the body, as shown at 4, 5, 6 of
figures 6 and 19, Plates XI. and XIV. This is necessary to convert the wing
from a long (Plate XI. figure 6, c d) into a short lever (Plate XI. figure 6, a 6),
and to destroy the momentum acquired by the wing during its more or less
vertical descent. While the wing is being shortened, the angles which the
several portions of its under surface make with the horizon are being diminished
(ed ef of figures 16 and 17, page 349); the angles made by the under surfaces
of the rowing feathers from within outwards being increased (123456789
of fig. 46, p. 378). These changes prepare the wing of the bird for making
an effective up or return stroke, and are necessitated by the more vertical
play of the bird’s wing, as compared with that of the msect. But for the
diminution of the actual area of the wing during the up stroke, the upper or
dorsal surface of the pinion would experience much resistance from the
air during its ascent. This difficulty is m a great measure obviated by the
wing being drawn close to the side of the body, and by its being made to
assume a somewhat crippled appearance, the tip of the wing folding upon
the root in a direction from below upwards, and in such a manner as to displace
comparatively little air (vide 4, 5, 6 of figure 6, Plate XI.) The pinion is
then, as a rule, elevated as a short lever (a 6 of figure 6, Plate XI.), until it
attains the position indicated at 1 of figures 5 and 18, Plates XI. and XIV.
In these situations the wing is for the most part deeply arched (vide figure 13,
Plate XIII.) When the wing has assumed the position indicated by 1 of

378 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

figures 5 and 18, Plates XI. and XIV., it is suddenly pushed away from the
body, extended and elevated, as shown at 2 and 3 of the same figures ; the angles
made by the several portions of its under surface with the horizon being in-
creased, while those formed by the under surfaces of the rowing feathers are
decreased (1234567 8 9 of fig. 47). The wing thus comes to form a kind
of natural parachute, as shown at 2, 2’ and 3, 3’ of figure 18, Plate XIV.
This completes the up or return stroke. While the wing is ascending, the
primary, secondary, and tertiary feathers rotate upon their long axes, and
present their thin margins to the air, into which they cut like so many knives.
The feathers are most widely separated at the beginning of the up stroke, and
least at the termination of that act, as they then flap together to make the
wing impervious, and prepare it for making the down stroke. The individual
primary, secondary, and tertiary feathers are so arranged and so rotated that they
open up, and close, and present the precise angles required for flight, whatever
the shape and whatever the position of the wing.

Figure 46 shows the tips of the primary (7) and secondary (s) feathers in the
wing of the piet during flexion, and it will be observed that the angles made

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 379

by the rowing feathers with the horizon (see straight dotted line) in a direction
from within outwards is greater than those made by the same feathers (7),
in extension, as represented at figure 47. In figure 46 the wing is folded upon
itself at x, and presents two arches, a larger (7) and a smaller (s), the numbers
1234567 8 9 giving the position of the primary feathers when counted from
without inwards, the arrows indicating the direction in which the primary and
secondary feathers open up and cut into the air from below upwards and from
within outwards during the up stroke. This figure shows that the primary and
secondary feathers (particularly the former), when viewed from the tip, or when
cut across, present a spiral contour (¢ g of figure 49). This arises from the
primary and secondary feathers being twisted upon themselves, as represented
at a 6b, c d of figure 50,

‘EE

77 i “
eo Ze
I b

Fig. 49. Primary Feather, showing double curves at anterior margin (c d), posterior margin (a b), and acrcss (c,).

Fig. 50. The same, seen from before edgeways. Bee Paes (c d) and posterior (@ 6) margins cross

Figure 47 shows that the primary and secondary feathers of the wing of
the piet are thrown into a beautiful groined arch in extension, preparatory
to the down stroke, the advantage in favour of a concave surface over a
convex one for seizing air or water being something like 2 to 1. It also
shows that the primary (7) and secondary (s) feathers in extension, and during
the down stroke, rotate upon their long axes in a direction from below
upwards, as indicated by the arrows abedefghijkimnopgq, so as to form
an arch which cannot be destroyed so long as the individual feathers remain
intact. In fact, the integral parts of the arch are so disposed that the greater
the pressure the greater the strength. Figure 48 shows a similar groined
arch formed by the roots of the primary and secondary feathers, the spirals
constituting the arch (abcdefghijkimnopq) running in an opposite direc-
tion to those seen in figure 47, from the fact of the primary and secondary
feathers being twisted upon themselves as already explained (compare 6 ¢ with
a d of figure 50). Fig. 20, Plate XIV. shows how the air is forced during
the down stroke in a spiral direction (vide arrows) from without inwards,

VOL. XXVI. PART II. 5D F

380 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

from before backwards, and from above downwards, the spiral currents
from the two wings impinging upon the sides of the bird, which is wedge
shaped, in such a manner as to force it upwards and forwards. The currents
also cross and neutralise each other below the body of the bird, and thus supply
additional buoyancy and propelling power. The arch made by the wing of the
gull, when fully extended and ready to give the down stroke, is seen at 3, 3’ of
figure 18, Plate XIV.; that made at the middle of the down stroke at figure
15, Plate XIII. ; and that made at the end of the down stroke at figure 13,
Plate XIII. The arch made by the wing of the gannet in extreme extension is
shown at figure 16, Plate XIII.

The Primary, Secondary, and Tertiary Feathers Imbricate or Overlap.—
Another point of interest in the bird’s wing is the manner in which the various
feathers (primary, secondary, and tertiary) overlap (fig. 20, Plate XIV.), and
the varying degrees of strength which they exhibit. Proceeding from the tip of
the wing towards the root we find as a rule that the first three primary feathers
are longer and stronger and overlap more than the second three—the second
three being longer, stronger, and overlapping more than the third three.
These points are well seen in the acuminate scythe-lke wing, of which that
of the gull (fig. 15, Plate XIII.) and gannet (fig. 16, Plate XIII.) are good
examples.* Similar remarks may be made of the secondary and tertiary feathers,
as areference to p q of fig. 16, Plate XIII., will show. Another not less inte-
resting feature is the varying position of the vanes of the primary, secondary, and
tertiary feathers. Thus, in the first primary the vane (¢ / of fig. 49, page 379),
is placed quite on the anterior margin (¢ d) the posterior margin (a 0) being three
or four times broader than the anterior one to admit of overlapping. The vane of
the feather occupies a more and more central position as we proceed from the tip
in the direction of the root of the wing, as shown at hi7k/ of fig. 45, page 376, and
also at 1, 2, 3, 4, 5,6, 7, 8,9, 7k im n, &c., of fig. 20, Plate XIV. The first
primary, as will be seen from this account, is eccentric in its nature. Itis more
eccentric than the second—the second bemg more eccentric than the third,
and so of all the primary and secondary feathers, until the stem of the feather
is found to occupy its centre. The posterior margin of the first primary, as a
consequence, rotates more than that of the second—the second than the third,
and so of the others—the valvular action of the wing being most marked at the
tip of the pinion, and gradually diminishing in the direction of the root. The
rowing feathers are necessarily eccentric. If the axis of each feather was not
placed nearer the anterior than the posterior margin the anterior margin would
rise as much as the posterior margin is depressed. This, however, is prevented

* In some cases, as for instance in the more rounded form of wing shown in fig. 20, Plate
XIV., the 4th, 5th, and 6th primaries are longer and stronger, and overlap more than the Ist, 2d, and
3d,

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 381

by the axis of the feather occupying an anterior position, the feather when it is

made to rotate causing the posterior margin (because of its greater breadth) to
move through a greater arc of a circle than the anterior margin. It is owing to
the greater travel of the posterior margin as compared with the anterior one, that
the feathers of the wing so readily open in flexion and close in extension. The
gradation in the length, and strength, and in the degree of overlapping is neces-
sitated by the fact that the feathers at the tip of the wing are exposed to a
much greater strain than those nearer the root—the former always travelling
through a much greater space in a given time than the latter.

The Wing of the Bird not always Opened Up to the same extent in the Up Stroke.
—The elaborate arrangements and adaptations just referred to for increasing
the area of the wing, and making it impervious to air during the down stroke,
and for decreasing the area and opening up the wing during the up stroke,
although necessary to the flight of the heavy-bodied, short-winged birds, as the
grouse, partridge, and pheasant, are by no means indispensable to the flight of
the long-winged oceanic birds, unless when in the act of rising from a level
surface ; neither do the short-winged heavy birds require to fold and open up
the wing during the up stroke to the same extent in all cases, less folding and
opening up being required when the birds fly against a breeze, and when they
have got fairly under weigh. All the oceanic birds, even the albatros, require to
fold and flap their wings vigorously when they rise from the surface of the
water. When, however, they have acquired a certain degree of momentum,
and are travelling at a tolerable horizontal speed, they can in a great measure
dispense with the opening up of the wing during the up stroke—nay, more, they
can in many instances dispense even with flapping. ‘This is particularly the case
with the albatros, which (if a tolerably stiff breeze be blowing) can sail about
for an hour at a time without once flapping its wings. In this case the wing is
wielded in one piece like the insect wing, the bird simply screwing and un-
screwing the pinion on and off the wind, and exercising a restraining influence
—the breeze doing the principal part of the work. In the bat the wing is
jointed as in the bird, and folded during the up stroke. As, however, the bat’s
wing, as has been already stated, is covered by a continuous and more or less
elastic membrane, it follows that it cannot be opened up to admit of the air
passing through it during the up stroke. Flight in the bat is therefore secured
by alternately diminishing and increasing the area of the wing during the up
and down strokes—the wing rotating upon its root and along its anterior
margin during its ascent and descent precisely as in the bird.

_ Analysis of the Movements of Extension and Flexion in the Wing of the Gannet.
—The changes which the wing undergoes in extension and flexion are seen to
great advantage in the gannet (figs. 9, 10, and 11, Pate XII.)

The pinion of this bird is remarkable for its great length as compared with

382 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

its breadth, and for the general elegance of its shape (vde figs. 11 and 16, Plates
XII. and XIII.) It is especially interesting from the fact that the wing
movements can be more readily and satisfactorily analysed by its aid than by
the aid of any other British wing with which I am acquainted. The following
account, taken from a perfectly fresh specimen, may prove interesting.

The joints of the gannet’s wing, particularly the shoulder joint (a of figs. 16
and 17, Plate XIII.) admit of very free movements. When the wing is slightly
flexed the under surface of the posterior margin of the pinion can be rotated
downwards and forwards until it makes a right angle with the horizon—the
greatest angle which it makes in extension amounting to something like 45°,
In flexion the elbow (s of figs. 9, 10, and 11, Plate XII.), wrist (¢), and meta-
carpal joints (v w) admit of a great variety of movements, the forearm (c d) moving
on the arm (ef), and the hand (a 6) upon the forearm (¢ d) in an oblique
spiral direction from above downwards and from below upwards. The whole
pinion, in fact, is flaccid, and the feathers opened up and thrown out of position
as shown more especially at figs. 9 and 10, Plate XII. The forearm is folded
upon the arm in nearly the same plane (vide x s ¢ of fig. 17, Plate XIII), the
secondary and tertiary feathers (c e g of fig. 9, Plate XII.) being inclined slightly
upwards and forwards, so that they form inclined surfaces with the horizon
—the secondaries forming an inclined surface which looks inwards and upwards
as indicated by the arrow marked c¢ d of fig. 9, Plate XII., the tertiary feathers
forming two inclined surfaces, one of which is directed upwards and outwards
as indicated by the arrow e fof fig. 9, Plate XII., the other inclining upwards
and inwards as shown at g h of fig. 9, Plate XII. The hand rotates upon the
wrist (¢ of fig. 9, Plate XII.,) as upon a hinge, the tip of the wing as it darts
out and in describing the segment of a circle (m n of fig. 9, Plate XII.) The
hand is folded upon the forearm in such a manner that the anterior margin of
the tip of the wing (v w 6 of fig. 9, Plate XII.) ascends, while the posterior
margin (a of fig. 9, Plate XII.) descends. As a consequence the hand and tip
of the wing are folded beneath the forearm or body of the wing as indicated by
the radius m 7 of fig. 9, Plate XII. The hand and tip of the wing form
with the horizon an inclined surface, which is directed outwards and upwards
as indicated by the arrows a 0 of fig. 9, Plate XII. The wpward and outward
inclination of the under surface of the outer portion of the wing of the gull is
well seen at a 6 of fig. 12, Plate XII. The tip of the wing, it will be observed,
acts during flexion as a true kite from below upwards and from within out-
wards.* We have in the flexed wing of the gannet four different sets of

* The same happens in the wings of all birds, and in the wing of the bat and insect. The out-
ward and upward inclination of the tip of the wing is well seen in the beetle. This portion of the wing
acts as a true kite, when the wing is being extended or thrust away from the body towards the termi-

nation of the up stroke. The under surface of the tip of the wing consequently contributes to flight —
during the up stroke.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 383

inclined surfaces, two directed upwards and outwards, viz., e fand a 6 of fig. 9,
Plate XII., and two directed upwards and znwards, viz.,c d and g h of fig. 9,
Plate XII. Those surfaces when the wing is moving are ever varying, and cause
the different portions of the pinion to act like so many kites. Thus, during
extension, the two portions of the wing marked a 6 and e/ (fig. 9, Plate XITI.,)
fly outwards and upwards, the two portions marked ¢ d and g h (fig. 9, Plate
XIL.,) flying inwards and upwards during flexion. As the two portions of the
wing, marked ad and ef, draw a current after them during extension, on which
the two portions marked ¢ d and g h operate during flexion, it follows that one
part of the wing, whatever its position in space, makes a current on which another
portion inevitably acts. This result is facilitated by the manner in which the
primary and secondary feathers rotate upon their long axes in flexion and
extension, and also by the ascent and descent of the wing, inasmuch as
flexion always occurs towards the end of the down stroke, and extension
towards the end of the up stroke. The wing, I may add, as a rule produces
a current during the up stroke on which it operates during the down
stroke and vice versa. ‘The inclined surfaces represented at fig. 9, Plate
XII., are reproduced in the partly extended wing at fig. 10, Plate XII., and a
comparison of the arrows marked by the same letters in the two figures will
show that the angles of inclination formed by the surfaces in question are some-
what changed. The wing when fully extended is seen at fig. 11, Plate XII.
Complete extension is followed by the obliteration of the inclined surfaces
indicated by the arrows a b,e f,cd,g h of figs. 9 and 10, Plate XII. The
obliteration of the inclined surfaces a b, ef, ¢ d, g h of figs. 9 and 10, Plate XIL.,
is followed by the production of other inclined surfaces, these being occasioned
by the rotation of the wimg upon its anterior margin (long axis) towards
the termination of extension. The angles of inclination formed by the under
surface of the wing in the extended condition are greatest towards the root and
least towards the tip of the wing, as shown at qg p o of fig. 16, Plate XIII.
When the gannet’s wing is extended and flexed by the aid of the hand, as repre-
sented at figs. 16 and 17, Plate XIII, it shows the screwing and unscrewing
action of the pinion to perfection ; the dorsal and ventral surfaces of the wing
oscillating on either side of a given line—the dorsal surface appearing above the
line in flexion (figs. 17, Plate XIII.,) and the ventral surface under the line in
extension (fig. 16, Plate XIII.) The upward and downward screwing of the
Wing in flexion and extension is also shown at fig. 8, Plate XII.—the wing to
the right of the observer being flexed, and having its anterior margin (d ¢/)
directed slightly downwards (vide arrow), the wing to the left being extended,
and having its anterior margin (d’ ef’) directed decidedly upwards (vide arrow.)

The Angles of Inclination which the Under Surface of the Gannet’s Wing
makes with the Horizon in Extension and Flexion vary.—When the wing of the

VOL. XXVI. PART II. DG

384 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

gannet is extended the angle which its under surface makes with the horizon,
especially the portion opposite the elbow joint (q of fig. 16, Plate XIII), is
much greater than one would anticipate—indeed, it is little short of 45°. The tip
of the wing (0p of fig. 16, Plate XIII.,) does not, however, make an angle of more
than 25° or 30°. This is a most interesting point, as it shows that the different
portions of the wing in extension make different angles with the horizon—that
made by the tip of the wing being the least, and that made by the root of the wing
the greatest. The inclined surfaces are no doubt adapted to suit the travel of
the wing, and to produce a uniform result as far as buoyancy is concerned.
Thus the wing acts with a gradually decreasing angle from the root towards
the tip—the speed of the wing increasing in the direction of its extremity. This
is important, as a surface with a small angle travelling at a high speed sup-
plies the same amount of buoying power as a surface with a greater angle movy-
ing at a lower speed. Indeed, on making a careful examination of the gannet’s
wing I have had no difficulty in determining that the different parts of the wing
not only make various angles of inclination with the horizon in an antero-
posterior direction at every stage of extension flexion in the down and up strokes,
but that they also make various angles of inclination with the horizon in a direc-
tion from within outwards. In other words, I find that in extension the wing
attacks the air from behind forwards and from within outwards at one and the
same instant—the different parts of the pinion tacking upon the air kite fashion,
precisely as a sailing vessel would. The same thing happens in the wing of the
insect. Here, as I have already pointed out, the posterior margin twists upon and
partially rotates round the anterior margin, so as to convert the wing into a
screw which moves in all its parts. This twisting and untwisting has the effect
of alternately producing a surface which attacks the air (at various angles of
inclination) from within outwards, and from behind forwards, and from without
inwards and from before backwards. Curiously enough, the inclined surfaces
formed by the different portions of the insect’s wing with the horizon vary to
accommodate themselves to the velocity acquired by its different parts—the
surfaces being least inclined where the speed is highest, and vzce versa. This,
therefore, is a fundamental point in the construction and application of all
wings, and affords the only rational solution of the involved problem of flight.
The various angles of inclination made by the wing with the horizon from
within outwards and the reverse, and from behind forwards and the reverse, are
all necessary to produce a perfect buoyancy.

When the wing of the gannet is fully extended it is also rendered more or
less rigid. The jomts, however, even the metacarpal ones, are free to move,
which shows that the wing, to be effective during the down stroke, must be
thoroughly under the control of the muscular and ligamentary system. This —
is all the more necessary, as the roots of the primary and secondary feathers

—-

DR PETTIGREW -ON THE PHYSIOLOGY OF WINGS. 385

have an inclination to move in an upward direction, and require to be re-
strained.

After carefully analysing the movements of the gannet’s wing in the dead
bird, I felt deeply impressed with the necessity of studying the same movements
in the living one. I therefore made an excursion to the Bass Rock (North
Berwick, Scotland) for this purpose, in July 1870. It was breeding season,
and the birds were in myriads, and so tame that they wheeled around and above
me at distances, in some cases, not-exceeding from six to eight yards. The
gannets which were hatching permitted me to approach within a yard
of them, and required to be driven from their nests by the aid of a stick. I
had, therefore, every facility for analysing the flight of this the most cherished
and beautiful of the British birds. Before proceeding to describe the results
of the expedition in question I may state, briefly, the measurement, weight,
&c., of the gannet, the movements of whose wings I have just recorded. For
the sake of comparison I will also give the weight and measurements of a
heron—this bird differing widely from the gannet in the configuration of its
wings.

Measurement, Weight, &c., of Gannet and Heron.—The following details of
weight, measurement, &c., of the gannet were supplied by an adult specimen
which I dissected during the winter of 1869. Entire weight, 7 lbs. (minus 3
ounces) ; length of body from tip of bill to tip of tail, 3 feet 4 inches; head
and neck, 1 foot 3 inches; tail, 12 inches; trunk, 13 inches; girth of trunk, 18
inches ; expanse of wing from tip to tip across body, 6 feet ; widest portion of
Wing across primary feathers, 6 inches ; across secondaries, 7 inches; across
tertiaries, 8 inches. Each wing, when carefully measured and squared, gave an
area of 194 square ches. The wings of the gannet, therefore, furnish a sup-

_ porting area of 3 feet 3 inches square. As the bird weighs close upon 7 lbs.,

this gives something like 138 square inches of wing for every 364 ounces of
body, 7.¢., 1 foot 1 square inch of wing for every 2 lbs. 44 ounces of body.

The heron, a specimen of which I dissected at the same time, gave a very
different result, as the subjoined particulars will show. Weight of body, 3 Ibs.
3 ounces ; length of body from tip of bill to tip of tail, 3 feet 4 inches; head
and neck, 2 feet; tail, 7 inches; trunk, 9 inches; girth of body, 12 inches;
expanse of wing from tip to tip across the body, 5 feet 9 inches ; widest portion
of wing across primary and tertiary feathers, 11 inches; across secondary
feathers, 12 inches.

Each wing, when carefully measured and squared, gave an area of 26 square
inches. The wings of the heron, consequently, furnish a supporting area of 4
feet 4 inches square. As the bird only weighs 3 Ibs. 3 ounces, this gives
something like 26 square inches of wing for every 254 ounces of bird, or 1 foot
54 inches square of wing for every 1 lb. 1 ounce of body.

386 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

In the gannet there is only 1 foot 1 square inch of wing for every 2 lbs.
4! ounces of body. The gannet has, consequently, less than half of the wing
area of the heron. The gannet’s wing is, however, a long narrow wing (that
of the heron is broad), extended transversely across the body in the direction
of its length; and this is found to be the most powerful form of wing—the
wings of the albatros, which measure 14 feet from tip to tip (and only one foot
across), elevating 18 lbs. without difficulty. If the wings of the gannet, which
have a superficial area of 3 feet 3 inches square, are capable of elevating 7 lbs.,
while the wings of the heron, which have a superficial area of 4 feet 4 inches,
can only elevate 3 Ibs., it is evident (seeing the wings of both are twisted
levers, and formed upon a common type) that the gannet’s wing must be vibrated
with greater energy than the heron’s wing; and this is actually the case.
The heron’s wing, as I have stated (foot note to page 392), makes 60 down and
60 up strokes every minute ; whereas the wing of the gannet, when the bird is
flying in a straight line to or from its fishing ground, makes close upon 150
up and 150 down strokes during the same period. The wings of the divers
and other short-winged, heavy-bodied birds are urged at a much higher speed,
so that a comparatively small wing can be made to elevate a comparatively heavy
body, if the speed with which the wing is driven only be increased sufficiently.*
Flight, therefore, is a question of power, speed, and small surfaces versus
weight. While there is apparently no fixed relation between the area of the
wing and the animal to be raised, there is (unless in the case of sailing birds,
which have acquired momentum) an unvarying relation as to the weight to be
elevated and the number of oscillations; so that the problem of flight would seem
to resolve itself into one of weight, power, velocity, and small surfaces, as against
comparative levity, debility, diminished speed, and extensive surfaces.t Ela-
borate measurements of wing area and minute calculations of speed can, con-
sequently, only determine the minimum of wing for elevating the maximum of
weight—flight being attainable within a comparatively wide range. That the
superficies of the wings destined to carry a certain weight may, and does vary,
is proved by the fact that large portions of the wings of insects and birds, as
I have pointed out,{ may be removed without destroying or even impairing the
function of flight. In such cases the speed with which the wings are driven is in-
creased in the direct ratio of the mutilation. It is further proved by the ingenious -
researches of M. pE Lucy, who has shown, by careful measurements, that the

* The grebes among birds and the beetles among insects furnish examples where small wings,
made to vibrate at high speeds, are capable of elevating great weights.

t “On the Mechanism of Flight,” by the Author, Trans. Linn. Soc., vol. xxvi. page 219.

t Vide page 326 and foot-note to pages 361 and 362 of the present memoir, and pages 219, 220,
221, and 222 of my memoir “On the Mechanical Appliances by which Flight is Attained in the
Animal Kingdom,” Trans. Linn. Society, vol. xxvi.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 387

area of the wings decreases as the size and weight of the body increase. M.
pE Lucy has tabulated his results, which I subjoin.*

INSECTS. BIrps.

Referred to the

Lucanus ) Stag- hectle (female),
cervus Stae-hectle ( ee
Rhinoceros- het, : ?

kilogramme. Y
Names. =2 Ibs. ot 02.2 hg 2 gr. Names. feieee
=2 Ibs. 3 oz. 4°428 dr.
vai, ft. in. sae ft. inch.
emai 3 eae LI S92 Swallow, 1 1042
Dragon-fly (anal), dy 2456 Sparrow, QO 5 1422
Coccinella (Lady-bird), Bah tire Turtle dove, Shay are 0 4 1003
Dragon-fly (common), pL V2 589 IE COMa My Ns Beret, Gs
Tipula, or Daddy long-egs Sou lel Stork, O52 20
Bee, . : 4 1 2 744 Vulture, 0 1 116
Meat-tfly, ; 1 3 543 Crane of Rasialia) 0 O 139
Drone (blue), . L220
Cockchafer, De 26550
Li
0 8
0 6

“It is easy, by the aid of this table, to follow the order, always decreasing,
of the surfaces, in proportion as the winged animal increases in size and weight.
Thus, in comparing the insects with one another, we find that the gnat, which
weighs 460 times less than the stag-beetle, has 14 times more of surface.
The lady-bird weighs 150 times less than the stag-beetle, and possesses 5 times
more of surface, &c. It is the same with the birds. The sparrow weighs about
10 times less than the pigeon, and has twice as much surface. The pigeon
weighs about 8 times less than the stork, and has twice as much surface.
The sparrow weighs 339 times less than the Australian crane, and possesses
7 times more surface, &c. If now we compare the insects and the birds,
the gradation will become even much more striking. The gnat, for example,
weighs 97,000 times less than the pigeon, and has 40 times more surface ;
it weighs three millions of times less than the crane of Australia, and possesses
140 times more of surface than this latter, the weight of which is about 9 kilo-
grammes 500 grammes (25 lbs. 5 oz. 9 eas troy, 20 lbs. 15 oz. 24 dr, avoirdu-
pois.

“ The Australian crane is the heaviest bird that I have weighed. It is that
which has the smallest amount of surface, for, referred to the kilogramme, it does
not give us a surface of more than 899 square centimetres (139 square inches),
that is to say, about an eleventh part of a square metre. But every one

* “On the Flight of Birds, of Bats, and of Insects, in reference to the subject of Aérial Locomo-
tion,” by M. pr Lucy, Paris.

VOL. XXVI. PART II. 5H

388 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

~ knows that these Grallatorial animals are excellent birds of flight. Of all travel-
ling birds they undertake the longest and most remote journeys. They are,
in addition, the eagle excepted, the birds which elevate themselves the highest,
and the flight of which is the longest maintained.”

Flight of Gannet as witnessed at the Bass Rock.—But to return to the gannet,
the flight of which, as witnessed from the Bass, I was about to describe.

The wings and body of the bird, as I fully satisfied myself, can be moved
in all their parts. The wings and body are, moreover, thoroughly under control.
The body can be twisted about in a remarkable manner—sideways and in
an upward and downward direction. The individual feathers of the wing are
likewise under control. In fact, the muscular movements can be seen extend-
ing along the pinion to the roots of the rowing feathers, the muscular influence
spreading thence to the tips. This could readily be ascertained, as the birds
wheeled round and round right overhead, and within a very few yards of where I
was standing.

When the gannet throws itself from a cliff it makes a large curve, the con-
vexity of which is directed downwards. It acquires speed and momentum by
a few gentle flappings of the wings, or it holds the wings comparatively motion-
less, and sails for a great distance without effort—the weight of the trunk domg
the principal portion of the work.* In the sailing movement the body is forced
into an upward or downward curve, according to circumstances.

When the bird has acquired momentum, either by flapping its wings or by
projecting itself from a cliff, it has the air perfectly under control. If it wishes
to turn to the right it elevates the left wing and depresses the right one, the
head and neck bending in the direction of the curve to be described. [If it
would turn to the left the movements are reversed.t If it desires to ascend,
the head, neck, body, and wings are elevated in an upward direction, so as to
increase the angle made by them with the horizon, the angle referred to being
decreased or reversed when the bird wishes to descend. If the bird aims at
horizontal flight, the head, neck, body, and wings are arranged so as to be nearly
parallel with the surface of the sea. The gannet wheels and skims about with
all imaginable ease and grace—now oscillating on the long axis of the body as
a centre, and now upon the long axes of the wings as a centre. In all these
movements the head, neck, tail, and body perform an important part.

When the gannet throws itself from a rock it rises to nearly the same level
as that from which it precipitated itself, without any apparent effort, thus showing
that the friction experienced in flight must be almost ni.

The neck, body, and tail, of the gannet are exceedingly flexible, and admit

* Compare with mechanical experiment described at pages 355 and 356.
} The swallow and crane, which dart along at a very high speed, tilt their bodies in turning; but,
in addition, flap their wings and fly round the curve they wish to describe.

—

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 389

of being curved in every direction. The feet are extended straight out behind
the bird, and appear on the under surface of the tail. The body forms an
elongated and very graceful ellipse, admirably adapted for cleaving the air and
eluding resistance.

When the gannet propels itself by the more or less vertical flappings of its
wings, the angles which the under surfaces of the wings and body make with the
horizon are very considerable—something like 25° or 30°. Of this I convinced
myself in a variety of ways.* When the bird has acquired speed and momentum,
and begins to sail, the angle made by the under surfaces of the body and
wings is reduced according to circumstances, and in some instances nearly
obliterated, the bird gliding along for long distances with its body and wings
apparently parallel to the surface of the ocean.

The wings of the gannet, when fully extended, are curved alternately for-
wards and backwards. Thus, the arm and hand are inclined backwards, and
the forearm forwards. When the wings are flexed in ordinary flight the move-
ment occurs principally at the wrist joint, the arm and forearm bending com-
paratively little, and affording a wide basis of support both during the down and
up strokes. In forced flight im flexion the wing bends perceptibly at the elbow
as well as the wrist, the wing during the up stroke forming a short lever, and
being thrown into a fine arch, the convexity of which is directed upwards. The
tip of the wing works out and in during the down and up strokes; and a close
examination satisfied me that the bird has the power of forcing the posterior
margin of its wings znto wave curves while the wings are rising and falling, the
air taking no part in the production of the waved movements.

The down stroke is delivered with perceptibly greater rapidity and energy
than the up stroke. Of this there can be no doubt whatever. This allows
the air, set in motion by the wing during its descent, time to re-act on the

under surface of the pinion so as to contribute to its elevation. This result is

facilitated by the wing striking very decidedly downwards and forwards.

When the gannet alights at its nest it delivers a few very energetic strokes
at right angles to the direction of its flight, and thus slows itself.

When the gannet plunges into the sea from a height it tilts its body until it
assumes a more or less perpendicular position, and descends with such impetuosity
as to displace the water in an upward direction, until it attains an altitude of
from 10 to 15 feet. It flies beneath the water with remarkable rapidity, and
emerges without difficulty, the momentum acquired during the descent assisting
it through and out of the water. In fact the gannet, when it stoops to pick up
a fish, simply describes a continuous downward curve, part of the curve being

* In the dragon-fly the anterior pair of wings make a smaller angle with the horizon than the pos-
terior pair. The first pair of wings are, consequently, more actively engaged as propellors—the second
pair as elevators.

390 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

formed in the air and part in the water. Those movements, so numerous, varied, |
and beautiful, are all the result of volition. It is impossible to resist this con-
clusion after deliberate and careful watching.

A Regulating Power necessary in Flight.—That the wing is propelled for the
most part by voluntary movements, may be ascertained in the following manner.

If the sentient nerve of a pigeon’s wing be divided (the motor nerve being
left intact) the bird flutters most energetically, but altogether fails to fly.* In
this experiment neither the flexibility, elasticity, nor the power which the wing
possesses of moving in all its parts, are tampered with. The guiding or con-
trolling power alone is impaired.

That the wing is vibrated intelligently admits of direct proof. Thus if we
hold a captured bird in the hand, we feel that it directs and controls the action
of its wings in such a manner that a tractile force is produced, now in one
direction now in another, in its efforts to escape ; nay more, that the force after
a brief fluttering is concentrated at that point where it is most loosely held, and
which offers the greatest chance of escape.

Second, The wings of birds, as any one may readily ascertain by watching
the flight of rooks, are visibly under control both during the down and up
strokes. They are, moreover, deliberate leisurely movements. By leisurely
movements, I mean such as are the result of design, and not such as would
be produced by the sudden recoil of a merely elastic apparatus. Those who have
watched, as I have frequently done, the rapid vibrations of natural and artificial
wings, will readily understand the difference here indicated. In the living wing
we have a smooth soft fanning continuous movement, quite devoid of dead
points ; whereas in artificial elastic wings, especially if worked vertically and
without elastic bands at their roots, we have a wavering, jerking, irregular
motion, particularly at the beginning of the up stroke.

Third, The blow-fly, as stated (p. 326), can fly with only one-third of its
original wing area, the two-thirds which represent the more highly elastic
portions of the wing being removed. In this case the wing is wielded intelli-
gently figure of 8 fashion, the mutilation not interfering either with the freedom
of motion enjoyed by the pinion at its root, or the power the insect possesses
of directing and controlling the wing throughout its entire vibration.

There are therefore at least five separate items to be considered in flight,
viz., intelligence and voluntary movements ; secondly, mobility or the power
which the wing possesses of moving its several parts ; thirdly, the flexibility and
elasticity of the wing; fourthly, the resistance and resiliency of the air upon
which the wing operates ; fifthly, the weight of the body of the flying animal,
which may be regarded as an independent moving power.

* “Experiments practically demonstrating the laws by which birds fly,” by Dr W. Suyru. Second
Annual Report of the Aéronautical Society of Great Britain for 1867.

ss.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 391

The wings of bats and birds are mobile because of their numerous joints
(shoulder, elbow, wrist, meta carpal, &c.), and because of the muscles and fibro-
elastic ligaments which operate upon these joints. They are also flexible and
elastic, the one (the bat) because of its long, thin tapering fingers and envelop-
ing membrane ; the other (the bird) because of its tapering, primary, secondary,
and tertiary feathers.

The insect wing is also mobile, the insect having the power not only of
moving the pinion in various directions at its root, but of causing the move-
ments generated at the roots to extend intelligently along the margins. The
insect wing is flexible and elastic in the same sense that the wing of the bat
and bird are flexible and elastic. The mobility, flexibility, and elasticity peculiar
to the living wing is more intimately blended in the wing of the insect than in
that of either the bat or bird. This arises from the fact that the wing of the
insect is usually in one piece, and jointed only at its root.

The Wing at all times thoroughly under Control.—The advantage which the
wing derives from being movable in all its parts, consists in this, that it can be
wielded intelligently even to its extremity. This enables the insect, bat, and
bird, to tread and rise upon the air as a master—to subjugate it in fact. The
wing, no doubt, abstracts an upward and onward recoil from the air, but in
doing this it exercises a selective and controlling power ; it seizes one current,
evades another, and creates a third; it feels and paws the air as a quadruped
would feel and paw a treacherous yielding surface. It is not difficult to com-
prehend why this should be so. If the flying creature is living, endowed with
volition, and capable of directing its own course, it is surely more reasonable
to suppose that it transmits to its travelling surfaces the peculiar movements
necessary to progression, than that those movements should be the result of
impact from fortuitous currents which it has no means of regulating. That the
bird requires to control the wing, and that the wing requires to be in a condition
to obey the behests of the will of the bird, is pretty evident from the fact that
most of our domestic fowls can fly for considerable distances when they are
young and when their wings are flexible ; whereas when they are old and the
wings stiff, they either do not fly at all or only for short distances, and with great
difficulty. This is particularly the case with tame swans. This remark also holds
true of the steamer or race-horse duck (Anas brachyptera), the younger speci-
mens of which only are volant. In the older birds the wings become too rigid
and the bodies too heavy for flight. Who that has watched a sea-mew struggling
bravely with the storm, could doubt for an instant that not only the wings but
every individual feather of the wing was perfectly under control? The whole
bird is an embodiment of animation and power. The intelligent active eye, the
easy graceful oscillation of the head and neck, the folding or partial folding of

one or both wings, nay more, the slight tremor or quiver of the individual
VOL. XXVI. PART II. 51

392 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

feathers of parts of the wings so rapid, that only an experienced eye can detect
it, all confirm the belief that the living wing has not only the power of directing,
controlling, and utilising natural currents, but of creating and utilising artificial
ones, which is not less important. But for this power, what would enable the
bat and bird to rise and fly in a calm, or steer their course in a gale? It is
erroneous to suppose that anything is left to chance where living organisms are
concerned, or that animals endowed with volition and travelling surfaces, should
be denied the privilege of controlling the movements of those surfaces quite
independently of the medium on or in which they are destined to operate. What
would we say of that quadruped or that fish which depended for the major
portion of its movements on the ground it trod or the water it navigated? I
will never forget the gratification afforded me on one occasion at Carlow
(Ireland) by the flight of a pair of magnificent swans. The birds flew towards
and past me, and I had my attention directed to their presence by a peculiarly
loud whistling noise made by their wings. They flew about fifteen yards from
the ground, and as their pinions were urged not much faster than those of the
heron,* I had abundant leisure for studying their movements. The sight was
very imposing, and as novel as it was grand. I had never seen anything before,
and certainly have seen nothing since that could in any way convey a more
adequate idea of the prowess and guiding power which a bird may exert.
What particularly struck me was the perfect mastery which they seemed to
possess over everything. They had their wings and bodies visibly under control,
and the air was attacked in a manner and with an energy which left little doubt
in my mind that it played quite a subordinate part in the great problem before
me. The necks of the birds were stretched out, and their bodies to a great
extent rigid. They advanced with a steady stately motion, and swept past with
a vigour and force which greatly impressed, and to a certain extent overawed,
me at the time.t Their flight was what one could imagine that of a flying
machine constructed in accordance with natural laws would be.

* T have frequently timed the beats of the wings of the common heron (Ardea cinerea) at Warren
Point (Ireland). In March 1869 I was placed under unusually favourable circumstances for obtain-
ing reliable results. I timed one bird high up over a lake for fifty seconds, and found that in that
period it made fifty down and fifty up strokes ; i.e., one down and one up stroke per second. I timed
another one in a heronry belonging to Major Hatt. It was snowing at the time (March 1869), but
the birds, notwithstanding the inclemency of the weather and the early time of the year, were actively
engaged in hatching, and required to be driven from their nests on the top of the larch trees by knock-
ing against the trunks thereof with large sticks. One unusually anxious mother refused to leave the
immediate neighbourhood of the tree containing her tender charge, and circled round and round it
right overhead. I timed this bird for ten seconds, and found that she made ten down and ten up
strokes ; 7.e., one down and one up stroke per second precisely as before. I have therefore no hesitation
in affirming that the heron, in ordinary flight, makes exactly sixty down and sixty up strokes per
minute. The heron, however, like all other birds when pursued or agitated, has the power of greatly
augmenting the number of its beats.

+ The above observation was made at Carlow on the Barrow in October 1867, and the account of
it is abstracted from my note-book.

—

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 395

How the Wing is Attached to the Body—Movements of the Shoulder, Elbow,
Wrist, and other Joints—Having endeavoured to prove, in a variety of ways,
that insects, bats, and birds have their wings thoroughly under control both
during the down and up strokes, I now proceed to show that the configuration
of the wing, its structure, its attachments to the body, its joints, its muscles
(voluntary in their nature), and its elastic ligaments, many of which have
muscular fibres running into them, all tend to confirm this belief.

While, however, saying so much, I take this opportunity of stating that the
structure of the living wing and its relations and attachments to the body are
such that if it moves at all it must move in such a manner as shall contribute
to flight. In other words, the wing is mechanically perfect ; and if it be made
to vibrate, even by artificial means, all its movements will tend in the direction
of flight. This, however, is a very different thing from asserting that the move-
ments of the living wing are purely mechanical in their nature. By mechanical
I mean such movements as would be produced by the elasticity of the wing
and the reaction of the air, minus volition, minus the voluntary muscles—mus-
culo-elastic ligaments and nerves of the wing. Flight is vito-mechanical in its
nature and intelligence, or that form of action which results from the habitual
use of intelligence, is necessary to its production.

All wings are constructed upon a common type. They are in every instance
carefully graduated, the wing tapering from the root towards the tip,.and from
the anterior margin in the direction of the posterior margin. They are ofa
generally triangular form, and twisted upon themselves in the direction of their
length, to form a helix or screw. They are convex above and concave below,
and more or less flexible and elastic throughout, the elasticity being greatest at
the tip and along the posterior margin. They are also movable in all their
parts. In all the wings which I have examined, whether in the insect, bat, or
bird, the wing is recovered, flexed, or drawn towards the body by the action of
elastic ligaments, these structures, by their mere contraction, causing the wing,
when fully extended and presenting its maximum of surface, to resume its posi-
tion of rest and plane of least resistance. The principal effort required in flight
is, therefore, made during extension and at the beginning of the down stroke.
The elastic ligaments are variously formed, and the amount of contraction which
they undergo is in all cases accurately adapted to the size and form of the wing
and the rapidity with which it is worked, the contraction being greatest in the
short-winged and heavy-bodied insects and birds, and least in the light-bodied
and ample-winged ones, particularly in such as skim or glide. The mechanical
action of the elastic ligaments, I need scarcely remark, ensures an additional
period of repose to the wing at each stroke; and this is a point of some im-
portance, as showing that the lengthened and laborious flights of insects and
birds are not without their stated intervals of rest.

394 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

The twisting of the wing upon itself during its action, to which I have fre-
quently directed attention, is occasioned in the bat and bird by the insertions
and direction of the muscles—by the spiral configuration of the articular sur-_
faces of the bones of the wing, and by the rotation of the bones of the arm,
forearm, and hand upon their long axes. In the insect it is due to the insertions
and direction of the muscles, and the conformation of the shoulder-joint, this
being furnished with a system of check-ligaments, and with horny prominences
or stops, set, as nearly as may be, at right angles to each other, and fashioned
so as to necessitate the wing acting in the manner specified.

To confer on the pinion the multiplicity of movement which it requires, it
is supplied with a double hinge or compound joint, which enables it to move
not only in an upward, downward, forward and backward direction, but also at
various intermediate degrees of obliquity. An insect furnished with wings thus
hinged may, as far as steadiness of body is concerned, be not inaptly compared
to a compass set upon gimbals, the universality of the wing-movements ren-
dering any elaborate attempt at balancing quite unnecessary.

In the bird the head of the humerus is convex and somewhat oval (not round),
the long axis of the oval being directed from above downwards, 7.¢., from the
dorsal towards the ventral aspect of the bird. The humerus can, therefore,
glide up and down in the facettes occurring on the articular ends of the coracoid
and scapular bones with great facility, much in the same way that the head of
the radius glides upon the distal end of the humerus. But the humerus has
another motion ; it moves like a hinge from before backwards, and vice versa.
The axis of the latter movement is almost at right angles to that of the former. —
As, however, the shoulder-joint is connected by long ligaments to the body, and
can be drawn away from it to the extent of one-eighth of an inch or more, it
follows that @ third and twisting movement can be performed, the twisting admit-_
ting of rotation to the extent of something like a quarter of aturn. In raising
and extending the wing preparatory to the downward stroke two opposite
movements are required, viz., one from before backwards, and another from
below upwards. As, however, the axes of these movements are at nearly
right angles to each other, a spiral or twisting movement is necessary to run
the one into the other—to turn the corner, in fact.

From what has been stated it will be evident that the movements of the
wing, particularly at the root, are remarkably free, and very varied. A directing
and restraining, as well as a propelling force, is therefore necessary.

Such complex force is to be found in the voluntary muscles which connect the
wing with the body in the insect, and which in the bat and bird, in addition to
connecting the wing with the body, extend along the pinion even to its tip.
It is also to be found in the musculo-elastic and other igaments. Ido not
propose entering upon a consideration of the muscular system of the wing of

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 399

the bat and bird, as this has been satisfactorily done already. I will, therefore,
confine the present remarks to the elastic ligaments, and more especially to those
of the bird, as being the most illustrative, alike from their size and situation.

The Wing Flexed and partly Elevated by the Action of Elastic Ligaments—
the Nature and Position of such Ligaments in the Pheasant, Snipe, Crested Crane,
Swan, &c—When the wing is drawn away from the body of the bird by the
hand the posterior margin of the pinion formed by the primary, secondary,
and tertiary feathers rolls down to make a variety of inclined surfaces with the
horizon. When, however, the hand is withdrawn, even in the dead bird, the wing
instantly folds up; and in doing so, reduces the amount of inclination in the several
surfaces referred to. This it does in virtue of certain elastic ligaments, which are
put upon the stretch in extension, and which recover their original form and posi-
tion in flexion. This simple experiment shows that the various inclined surfaces
requisite for flight are produced by the mere act of extension and flexion in
thedead bird. It is not, however, to be inferred from this circumstance that
flight in the animal kingdom is a purely mechanical act any more than ordi-
nary walking is. The muscles, bones, ligaments, feathers, &c. are so adjusted
with reference to each other that if the wing is moved at all, it must be moved
in the proper direction—an arrangement which enables the bird to fly without
thinking just as we can walk without thinking. There cannot, however, be a
shadow of a doubt that the bird has the power of controlling its wings both
during the down and up strokes; for how otherwise could it steer and direct
its course with such precision in obtaining its food ? how fix its wings on a level
with or above its body for skimming purposes ? how forma curve ? how fly with,
against, or across a breeze? how project itself from a rock directly into space,
or how elevate itself from a level surface by the laboured action of its wings ?

The wing of the bird is elevated to a certain extent in flight by the reac-
tion of the air upon its under surface; but it is also elevated by muscular
action—by the contraction of the elastic ligaments, and by the body falling
downwards and forwards in a curve.

That muscular action is necessary is proved by the fact that the pinion
is supplied with distinct elevator muscles*—nay, more, that the bird can, and
always does, elevate its wing prior to flight, quite independently of the air.
When the bird is fairly launched into space the elevator muscles are assisted

* C. J. L. Krarup, a Danish author, gives it as his opinion that the wing is elevated by a vital
force, viz., by the contraction of the pectoralis minor; this muscle, according to him, acting with }th
the intensity of the pectoralis major (the depressor of the wing). He bases his statement upon the
fact that in the pigeon the pectoralis minor or elevator of the wing weighs 3th of an ounce, whereas
the pectoralis major or depressor of the wing weighs ths of an ounce. It ought, however, to be
borne in mind that the volume of a muscle does not necessarily determine the precise influence exerted by
its action ; for the tendon of one muscle may be made to act upon a long lever, and, under favourable con-
ditions, for developing its powers, while that of another muscle may be made to act upon a short lever,
and, consequently, under unfavourable conditions.—On the Flight of Birds, p. 30. Copenhagen, 1869.

VOL. XXVI. PART II. aya

396 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

by the falling forward of the body, by the reaction of the air, and by the con-
traction of the elastic ligaments. The air and the elastic ligaments contribute
to the elevation of the wing, but both are obviously under control—they, in
fact, form links in a chain of motion which at once begins and terminates in
the muscular system.

That the elastic ligaments are subsidiary and to a certain extent under the
control of the muscular system in the same sense that the air is, is evident from
the fact that voluntary muscular fibres run into the ligaments in question at
various points. Thus, in the pheasant, as shown at a 0 of figure 23, Plate XV.,
red muscular fibres are seen terminating in the fibrous and elastic tissues
cand. These structures act in conjunction, and fold or flex the forearm on
the arm. At // voluntary muscle is seen acting in concert with the elastic
ligament gz to flex the hand upon the forearm. The arm is drawn towards
the body by the elastic igament d and by the muscles v w.

The elastic ligaments, while occupying a similar position in the wings of all
birds, are variously constructed in the several species. In the common snipe,
for example, as represented at figure 21, Plate XV., the voluntary muscular
slip a terminates in the fibro-elastic band 4; this again being geared to volun-
tary muscle x, and to certain musculo-fibrous bands 7. Their conjoined action
is to flex the forearm upon the arm, the arm being drawn towards the body by
a musculo-fibrous ligament d, e. The elastic ligament g 2 flexes the hand upon
the forearm, and the ligament 7 the fingers upon the hand. A somewhat
similar arrangement is formed in the wing of the crested crane, as shown at
figure 24, Plate XVI. Thus, at a, 6, voluntary muscular slips are seen termi-
nating in the elastic band 4, this splitting up into two portions at 4, m. A some-
what similar band is seen at 7, and all three are united to, and act in conjunc-
tion with, the great fibro-elastic web ¢ to flex the elbow. The musculo-fibro-
elastic ligament / g, h 7, as already explained, envelopes the root of each
primary, secondary, and tertiary feather. It also forms a symmetrical network,
so that it at once supports the feathers and limits their peculiar actions. In
the swan the muscular slip which corresponds to a of figure 24, Plate XVI.
(crested crane), terminates in a fibrous band, which corresponds to 4; but the
muscular slip corresponding to } terminates in a well-defined tendon, not in the
fibrous band m, but in a distinct muscle, 5 inches in length and + of an inch
in breadth. This muscle is situated in the anterior margin of the wing, mid-
way between the shoulder and wrist joints, and exercises a most potent influence
in folding the elbow. The band marked 7 in the crane’s wing is at least four
times broader in the swan’s wing.

The Elastic Ligaments more Highly Differentiated in Wings which Vibrate
Rapidly.—From what has been stated, it will be evident that the elastic liga-
ments of the swan are more complicated and more liberally supplied with

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 397

voluntary muscle than those of the crane, and this is no doubt owing to the
fact that the wings of the swan are driven at a much higher speed than those
of the crane. In the snipe the wings are vibrated very much more rapidly than
in the swan, and as a consequence we find that the fibro-elastic bands are not
only greatly increased, but they are also geared to a much greater number of
voluntary muscles, all which seems to prove that the elastic apparatus employed
by nature for recovering or flexing the wing towards the end of the down stroke
become more and more highly differentiated in proportion to the rapidity with
which the wing is moved.* The reason for this is obvious. If the wing is to
be worked at a higher speed, it must, as a consequence, be more rapidly flexed
and extended. The rapidity with which the wing of the bird is extended and
flexed is in some instances exceedingly great ; so great, in fact, that it escapes
the eye of the ordinary observer. The rapidity with which the wing darts in
and out in flexion and extension would be quite inexplicable, but for a know-
ledge of the circumstance that the different portions of the pinion are disposed
at various angles of inclination (vde 2, s, t, w of figures 9 and 10, Plate XIT.),
these angles being instantly increased or diminished by the slightest quiver of
the muscular and fibro-elastic systems. If we take into account the fact that
the wing of the bird is recovered or flexed by the combined action of voluntary
muscles and elastic ligaments; that it is elevated to a considerable extent by
voluntary muscular effort; and that it is extended and depressed entirely by
muscular exertion, we shall have difficulty in avoiding the conclusion that the
wing is thoroughly under the control of the muscular system, not only in flexion
and extension, but also throughout the entire down and up strokes.

An arrangement in every respect analogous to that just described is found
in the wing of the bat, the covering or web of the wing in this instance forming
the principal elastic ligament. In fact, the bones and muscles of the bat’s
wing, and the inclined surfaces made by its different portions with each other
and with the horizon during flexion and extension, and during the down and up
strokes, so closely resemble those of the bird that a separate description is un-
necessary. From the foregoing description it will be obvious that the wing of
the bird and bat is a highly differentiated organ, endowed with independent
movements, which enable it to direct and control the air for a purpose.

How Balancing is Effected in Flight—The manner in which insects, bats,
and birds balance themselves in the air has hitherto, and with reason, been
regarded a mystery, for it is difficult to understand how they maintain their
equilibrium when the wings are beneath their bodies. Figures 3 and 4, page 338,
throw considerable light on the subject in the case of the insect. In those
figures the space (a, g) mapped out by the wing during its vibrations is entirely

* A careful account of the musculo-elastic structures occurring in the wing of the pigeon is given
by Mr Macemuivray in his admirable “ History of British Birds,” pages 37 and 38. Lond. 1837.

398 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

occupied by it ; @.e., the wing (such is its speed) is in every portion of the space
at nearly the same instant, the space representing what is practically a solid
basis of support. As, moreover, the wing is jointed to the upper part of the
body (thorax) by a universal joint, which admits of every variety of motion,
the insect is always suspended (very much as a compass set upon gimbals is
suspended), the wings, when on a level with the body, vibrating in such a manner
as to occupy a circular area, in the centre of which the body is placed (vide
rdbf of fig. 51, page 399). The wings, when vibrating above and beneath the
body occupy a conical area, the apex of the cone being directed upwards when
the wings are below the body, and downwards when beneath it. Those points
are well seen in the bird at figures 18 and 19, Plate XIV. In figure 18
the inverted cone formed by the wings when above the body is represented,
and in figure 19 that formed by the wings when below the body is given. In
these figures it will be observed that the body, from the insertion of the roots of
the wings into its upper portion, is always suspended, and this, of course, is
equivalent to suspending the centre of gravity. In the bird and bat, where the
stroke is delivered more vertically than in the insect, the basis of support is
increased by the tip of the wing folding inwards and backwards in a more or
less horizontal direction at the end of the down stroke ; and outwards and for-
wards at the end of the up stroke. This is accompanied by the rotation of the
outer portion of the wing upon the wrist as a centre (vide ¢ of figures 9 and 10,
Plate XII.), the tip of the wing, because of the ever varying position of the
wrist, describing an ellipse. In insects whose wings are broad and large
(butterfly), and which are driven at a comparatively low speed, the balancing
power is diminished. In insects whose wings, on the contrary, are long and
narrow (blow-fly), and which are driven at a high speed, the balancing power
is increased. It is the same with short and long winged birds, so that the
function of balancing is in some measure due to the form of the wing, and the
speed with which it is driven, the long wing and the wing vibrated with great
energy increasing the capacity for balancing. When the body is light and
the wings very ample (butterfly and heron), the descent of the wing and the
reaction of the air during the up stroke displaces the body to a marked
extent. When, on the other hand, the wings are small and the body large, the
reaction produced on the trunk by the vibration of the wing is scarcely per-
ceptible. Apart, however, from the shape and dimensions of the wing, and the
rapidity with which it is urged, it must never be overlooked that all wings (as
has been pointed out) are attached to the bodies of the animals bearing them
by some form of universal joint, and in such a manner that the bodies, whatever
the position of the wings, are accurately balanced, and swim about precisely
after the fashion of a compass set upon gimbals. To such an extent is this true,
that the position of the wing isa matter of indifference. Thus the pinion may be

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 399

above, beneath, or on a level with the body; or it may be directed forwards,
backwards, or at right angles to the body. In either case the body is balanced
mechanically and without effort. To prove this point, I made an artificial wing
and body, and united the one to the other by a universal joint. I found,as I had
anticipated, that place the wing in whatever position I chose, whether above,
beneath, or on a level with the body, or at either side of it, the body almost
instantly attained a position of rest. The body was, in fact, equally suspended
and balanced from all points.

Rapidity of Wing Movements partly Accounted for.—Much surprise has
been expressed at the enormous rapidity with which some wings are made

ot Os
\

Fig. 51.*

to vibrate. The wing of the insect is, as a rule, very long and narrow. Asa
consequence, a comparatively slow and very limited movement at the root
confers great range and immense speed at the tip, the speed of each portion
of the wing increasing as the root of the wing is receded from. This is
explained on a principle well understood in mechanics, viz., that when a rod
hinged at one end is made to move in a circle, the tip or free end of the rod
describes a much wider circle ¢n a given time than a portion of the rod nearer the

* In this diagram I have represented the wing by a straight rigid rod. The natural wing, how-
ever, is curved, flexible, and elastic. It likewise moves in curves, the curves being most marked towards
the end of the down and up strokes, as shown at m, n,0,p. The curves, which are double figure
of 8 curves, are obliterated towards the middle of the strokes (r, a). This remark holds true of all
natural wings, and of all artificial wings properly constructed. The curves and the reversal thereof are
hecessary to give continuity of motion to the wing during its vibrations, and what is not less important,
to enable the wing alternately to seize and dismiss the air.

VOL. XXVIL. PART. I. 5.1L

400 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

hinge. Thus if a 0 of figure 51 (p. 399) be made to represent the rod hinged at a,
it travels through the space db fin the same time that it travels through 7 £/;
and through the space 7 // in the same time that it travels through the space
g hi; and through the space g / 7 in the same time it travels through e @ ¢, which
is the area occupied by the thorax of the insect. If, however, the rod a 0 travels
through the space dbf in the same time that it travels through the space eae,
it follows of necessity that the portion of the rod marked a moves very much
slower than that marked 6. The muscles of the insect are applied at the point
a, as short levers (the point referred to corresponding to the thorax of the
insect), so that a comparatively slow and limited movement at the root of the
wing produces the marvellous speed observed at the tip, the tip and body of
the wing being those portions which occasion the blur or impression produced on
the eye by the rapidly oscillating pinion. But for this mode of augmenting the
speed originally inaugurated by the muscular system, it is difficult to comprehend
how the wings could be driven at the velocity attributed to them. The wing of
the blow-fly is said to make 300 strokes per second, 2.¢., 18,000 strokes per minute.
Now it appears to me that muscles to contract at the rate of 18,000 times in the
minute would be exhausted in a very few seconds, a state of matters which
would render the continuous flight of insects impossible. (The heart contracts
only between 60 and 70 times in a minute.) I am therefore disposed to
believe that the number of contractions made by the thoracic muscles of insects
has been greatly overstated, the high speed at which the wing is made to vibrate
being due less to the separate and sudden contractions of the muscles at its
roots than to the fact that the speed of the different parts of the wing is increased
in a direct ratio as the portions in question are removed from the driving point,
as already explained. Speed is certainly a matter of great importance in wing
movements, as the elevating and propelling power of the pinion depends to a
great extent upon this condition. Speed, however, may be produced in two ways
—either by a series of separate and opposite movements, such as is witnessed im
the action of a piston, or by a series of separate and opposite movements, acting
upon an instrument so designed that a movement applied at one part increases in
rapidity as the point of contact is receded from, as happens in the wimg. In the
piston movement the motion is uniform, or nearly so, all parts of the piston
travelling at very much the same speed. In the wing movements, on the con-
irary, the motion is gradually accelerated towards the tip of the pinion, where
the pinion is most effective as an elevator, and decreased towards the root,
where it is least effective ; an arrangement calculated to reduce the number
of muscular contractions, while it contributes to the actual power of the wing.
This hypothesis, it will be observed, guarantees to the wing a very high speed,
with comparatively few reversals and comparatively few muscular contractions.

In the bat and bird the wings do not vibrate with the same rapidity as m

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 401

the insect, and this is accounted for by the circumstance, that in them the
muscles do not act exclusively at the root of the wing. In the bird and bat
the muscles run along the wing towards the tip for the purpose of flexing or
folding the wing prior to the up stroke, and for opening out or expanding it
prior to the down stroke.

As the wing must be folded or flexed and opened out or expanded every
time the wing rises and falls, and as the muscles producing flexion and extension
are long muscles with long tendons, which act at long distances as long levers,
and comparatively slowly, it follows that the great short muscles (pectorals, &c.)
situated at the root of the wing must act slowly likewise, as the muscles of the
thorax and wing of necessity act together to produce one pulsation or vibration
of the wing. What the wing of the bat and bird loses‘in speed it gains in
power, the muscles of the bird and bat’s wing acting directly upon the points
to be moved, and under the most favourable conditions. In the insect, on the
contrary, the muscles act indirectly, and consequently at a disadvantage. If the
pectorals only acted, they would act as short levers, and confer on the wing of
the bat and bird the rapidity peculiar to the wing of the insect. The tones
produced by the bird’s wing would in this case be heightened. The swan in
flying produces a loud whistling sound, and the pheasant, partridge, and grouse
a sharp whirring noise like the stone of a knife-grinder.

It is a mistake to suppose, as many do, that the tone or note produced by
the wing during its vibrations is a true indication of the number of beats made
by it in any given time. This will be at once understood, when I state that a
long wing will produce a higher note than a shorter one driven at the same
speed and having the same superficial area, from the fact that the tip and body
of the long wing will move through a greater space in a given time than the
tip and body of the shorter wing. This is occasioned by all wings being jointed
at their roots, the sweep made by the different parts of the wing in a given time
being longer or shorter in proportion to the length of the pinion. It ought,
moreover, not to be overlooked that in insects the notes produced are not
always referrible to the action of the wings, these, in many cases, being trace-
able to movements induced in the legs and other parts of the body.

It is a curious circumstance that if portions be removed from the posterior
margins of the wings of a buzzing insect, such as the wasp, bee, blue-bottle fly,
&c., the note produced by the vibration of the pinions is raised in pitch. This
is explained by the fact that an insect, whose wings are curtailed, requires to
drive them at a much higher speed in order to sustain itself in the air. That
the velocity at which the wing is urged is instrumental in causing the sound, is
proved by the fact that in slow flying insects and birds no note is produced ;
whereas in those which urge the wing at a high speed, a note is elicited which
corresponds within certain limits to the number of vibrations and the form of

402 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

the wing. It is the posterior or thin flexible margin of the wing which is more
especially engaged in producing the sound, and if this be removed, or if this
portion of the wing, as is the case in the bat and owl, be constructed of very
soft materials, the character of the note is altered. An artificial wing, if pro-
perly constructed and impelled at a sufficiently high speed, emits a drumming
noise, which closely resembles the note produced by the vibration of short-
winged, heavy-bodied birds, all which goes to prove that sound is a concomitant
of rapidly vibrating wings.

ARTIFICIAL FLIGHT.

The subject of artificial flight, notwithstanding the large share of attention
bestowed upon it, has been particularly barren of results. This is the more to
be regretted, as the interest which has been taken in it from early Greek and
Roman times has been universal. The unsatisfactory state of the question is
to be traced to a variety of causes, the most prominent of which are—

1st, The extreme difficulty of the problem.

2d, The incapacity or theoretical tendencies of those who have devoted
themselves to its elucidation.

3d, The great rapidity with which wings, especially insect wings, vibrate,
and the difficulty experienced in analysing their movements.

4th, The great weight of all flying things when compared with a correspond-
ing volume of air.

5th, The discovery of the balloon, which has retarded the science of aérosta-
tion, by misleading men’s minds and causing them to look for a solution of the
problem by the aid of a machine lighter than the air, and which has no analogue
in nature. Flight has been unusually unfortunate in its votaries. It has been
cultivated by profound thinkers, especially mathematicians, who have worked
out innumerable theorems, but who, it would appear, never bethought them of
verifying their results by experiment; and by uneducated charlatans who,
despising the abstractions of science, have made the most ridiculous attempts —
at a practical solution of the problem. Thus bandied about, artificial flight has
become the idol of a few and the jest of the many. The term has been employed,
on the one hand, to represent the highest soarings of the human mind, and on
the other, to typify the extinction or aberration of intellect, the word flighty
signifying whatever is utopian or foolish.

Flight, as the question stands at present, may be divided into two principal
varieties which represent two great sects or schools—

1st, The Balloonists, or those who advocate the employment of a machine
specifically lighter than the air.

2d, Those who believe that weight is necessary to flight.

The second school may be subdivided into (a) those who advocate the

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 403

employment of rigid inclined planes driven forward in a straight line, or revolving
planes (aérial screws); and (>) such as trust for elevation and propulsion
to the vertical flapping of wings.

Balloon.—The balloon, as all are aware, is constructed on the obvious prin-
ciple that a machine lighter than the air must necessarily rise through it. The
MonrTco.rier brothers invented such a machine in 1782. Their balloon con-
sisted of a paper globe or cylinder, the motor power being super-heated air
supplied by the burning of vine twigs under it. The Montgolfier or fire
balloons, as they were called, were superseded by the hydrogen gas balloon of
MM. Cuartes and Rosert, this being in turn supplanted by the ordinary gas
balloon of Mr Grern. Since the introduction of coal gas in the place of
hydrogen gas, no radical improvement has been effected, all attempts at guiding
balloons having signally failed. This arises from the vast extent of surface
which they necessarily present, rendering them a fair conquest to every breeze
that blows, and because the power which animates them is a mere lifting power
which, in the absence of wind, must act in a vertical line, all other motion being
extraneous and foreign to it. It consequently rises through the air in opposi-
tion to the law of gravity, very much as a dead bird falls in a downward
direction in accordance with it. Having no hold upon the air, this cannot be
employed as a fulcrum for regulating its movements, and hence the cardinal
difficulty of ballooning as an art.

Finding that no marked improvement has been made in the balloon since
its introduction in 1782, the more advanced thinkers have within the last
quarter of a century turned their attention in an opposite direction, and have
come to regard flying creatures, all of which are much heavier than the air, as
the true models for flying machines. An old doctrine is more readily assailed
than uprooted, and accordingly we find the followers of the new faith met by
the assertion that insects and birds have large air cavities in their interior, that
those cavities contain heated air, and this heated air in some mysterious manner
contributes to, if it does not actually produce, flight. No argument could be
more fallacious. To render a flying creature buoyant by means of air-cells, it
would require to have its superficial area increased a thousand fold (would, in
fact, require to be converted into a balloon); and, besides, many admirable fliers,
such as the bats, have no air-cells, while many birds, the apteryx for example,
and many animals never intended to fly, such as the orang-outang and a large
number of fishes, are provided with them. It may therefore be reasonably
concluded that flight is in no way connected with air-cells, and the best proof
that can be adduced is to be found in the fact that it can be performed to
perfection in their absence.

The Inclined Plane.—The modern school of flying is in some respects quite
as irrational as the ballooning school.

VOL, XXVI. PART II. OM

404 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

The favourite idea with most is the wedging forward of an inclined plane
upon the air by means of a “wis a tergo.”

The inclined plane may be made to advance in a horizontal line or made
to rotate in the form of a screw. Both plans have their adherents. The one
recommends a large supporting area extending on either side of the weight to
be elevated, the surface of the supporting area making an all but inappreciable
angle with the horizon, the whole being wedged forward by the action of vertical
screw propellers. This was the plan suggested by HENSon and STRINGFELLOW,
and partly carried out by the latter.

WENHAM* has advocated the employment of superimposed planes, with a
view to augmenting the support furnished while it diminishes the horizontal
space occupied by the planes. These planes WENHAM designates Acroplanes.
They are inclined at a very slight angle to the horizon, and are wedged forward
either by the weight to be elevated or by the employment of vertical screws.
WENHAM’ plan was adopted by STRINGFELLowt in a model which he exhibited
at the Aéronautical Society’s Exhibition, held at the Crystal Palace in the
summer of 1868.

The subjoined woodcut (fig. 52), taken from a photograph, gives a very good

S ead
| @]
AS , SZ

idea of the model in question, a 6 ¢ representing the superimposed planes, d the
tail, and e / the vertical screw propellers.

The superimposed planes (a 6 ¢) in this machine contained a sustaining area
of 28 square feet in addition to the tail (d). :

Its engine represented a third of a horse power, and the weight of the whole
(engine, boiler, water, fuel, superimposed planes, and propellers) was under 12
‘ Ibs. Its sustaining area, if that of the tail (d) be included, was something like
36 square feet, z.¢., 3 square feet for every pound—the sustaining area of the
gannet (p. 385), it will be remembered, being less than one foot of wing for
every two pounds of body.

* On Aérial Locomotion, by F. H. Wenuam, Esq., World of Science for June, 1867.
+ Flying Machines, by F. W. Bruary, Esq., Popular Science Review for January, 1869.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 405

The model was forced by its propellers along a wire at a great speed, but, so
faras I could determine, failed to lift itself notwithstanding its extreme lightness
and the comparatively very great power employed.*

Mr Henson’st aérial machine was very similar in principle to Mr
STRINGFELLOW’S. “The chief feature of the invention was the very great
expanse of its sustaining planes, which were larger in proportion to the weight

‘it had to carry than those of many birds. The machine advanced with its front
edge a little raised, the effect of which was to present its under surface to the
air over which it passed, the resistance of which, acting upon it like a strong
wind on the sails of a windmill, prevented the descent of the machine and its
burden. The sustaining of the whole, therefore, depended upon the speed at which
it travelled through the air, and the angle at which its under surface impinged on
the air in its front. . . . The machine, fully prepared for flight, was started
from the top of an inclined plane, in descending which it attained a velocity
necessary to sustain it in its further progress. That velocity would be gradually
destroyed by the resistance of the air to the forward flight ; it was, therefore,
the office of the steam engine and the vanes it actuated simply to repair the loss
of velocity; it was made therefore only of the power and weight necessary for that
small effect. ” The editor of “ Newton’s Journal of Arts and Science”
speaks of it thus—‘ The apparatus consists of a car containing the goods, passen-
gers, engines, fuel, &c., to which a rectangular frame, made of wood or bamboo
cane, and covered with canvas or oiled silk, is attached. This frame extends on
either side of the car in a similar manner to the outstretched wings of a bird; but
with this difference, that the frameis immovable. Behind the wings are two vertical
fan wheels, furnished with oblique vanes, which are intended to propel the
apparatus through the air. The rainbow-lke circular wheels are the propellers,
answering to the wheels of a steam-boat, and acting upon the air after the
manner of a windmill. These wheels receive motion from bands and pulleys
from a steam or other engine contained in the car. To an axis at the stern of
the car a triangular frame is attached, resembling the tail of a bird, which is
also covered with canvas or oiled silk. This may be expanded or contracted
at pleasure, and is moved up and down for the purpose of causing the machine
to ascend or descend. Beneath the tail is a rudder for directing the course of
the machine to the right or to the left ; and to facilitate the steering a sail is
stretched between two masts which rise from the car. The amount of canvass
or oiled silk necessary for buoying up the machine is stated to be equal to one
square foot for each half pound of weight.{

* Mr Srriverstiow stated that his machine occasionally left the wire, and was sustained by its
superimposed planes alone.

+ Mr Henson designed his aérostat in 1843.

} Astra Castra, by Harron Turner, Esq. London, 1865, pages 311 and 312.

7 7

406 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

The idea embodied by HENSON, STRINGFELLOW, and WENHAM is plainly that
of a boy’s kite sailing upon the wind. The kite, however, is a more perfect
flying apparatus than that furnished by HEnson, SrrINGrELLow, and WENHAM,
inasmuch as the inclined plane formed by its body strikes the air at various
angles—the angles varying according to the length of string, strength of breeze,
length and weight of tail, &c. HENson’s, STRINGFELLOW’s, and WENHAM’S
methods, although carefully tried, have hitherto failed. The objections are
numerous. In the first place, the supporting planes (aéroplanes or otherwise)
are rigid. This is a point to which I wish particularly to direct attention.
Second, They stroke the air at a given angle. Here again, there is a departure
from nature. Third, A machine so constructed must be precipitated from a
height or driven along the surface of the land or water at a high-speed to supply
it with initial velocity. Fourth, It is unfitted for flying with the wind unless its
speed greatly exceeds that of the wind. Fifth, It would have considerable
difficulty in flying across the wind, and considerable risk would be incurred in
landing because of the velocity attained. Sixth, The sustaining surfaces are
comparatively very large. They are, moreover, passive or dead surfaces, 7.¢.,
they have no power of moving or accommodating themselves to altered circum-
stances. In this respect they somewhat resemble the surfaces presented by a
balloon—their great extent rendering them liable to be seized and tossed by the
wind.

The Aérial Screw.—Our countryman, Sir GrorGE CAyLey, gave the first
practical illustration of the efficacy of the screw as applied to the air in 1796.
In that year he constructed a small machine consisting of two screws made of
quill feathers. The screws were each composed of four feathers stuck in a piece
of cork, the corks being drilled in the centre to receive a driving shaft or axis.
To the shaft a whalebone spring, with a string which coiled round the shaft (and
by which the spring was wound up), was affixed. By turning the upper screw
(the lower one being secured) a sufficient number of times, the proper degree of
tension was conferred on the spring; and the instant the apparatus was
liberated it flew into the air. CAyYLEy’s screws were peculiar, inasmuch as they
were superimposed and rotated in opposite directions. He estimated that if
the area of the screws was increased to 200 square feet, and moved by a man,
they would elevate him. CAyLey’s interesting experiment is described at length,
and the apparatus figured in ‘ Nicholson’s Journal” for 1809, p. 175. In 1842
Mr PuItuirs also succeeded in elevating a model by means of revolving fans. Mr
PuHILiies’s model was made entirely of metal, and when complete and charged
weighed 2 lbs. It consisted of a boiler or steam generator and four fans
supported between eight arms. The fans were inclined to the horizon at an
angle of 20°, and through the arms the steam rushed on the principle discovered
by Hero of Alexandria. By the escape of steam from the arms, the fans were

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 407

made to revolve with immense energy, so much so that the model rose to a great
altitude, and flew across two fields before it alighted. The motive power
employed in the present instance was obtained from the combustion of charcoal,
nitre, and gypsum, as used in the original fire annihilator, the products of
combustion mixing with water in the boiler, and forming gas charged steam,
which was delivered at a high pressure from the extremities of the eight arms.
This model is remarkable as being probably the first which actuated by steam
has flown to any considerable distance.* The French have espoused the aérial
screw with great enthusiasm, and within the last few years (1863) M. M. Napar,t
DE PontTIN D’AMECOURT, and DE LA LANDELLE have constructed clockwork
models (orthopteres), which not only raise themselves into the air, but carry a

Fig. 53. Flying Machine designed by M. pz LA LANDELLE.

certain amount of freight. These models are exceedingly fragile, and because of
the prodigious force required to propel them usually break after a few trials.
The above woodcut (figure 53) embodies M. pE LA LANDELLE’s ideas.

* Report on the First Exhibition of the Aéronautical Societ of Great Britai
Palace, London, in June 1868, page 10. cig sadeneiialain
. + Mons. Napar, in a paper written in 1863, enters very fully into the subject of artificial
fight, as performed by the aid of the screw. Liberal extracts are given from Napar’s paper in
Astra Castra,” by Captain Harton Turner. London, 1865, page 340. To Turnur’s handsome
volume the reader is referred for much curious and interesting information on the subject of
Aérostation,

VOL. XXVI. PART II. 5N

408 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

In the helecopteric models made by M. M. Napar, Pontin p’AmEcoURT, and
DE LA LANDELLE, the screws (mnopqr st, figure 53, p. 407) are arranged in
tiers, 7.¢., the one screw is placed above the other. In this respect they resemble
the aéro-planes recommended by Mr WenuAM, and tested. by Mr StTRINGFELLOw,
(compare mnopgqrstof fig. 53. p. 407, with a b ¢ of fig. 52, p. 404). The
superimposed screws, as already explained, were first figured and described
by Sir Grorce CaAyLey. The French screws, and that employed by Mr
PuHILuirs, are rigid or unyielding, and strike the air at a given angle, and herein,
I believe, consist their principal defect. This arrangement results in a ruinous
expenditure of power, and is accompanied by a great amount of slip. The
aérial screw, and the machine to be elevated by it, can be set in motion without
a preliminary run, and in this respect it has the advantage over the machine
supported by sustaining planes. It has, in fact, a certain amount of inherent
motion, its sustaining surfaces being active or moving surfaces. It is accordingly
more independent than the machine designed by HENSON, STRINGFELLOW, and
WENHAM.

I may observe with regard to the system of rigid inclined planes wedged
forward at a given angle in a line or in a circle, that it does not embody the
principle carried out in nature.

The wing of a flying creature, as I have taken pains to show, is not rigid; —
neither does it always attack the air at one angle. On the contrary, it is capable
of moving in all its parts, and attacks the air at an injinite variety of angles.
Above all, the surface exposed by a natural wing, when compared with the
great weight it is capable of elevating, is remarkably small. This is accounted
for by the length and the great range of motion of natural wings, the latter
enabling the wings to convert large tracts of air into supporting areas. It is
also accounted for by the multiplicity of the movements of natural wings, these
enabling the pinions to create and rise upon currents of their own forming, and
to select and utilise existing currents.

If any one watches an insect, a bat, or a bird when dressing its wings, he
will observe that it can incline the under surface of the wing at a great variety
of angles to the horizon. This it does by causing the wing to rotate along its
anterior or thick margin, or by twisting the posterior or thin yieldmg margin
around the anterior or thick margin. As a result of this movement, the two
margins are forced into double and opposite curves, and the wing converted
into a plastic helix or screw. We will further observe that the bat and bird, and
some insects, have, in addition, the power of folding and drawing the wing
towards the body during the up stroke, and of pushing it away from the body
and extending it during the down stroke, so as alternately to diminish and
increase its area, arrangements necessary to decrease the amount of resistance
experienced by the wing during its ascent, and increase it during its descent. It

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 409

is scarcely necessary to add, that in the aéro-planes and aérial screws, as at
present constructed, no provision whatever is made for suddenly increasing or
diminishing the sustaining area, of conferring elasticity upon it, or of giving to the
supporting surfaces that infinite variety of angles which would enable them to seize
and disentangle themselves from the air with rapidity. Many investigators are
of opinion that flight is a question of mere levity and power, and that if a
machine could only be made light enough and powerful enough, it must of
necessity fly, whatever the nature of its flying surfaces. A grave fallacy lurks
here. Birds are not more powerful than quadrupeds of equal size, and Strinc-
FELLOW’S machine, which, as we have seen, only weighed 12 /bs., exerted one-third
of a horse power. The probabilities therefore, are, that flight is dependent to a
ereat extent on the nature of the flying surfaces, and the mode of applying those
surfaces to the air.

Artyicial Wings (BorELLI’s Views).—With regard to the production of fight
by the flapping of wings, much may and has been said. Of all the methods yet
proposed, it is unquestionably by far the most ancient. Discrediting as apocry-
phal the famous story of Dapatus and his waxen wings, we certainly have a
very graphic account of artificial wings in the “De Motu Animalium” of
BorRE.uI, published as far back as 1680, 7.¢., nearly two centuries ago.*

Indeed it will not be too much to affirm, that to this distinguished physiologist
and mathematician belongs almost all the knowledge we at present possess of
artificial wings and their actions. He was well acquainted with the properties
of the wedge, as applied to flight, and he was likewise cognisant of the flexible
and elastic properties of the wing. To him is to be traced the purely mechanical
theory of the wing’s action. He figured a bird
with artificial wings, each wing consisting of a rigid
rod in front and flexible feathers behind. I have
thought fit to reproduce Boretwi’s figure, both be-
cause of its great antiquity, and because it is emi-
nently illustrative of his text.t

The wings, as a reference to fig. 54 will show,
are represented as striking vertically downwards

(gh). They remarkably accord with those describ-
_ ed by Straus-DurckHEmM, GirarpD, and quite recently by Professor Marry.{

BorE11! was of opinion that flight resulted from the application of an inclined
plane, which beats the air, and which has a wedge action. He, in fact, endeavours
to prove that a bird wedges itself forward upon the air by the perpendicular

* Borevut. De Motu Animalium. Sm. 4to. 2 vols. Rome 1680.

+ “De Motu Animalium,” Lugduni Batavorum apud Petrum Vander. Anno mpcnxxxy. Tab.
XIII. figure 2. (New edition.)

+ Revue des Cours Scientifiques de la France et de lEtranger. Mars 1869.

410 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

vibration of its wings, the wings during their action forming a wedge, the base
of which (cde) is directed towards the head of the bird, the apex (a/) being
directed towards the tail (d). This idea is worked out in propositions 195 and
196 of the first part of BorELuI’s book. In proposition 195 he explains how,
if a wedge be driven into a body, the wedge will tend to separate that body into
two portions ; but that if the two portions of the body be permitted to react
upon the wedge, they will communicate oblique impulses to the sides of the
wedge, and expel it, base first, in a straight line.

Following up the analogy, BoreLu endeavours to show in his 196th pro-
position, “that if the air acts obliquely upon the wings, or the wings obliquely
upon the air (which is, of course, a wedge action), the result will be a horizontal
transference of the body of the bird.” In the proposition referred to (196)
BorELLI states—“ If the expanded wings of a bird suspended in the air shall
strike the undisturbed air beneath it with a motion perpendicular to the horizon,
the bird will fly with a transverse motion in a plane parallel with the horizon.”
In other words, if the wings strike vertically downwards, the bird will fly horizon-
tally forwards. He bases his argument upon the belief that the anterior
margins of the wings are rigid and unyielding, whereas the posterior and after
parts of the wings are more or less flexible, and readily give way under pres-
sure. If, he adds, the wings of the bird be expanded, and the under surfaces
of the wings be struck by the air ascending perpendicularly to the horizon, with
such a force as shall prevent the bird gliding downwards (2.¢., with a tendency
to glide downwards) from falling, it will be urged im a horizontal direction.
This follows because, in BoRELLI’s opinion, the two osseous rods (vi7g@) forming
the anterior margins of the wings resist the upward pressure of the air, and so
retain their original form (literally extent or expansion), whereas the flexible after
parts of the wings (posterior margins) are pushed up and approximated to form a
cone, the apex of which (vide af of figure 54, p. 409) is directed towards the
tail of the bird. In virtue of the air playing upon and compressing the sides of
the wedge formed by the wings, the wedge is driven forwards in the direction of
its base (c, b, e), which is equivalent to saying that the wings carry the body of
the bird to which they are attached in a horizontal direction.” Bore ut restates
the same argument in different words, as follows :—

“Tf,” he says, “the air under the wings be struck by the flexible portions of
the wings (/labella, literally fly flaps or small fans) with a motion perpendicular
to the horizon, the sails (vela) and flexible portions of the wings (/labella) will
yield in an upward direction, and form a wedge, the point of which is directed
towards the tail. Whether, therefore, the air strikes the wings from below, or
the wings strike the air from above, the result is the same—the posterior or
flexible margins of the wings yield in an upward direction, and in so doing urge
the bird in a horizontal direction.”

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 411

In his 197th proposition, Bore follows up and amplifies the arguments
contained in propositions 195 and 196. Thus, he observes, “It is evident that
the object of flight is to impel birds upwards, and keep them suspended in the
air, and also to enable them to wheel round in a plane parallel to the horizon.
The first (or upward flight) could not be accomplished unless the bird were
impelled upwards by frequent leaps or vibrations of the wings, and its descent
prevented. And because the downward tendency of heavy bodies is perpen-
dicular to the horizon, the vibration of the plain surfaces of the wings must be
made by striking the air beneath them im a direction perpendicular to the
horizon, and in this manner nature produces the suspension of birds in the air.

With regard to the second or transverse motion of birds (7.e., horizontal
flight) some authors have strangely blundered ; for they hold that it is like that
of boats, which, being impelled by oars, moved horizontally in the direction of
the stern, and pressing on the resisting water behind, leaps with a contrary
motion, and so are carried forward. In the same manner, say they, the wings
vibrate towards the tail with a horizontal motion, and likewise strike against
the undisturbed air, by the resistance of which they are moved forward by a
reflex motion. But this is contrary to the evidence of our sight as well as to
reason ; for we see that the larger kinds of birds, such as swans, geese, &c.,
never vibrate their wings, when flying, towards the tail with a horizontal
motion like that of oars, but always bend them downwards, and so describe
circles raised perpendicularly to the horizon.*

Besides, in boats the horizontal motion of the oars is easily made, and a
perpendicular stroke on the water would be perfectly useless, inasmuch as their
descent would be impeded by the density of the water. But in birds such a
horizontal motion (which indeed would rather hinder flight) would be absurd,
since it would cause the ponderous bird to fall headlong to the earth ; whereas
it can only be suspended in the air by constant vibration of the wings perpen-
dicular to the horizon. Nature was thus forced to show her marvellous skill
im producing a motion which, by one and the same action, should suspend the
bird in the air, and carry it forward in a horizontal direction. This is effected
by striking the air below perpendicularly to the horizon, but with oblique
strokes—an action which is rendered possible only by the flexibility of the
feathers, for the fans of the wings in the act of striking acquire the form of a
wedge, by the forcing out of which, the bird is necessarily moved forwards in a
horizontal direction.”

The points which BorEtii endeavours to establish are these :—

First, That the action of the wing is a wedge action.

* It is clear from the above that Bornrxt did not know that the wings of birds strike forwards
as well as downwards during the down stroke. He seems to have been equally ignorant of the fact
that the wings of insects vibrate in a more or less horizontal direction.

VOL. XXXVI. PART IT. 50

412 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

Second, That the wing consists of two portions—a rigid anterior portion,
and a non-rigid flexible portion. The rigid portion he represents in his artificial
bird (figure 54, page 409) as consisting of a rod (e,r), the yielding portion of
Seathers (a, 0).

Third, That if the air strikes the under surface of the wing perpendicularly
in a direction from below upwards, the flexible portion of the wing will yield in
an upward direction, and form a wedge with its neighbour.

Fourth, Similarly and conversely, if the wing strikes the air perpendicularly
from above, the posterior and flexible portion of the wing will yield and be forced
in an upward dirction.

Fifth, That this upward yielding of the posterior or flexible margin of the
wing results in and necessitates a horizontal transference of the body of the bird.

Sixth, That to sustain a bird in the air the wings must strike vertically
downwards, as this is the direction in which a heavy body, if left to itself, would
fall.

Seventh, That to propel the bird in a horizontal direction, the wings must
descend in a perpendicular direction, and the posterior or flexible portions of
the wings yield in an upward direction, and in such a manner as virtually
to communicate an oblique action to them.

Eighth, That the feathers of the wing are bent in an upward direction when
the wing descends, the upward bending of the elastic feathers contributing to
the horizontal travel of the body of the bird.

I have been careful to expound Bore 11's views for several reasons :—

1st, Because the purely mechanical theory of the wing’s action is to be
traced to him.

2d, Because his doctrines have remained unquestioned for nearly two
centuries, and have been adopted by all the writers since his time, without, I
regret to say in the great majority of cases, any acknowledgment whatever.

3d, Because his views have been revived by the modern French school, and

4th, Because in commenting upon and differmg from BoreE.it I will neces-
sarily comment upon and differ from all his successors.

The Duke of ARGYLL agrees with Bore ui in believmg that the wing
invariably strikes perpendicularly downwards. His words are—-‘‘ Except
for the purpose of arresting their flight birds can never strike except directly
downwards ; that is, against the opposing force of gravity.” Professor OWEN
in his “ Comparative Anatomy,” Mr M‘Griitvray in his “ British Birds,” Mr
Brsuop in his article Motion in the “ Cyclopedia of Anatomy and Physiology,”
and M. Liars “on the flight of birds and insects” in the “ Annals of Natural
History,” all assert that the stroke is delivered downwards and more or less
backwards. To obtain an upward recoil, one would naturally think all that is
required is a downward stroke, and to obtain an upward and forward recoil,

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 413

one would naturally conclude a downward and backward stroke alone is requisite.
This reasoning is true of water and wings, but it is not true of air and wings.

In the first place, a natural wing, or a properly constructed artificial one,
cannot be depressed either vertically downwards, or downwards and backwards.
It will of necessity descend downwards and forwards in a curve. This arises
from its being flexible and elastic throughout, and in especial from its being
carefully graduated as regards thickness, the tip being thinner and more elastic
than the root, and the posterior margin than the anterior margin.

In the second place, there is only one direction in which the wing could
strike so as at once to support and carry the bird forward. The bird, when
flying, is a body in motion. It has therefore acquired momentum. Ifa grouse
is shot on the wing 7¢ does not fall vertically downwards, as BoRELLI and his
successors assume, but downwards and forwards. The flat surfaces of the
wings are consequently made to strike downwards and forwards, as they in this
manner act as kites to the falling body, which they bear, or tend to bear, wpwards
and forwards. So much for the direction of the stroke during the descent of
the wing. Let us now consider to what extent the posterior margin of the wing
yields in an upward direction when the wing descends. BorELLI does not
state the exact amount. The Duke of ArGyLi, who agrees with Bore.
that the posterior margin of the wing is elevated during the down stroke, avers
that, whereas the air compressed in the hollow of the wing cannot pass through
the wing owing to the closing upwards of the feathers against each other, or
escape forwards because of the rigidity of the bones and of the quills in this
direction, it passes backwards, and in so doing lifts by its force the elastic ends
of the feathers. In passing backwards it communicates to the whole line of
both wings a corresponding push forwards to the body of the bird. The same
volume of air is thus made, in accordance with the law of action and reaction,
to sustain the bird and carry it forward.* Mr M‘GiItiivray observes that “to
progress in a horizontal direction it is necessary that the downward stroke
should be modified by the elevation in a certain degree of the free extremities of
the quills.t

Marey’s Views.—Professor Marey states that during the down stroke the
posterior or flexible margin of the wing yields in an upward direction, to such
an extent as to cause the under surface of the wing to look backwards, and make
a backward angle with the horizon of 45° plus or minus according to circum-
stances.{ That the posterior margin of the wing yields in a slightly upward
direction during the down stroke to prevent shock, I admit. The amount of

* Reign of Law. “Good Words,” February 1865, p. 128.

+ History of British Birds. Lond. 1837, p. 43.

{ Méchanisme du vol chez les insectes. Comment se fait la propulsion, by Professor E. J. Marry.
Revue des Cours Scientifiques de la France et de l’Etranger for 20th March 1869, p. 254.

414 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

yielding is, however, in all cases very slight, and the little upward movement
there is, is in part the result of the posterior margin of the wing rotating
around the anterior margin as an axis. That the posterior margin of the wing —
never yields in an upward direction until the under surface of the pinion makes
a backward angle of 45° with the horizon, as MArery remarks, is a matter of
absolute certainty. This statement admits of direct proof. If any one watches
the horizontal or upward flight of a large bird, he will observe that the pos-
terior or flexible margin of the wing never rises during the down stroke to a
perceptible extent, so that the under surface of the wing never looks backwards.
On the contrary, he will find that the under surface of the wing (during the
down stroke) invariably looks forwards—the posterior margin of the wing being”
inclined downwards and backwards, the anterior one upwards and forwards, as —
shown atcdef, jkim of fig. 15, page 345; h7 of fig. 38, page 361; 1, 2,357
4,5, 6 of figs. 18 and 19, Plate XIV. ; and q po of figs. 14 and 15, Plate XIII.

The under surface of the wing, as will be seen from this account, not only
looks forwards, but it forms a true kite with the horizon, the angles made by
the kite varying at every part of the down stroke, as shown more particularly
at c,d,e,f; j, k,l, m of fig. 15, page 345.

Professor Margy goes on to state that not only does the posterior margin of
the wing yield ix an upward direction during the down stroke until the under —
surface of the pinion makes a backward angle of 45° with the horizon (page
415, fig. 55, ac; ab), but that during the up stroke it yields to the same extent ~
in an opposite direction (xd; ab). The posterior flexible margin of the wing,
according to Margy, thus passes through a space of 90° every time the wing”
reverses its course, this space being dedicated to the mere adjusting of the
planes of the wing for the purposes of flight. The planes, moreover, he
asserts, are adjusted not by vital and vito-mechanical acts but by the action of
the air alone; this operating on the under surface of the wing and forcing its
posterior margin upwards during the down stroke; the air during the up stroke
acting upon the posterior margin of the upper surface of the wing, which it
forces downwards. Marry thus delegates to the air, the difficult and delicate
task of arranging the details of flight. The time, power, and space occupied”
in reversing the wing alone, according to this view, are such as to render flight
impossible. That the wing does not act as stated by Margy, may be readily
proved by experiment. It may also be proved diagrammatically, as a reference .
to fig. 55, page 415, will show. a

Let a, b of fig 55 represent the horizon; m, n the line of vibration ; 2, ¢ the
wing inclined at an upward backward angle of 45° in the act of making the —
down stroke, and a, d the wing inclined at a downward backward angle of 45° |
and in the act of making the up stroke. When the wing a ¢ descends it will tend
to dive downwards in the direction / (giving very little of any horizontal sup-

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 415

port) ; when the wing 2 d ascends it will endeavour to rise in the direction g,
as it darts up like a kite (the body bearing it being in motion). If we take the
resultant of these two forces, we have at most propulsion in the direction a d.
This, moreover, would only hold true if

the bird was as light as air. As, how- a

ever, gravity tends to pull the bird down-

wards as it advances, the real flight of ;

the bird, according to this explanation, j <-—-—_-_- Y LSE ee
would fall in a line between 6 and f ee iy
probably in ah. It could not possibly i; LR BSA

be otherwise ; the wing described and Wh

figured by Margy is in one piece, and a

vibrated vertically on either side of a Fig. 55.

given line. If, however, a wing in one piece is elevated and depressed ina
strictly perpendicular direction, it is evident that the wing will experience a
greater resistance during the up stroke, when it is acting against gravity, than
during the down stroke, when it is acting with gravity. As a consequence, the
bird will be more vigorously depressed during the ascent of the wing than it
will be elevated during its descent. That the mechanical wing referred to
by Marey is not a flying wing, but a mere propelling apparatus, seems evident
to himself, for he states that “the winged machine designed by him has un-
questionably not motor power enough to support its own weight.”*

The manner in which the natural wing (and the artificial wing properly con-
structed and propelled) evades the resistance of the air during the up stroke,
and gives continuous support and propulsion, is very remarkable. Fig. 56, page
416, willillustrate the principle. Let a) represent the horizon ; m7 the direction
of vibration ; # s the wing ready to make the down stroke, and #¢ the wing ready
to make the up stroke. When the wing 2s descends, the posterior margin (s)
is screwed downwards and forwards in the direction s,¢, the forward angle
which it makes with the horizon increasing as the wing is lowered. The air is
thus seized by a great variety of inclined surfaces, and as the under surface of
the wing, which is a true kite, looks upwards and forwards, it tends to carry the
body of the bird upwards and forwards in the direction x w. When the wing
a,t makes the up stroke, it rotates from below upwards to prepare for the second
down stroke. The wing does not, however, ascend in the direction ¢, s.. On the
contrary, it darts up like a true kite, which it is, in the direction 2, v, in virtue of
the reaction of the air, and because the body of the bird, to which it is attached,
had a forward motion communicated to it by the wing during the down stroke.
The resultant of the forces acting in the lines 2v and #8, is one acting in the

* Revue des Cours Scientifiques de la France et de lEtranger. 8vo. March 20, 1869.
VOL. XXVI. PART II. a

416 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

direction xw, and if allowance be made for the operation of gravity, the
flight of the bird will fall somewhere between w and 6, probably in the line a, r.
This arises from the wing acting as an eccentric—from the upper concave sur-
face of the pinion being always directed
upwards, the under concave surface
downwards—from the under surface,
which is a true kite, darting forwards in
wave curves both during the down and
up strokes, and never making a back-
! ward angle with the horizon ; and lastly,
rule from the wing employing the air under

it as a fulcrum during the down stroke
the air, on its part, reacting on the under
surface of the pinion, and when the proper time arrives, contributing to the
elevation of the wing.

If, as Boretir and his successors believe, the posterior margin of the wing
yielded to any marked extent in an upward direction during the down stroke,
and more especially if it yielded to such an extent as to cause the under surface
of the wing to make a backward angle with the horizon of 45°, one of two things
would inevitably follow—either the air on which the wing depended for support
and propulsion would be permitted to escape before it was utilised, or the wing
would dart rapidly downward, and carry the body of the bird with it. If
the posterior margin of the wing yielded in an upward direction to any
marked extent during the down stroke it would be tantamount to removing
the fulcrum (the air) on which the lever formed by the wing operates. The
wing of the bird, as I have fully explained (see pages from 366 to 384 inclu-
sive), acts as a kite both during the down and up strokes, the ventral aspect
of the kite being always directed forwards (vide from c to m of fig. 15,
page 345).

If a bird flies in a horizontal direction the angles made by the under sur-
face of the wing with the horizon are very slight, but they always look forwards.
If a bird flies upwards the angles in question are increased. In no instance, |
however, unless when the bird is everted and flymg downwards, is the posterior
margin of the wing on a higher level than the anterior one. This holds true of
natural flight, and, consequently, ought to hold true of artificial flight.

With regard to the cone formed, according to Boretu, by the vertical
descent of the two wings, or what, in his opinion, is the same thing, the per-
pendicular ascent of the air, and which is represented at fe ¢ of figure 54, page
409; I think it would be more accurate to state that, instead of the two wings
taken together forming one cone, that each wing by itself forms two cones.

The base of BoreEtu’s cone (e 0 c, figure 54, p. 409), it will be remembered,

——

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 417

is inclined forwards in the direction of the head of the bird—the bases of the
cones formed by each natural wing being, on the contrary, directed outwards
(vide « bd of figure 12, page 342) and backwards (see ¢ pn of same figure).
This arises from the fact that the wing rotates upon two axes (ab and cd of
figure 45, page 376); because it rotates on its root (a of figure 19, Plate XIV.)
to form one cone (af ¢ of same figure), and because, while it is rotating on
its root, it is also rotating along its anterior margin (a0) to form a second
cone, chg. The wing, in forming the cone a fe elevates, and in forming
the cone chg propels. The base of the wedge which furnishes the horizontal
transference is, therefore, turned in the direction of the tail of the bird,
which is just the opposite of what BoreLii maintains, the base of his wedge
being turned in the direction of the head.

BorELLI, and all who have written since his time, are unanimous in affirming
that the horizontal transference of the body of the bird is due to the perpen-
dicular vibration of the wings, and to the yielding of the posterior or flexible
margins of the wings in an upward direction as the wings descend. I am, how-
ever, disposed to attribute it to the fact (1st), that the wings, both when elevated
and depressed, leap forwards in curves, those curves uniting to form a con-
tinuous waved track ; (2d), to the tendency which the body of the bird has to
swing forwards, in a more or less horizontal direction, when once set in motion;
(3d), to the construction of the wings (they are elastic helices or screws, which
twist and untwist while they vibrate, and tend to bear upwards and onwards any
weight suspended from them); (4th), to the reaction of the air on the under
surfaces of the wings; (5th), to the ever-varying power with which the wings
are urged, this being greatest at the beginning of the down stroke, and least at
the end of the up one; (6th), to the contraction of the voluntary muscles and
elastic ligaments, and to the effect produced by the various inclined surfaces
formed by the wings during their oscillations ; (7th), to the weight of the bird
—weight itself, when acting upon wings, becoming a propelling power, and so
contributing to horizontal motion. This is proved by the fact that if a sea
bird launches itself from a cliff with expanded motionless wings, it sails along
for an incredible distance before it reaches the water.

The authors who have adopted Bore.tr’s plan of artificial wing, and who
have indorsed his mechanical views of the wing’s action most fully, are CHABRIER,
STRAUS-DURCKHEIM, GIRARD, and Margy. Bors.u1’s artificial wing, as a

reference to fig 54, page 409, will show, consists of a rigid rod in front, and a

flexible sail, composed of feathers, behind. It acts upon the air, and the air
acts upon it, as occasion demands.

CHasrier’s Views.—CHABRIER states that the wing has only one period of
activity—that, in fact, if the wing be suddenly lowered by the depressor muscles,
it is elevated solely by the reaction of the air. There is one unanswerable objec-

418 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

tion to this theory—the bats and birds, and some, if not all, the insects have
distinct elevator muscles. The presence of well-developed elevator muscles
implies an elevating function ; and, besides, we know that the insect, bat, and
bird can elevate their wings when they are not flying, and when, consequently,
no reaction of the air is induced (pages 364, 365, 395, 396, and 397).

SrrRAvUSs-DURCKHEIM’S Views.—DuRCKHEIM believes that the insect abstracts
from the air by means of the inclined plane a component force (composant)
which it employs to support and direct itself. In his Theology of Nature he
describes a schematic wing as follows :—It consists of a rigid ribbing in front,
and a flexible sail behind. A membrane so constructed will, according to him,
be fit for flight. It will suffice if such a sail elevates and lowers itself successively.
It will, of its own accord, dispose itself as an inclined plane, and receiving
obliquely the reaction of the air, it transfers into tractile force a part of the
vertical impulsion it has received. These two parts of the wing are moreover
equally indispensable to each other. If we compare the schematic wing of
DvRCKHEIM with that of Bore they will be found to be identical, both as
regards their construction and the manner of their application.

Marey’s Views continued.—Professor Margy, so late as 1869, repeats
BorEL.i’s arguments and views with very trifling alterations. Margy describes
two artificial wings, the one composed of a rigid rod and sai/—the rod repre-
senting the stiff anterior margin of the wing; the sail, which is made of paper
bordered with card board, the flexible posterior portion. The other wing
consists of a rigid nervure in front and behind of thin parchment which sup-
ports jine rods of steel. He states, that if the wing only elevates and depresses
itself, “the resistance of the air is sufficient to produce all the other move-
ments. In effect the wing of an insect has not the power of equal resistance
in every part. On the anterior margin the extended nervures make it raged,
while behind it is fine and flexible. During the vigorous depression of the
wing the nervure has the power of remaining rigid, whereas the flexible
portion, being pushed in an upward direction on account of the resistance it
experiences from the air, assumes an oblique position which causes the upper
surface of the wing to look forwards.” ‘The reverse of this takes place during
the elevation of the wing—the resistance of the air from above causing the
upper surface of the wing to look backwards. . . . “At first the plane of
the wing is parallel with the body of the animal. It lowers itself—the /ront
part of the wing strongly resists, the sail which follows it being flexible yields.
Carried by the ribbing (the anterior margin of the wing) which lowers itself, the
sail or posterior margin of the wing being raised meanwhile by the air, which sets
it straight again, the sail will take an intermediate poset and incline itself
about 45° plus or minus according to circumstances.”

“ The wing continues its movements of depression inclined to the horizon, but

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 419

the impulse of the air which continues its effect, and naturally acts upon the
surface which it strikes, has the power of resolving itself into two forces, a
vertical and a horizontal force, the first suffices to raise the animal, the second to
move it along.”*

I have already adverted at considerable length (pages 413, 414, and 415) to
the movements and peculiarities of Professor MArery’s artificial wing, and need
not again return to it. I will only observe, in passing, that it is not a little
curious that BoreELLr’s artificial wing should have been reproduced at a distance
of nearly two centuries.

The Author's Views :—his Method of Constructing and Applying Artificial
Wings as Contradistinguished from that of Borel, CHABRIER, DURCKHEIM,
Marey, &¢.—The artificial wings which I have been in the habit of making for
several years differ from those recommended by Boreiui, DurckHEm, and
Marey in four essential points :—

1st, The mode of construction.

2d, The manner in which they are applied to the air.

3d, The nature of the power employed.

4th, The necessity of adopting certain elastic substances at the root of the
wing if in one piece, and at the root and in the body of the wing if in several
pieces.

And, first, as to the manner of construction.

BoreE.ui, DurcKHEIM, and MArEyY maintain that the anterior margin of the
wing should be rigid; I, on the other hand, believe that no part of the wing
whatever should be rigid, not even the anterior margin, and that the pinion
should be flexible and elastic throughout.

That the anterior margin of the wing should not be composed of a rigid rod

* Compare Marey’s description with that of Boreut, a translation of which I subjoin. ‘ Let
a bird be suspended in the air with its wings expanded, and first let the under surfaces (of the wings)
be struck by the air ascending perpendicularly to the horizon with such a force that the bird gliding
down is prevented from falling: I say that it (the bird) will be impelled with a horizontal forward
motion, because the two osseous rods of the wings are able, owing to the strength of the muscles, and
because of their hardness, to resist the force of the air, and therefore to retain the same form (literally
extent, expansion), but the total breadth of the fan of each wing yields to the impulse of the air when
the flexible feathers are permitted to rotate around the “manubria” or osseous axis, and hence it is
necessary that the extremities of the wings approximate each other: wherefore the wings acquire the
form of a wedge whose point is directed towards the tail of the bird, but whose surfaces are compressed
on either side by the ascending air in such a manner that it is driven out in the direction of its base.
Since, however, the wedge formed by the wings cannot move forward unless it carry the body of the
bird along with it, it is evident that it (the wedge) gives place to the air impelliny it, and therefore
the bird flies forward in a horizontal direction. But now let the substratum of still air be struck
by the fans (feathers) of the wings with a motion perpendicular to the horizon. Since the fans and
sails of the wings acquire the form of a wedge, the point of which is turned towards the tail (of the
bird), and since they suffer the same force and compression from the air, whether the vibrating wings
strike the undisturbed air beneath, or whether, on the other hand, the. expanded wings (the osseous
axis remaining rigid) receive the percussion of the ascending air; in either case the flexible feathers

yield to the impulse, and hence approximate each other, and thus the bird moves in a forward direc-
tion.” —De Motu Animalium, pars prima, prop. 196, 1685.

VOL. XXVI. PART II. 5 Q

420 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

may, I think, be demonstrated in a variety of ways. Ifa rigid rod be made to
vibrate by the hand the vibration is not smooth and continuous; on the con-
trary, it is irregular and jerky, and characterised by two halts or pauses (dead
points), the one occurring at the end of the wp stroke, the other at the end
of the down stroke. This mechanical impediment is followed by serious con-
sequences as far as power and speed are concerned—the slowing of the wing at
the end of the down and up strokes involving a great expenditure of power and a
disastrous waste of time. The wing, to be effective as an elevating and pro-
pelling organ, should have no dead points, and should be characterised by a
rapid winnowing or fanning motion. It should reverse and reciprocate with
the utmost steadiness and smoothness—in fact, the motions should appear as
continuous as those of a fly-wheel in rapid motion : they are so in the insect.

To obviate the difficulty in question, it is necessary, in my opinion, to employ
a tapering elastic rod or series of rods bound together for the anterior margin
of the wing.

If a longitudinal section of bamboo cane, 10 feet in length, and 1 inch in
breadth (vide fig. 57, p. 421), be taken by the extremity and made to vibrate, it
will be found that a wavy serpentine motion is produced, the waves being greatest
when the vibration is slowest (fig. 58, p. 421), and least when it is most rapid (fig.
59, p. 421). It will further be found that at the extremity of the section where
the impulse is communicated there is a steady reciprocating movement devoid of
dead points. The continuous movement in question is no doubt due to the fact
that the different portions of the reed reverse at different periods—the undula-
tions Induced being to an interrupted or vibratory movement very much what —
the continuous play of a fly-wheel is to a rotatory motion.

The Wave Wing of the Author.—If a similar reed has added to it, tapering
rods of whalebone, which radiate in an outward direction to the extent of a foot
or so, and the whalebones be covered by a thin sheet of india-rubber, an artificial
wing, resembling the natural one in all its essential points, is at once produced
(vide fig. 60, p. 421). I propose to designate this wing, from the peculiarities of
its movements, the wave wing (fig. 61, p. 421). If the wing referred to (fig. 61)
be made to vibrate at its root, a series of longitudinal (¢ d e) and transverse
(fg h) waves are at once produced, the one series running in the direction of
the length of the wing, the other in the direction of its breadth (vide p. 330).
This wing further twists and wntwists, figure of 8 fashion, during the down and
up strokes, as shown at figure 62, page 423 (compare with figure 2, p. 336).
There is, moreover, a continuous play of the wing, the down stroke gliding
into the up one, and vice versa, which clearly shows that the down and up
strokes are parts of one whole, and that neither is perfect without the other.

This wing is endowed with the very remarkable property that it will fly m
any direction, demonstrating more or less clearly that flight is essentially a pro-

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 421

gressive movement, 7.¢., a horizontal rather than a vertical movement. Thus, if
the anterior or thick margin of the wing be directed upwards, and the angle
which the under surface of the wing makes with the horizon be something like

a b
Sig se EE BS OE SEE EAS 2 SS Se Se Se SET |

Fig. 61.||

45°, the wing will, when made to vibrate by the hand, fly with an undulating
motion in an upward direction, like a pigeon to its dovecot. If the under sur-
face of the wing makes no angle, or a very small angle, with the horizon, it will

* Fig. 57 represents a longitudinal section of bamboo reed 10 feet long, and 1 inch wide.

+ Fig. 58. The appearance presented by the same reed when made to vibrate by the hand. The reed vibrates on
either side of a given line (x x), and appears as if in two places at the same time, viz., cand f, gandd, eandh,. Itis
thus during its vibration thrown into figures of 8 or opposite curves.

{ Fig. 59. The appearance presented by the same reed when made to vibrate more rapidly. In this case the waves
made by the reed are less in size, but more numerous than in fig. 58. The reed vibrates alternately on either side of the
line # x, being now at 7 now at m, now at 7 now at j, now at & now at 0, now at p now atl. This reed, when made to
vibrate by the hand, has no dead points, a circumstance due to the fact that no two parts of it reverse or change their
curves at precisely the same instant. It is because of this curious reciprocating motion that the wing can seize and dis-
engage itself from the air with such rapidity.

§ Fig. 60. The same reed with a flexible elastic curtain or fringe added to it. The curtain consists of tapering
whalebone rods covered with a thin layer of india-rubber. «@ } anterior margin of wing. ¢ d posterior ditto.

|| Fig. 61 gives the appearance presented by the artificial wing (fig. 60) when made to vibrate by the hand. It is
thrown into longitudinal and transverse waves. The longitudinal waves are represented by the arrows cde, and the
transverse by the arrows fy. A wing constructed on this principle gives a continuous elevating and propelling power.
Tt developes figure of 8 curves during its action in longitudinal, transverse, and oblique directions. It literally floats upon
the air. It has no dead points—is vibrated with amazingly little power, and has apparently no slip. It can fly in an
upward, downward, or horizontal direction by merely altering its angle of inclination to the horizon. It must be
applied to the air by an irregular motion—the movement being most sudden and vigorous always at the beginning of
the down stroke.

422 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

dart forward in a series of curves in a horizontal direction, like a crow in rapid
horizontal flight. Ifthe angle made by the under surface of the wing be reversed,
so that the thick margin of the wing be directed downwards, the wing will
describe a waved track, and jy downwards, as a sparrow from a house-top or
from a tree. In all those movements progression is a necessity. The move-
ments are continuous gliding forward movements. There is no halt or pause
between the strokes, and if the angle which the under surface of the wing makes
with the horizon be properly regulated, the amount of steady tractile and
buoying power developed is truly astonishing. This form of wing, which may
be regarded as the realisation of the figure of 8 theory of flight, elevates and
propels both during the down and up strokes, and its working is accompanied
with almost no slip. It seems literally to float upon the air. No wing that is ~
rigid in the anterior margin can twist and untwist during its action, and produce
the figure of 8 curves generated by the living wing. To produce the curves in
question, the wing must be flexible, elastic, and capable of change of form in
all its parts. The curves made by the artificial wing, as has been stated (p. 420),
are largest when the vibration is slow, and least when it is quick. In like
manner, the air is thrown into large waves by the slow movement of a large
wing, and into small waves by the rapid movement of a smaller wing. The size
of the wing curves and air waves bear a fixed relation to each other, and both
are dependent on the rapidity with which the wing is made to vibrate. This is
proved by the fact that insects, in order to fly, require, as a rule, to drive
their small wings with immense velocity. It is further proved by the
fact that the small humming bird, in order to keep itself stationary before
a flower, requires to oscillate its tiny wings with great rapidity, whereas the
large humming bird (Patagona gigas), as was pointed out by Darwin, can
attain the same object by flapping its large wings with a very slow and powerful
movement. In the larger birds the movements are slowed in proportion to the
size, and more especially in proportion to the length of the wing, the cranes and
vultures moving the wings very leisurely, and the large oceanic birds dispensing
in a great measure with the flapping of the wings, and trusting for progression
and support to the wings in the expanded position.

This leads me to conclude that very large wings may be driven with a com-
paratively slow motion, a matter of some inportance in artificial flight secured
by the flapping of wings.

How to Construct an Artificial Wave Wing on the Insect Type.—The follow-
ing appear to me to be essential features in the construction of an artificial
wing :—

The wing should be of a generally triangular shape.

It should taper from the root towards the tip, and from the anterior margin
in the direction of the posterior margin.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 423

It should be convex above and concave below, and slightly twisted upon
itself.

It should be flexible and elastic throughout, and should twist and untwist
during its vibration, to produce figure of 8 curves along its margins and through-
out its substance.

Such a wing is represented at figure 62.

Tig. 62.*

If the wing is in more than one piece, joints and springs require to be added
to the body of the pinion.

In making a wing in one piece on the model of the insect wing, such as that
shown at figure 62, I employ one or more tapering elastic reeds, which
arch from above downwards (a 6) for the anterior margin. To this I add
tapering elastic reeds, which radiate towards the tip of the wing, and which
also arch from above downwards (g,,7). These latter are so arranged that
they confer a certain amount of spirality upon the wing, the anterior (a 6) and
posterior (cd) margins being arranged in different planes, so that they appear to
cross each other. I then add the covering of the wing, which may consist of
india-rubber, silk, tracing cloth, linen, or any similar substance.

If the wing is large, I employ steel tubes, bent to the proper shape. In
some cases I secure additional strength by adding to the oblique ribs or stays
(gh 7 of figure 62) a series of very oblique stays, and another series of cross
stays, as shown at m and a, n, 0, p, q of fig. 63, page 424.

_ ~ Fig. 62. Elastic spiral wing, which twists and untwists during its action, to form a mobile helix or screw. This
Wing is made to vibrate by a direct piston action, and by a slight adjustment can be propelled vertically, horizontally,
or at any degree of obliquity.

a, b, Anterior margin of wing, to which the neure or ribs are affixed. c, d, Posterior margin of wing crossing
anterior one. 2, Ball and socket joint at root of wing, the wing being attached to the side of the cylinder by the socket.
t, Cylinder. r, 7, Piston, with cross heads (1, w) and piston head (s). 0, 0, Stuffing boxes. e, 7, Driving chains. m,
Superior elastic band, which assists in elevating the wing. 7, Inferior elastic band, which antagonises m. The alternate

stretching of the superior and inferior elastic bands contributes to the continuous play of the wing, by preventing dead
points at the end of the down and up strokes.

VOL. XXVI. PART II. oR

424 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

This form of wing is made to oscillate upon two centres (x and / of fig. 63), to
bring out the peculiar eccentric action of the pinion.

If I wish to produce a very delicate light wing, I do so by selecting a fine
tapering elastic reed, as represented at a b of figure 64, p. 425.
' To this I add successive layers (,h, 9,F, e,) of some flexible
material, such as parchment, buckram, tracing cloth, or even
paper. As the layers overlap each other, it follows that there
are five layers at the anterior margin (a 0), and only one at the
posterior (cd). This form of wing is not twisted upon itself
structurally, but it twists and untwists, and becomes a true screw
during its action.
How to Construct a Wave Wing which shall evade the superim-

> ~—

Fig. 63.*

posed Air during the Up Stroke-—To construct a wing which shall elude the air
during the up stroke, it is necessary to make it valvular, as shown at fig. 65, p. 425.

* Fig. 68. Artificial Wing with Driving Apparatus.

a 6, Strong elastic rod, which tapers towards the tip of the wing.

d, ¢, f, g, h, i,j, k, Tapering curved reeds, which run obliquelyf rom the anterior to the posterior margin of the wing,
and whch radiate towards the tip.

m, Similar curved reeds, which run still more obliquely. :

a, n, 0, p, g, Tapering curved reeds, which run from the anterior margin of the wing, and at right angles to it. These
support the two sets of oblique reeds, and give additional strength to the anterior margin.

x, Ball and socket joint, by which the root of the wing is attached to the cylinder.

s, Steam cylinder.

r, Piston, with cross bar, with which driving gear (¢) is connected by ball and socket joint (7), and by a hinge
joint (m). The hinge joint is mounted on a tube, through which the root of the wing passes, and within which it ean
rotate in the direction of its length (long axis). The hinge joint and the tube on which it is mounted can be moved out
and in upon the root of the wing, and fixed by the aid of pins. By this means the range of the wing, 7.¢., the length of
the stroke, can be increased or diminished. The driving gear is arranged on a similar principle. Thus, by causing the
portion marked w to move within the tube (¢) in an upward direction, the wing vibrates on a higher level than natural.
If, on the other hand, the portion marked u be moved in a downward direction, the wing vibrates on a lower level. The
range of the wing and its are of vibration are thus easily regulated.

1, 2, Cross bar attached to steam chest (7) and to cylinder (s). To this anterior (v) and posterior (w) elastic
bands are affixed. Those elastic bands (anterior and posterior) are bound to the anterior and posterior portions of the
ring c; y, superior elastic band ; 2, inferior ditto.

3, 4, Steel springs running at right angles to each other, and attached respectively to the cross bar and the root of the
wing anteriorly. They come in contact when the wing descends, and prevent the anterior margin of the wing from dipping,
i.e., from diving downwards during the down stroke. This result is also secured by inserting the superior elastic band (y)
into the upper and anterior portion of the ring c. Indeed, by employing a cross bar or lever, similar to that marked 4,
in place of the ring c, the amount of rotation of the posterior margin round the anterior one can be regulated both
during the down and up strokes. If the superior elastic band (y) be moved towards the tip of the lever, the degree of
rotation is increased ; if it be moved towards the root of the lever, it is diminished.

5. Rod fixed to posterior of cylinder, and bearing cross bar (6), to which the superior elastic band (zy) is attached.

Norr.—In the present arrangement the steam chest (7) and valve occupy the ceutre of the cylinder posteriorly, the
valve being opened and closed by the aid of an idle rod (furnished with two kickers), which passes through a loop pro-
jecting from the piston anteriorly. The idle rod and kickers move a small lever (9), which in turn moves the spindle
(8), to which the steam valve is attached. The cylinder is fixed to the top of the boiler, and the ports for the admission of
steam to the cylinder are unequal in size, the woper port being larger than the under one. Unequal quantities of steam
are thus admitted to the top and bottom of the cylinder respectively, the greater quantity admitted to the top causing
the wing to descend much more quickly than it ascends. From the above figure it will be seen that the movements of
the wing are communicated directly from the piston, a great saving in weight and power being thus effected.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 425

This wing, as the figure indicates, is composed of numerous narrow segments
(777,999), 80 arranged that the air, when the wing is made to vibrate, opens
or separates them at the beginning of the up stroke, and closes or brings them
together at the beginning of the down stroke.

z Spent Hd b

Fig. 64.*

The time and power required for opening and closing the segments is com-
paratively trifling, owing to their extreme narrowness and extreme lightness.
The space, moreover, through which they pass in performing their valvular

Ape PR

Fig. 65.t

action is exceedingly small. The wing under observation is flexible and elastic
throughout, and resembles in its general features the other wings described.

I have also constructed a wing which is self-acting in another sense. This
consisted of two parts—the one part being made of an elastic reed, which
tapered towards its extremity, the other of a flexible sail. To the reed, which
corresponded to the anterior margin of the wing, delicate tapering reeds were
fixed at right angles, the principal and subordinate reeds being arranged on the
same plane. The flexible sail was attached to the under surface of the principal
reed, and was stiffer at its insertion than towards its free margin. When the
wing was made to ascend, the sail, because of the pressure exercised upon its
upper surface by the air, assumed a very oblique position, so that the resistance
experienced by it during the wp stroke was very slight. When, however,
the wing descended, the sail instantly flapped in an upward direction, the

* Fig. 64. x, Ball and socket joint at root of wing. a, b, Anterior margin of wing. c, d, Posterior margin of wing.

?, Portion of wing composed of one layer of flexible material. h, Portion of wing composed of two layers. 4g, Portion
a wing composed of three layers. jf, Portion of wing composed of four layers. ¢, Portion of wing composed of five
ayers.

+ Fig. 65. Flexible valvular wing with India-rubber springs attached to its root.

_%,, Anterior margin of wing, tapering and elastic. c,d, Posterior margin of wing, elastic. /, f,f, Segments
which open during the up stroke and close during the down, after the manner of valves. These are very narrow,
and open and close instantly. g, g, g, The same segments magnified. «a, Universal joint. m, Superior elastic band.
n, Dittoinferior. 0, Ditto anterior. p, ¢g, Ditto oblique. 7, Ring into which the elastic bands are fixed.

426 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

subordinate reeds never permitting its posterior or free margin to rise above its
anterior or fixed margin. The under surface of the wing consequently descended
so as to present a flat surface to the earth. It experienced much resistance
from the air during the down stroke, the amount of buoyancy thus furnished being
very considerable. The above form of wing is more effective during the down
stroke than during the up. It, however, elevates and propels during both, the
forward travel being greatest during the down stroke.

Compound Wave Wing.—In order to render the movements of the wing as
simple as possible, I was induced to devise a form of pinion, which for the sake
of distinction I shall designate the Compound Wave Wing. This wing consists
of two wave wings united at their roots, as represented at b,c, (A, A’) of fig.
66. It is attached by its centre to the head of the piston by a compound joint

/
= TUN
ara a ~SEAMINMN
Nifii/, y SA NA AY
/ rid
i 1 f

a ~. : a

o
i) Ll

Wine

\
\

r) ge

Fig. 66.

(x), which enables it to move in a circle, and to rotate along its anterior margin
(a, b, c,d, A, A’) in the direction of its length. The circular motion is for steer-
ing purposes only. The wing rises and falls with every stroke of the piston,
and the movements of the piston are quickened during the down stroke, and
slowed during the up one (vide note to fig. 63, p. 424, also pp. 432 and 433).
During the up stroke of the piston the wing is very decidedly convex on its
upper surface (1, b,c, d, A, A’), its under surface being deeply concave and inclined
obliquely upwards and forwards. It thus evades the air during the up stroke.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 427

During the down stroke of the piston the wing is flattened out im every direction,
and its extremities twisted in such a manner as to form two screws, as shown
ata’ bc’ d; ef gh’; B, B’ of figure. The active area of the wing is by this
means augmented, so that it seizes the air with great avidity during the down
stroke. The area of the wing may be still further increased and diminished
during the down and up strokes by adding joints to the body of the wing on the
principle recommended at pages 428, 429, 430, and 431, figs. 67, 68, and 69.
The degree of convexity given to the upper surface of the wing can be increased
or diminished at pleasure by causing a cord (¢7; A, A’) and elastic band
(£4) to extend between two points, which may vary according to circumstances.
The wing is supplied with vertical springs, which assist in slowing and reversing
it towards the end of the down and up strokes, and these, in conjunction
with the elastic properties of the wing itself, contribute powerfully to its con-
tinued play. The compound wave wing produces the currents on which it
rises. Thus during the up stroke it draws after it a current, which being met
by the wing during its descent, confers additional elevating and propelling
power. During the down stroke the wing in like manner draws after it a cur-
rent which forms an eddy, and on this eddy the wing rises, as explained at page
438, fig. 72. The ascent of the wing is favoured by the superimposed air play-
ing on the upper surface of the posterior margin of the organ, in such a manner
as to cause the wing to assume a more and more oblique position with reference
to the horizon. This change in the plane of the wing enables its upper surface
to avoid the superincumbent air during the up stroke, while it confers upon its
under surface a combined kite and parachute action. The compound wave
wing leaps forward in a curve both during the down and up strokes, so that the
wing during its vibration describes a waved track, as shown at a, ¢, é, g, @ of fig.
14, page 344. The compound wave wing possesses most of the peculiarities of
single wings when made to vibrate simultaneously. It forms a most admirable
elevator and propeller, and has this advantage over ordinary wings, that it can be
worked without injury to itself, when the machine which it is intended to elevate
is resting on the ground. Two or more compound wave wings may be arranged
on the same plane, or superimposed, and made to act in concert. They may also
by a slight modification be made to act horizontally instead of vertically. The
length of the stroke of the compound wave wing is determined in part, though
not entirely, by the stroke of the piston—the extremities of the wing, because of
their elasticity, moving through a greater space than the centre of the wing. By
fixing the wing to the head of the piston all gearing apparatus is avoided, and
the number of joints and working points reduced—a matter of no small import-
ance when it is desirable to conserve the motor power and keep down the

weight.
How to Construct a Wave Wing on the Bat and Bird type.—In order to

VOL, XXVI. PART II. aS

428 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

imitate the bat and bird’s wing successfully it is necessary to introduce joints :
the artificial wing, in fact, requires to be composed of several pieces, so that
it will flex or fold towards the end of the down stroke, and open out or expand
towards the end of the up stroke. This is requisite for several reasons. In
the first place, the wing of the bat and bird is made to vibrate in a much more
vertical direction (figs. 5 and 6, Plate XI., figs. 18 and 19, Plate XIV.,) than
that of the insect (fig. 4, Plate XI.) They have therefore to contend directly
with the resistance furnished by the superimposed air. As a consequence, the
wing in such of the bats and birds as do not sail or skim must be folded more
or less completely during the up stroke to diminish the wing area, so as to
elude the resistance offered by the air when the wing is being elevated. It is
for this reason too, that in the bird the rowing feathers open up or separate
during the up stroke. As the wings of the bat and bird afford comparatively
little support during the up stroke, it follows that the wing area must be
increased to its utmost during the down stroke. But for the folding or clos-
ing of the wing towards the termination of the down stroke, the downward
passage of the pinion, as I have repeatedly satisfied myself by experiment,
could not be suddenly arrested and a new upward passage commenced. In
other words, the wing could not be reversed. At the beginning of the down
stroke the wing is a long lever, and acts as such, (vide ¢ d of fig. 6, Plate XI.)
It is depressed with extreme energy and acquires during its descent a degree
of momentum which could not possibly be checked if the wing was not sud-

’

denly flexed and instantly converted from a long into a short lever, (vide a b of —

fig. 6, Plate XI.) The wing is therefore by this very simple contrivance, not
only robbed of its momentum, but what is quite as important, it is prepared
for making the up or return stroke. Ifthe wing of a gull just dead be taken,
and the air winnowed by it in a more or less vertical direction, it will be found
to fly open and to extend itself during the down stroke, and to fold up or close
during the up stroke. The quicker the wing is made to vibrate, the more
admirable is the result. Indeed, the gull’s wing, when made to oscillate as
recommended, reverses perfectly and has no dead points. It moreover furnishes
a steady persistent buoying power which is quite remarkable when the limited
dimensions of the pinion are taken into account.

To construct a bat or bird’s wing, I take a tapering flexible reed, and cut
it into three pieces, each piece varying in length. These I bend to the shape
required as shown at a, d, and g of figs. 67 and 68, page 429. .

The shortest and thickest piece (a) I furnish with a ball and socket joint at

one end (z), and a hinge joint (6) at the other. The second shortest and —

strongest piece (d) I supply with a hinge joint at either end (0 and e) ; and the
- third piece, which is the longest and weakest, I provide with a hinge joint at its
thicker or proximal end (¢). When the three pieces are joined together as

sé.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 429

shown in the figures, I apply to each of the pieces at intervals tapering elastic
curved reeds (0, p, q, fig. 68), the reeds radiating in the direction of the tip of
the wing, and in such a manner as to confer a certain degree of spirality upon
it. Ithen cause elastic substances (h 7, 7 4, of figs. 67 and 68,) to extend
between the pieces (a, d,g). The covering is then added in one piece if a bat’s
wing is desired, and in several (see valvular wing, fig. 65, page 425,) if the more
highly differentiated wing of the bird is aimed at.

fig 63:

The covering may consist of a thin layer of india-rubber, silk, linen, or any
other flexible material.

To the inner extremity of the distal reed (7) I attach a cord or wire, and
this cord or wire (/, m, 7,) I pass through an aperture in the outer extremity (c)
of the proximal reed. I then fix the free end of the cord to a loop in the
cylinder (vide q of fig. 69, p. 430), from which the wing receives its movements
by a direct piston action.

The arrangement is represented at fig. 69, page 430.

When the wing is elevated from B to A (fig. 69) by the direct action of the

; i. Figures 67 and 68. Wing made to close or fold during the up stroke, and to open out or expand during the down
stroke.

At fig. 67, the wing is represented as folded upon itself. a Universal joint at root of wing. a Proximal portion
of wing. d Central portion of wing. g Distal portion of wing. & Joint uniting proximal and central portions of wing.
e Joint uniting central and distal portions. 7, 7 & Sheet of elastic substance which when contracted as represented,
tends to approximate the proximal (a), central (d), and distal (g), portions of wing. 7, m,n A cord or wire fixed at
fand running through an aperture atc. If this cord be rendered taught (provided the root of the wing (2) is fixed in
its socket), it causes the proximal (a), central (d), and distal (g) portions of the wing suddenly to dart out and arrange
themselves in a nearly straight line as shown at a, d, g of fig. 68.

__ At fig. 68 the wing is represented as fully extended or spread out. The lettering is the same as in fig. 67. 0, p, q
Ribs or stays of the wing which support the covering or curtain.

430

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

piston (7 7), and the gearing apparatus (y 2), it is likewise extended or spread
out, the mere elevation of the piston rendering the cord or wire (/, n,) taught—

Fig. 69.*

the taughtness of the cord causing the different parts of the wing to fly out-
wards, while it at the same time puts the elastic substances (4 4) on the stretch. .

* Fig. 69. Wing which folds upon itself during the up stroke, and expands during the down one, made to vibrate
by a direct piston action. At A the wing is fully expanded and in the act of commencing the down stroke. At B the
wing is at mid stroke and very slightly folded. At C the wing is fully folded, and ready to begin the up stroke. It is
thus that the wing acts as a Jong lever at the beginning of the down stroke, and a short one at the beginning of the up one.
Compare with figs. 18 and 19, Plate XIV., and also with figs. 9, 10, and 11, Plate XII. The lettering of the wing in
the present fig. is the same as in fig. 68, p. 429.

x Universal joint at root of wing received into cup-shaped cavity (v) of cylinder (¢).

a Proximal, d central, and g distal portions of wing.

b, e, Joints which unite the three portions of the wing to each other.

J, 7, Points to which the cord or wire of wing is fixed.

ce, Aperture through which cord or wire of wing glides as the wing ascends and descends. When the piston ascends
it elevates the wing by its gearing yz. It also renders the cord 7 » taught, the cord in its turn extending the wing (A)
and the elastic substance & When the piston descends to mid stroke the wing is very slightly folded (B) and the
cord U' n' somewhat relaxed. When the piston has quite descended the cord 2’ n” is very much relaxed, and as a conse-
quence the elastic substance extending between the different portions of the wing has contracted, the wing being thereby
folded upon itself (C). The elastic substance may be dispensed with, if a strong elastic cord be employed instead of the
non-elastic one J, n. _If two cords be fixed to two points on the cylinder as at p and q, and the one cord be passed on
the upper surface of the wing, and the remaining one on the under surface, the wing will be under control during the _
whole of the down and up strokes, the one cord extending the wing, the other flexing it.

t, t, Cylinder. 0, 0, Stuffing boxes. r7, Piston. w, w, Cross heads for driving gear. yz Driving gear. $ Piston
head, v, Cup-shaped cavity for receiving root of wing.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 431

The instant the wing begins to descend, the cord is more or less relaxed, and a
struggle ensues between the air, which endeavours to keep the wing open, and
the elastic substances (4) which endeavour to close it. If the wing is very
forcibly depressed, it is kept open till quite near the end of the down stroke,
when the elastic bands close it (C), destroy its momentum, and prepare it for
the up stroke. This form of wing acts as a short lever (C) during the up
stroke, and a long one (A) during the down stroke. It therefore eludes the
superimposed air to a great extent when it is being elevated. If it is thought
desirable to differentiate the wing still further in imitation of the bird’s wing,
it is only necessary to add a series of segments similar to those represented at
fig. 65, page 425, these segments representing the individual rowing feathers.
What especially struck me on analysing the movements of the artificial bat and
bird’s wing, was the fact, that during their vibrations figure of 8 curves were
developed along their anterior and posterior margins similar to those found in
the living wings ; that the under surfaces of the pinions made various angles of
inclination with the horizon analogous to those made by the natural wings ;
these changes being induced in a great measure independently of the air, in
virtue apparently of inherent structural peculiarities. This I regard as a very
remarkable circumstance, and one well worthy the attention of the physiologist
and mechanician.

How to apply Artyicial Wings to the Air.—BoreE ui, DuRCKHEIM, Marey, and
all the writers with which I am acquainted, assert that the wing should be
made to vibrate vertically. I believe that if the wing be in one piece it should
be made to vibrate obliquely and more or less horizontally. If, however, the wing
be made to vibrate vertically, it is necessary to supply it with a ball and socket
joint, and with springs at its root (mn of fig. 62, page 423), to enable it do
leap forward in a curve when it descends, and in another and opposite curve
when it ascends (vde a, ¢, e, g,7 of fig. 14, page 344). This arrangement practi-
cally converts the vertical vibration into an oblique one. If this plan be not
adopted the wing is apt to foul at its tip. In applying the wing to the air it
ought to have a figure of 8 movement communicated to it either directly or
indirectly. It is a peculiarity of the artificial wing properly constructed, (as it
is of the natural wing), to twist and untwist and make figure of 8 curves during
its action (see a b, cd of fig. 62, page 423), this enabling it to seize and let go
the air with wonderful rapidity, and in such a manner as to avoid dead points.
If the wing be in several pieces it may be made to vibrate more vertically than
a wing in one piece, from the fact that the outer half of the pinion moves for-
wards and backwards when the wing ascends and descends so as alternately to
become a short and long lever. (Compare C with A of fig. 69, page 480), this
arrangement permitting the wing to avoid the resistance experienced from the air
during the up stroke, while it vigorously seizes the air during the down stroke.

VOL. XXVI. PART II. oT

432 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

If the body of a flying animal be in a horizontal position, a wing attached to
it in such a manner that its under surface shall look forwards, and make an up-
ward angle of 45° with the horizon, is in a position to be applied either verti-
cally (figs. 5 and 6, Plate XI.), or horizontally (figs. 3, 4, 5, and 6, page 338).
Such, moreover, is the conformation of the shoulder joint in insects, bats, and
birds, that the wing can be applied vertically, horizontally, or at any degree of
obliquity without inconvenience.* It is in this way that an insect which may
begin its flight by causing its wings to make figure of eight horizontal loops
(vide fig. 8, page 340), may gradually change the direction of the loops, and make
them more and more oblique until they are nearly vertical (see fig. 13, page 342).
In the beginning of such flight the insect is screwed nearly vertically upwards ;
in the middle of it, it is screwed upwards and forwards; whereas, towards the
end of it, the insect advances in a waved line almost horizontally (see q, 7, s, t, of
fig. 11, page 341). The muscles of the wing are so arranged that they can propel
it in a horizontal, vertical, or oblique direction. It is a matter of the utmost
importance that the direction of the stroke and the angles made by the surfaces
of the wing during its vibration with the horizon should be distinctly under-
stood, as it is on these that all flymg creatures depend when they seek to elude
the upward resistance of the air, and secure a maximum of elevating and _ pro-
pelling power with a minimum of slip.

Nature of the Forces required for Propelling artificial wings.—BoRELLuI,
DurckHem, and Margy affirm that it suffices if the wing merely elevates
and depresses itself by a rythmical movement in a perpendicular direction,
while CHABRIER is of opinion that a movement of depression only is required.
All those observers agree in believing that the details of flight are due to the
reaction of the air on the surface of the wing. Repeated experiment has, how-
ever, convinced me that the artificial wing must be thoroughly under control,
both during the down and up strokes—the details of flight being in great measure
due to the movements communicated to the wing by an intelligent agent. In order
to reproduce flight by the aid of artificial wings I find it necessary to employ a.
power which varies in intensity at every stage of the down and up strokes. The
power which I find suits best is one which is made to act very suddenly and
forcibly at the beginning of the down stroke, which gradually abates in intensity
until the end of the down stroke where it ceases to act in a downward direction.
The power is then made to act inan upward direction, and gradually to decrease
until the end of the up stroke. The force is thus applied more or less con-
tinuously, its energy being increased and diminished according to the position

* The human wrist is so formed that if a wing be held in the hand at an upward angle of 4am
the hand can apply it to the air in a vertical or horizontal direction without difficulty. This arises
from the power which the hand has of moving in an upward and downward direction, and from side
to side with equal facility. The hand can also rotate on its long axis, so that it virtually represents all
the movements of the wing at its root.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 433

of the wing, and the amount of resistance which it experiences from the air.
The flexible and elastic nature of my peculiar form of wing (wave-wing), as-
sisted by certain springs to be presently explained, ensure a continuous vibra-
tion where neither halts nor dead points are observable. I obtain the varying
power required by a direct piston action, and by working the steam expansively
(vide figs. 62, 63, and 69, pages 423, 424, and 430). The power employed is
materially assisted, particularly during the up stroke, by the reaction of the air
and the elastic structures about to be described. An artificial wing, propelled
and regulated by the forces recommended, is in some respects as completely
under control as the wing of the insect, bat or bird.

Necessity for supplying the root of artificial wings with elastic structures in
imitation of the muscles and elastic ligaments of flying animals.— Bore, DuRCK-
HEIM, and Margy, who advocate the perpendicular vibration of the wing, make
no allowance, so far as I am aware, for the wing leaping forward in curves dur-
ing the down and up strokes. As a consequence, the wing is jointed in their
models to the frame by a simple joint which moves only in one direction, viz.,
from above downwards, and vice versa. Observation and experiment have,
however, convinced me that an artificial wing, to be effective as an elevator and
propeller, ought to be able to move not only in an upward and downward direc-
tion, but also ina forward, backward, and oblique direction, nay, more, that it
should be free to rotate along its anterior margin in the direction of its length :
in fact, that its movements should be universal. Thus it must be able to rise
or fall, to advance or retire, to move at any degree of obliquity, and to rotate
along its anterior margin. To secure the several movements in question I fur-
nish the root of the wing with a ball and socket-joint, ¢.¢., a universal joint (see
@ of fig. 62, page 423 ; and 2 of fig. 68, page 424). To regulate the several move-
ments when the wing is vibrating, and to confer on the wing the various in-
clined surfaces requisite for flight, as well as to delegate as little as possible to

the air, I employ a cross system of elastic bands. These bands vary in length,
Strength, and direction, and are attached to the anterior margin of the wing
(near its root), and to the cylinder (or a rod extending from the cylinder) of the
model respectively (vide m, n of fig. 62, page 423). The principal bands are four
in number: a superior (fig. 63, page 424, y), inferior (z), anterior (v), and posterior
(w). The superior band extends between a rod proceeding from the upper part
of the cylinder (5) of the model, and the upper surface of the anterior margin
(a, b,) of the wing; the inferior band (z), extending between the under part of
the cylinder or boiler and the inferior surface of the anterior margin (d, ¢, /,) of
the pinion. The anterior (v), and posterior (w), bands are attached to the anterior
and posterior portions of the wing and to rods extending from the centre of the
anterior and posterior portions of the cylinder. Oblique bands are added (vide
P, 7 of fig. 65, page 425), and these are so arranged that they give to the wing

434 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

during its descent and ascent the precise angles made by the wing with the
horizon in natural flight. The superior bands are stronger than the inferior
ones, and are put upon the stretch during the down stroke. They thus help
the wing over the dead point at the end of the down stroke, and assist, in con-
Junction with the reaction obtained from the air, in elevating it. The posterior
bands are stronger than the anterior ones to restrain within certain limits the
strong tendency which the wing has to leap forward in curves towards the end
of the down and up strokes. The oblique bands, aided by the air, give the
necessary degree of rotation to the wing in the direction of its length. This effect
can, however, also be produced independently by the four principal bands. From
what has been stated it will be evident that the elastic bands exercise a restrain-
ing influence, and that they act in unison with the driving power and with the
reaction supplied by the air. They powerfully contribute to the continuous vibra-
tion of the wing, the vibration being peculiar in this that it varies in rapidity at
every successive stage. I derive the motor power, as has been stated, from a
direct piston action, the piston being urged either by steam worked expansively
or by the hand, if it is merely a question of illustration. In the hand models the
“muscular sense” at once informs the operator as to what is being done. Thus
if one of the wave wings supplied with a ball and socket joint, and a cross
system of elastic bands as explained, has a sudden vertical impulse communi-
cated to it at the beginning of the down stroke, the wing darts downwards and
Jorwards in a curve (vide a, ¢, of fig. 14, page 344), and in doing so zt elevates and
carries the piston and cylinder forwards. The force employed in depressing the
wing is partly expended in stretching the superior elastic band (y of fig. 63, page
424), the wing being slowed towards the end of the down stroke. The instant
the depressing force ceases to act the superior elastic band (vy) contracts, and
the air reacts ; the two together, coupled with the tendency which the model has
to fall downwards and forwards during the up stroke, elevating the wing. The
wing when it ascends describes an upward and forward curve, as shown at ¢ é
of fig. 14, page 344. The ascent of the wing stretches the inferior elastic band
(z of fig. 63, page 424) in the same way that the descent of the wing stretched
the superior band. The superior and inferior elastic bands antagonise each other
and reciprocate with vivacity. While those changes are occurring the wing is’
twisting and untwisting in the direction of its length and developing figure of eight
curves along its margins (page 423, fig. 62, a b, ed), and throughout its sub-
stance similar to what are observed under like circumstances in the natural wing
(vide figs. 39, 40, 41, 42, and 43, page 362). The angles, moreover, made by the
under surface of the wing with the horizon during the down and up strokes are
continually varying—the wing all the while acting as a kite, which flies steadily
upwards and forwards (fig. 15, page 345). As the elastic bands, as has been
partly explained, are antagonistic in their action the wing is constantly oscillating

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 435

in some direction, there being no dead point either at the end of the down or
up strokes. Asa consequence, the curves made by the wing during the down
and up strokes respectively, run into each other to form a continuous waved
track, as represented at figs. 13,14, and 15, pages 342, 344, and 345. <A con-
tinuous movement begets a continuous buoyancy, and it is quite remarkable to
what an extent, wings constructed and applied to the air on the principles
explained, elevate and propel—how little power is required, and how little of
that power is wasted in slip.

If the piston, which in the experiment described has been working vertically,
be made to work horizontally, a series of essentially similar results are obtained.
When the piston is worked horizontally, the anterior and posterior elastic bands
require to be of nearly the same strength, whereas the inferior elastic band
requires to be much stronger than the superior one, to counteract the very
decided tendency the wing has to fly upwards. The power also requires to be
somewhat differently applied. Thus the wing must have a violent impulse
communicated to it when it begins the stroke from right to left, and also when
it begins the stroke from left to right (the heavy parts of the spiral line repre-
sented at fig. 8, page 340, indicate the points where the impulse is communi-
cated). The wing is then left to itself, the elastic bands and the reaction
of the air doing the remainder of the work. When the wing is forced by the
piston from right to left, it darts forwards ina double curve, as shown at fig.
70, the various inclined surfaces made by the wing with the horizon changing
at every stage of the stroke.

Fig. 70.* : Fig. 71.+

At the beginning of the stroke from right to left, the angle made by the
under surface of the wing with the horizon (a 2’) is something like 45°, whereas
at the middle of the stroke it is reduced to 20° or 25°. At the end of the stroke
the angle gradually increases to 45°, then to 90°, after which the wing suddenly
turns a somersault, and reverses precisely as the natural wing does at e, f, g of
figs. 3 and 5, page 338. The artificial wing reverses with amazing facility, and
in the most natural manner possible. The angles made by its under surface

* Fig. 70. Stroke of artificial wave wing from right to left. x, 2, Horizon. m,n, 0, Wave track described by
wing from right to left. p, Angle made by wing at beginning of stroke. gq, Ditto, made at middle of stroke. 0, Ditto,
towards end of stroke. c, Wing in the act of reversing ; at this stage the wing makes an angle of 90° with the horizon,
and its speed is less than at any other part of its course. d, Wing reversed, and in the act of darting up to w, to begin
the stroke from left to right (vide w of fig. 71).

+ Fig. 71. Stroke of artificial wave wing from left to right. 2, a, Horizon. w, v, w, Wave track described by
wing from left to right. ¢, Angle made by the wing with the horizon at beginning of stroke. y, Ditto, at middle of
stroke. 2, Ditto, towards end of stroke. 7, Wing in the act of reversing ; at this stage the wing makes an angle of 90°
with the horizon, and its speed is less than at any other part of its course. s, Wing reversed, and in the act of darting
up to m, to begin the stroke from right to left (vide sn of fig. 70).

VOL. XXVI. PART II. dU

436 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

with the horizon, depend chiefly upon the speed with which the wing is urged
at different stages of the stroke, the angle always decreasing as the speed
increases, and vice versa. As a consequence, the angle is greatest when the
speed is least.

The course described, and the angles made by the artificial wave wing with the
horizon during the stroke from right to left, are represented at fig. 70, page 435.

When the wing reaches the point }, its speed is much less than it was at gq.
The wing is, in fact, preparing to reverse. At c the wing is in the act of revers-
ing (compare with ¢ of figs. 16 and 17, page 349), and, as a consequence, its
speed is at its minimum, and the angle which it makes with the horizon at its
maximum. At d the wing is reversed, its speed being increased, and the angle
which it makes with the horizon diminished. Between the letters d and wu the
wing darts suddenly up like a kite, and at w it is in a position to commence the
stroke from left to right, as indicated at w of fig. 71 p. 435. The course described,
and the angles made by the wing with the horizon during the stroke from left
to right, are represented at fig. 71 (compare with figs. 4 and 6, page 338), The
stroke from left to right is in every respect the converse of the stroke from
right to left, so that a separate description is unnecessary.

The Artificial Wave Wing can be driven at any speed—it can make tts own
currents, or utilise existing ones,—The remarkable feature in the artificial wave
wing is its adaptability. It can be driven slowly, or with astonishing rapidity.
It has no dead points. It reverses instantly, and in such a manner as to dissi-
pate neither time nor power. It alternately seizes and evades the air so as to
extract a maximum amount of support with a minimum of slip, and with a
minimum expenditure of power, It supplies a degree of buoying and propelling
power which is truly remarkable. Its buoying area is nearly equal to half a
circle. It can act upon still air, and it can create and utilise its own currents.
I proved this in the following manner. I caused the wing to make a horizontal
sweep from right to left over a candle ; the wing rose steadily as a kite would,
and after a brief interval, the flame of the candle was persistently blown from
right to left. I then waited until the flame of the candle assumed its normal
perpendicular position, after which I caused the wing to make another and
opposite sweep from left to right. The wing again rose kite fashion, and the
flame was a second time affected, being blown in this case from left to right. I
now caused the wing to vibrate steadily and rapidly above the candle, with this
curious result, that the flame did not incline alternately from right to left and
from left to right. On the contrary, it was blowu steadily away from me, 2.¢.,
in the direction of the tip of the wing, thus showing that the artificial currents
produced, met and neutralised each other always at mid stroke. I also found
that under these circumstances the buoying power of the wing was remarkably
increased.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 437

Compound rotation of the Artificial Wave Wing: the different parts of the
Wing travel at different speeds.—The artificial wave wing, like the natural wing,
revolves upon two centres (a b, ¢ d of fig. 45, page 376, and a, / of fig. 63, page
494) and owes much of its elevating and propelling, seizing and disentangling
power to its different portions travelling at different rates of speed (see fig. 51,
page 399), and to its storing up and giving off energy as it hastens to and fro.
Thus the tip of the wing moves through a very much greater space in a given
time than the root, and so also of the posterior margin as compared with the
anterior, This is readily understood, by bearing in mind that the root of the
wing forms the centre or axis of rotation for the tip ; while the anterior margin
is the centre or axis of rotation for the posterior margin. The momentum,
moreover, acquired by the wing during the stroke from right to left ¢s expended
in reversing the wing, and in preparing it for the stroke from left to right, and
vice versa; a continuous to and fro movement devoid of dead points being thus
established. If the artificial wave wing be taken in the hand and suddenly
depressed in a more or less vertical direction, it immediately springs up again,
and carries the hand with it. It, in fact, describes a curve whose convexity is
directed downwards, and in doing so, carries the hand upwards and forwards.
If a second down stroke be added, a second curve is formed; the curves
running into each other, and producing a progressive waved track similar to
what is represented at a, ¢, @, 7, 7 of fig. 14, page 344. This result is favoured if
the operator runs forward so as not to impede or limit the action of the wing.

How the Wave Wing creates currents, and rises upon them, and how the air
assists in elevating the Wing.—In order to ascertain in what way the air contri-
butes to the elevation of the wing, I made a series of experiments with natural
and artificial wings. On concluding these experiments, I felt convinced that
when the wing descends it compresses and pushes before it, in a downward and
forward direction, a column of air represented by a, b, ¢ of fig. 72, p. 438.* The
air rushes in from all sides to replace the displaced air, as shown at d, e, f, g, h, 7,
and so produces a circle of motion indicated by the dotted line s, ¢, v7, w. The
Wing rises upon the outside of the circle referred to, as more particularly seen
at d, ¢, v, w. The arrows, it will be observed, are all pointing upwards, and as
these arrows indicate the direction of the reflex or back current, it is not diffi-
cult to comprehend how the air comes indirectly to assist in elevating the wing.
A similar current is produced to the right of the figure, as indicated by Z, m,
0, P, Y, 7, but seeing the wing is always advancing, this need not be taken into
account.

* The artificial currents produced by the wing during its descent may be readily seen by partially
filling a chamber with steam, smoke, or some impalpable white powder, and causing the wing to
descend in its midst. By a little practice, the eye will not fail to detect the currents represented at
d, ¢, f, g, h, 1, l, m, n, 0, p, q, 7 of fig. 72, p. 438.

438 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

If fig. 72 be made to assume a horizontal position, instead of the oblique
position which it at present occupies, the manner in which an artijicial current
is produced by one sweep of the wing from right to left, and utilised byit ina _
subsequent sweep from left to right, will be readily understood. The artificial
wave wing makes a horizontal sweep from right to left, 7.¢., it passes from the
point a to the point ¢ of fig. 72. During its passage it has displaced a column
of air. To fill the void so created, the air rushes in from all sides, viz., from
d, ¢,f,9, h,t; 1, m, 0,p,¢,7r. The currents marked g, h,1; p, g, 7, Tepre
sent the reflex or artificial currents. These are the currents which, after a
brief interval, force the flame of the candle from right to left. It is those
same currents which encounter the wing, and contribute so powerfully to its

Fig. 72.

elevation, when it sweeps from left to right. The wing, when it rushes from
left to right, produces a new series of artificial currents, which are equally
powerful in elevating the wing when it passes a second time from right to
left, and thus the process of making and utilising currents goes on so long
as the wing is made to oscillate. In waving the artificial wing to and fro,
I found the best results were obtained when the range of the wing and the
speed with which it was urged were so regulated as to produce a perfect
reciprocation. Thus, if the range of the wing be great, the speed should also
be high, otherwise the air set in motion by the right stroke will not be utilised —
by the left stroke, and vice versa. If, on the other hand, the range of the
wing be small, the speed should also be low, as the short stroke will ena
the wing to reciprocate as perfectly as when the stroke is longer and ~

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 439

speed quicker. When the speed attained is high, the angles made by the
under surface of the wing with the horizon are diminished ; when it is low,
the angles are increased. From these remarks it will be evident that the
artificial wave wing reciprocates in the same way that the natural wing recipro-
cates, the reciprocation being most perfect when the wing is vibrating in a
given spot, and least perfect when it is travelling at a high horizontal speed.

The Artificial Wing propelled at various degrees of speed during the down
and up strokes.—The tendency which the artificial wave wing has to rise again
when suddenly and vigorously depressed, explains why the e/evator muscles of the
wing should be so small when compared with the depressor muscles—the latter
being something like seven times larger than the former. ‘That the contraction
of the elevator muscles is necessary to the elevation of the wing, is abundantly
proved by their presence, and that there should be so great a difference between
the volume of the elevator and depressor muscles is not to be wondered at, when
we remember that the whole weight of the body is to be elevated by the rapid
descent of the wings—the descent in question being entirely due to the vigorous
contraction of the pectoralis major. If, however, the wing was elevated with
as great a force as it is depressed, it is plain that the good effected during the
descent would be utterly undone, as the wing, during its ascent, would experience
a much greater resistance from the air than it did during its descent. The
wing is consequently elevated more slowly than it is depressed, the elevator
muscles exercising a controlling and restraining influence. By slowing the
wing during the up stroke, the air has an opportunity of reacting on its under
surface, as explained at page 351.

The Artificial Wave Wing as a Propeller.—The wave wing makes an
admirable propeller if its tip be directed vertically downwards, and the wing
lashed from side to side with a sculling figure of 8 motion, similar to that executed
by the tail of the fish. Three wave wings may. be made to act in concert and
with a very good result ; two of them being made to vibrate figure of 8 fashion
in a more or less horizontal direction with a view to elevating, the third being
turned in a downward direction, and made to act vertically for the purpose of
propelling.

A New Form of Aerial Screw.—If two of the. wave wings represented at
fig. 62, page 423, be placed end to end, and united to a vertical portion of tube
to form a two-bladed screw, similar to that employed in navigation, a most
powerful elastic aerial screw is at once produced, as seen at fig. 73, page 440.

This screw, which for the sake of uniformity I denominate the aerial ware
screw, possesses advantages for aerial purposes to which no form of rigid screw
yet devised can lay claim. The way in which it clings to the air during its
revolution, and the degree of buoying power it possesses are quite astonishing.
It is a self-adjusting, self-regulating screw, and as its component parts are

VOL. XXVI. PART II. DX

440 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

flexible and elastic, it accommodates itself to the speed at which it is driven, and
gives a uniform buoyancy. The slip I may add is nominal in amount. This
screw is exceedingly light, and owes its efficacy te its shape and the graduated
nature of its blades, the anterior margin of each blade being comparatively
rigid, the posterior margin being comparatively flexible and more or less elastic.
The blades are kites in the same sense that natural wings are kites, and are
flown as such when the screw revolves. I find the aerial wave screw flies best
and elevates most when its blades are inclined at a certain upward angle as
indicated in the figure (73). The aerial wave screw may have the numbers of
its blades increased by placing the one above the other, and two or more screws
may be combined and made to revolve in opposite directions so as to make _
them reciprocate, the one screw producing the current on which the other
rises, aS happens in natural wings.

4 if :
Hf}\
c i of IJ

ee
. V WwW y y

iTS eS

The Aerial Wave Screw operates also upon Water.—The form of screw just
described is adapted in a marked manner for water, if the blades be made of
carefully tempered finely graduated steel plates and reduced in size. It bears”
the same relation to, and produces the same results upon, water as the tail and
fin of the fish. It throws its blades during its action into double figure of 8°
curves, similar in all respects to those produced on the anterior and posterior
margins of the natural and artificial flymg wing. As the speed attained by the
several portions of each blade varies, so the angle at which each part of the

* Fig. 72. Aerial wave screw whose blades are slightly twisted upon themselves (a b, cd; e f, g h), so that those
portions nearest the root (d h) make a greater angle with the horizon than those parts nearer the tip (bf). The angle
is thus adjusted to the speed attained by the different portions of the screw. The angle admits of further adjustment
by means of the steel springs z, s, these exercising a restraining, and to a certain extent a regulating influence which
effectually prevents shock. _

It will be at once perceived from this figure that the portions of the screw marked m and 7 travel at a much low
speed than those portions marked 0 and p, and these again more slowly than those marked g and7. As however tl
angle which a wing or a portion of a wing, as I have pointed out, varies to accommodate itself to the speed attained I
the wing, or a portion thereof, it follows, that to make the wave screw mechanically perfect, the angles made by t
several portions must be accurately adapted to the travel of its several parts as indicated above.

x, Vertical tube for receiving driving shaft. v, w, Sockets in which the roots of the blades of the serew ro
the degree of rotation being limited by steel springs z, s. a b, e f, Tapering elastic reeds forming anterior or thi
margins of blades of screw. dc, hg, Posterior or thin elastic margins of blades of screw. m n, 0p, ¢ 7, Radii fo
by the different portions of the blades of the screw when in operation. The arrows indicate the direction of travel.

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS, 441

blade strikes varies, the angles being always greatest towards the root of the
blade and least towards the tip. The angles made by the different portions of the
blade are diminished in proportion as the speed with which the screw is driven
is increased. The screw in this manner is self-adjusting, and extracts a large
percentage of propelling power with very little force and surprisingly little ‘slip.
A similar result is obtained, if two finely graduated angular-shaped steel
plates be placed end to end and applied to the water (vertically or horizontally
matters little), with a slight sculling figure of 8 motion, analogous to that
performed by the tail of the fish, porpoise, or whale. If the thick margin of the
plates be directed forwards, and the thin ones backwards, an unusually effective
propellor is produced. This form of propellor is likewise very effective in air.

EXPLANATION OF THE PLATES.
Puate XI.

Figures 1, 2, and 3 show how the wing of the gull is elevated and extended towards the termination
of the up stroke to prepare it for making the down stroke. At figure 3 the wing is repre-
sented as folded upon itself, and in the act of being elevated. It is, therefore, elevated as
a short lever, the resistance experienced from the superimposed air being thus greatly
diminished, The wing acts as a short lever from the time it leaves the position indicated
by 6 of figure 6 until it assumes the position indicated by o of figure 3. At figure 2 the
wing is raised higher than in figure 3, and partly extended—the elevation and extension
of the wing occurring simultaneously. At figure 1 the wing is fully elevated and fully
extended, and, consequently, ready to make the down stroke. It descends as a long lever,
with great energy, until it assumes the position indicated by 4 of figure 6. The resistance
which the wing experiences from the air beneath, is consequently, very great, the buoying
power of the wing bearing a fixed relation to the resistance in question, The under sur-
face of the wing, when in the position represented at figure 3, makes a very slight angle
with the horizon bd. This arises from the fact that the different portions of the wing,
when the wing is folded upon itself, are on nearly the same plane. The angle or angles
—for they are numerous—made by the under surface of the wing with the horizon become
larger when the wing is partly extended, as shown at figure 2 : bd, representing the horizon,
and ¢ 6 d the angle which the root of the wing makes with it. The angles become still
larger when the wing is fully extended, as a comparison of ¢ b d of figure 1 with c b d of
figure 2 will show. ‘The under surface of the wing, it will be observed, makes a variety
of inclined surfaces with the horizon while the pinion is being extended. The angles of
inclination made by the inclined surfaces in question are increased and diminished by the
ascent or descent of the posterior margin of the wing, o p q (the anterior margin acts as an
axis to the posterior one), the angles being always greatest when the wing is extended,
and least when it is flexed. The angles, moreover, made by the root of the wing are
always greater than those made by the tip. The various inclined surfaces made by the
under surface of the wing are intimately associated with the power the wing possesses of
alternately seizing and evading the air. The angles are greater at the root of the wing
than at the tip, because the portions of the pinion nearer the root travel at a lower speed
than portions nearer the tip. The various inclined surfaces made by the wing in flexion
and extension are well seen at figures 16 and 17, Plate XIII. At figure 17 the anterior
margin of the wing (ws ¢ vw) is nearly on a level with the posterior margin (0, p,q). At figure
16, on the other hand, the anterior margin (2, s, ¢, v, w) is elevated and the posterior margin
(op q) depressed. A careful examination of those figures (particularly figure 16) will also
show that the angles of inclination made by the several portions of the under surface

442 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

of the wing vary—the angle made by the part g of figure 16 with the horizon being
greater than that made by the part p—this, again, being greater than that made by the tip
of the wing 0. Those points are also illustrated at figure 8, Plate XII. The letters in
figures 1, 2, and 3 (Plate XI.) represent the same parts of the wing—z, shoulder joint ;
s, elbow joint; ¢, wrist joint; v, w, hand and finger joints; x s ¢ v w, anterior margin of
wing ; 0 p q, posterior margin.
Figure 4 represents the very oblique and almost horizontal direction of the stroke of the wing in the
flight of the insect (wasp)—how the wing is twisted upon itself at the end of the up (a)
and down (6) strokes, and how the tip of the wing, during its vibration, describes a figure
of 8 track in space (a, ¢, d).
Figures 5 and 6 show the more or less perpendicular direction of the stroke of the wing in the flight of
the bird (gull) —how the wing is gradually extended as it is elevated (1, 2, 3 of figure 5)—
how it descends as a long lever until it assumes the position indicated by 4 of figure 6—
how it is flexed towards the termination of the down stroke, as shown at 4, 5, 6 of figure
6, to convert it into a short lever (a b), and prepare it for making the up stroke. The dif-
ference in the length of the wing during flexion and extension is indicated by the short
and long levers w 6 and ¢ d of figure 6. The sudden conversion of the wing from a long into a
short lever at the end of the down stroke is of great importance, as it robs the wing of its
momentum, and prepares it for reversing its movements. Those same points are illus-
trated at figures 18 and 19, Plate XIV. At 4 of figure 19 the wing is represented as
fully extended, and in the middle of the down stroke. At 5 of the same figure the wing
is being flexed and slowed, and at 6 it is fully flexed, and its momentum destroyed. The
wing is then elevated as a short lever until it assumes the position indicated at 1’ of figure
18. It is subsequently elevated and extended, as shown at 2’ and 3’ (fig. 18). At 3” it i
transformed into a long lever, and in a condition to make a second down stroke. Figure 19
also shows the compound rotation of the wing—the tip of the wing rotating upon the axis
e d, and describing an arc of a circle, e f—the posterior margin of the wing rotating upon
the axis (a b), and describing the arc g h. The compound rotation of the wing occurs —
simultaneously with the down and up strokes, and it is to it that the great variety of ©
inclined surfaces made by the under surface of the wing is principally due.

Pruate XII.

Figure 7 is designed to show that the angle made by the under surface of the wing (more particularly j
at its root) with the lee is much greater than is generally sanpaseds This arises from
the fact that the body of the bird is inclined in an upward direction in flight, and
because the anterior margin of the wing (a) curves in a downward direction in such a
manner as to conceal the actual angle made. Thus, if e f be taken to represent the horizon,
the angle apparently made by the under surface of the wing with it isa@b d. The real
angle, however, isc bd.

Figure 8. The ine, or green plover (Vanellus cristatus, Meyer), with one wing fully extended (¢ b,
a6 fF) the other being in a semiflexed condition (d ef, ¢ 6). In the extended wing the
anterior or thick sane (d’ é f’) of the pinion is directed upwards and forwards can
arrow), the posterior or thin margin (¢ 6) downwards and backwards. The reverse of this
happens during flexion, the anterior or thick margin (d e f) of the pinion being directed
slightly downwards and forwards (vide arrow), the posterior or thin margin bearing the
rowing feathers slightly upwards and backwards. The wings, therefore, twist in opposite —
directions during extension and flexion. In the flexed condition of the wing the anterior
(def) and posterior (bc) margins are nearly on the same level, and the wing acts as
a short lever. In this condition of the pinion the primary or rowing feathers (0) are
separated from each other, and inclined obliquely upwards and outwards. (These feathers ©
are also shown at 1, 2, 3, 4, 5, 6, 7, 8, 0, 9 of figure 46, page 378.) When, therefore,
the wing ascends, the feathers in question (as well as the secondary feathers) cut into
the air like so many knives. They thus diminish the resistance experienced from the
superimposed air during the up stroke, a result to which the flexing or folding of the
wing and its conversion into a short lever contributes. From this account it will be
seen that when the wing is flexed the angles made by its under surface with the horizon
are diminished, whereas those made by the individual primary and secondary feathers are
increased. When the wing is extended it rotates in the direction of its length, the
anterior margin (de /f) being gradually inclined upwards and backwards, the posterio

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 445

one downwards and forwards. The rotation of the wing on its long axis during exten-
sion increases the angles made by the under surface of the wing with the horizon,
but decreases the angles made by the individual primary and secondary feathers, these
being made to flap together, and to assume amore or less horizontal position, as is well
shown at abcdefghijklmnopq of figure 48, page 378. This flapping together of the
primary and secondary feathers during extension effectually prevents the air from passing
between them. The power of the wing is greatly augmented during the down stroke—
Ist, by its being converted into a long lever; 2d, by the flapping of the feathers together ;
3d, by its under surface being rendered deeply concave (page 378, figure 48); and 4th,
by the various angles of inclination made by the several portions of the under surface
of the wing with the horizon being increased. These points are further illustrated at
ficures 16 and 17, Plate XIII. At figure 17 the margins of the primary (0 p) and
secondary (q) feathers, as seen in flexion, are given; whereas in figure 16 the flat of the
feathers (0 p qg), as seen in extension, are shown. These figures also show that, as the
angles made by the under surface of the wing with the horizon increase, the angles
made by the individual primary and secondary feathers (0 p q) decrease, and vice versa.
The angles made by the primary and secondary feathers are increased during the up
stroke, when the speed of the wing is slowed, and decreased during the down stroke,
when the speed is increased, an inclined surface, which forms a large angle with the horizon,
giving, when forced against the air at a low speed, the same amount of buoying power
as an inclined surface, which forms a smaller angle when urged at a lower speed.

Figures 9, 10, and 11 (Plate XIT.) show the wing of the gannet in the flexed, semiflexed, and extended
condition. Those figures are also intended to illustrate how the various inclined surfaces
made by different portions of the under surface of the wing in extension and flexion are
directed forwards, backwards, outwards, and inwards. ‘Thus in flexion and semiflexion
(figures 9 and 10), the portions of the wing marked g h and ¢ d, are inclined upwards and
inwards (vide arrows), whereas the portions marked e f and a b are inclined upwards and
outwards. When the wing is being extended, as in figure 10, the portions marked ef an
a b produce or draw after them a current, on which the portions marked ¢ d and g h operate
when the wing is being flexed, and vice versa. When the wing is fully extended, as at figure
11, the inclined surfaces indicated by g h, c d, e f, and a b of figures 9 and 10 disappear,
the under surface of the wing making a variety of inclined surfaces, which are directed
principally upwards and forwards, as shown at figure 16, Plate XIII. It is in this way
that the wing is capable of change of form in all its parts, and it will be observed that
those changes are induced irrespectively of any resistance experienced from the air.
When the wing ascends, it draws after it a current on which it operates when it descends;
and when the wing descends, it produces a current which assists in the elevation of the
wing. By the acts of flexion and extension, and by the down and up strokes, the wing of the
bird and bat produces the whirlwind on which it depends for support and progress. The
tip of the wing rotates upon ¢ of figures 9 and 10 (Plate XII.) as a centre, and by its alter-
nately darting in and out in flexion and extension, it describes the segment of a circle
(m n), and contributes to the stability of the bird by increasing the area of support.

The letter x in figures 9, 10, and 11 indicates the shoulder joint; s, the elbow joint; ¢, the wrist
joimt; v and w, the hand and finger joints; o p (fig. 11), the primary feathers; p gq,
the secondary feathers ; 7 the tertiary feathers ; xs tv w, the anterior margin of the wing;

: and o p qr the posterior margin.

Figure 12 shows how the wing is twisted upon itself structurally, and how the tip of the wing forms
an inclined surface, which is directed upwards and outwards (see arrows marked a and ).
x, mM, n, anterior margin of wing; 0 p q, posterior margin.

Puate XIII.

Figures 13, 14, and 15 represent the flight of the gull with the wings in the flexed, semi-flexed, and
extended conditions. The letters indicate the same parts of the wing in all the figures,
x representing the shoulder joint, s the elbow joint, ¢ the wrist joint, and v and w the
hand and finger joints ; o p the primary feathers, and q the secondary ones. At figure 15
the wings are fairly twisted upon themselves, and form true screws. In this figure the
pinions are extended to their utmost, and affording their maximum of support. They
are represented as they are seen at the middle of the down stroke. At figure 14 the

VOL. XXVI. PART II. on¥

444 DR PETTIGREW ON THE PHYSIOLOGY OF WINGS.

wings are slightly flexed and deeply concave on their under surfaces, the greater
concavity of the wings compensating in part for the diminution in their length. They
are also further depressed than in figure 15. At figure 13 the wings are represented
as seen at the end of the down stroke, the concavity of their under surfaces being still
more increased, and their length still more diminished. The wings are now short
levers, and prepared to make the up stroke, the great convexity of their upper sur-
faces diminishing the resistance which they experience from the superimposed air during
their ascent. Figures 13, 14, and 15 illustrate very clearly how the downward and for-
ward fall of the body during the up stroke contributes to the elevation of the wings.
Thus in figure 13 the body is up and the wings down. At figure 14 the body has fallen ‘
a little, and the wings are elevated and spread out more than in fig. 13. At figure 15

the body has fallen further, and the wings are spread out to their utmost, and on a level ¥
with the body. If we now turn to figure 18 of Plate XIV. we will see that the body
continues to fall and the wings to rise, as shown at 1, 2,3; 1’ 2’ 3’. At 3, 3’ of this
figure the wings are elevated to their utmost, and the body depressed to its utmost. The
wings are consequently in a position to make a new down stroke. From these figures it

will be evident that the wings and body rise and fall alternately, the fall of the body con-
tributing to the elevation of the wings, and the descent of the wings necessitating the
ascent of the body. It is in this way that the weight of the body comes to play an
important part in flight. The alternate waved tracks described by the wings and body in 44
flight are given at figure 14, page 344; a, ¢, e, g, i giving the undulaticns made by the f
wings; 1, 2, 3, 4, 5, those made by the body.

Figures 16 and 17 (Plate XIII.) show the wing in the extended and flexed condition in the gannet. :
In these figures the body of the bird is exactly in the same position. When the wing is
flexed, as in figure 17, it is crushed together, the tip of the wing (s, p, v, w) folding
beneath the central portion (p, q, t), the central portion and root (ws7) flapping together
on nearly the same plane. It is by this means that the wing is converted from a long
into a short lever. The flexing of the wing reduces the angles of inclination formed by
the several portions of the under surface of the wing with the horizon, and causes the
anterior (a, s, t, v, w) and posterior (0 p q) margins of the pinion to occupy nearly the
same level. It, however, increases the angles of inclination made by the primary and
secondary feathers, these changes being necessary to reduce the resistance experienced
from the air during the up stroke. When the wing is flexed, all its parts areina lax
condition, the wing being principally under the control of the elastic ligaments, the muscles
acting more especially during extension. When the wing is pushed away from the side of
the body, and extended as represented at flgure 16, the angles of inclination made by the
several portions of the under surface of the pinion with the horizon are increased, while
those made by the primary (0 p) and secondary (p q) feathers are diminished. The
pinion, moreover, is rendered more or less rigid. When the wing is fully extended, it
acts as a long lever (compare length of wing in figures 16 and 17). By increasing its
length, the wing also increases its power and speed towards the tip. It therefore attacks
the air with great violence during the down stroke, and insures a corresponding upward
recoil of the body. The angles of inclination made by the several portions of the under
surface of the wing with the horizon vary. Thus the angle made by the portion qs is
greater than that made by the portion p v, and thit made by p v greater than that made
by ow. The diminution and increase of the angles bears a fixed relation to the speed at
which the different portions of the wing travel, the angle always being greatest when the
speed is lowest, and vice versa. The change in the angles is principally due to the rota-
tion of the wing in the direction of its length, the posterior margin of the pinion rotating
round the anterior one in a downward direction during extension (figure 16, vide arrows),
and in an upward direction during flexion (figure 17, vide arrows). It is this rotation of
the wing upon its long axis which presents the upper or dorsal surface of the pinion to
the spectator in flexion (figure 17), and the under or ventral surface in extension (figure
16.) These points are further illustrated at figure 8, Plate XII. (see description of figure
8.) In figures 16 and 17 the same letters are affixed to the same portions of the wing
in both ; w representing the shoulder joints; s, the elbow joint ; ¢ the wrist joint; %
the hand and finger joints; o p, the primary feathers; p, g, the secondary feathers; 7,
the tertiary feathers ; «, s, ¢, v, w, the anterior margin of the pinion; 0 p q, the posterior
margin.

o icf

DR PETTIGREW ON THE PHYSIOLOGY OF WINGS. 445

Puate XIV-

Figures 18 and 19 represent the several positions assumed by the wing of the gull during extension
and flexion, and during the down and up strokes. Figure 19 also shows how the wing
during its ascent and descent rotates upon two axes. At 4 of figure 19 the wings are
represented as they appear at the middle of the down stroke. They are fully extended, and
affording their maximum of support. At 5 of this figure the wings are slightly flexed, ana
more deeply arched than at 4. They arealso ona lower level. At 6 the wings are represented
as they appear at the end of the down stroke. They are now fully flexed and form short levers.
They are also more deeply arched than at 5, a circumstance which prepares them for making
the up stroke, as the arching renders the upper or dorsal surfaces of the wings very markedty
convex. The wings, when in the positions indicated by 6 of figure 19, are elevated as
short levers, until they assume the positions indicated by 1, 1’ of figure 18. The wings
are then pushed away from the body, and extended and elevated, as shown at 2, 2’
and 3, 3’ (fig 18). At 3 3’ the wings are fully extended and fully elevated, and ready to make
the down stroke. They descend as long levers, until they assume the positions indicated
by 4 of figure 19, the changes in position just described being repeated in rapid succession
as the wings vibrate. The wings are flexed towards the termination of the down stroke
(5 and 6 of figure 19) to convert them into short levers, to destroy the momentum acquired
by them during their descent, and to prepare them for making the up stroke. They are
extended towards the termination of the up stroke (2, 2’; 3, 3’ of figure 18) to convert
them into long levers, and to prepare them for making the down stroke. Figure 18 repre-
sents the bird when it is flying vigorously, or when it is rising or picking up garbage
from the surface of the sea. In leisurely flight the wings do not rise much above the level
of the body, as shown at figure 19. In this case the wings are made to play rather under
than above the body (vide p. 374). The compound rotation of the wing is shown at figure
19, the wing rotating at its root (a) and along its anterior margin (c 5), the tip of the
wing describing the are of one circle (e bf), and the posterior margin of the wing the are
of another circle (gdh). The compound rotation of the wing is further illustrated at
figure 45, page 376.

Figure 20. Wing of the piet in the extended position.—In this figure the under lapping of the
primary (1 2345678 9) and secondary (jk2mnopqrs) feathers are shown, and how
the axis of each primary feather occupies a more and more central position in proportion
as it is placed nearer the secondary feathers. This want of symmetry in the primary
feathers is necessary to their valvular action during flexion and extension. The wing
during its vibration forces a certain portion of the air in waves along its under surface in
the direction of its root, as indicated by the arrows and dotted lines; the greater portion
of the air, however, is urged from the tip and posterior margin of the wing in a backward
and downward direction, the reaction propelling the body upwards and forwards. The
commotion produced in the air by the tip and posterior margin of the wing is on all
occasions very great, as the exposure of a flame behind or to the outside of the wing will
readily satisfy.

Prats XV.

Figures 21 and 22 represent the muscles and elastic ligaments of the wings of the snipe, as seen on
the ventral and dorsal aspects. In figure 21 (ventral aspect) the wing to the right of the
observer is fully extended, and the elastic ligaments put upon the stretch. The wing to

-the left of the observer is represented as flexed, the elastic ligaments being in a state of
contraction. The same points are illustrated at figure 22, which represents the dorsal
aspect of the bird. The wing is flexed principally by the action of the elastic ligaments.
It is extended chiefly by voluntary muscular efforts. Those figures show the difference in
the length of the wing in the extended and flexed condition, the pinion being a long lever
in extension, and a short one in flexion. That the elastic ligaments are subsidiary, and to
a certain extent under the control of the muscular system, is evident from the fact that
voluntary muscular fibres run into the ligaments in question. Thus the voluntary muscular
slip marked a in figure 21 terminates in the fibro-elastic band & ; this, again, being geared
to voluntary muscle x, and to certain musculo-fibrous bands 7. Their conjoined action is
to flex the forearm upon the arm, the arm being drawn towards the body by a musculo-

446 DR PETIGREW ON THE PHYSIOLOGY OF WINGS.

fibrous igament d, e. The elastic ligament g, 7 flexes the hand upon the forearm, and the
ligament 7 the fingers upon the hand.

Figure 23 shows the muscles and elastic ligaments in the wings of the pheasant, as seen on the dorsal
aspect, the wing to the right of the observer being fully extended, that to his left being
fully flexed. In the former the elastic ligaments are put upon the stretch ; in the latter,
they are in a state of contraction.

a, b, Voluntary muscular fibres, terminating in fibrous and elastic tissues ¢ and k. These structures
act in conjunction, and fold or flex the forearm on the arm.

Jf h, Voluntary muscular fibres, sending processes into elastic ligament g 7, to flex the hand upon the
forearm. The arm is drawn towards the body by the elastic ligament d, and by the
muscles v, w.

Puate XVI.

Figures 24 and 28 show the muscles and elastic ligaments, and the arrangement of the primary and
secondary feathers on the ventral and dorsal aspects of the wing of the crested crane.
The wing is in the extended condition in both cases.

a b, Voluntary muscular fibres terminating in elastic band &. This band splits up into two portions
(x, m, figure 24). A somewhat similar band is seen at j (figure 24). These three bands
are united to, and act in conjunction with, the great fibro elastic web c, to flex the fore-
arm on the arm.

tg,h, i, Musculo-fibro-elastic ligament, which envelopes the roots of the primary and secondary feathers.
The muscnlo- fibro- elastic ligament forms a symmetrical network of great strength and
beauty, its component parts being arranged in such a manner as to envelope the root of
each individual feather. The network in question supports the feathers, and limits their —
peculiar valvular action. It is enlarged at figures 25 and 27, and consists of three longi-
tudinal bands, 7 s,¢u,vw. Between these bands two oblique bands, g and h, run; the
oblique bands occurring between every two feathers. The marginal longitudinal band
(v, w) splits up into two processes, one of which curves round the root of each feather (2)
in a direction from right to left (a,b, c), the other in a direction from left to right (d,e, f).
These processes are also seen at m,n of figure 26. They have the root of each feather
completely under control, and their function, in conjunction with the oblique bands, is to
rotate the feathers from right to left during flexion, and from left to right during exten-
sion. The longitudinal and oblique bands are so geared together that they work in har-
mony, all the feathers enveloped by them being made to rotate in the same direction at
exactly the same instant of time. It is in virtue of the rotation of the individual primary
and secondary feathers at their roots that the feathers are separated from each other during
flexion, and brought into close contact during extension ; and thus it is that the air is
avoided during the up stroke, and seized during the down one. The primary and
secondary feathers are supported on their dorsal aspects by a series of subsidiary feathers
(mnop of figure 28), which are placed obliquely across their roots, and act as buffers.
The subsidiary feathers prevent the primary and secondary feathers from rising too far
during the down stroke.

Figures 25, 26, and 27. See under figures 24 and 28

CONTENTS.
NATURAL FLIGHT.

Introductory Remarks,

History of the figure of 8 theory o1 Wi ing Toye ment.

The Wing a Pricced lever or helix, .

The Wing twists and untwists during its action,

The image produced on the eye by the Wing in motion is concavo-convex wad fisted,
The iyade rotates on its long axis,

Gormpound rotation of the Wane) .
The Wing dnring its action reverses its see and deeenibes a ages ot 8 ies in space,

CONTENTS. 447

PAGE
The figure of 8 in rapid horizontal flight is opened out so that the Wing describes a looped
and then a waved track, ; 328, 340, 341, 342
Method of testing the eccrine of the Gaare of 8 sheory of Wine movements, : 334
Mode of Investigation pursued by the ation, é : : : : : 332
The Wing eapable of change of form in all its parts, 329
The Wing mobile and flexible as well as elastic—mobility ail Rerinilites Ssoseieu to flight, 365
The margins of the Wing thrown into opposite curves during extension and flexion, 328
The ine inclined pede at the end of the down stroke, aad backwards at the end of ote
up Boke : 335
The rotation of the sisheten margin of ae ae ma iguaman dhivecine increases the
elevating, but decreases the propelline: power, . : é : : : 368
The Wing attacks the air at various angles, . ; ; : : . . 33/, 383
The iyane during its vibration produces a cross pulsation, . ; 330
Analogy linea the Wing in motion and the sounding of sonorous aailee, ; . 330
The Wing during its oilina dene moves on the surface of an imaginary sphere, : : 343
The tip of the Wing describes an ellipse, . . : : . ° 330
The Wing vibrates Sneeneiley on either side of a nen line, . : : : 5 374
The natural Wing, when elevated and depressed, must move forwards, : : : 344
The Wing acts as a kite both during the down and up strokes, : : ; : 345
Where the kite formed by the Wing differs from the Boy’s kite, . : ; ; 346
Points wherein the Wing differs from the scull of the Boatman, ; : ; . 339, 340
A regulating power necessary in Flight, : ; : : 3 , : 390
_ The Wing at all times thoroughly under control, .. : : 3 : ; 391
Rapidity of Wing movements partly accounted for, . f . : ; A 399
How balancing is effected in flight, . ; . : ; ; : : 397
The Body and Wing move in opposite curves, ‘ : : : : ; 347
The Body ascends when the Wing descends, and vice versa, : ; : . 343, 352
Weight contributes to horizontal fli ght, 3 395
Weight necessary to flying neal as at present cadena: Weight and ler ity soli
considered with regard to aériel and subaquatic Flight ine ; 5 é 371
The Wing elevated indirectly. by the operation of one 3 ‘ : ; : 370
The Wing acts upon yielding fulera, i : : ; : 355
Bencideration of the forces which propel the Wing a the ieee , ‘ : ‘ 363
Analysis of the down and up strokes of the Wing of the Insect, . 347
The direction of the stroke of the Wing of the Insect, what effective, what non- Anni,
the kite-like action of the Wing, : : : 337
Mechanical theory of the action of the Insect’s Wine as stated by Chabrier, : 5 357
_ Objections thereto, . . 358, 364
Analysis of the movements of the Wie of the Wospanorecel of the Planes of the Wine
1” reciprocating action, &c., 338
The down and up pales of the ee of the Butterfly ; 2 merece at. destiomrten of the aya
area; development of figure of 8 curves on the margins of the Wing, . ; : 359
Analysis as the down and up strokes of the Wing of ae Bird and Bat, ; : 366
The Wing of the Bird descends as a long lever G 368), and ascends as a short lever, : 373
The importance to be attached to the concavo-convex form of Wing in Birds, : 369
The under or concave surface of the Wing of the Bird effective both during the down and a
strokes, 369
The Wing af the Bird Rane a nabaeal seneclues ‘pon ies the body depends both qunee
the aa and up strokes, : Sil
The Wing of the Bird cranked slightly Feapanndl compen rotation of the Quill F chen : 375
The primary, secondary, and tertiary Feathers are geared to each other, and-act in concert, . 376
They overlap or imbricate, . ; ; 380
The up or return stroke of the Wave of the Bad: sieve Aeon of Wing, : : 377
The Wing not always opened up to Hite same een in the up stroke, ; 381
Lax condition of the Shoulder Joint in Birds (p. 370); how the Wing is attached to the body ; :
movements of the Shoulder, Elbow, Wrist, and other Joints, . : : 393

VOL. XXVI. PART IL. Bz

448 | CONTENTS. |

The Wing flexed and partly elevated by the action of elastic ligaments—the nature and posi-
tion of said ligaments in the Pheasant, Snipe, Crested Crane, Swan, &c.,

The elastic ligaments more highly differentiated in Wings which vibrate rapidly,

Analysis of the movements of extension and flexion in the Wing of the Gannet,

Measurement, weight, &c., of Gannet and Heron,

Flight of Gannet as witnessed from the Bass Rock, .

ARTIFICIAL FLIGHT.

The Balloon, :

The Inclined Plane,

The Aérial Screw, :

Artificial Wings ; Borelli’s views,

Professor Marey’s views,

Chabrier’s views,

Straus-Durckheim’s views, E

The Author’s views—his method of consauciite and applyiue acacia Wines as coutmaae”
tinguished from that of Borelli, Durckheim, and nat aes

The Wave Wing of the Author, :

Compound Wave Wing ditto,

How to construct an artificial Wave Wing on une Tmeect oe

How to construct an artificial Wave Wing on the type of the bat and ah ;

How to construct an artificial Wave Ware which shall evade the cess of the air during
the up stroke,

Compound rotation of the paced Ways tne cane Paes ere of ie Wing fave ot
different speeds, }

How the Wave Wing creates onmrants and rises upon em oad how the air ace in elevating
the Wing, :

The Aérial Wave-screw of the Sexinon

How to apply artificial Wings to the air,

As to the nature of the forces required for propellfae artificial Wines :

Artificial Wings propelled at various degrees of speed during the down and up strokes,

Necessity for supplying the roots of artificial Wings with eee structures in imitation of the
muscles and elastic ligaments of flying animals,

XVI.—Additional Note on the Motion of a Heavy Body along the Circum-
Jerence of a Circle. By Epwarp Sane, Esq., F.R.S.E.

(Read 19th December 1870.)

In the twenty-fourth volume of the Society’s Transactions, a very convenient
formula is given for computing the time of oscillation in a circular arc ; and the
investigation of that formula is conducted by an appeal to the actual pheno-
mena. It is defective in so far that it contemplates chiefly the time of oscilla-
tion over the whole arc, and does not enable us conveniently to compute the
time in which a part of that arc is described.

The object of the present note is to supply that defect, and to present the
whole subject in a new aspect remarkable alike for its generality and for its
simplicity.

Referring to the first figure given in the paper cited, let N be the nadir and
Z the zenith point of a circle placed upright; and let us suppose a heavy
physical point to be projected from N along the circumference with a known
initial velocity, the object of our inquiry is to ascertain the law of its motion,
and to compute the time in which it describes a given arc.

If the initial velocity be due to a fall through the height NA greater than
the diameter, the body will reach the zenith point Z with a velocity due to a
descent through ZA, and will continue its motion along the other semicircum-
ference, reaching N with the same velocity as at first; thus it will circulate
along the whole circumference with a variable speed.

But if the initial velocity be that due to a fall through NB less than the
diameter, the body launched in the direction Nf will gradually lose speed until
it come to rest at the point F on a level with B. Thereafter it will descend
along FN, pass to the other side, and again return to N, repeating over and
over again its oscillation.

In the original paper a connection was established between cases of con-
tinuous and reciprocating motion. This connec-
tion may be more neatly traced by the following , ee a
scheme :—

Let a trigon have two sides AC, CB of

definite length jointed together at C, and let ¢
the angle ACB gradually change. Beginning
with AC, CB in a straight line, the angles A S

at A and C are each zero. As CAB in- Fig. 3.
creases, ABC also increases, until, if CA be the shorter leg, CAB becomes
VOL. XXVI. PART II. ak

450 MR EDWARD SANG ON THE MOTION OF A HEAVY BODY

a right angle, at which instant ABC has reached a maximum value. After
that, CAB still continuing to increase, ABC must decrease and become zero
Just when CAB becomes 180°. Afterwards, when
x CAB has become reverse, or greater than 180°,
one ABC appears on the other side and reaches its
A maximum value on that side when CAB = 270°.
Thereafter ABC decreases to become zero just
DS ee rar sae when CAB = 360°.
Thus the continuous growth of the angle
| CAB may typify the continuous motion, while the
reciprocating angle ABC typifies the oscillatory
ei motion of the heavy body. Seeing, then, that the
4 general phases of this arrangement represent the
leading characters of the two kinds of motion, —
aces we may inquire somewhat more narrowly into the
c

resemblance. For this purpose we shall put ACB,

Fig. 7 ;
= ree Jig. 9, for one form of the changeable trigon, and —
G ane: z imagine its form to be altered by turning the arm
ee CA into the closely approximate position Ca, so

that ACa becomes the decrement of the angle —
ACB, while ABa is the increment of ABC. These —
changes bemg supposed to be infinitesimally
minute, the arc Aa may be regarded as a short
straight line perpendicular to CA. Draw CP and
ae perpendicular to AB; then the minute trigon
Aea is similar to CPA, whence Aa:ae::AC:AP. Now the angle ACa is —

His. 9!

expressed by 34 , and ABa by aR wherefore

Ag 025 HAC AG.

ACa : ABa :: 5G ' api! aa: gp: AB: AP.

Hence, if we regard ACa as the differential of the exterior angle ECB, we
have

d@.ECB:d.ABC:: AB: AP, and consequently
@. ECB ad ACAB ? AB {PBs

so that the differentials of the three angles ECB, CAB, and ABC are propor —
tional respectively to AB, BP, and PA. If then we suppose the angle at A to
be generated with a velocity varying as the distance PB, that at B will be
generated with a velocity proportional to AP, and the exterior angle at C with

ALONG THE CIRCUMFERENCE OF A CIRCLE. 451
a velocity proportional to the whole subtense AB. Putting, for shortness’ sake,
BC = a, CA = 3b, we have CP = Od sina, and PB’ = @ — 0’ sin A’, that is,

dA
dt

dB
dt

a ./(a@ — 6 sin A’*), and similarly
a ,/(6 — @ sin B’).

When a body projected from N, with the
velocity due to a descent through AN, has
reached the point a, its velocity there is that
which is due to a descent from A to G on
the same level with a. Now, if we put A
for the diameter NZ, H for the whole height
NA, and A for the angle NZa, we have
ia = A.sin A, NG = A.sin A’, AG = H—
Asin A’. Wherefore, if g be the intensity of
gravitation as measured by the velocity which
a falling body acquires in one second,

—— = = /29-n/(H—A. sin A’), 01

yey = Nia xen’).

Hence, if we assume in our trigon ABC,

Wifes PB /29 ?

BA pred iad Ca Lawsolte tet
the expression = = /29. /(a —?’ sin A’) becomes identic with
dA :

GT = V2 (ie -3 —-—> x sin A®)on putting
Lola N ER ater Bloay. 5 err Gal: bas a?
ea Ne ee ES ge Oe

In order, then, to obtain the mechanical arrangement typified by the
variable trigon ABC, we must describe a circle having its diameter inversely
proportional to the square of AC, and suppose the initial velocity to be that
due to a descent through a height exceeding this diameter in the ratio a’: 6’.
This gives us the motion represented by the variable angle A. Again, we make
another circle having its diameter inversely proportional to a’, and take a height
less than this in the ratio of }?: a’; the oscillatory motion in this arc is repre-
sented by the variations of the smaller angle B; and the time of describing

452 MR EDWARD SANG ON THE MOTION OF A HEAVY BODY

the whole circumference in the one case is equal to the time of an oscillation
in the other case.

Farther, if we put C for half the exterior angle at c, that is, for the half
sum of A and B, we have

AB? = @ +8 + 2ab cos 2C
a + 2ab + & — 4ab sin C’;

now

= ,/2g.AB = ./2g ./ {(a + 6) — 4ab sin Ct, or

= = V/2%9 NEG 5 “y — ab pe cr,

wherefore if we put C for the half sum of a and 6, d for the mean proportional

between them, we have = = /29.,/(¢ —d@ sin C’), an equation identic in

form with that for the variation of A. Hence if we produce AC, draw CD,
jig. 10, bisecting the angle BCE, lay off CE a mean proportional between AC
and CB, and then inflect CD equal to the half sum of these same, the distance
QD, which is just half of AB, will represent the velocity with which the angle
ECD changes. At the same time CQ will represent the rate of change of the

angle at D, or
= = ,/29 /(@ —eé sm D’),

and thus we can obtain another pair of motions, one continuous, the other
alternate, synchronous with each other and with the preceding pair.

From this trigon CED we can, in the same way, that is, by making

ALONG THE CIRCUMFERENCE OF A CIRCLE. 453

EG = (CE. ED), GF = 4 (CE + ED), obtain a third trigon EGF ; and we
can continue this series of trigons indefinitely. The ratio of CE to ED is much
nearer to a ratio of equality than is AC: CB; EG: GF is still nearer, and after
a very few steps the ratio of say GI to IH becomes, sensibly, an equality. By
continuing the progression in the opposite direction, that is, by making
Ad = CB + ,/(CB’? — CA’), AC = CB — ,/(CB’ — CA’), we obtain trigons
more and more scalene, the ratio of disparity of the two sides increasing with
greater rapidity at each step. Hence of the general formula

= ds
dt G r/29 ae a (s? = yt sin s*) y)

we can by continuing this series either way, render s equal to 7, or greater or
less than 7 in any enormous degree. The integration in these three cases must
be considered separately.

In the first place, let us suppose that s, represented by cA, is infinitely small
in comparison with 7, represented by Ad. Here the arc S can never exceed a
certain infinitesimally small limit, so that it may be held to be equal to its sine,
and thus the formula becomes

dt,/(2g) = NCES

which is easily integrated, as in the familiar case of isochronous motions.
In the second place, when s, represented by Ad, is many times longer than
r, or Ac, the velocity of the moving point being proportional to gd, may be

held as constant; in which case the integral becomes ¢ ,/ (29) = - ; this limit

has been used in the previous paper. Both of these limits belong to the
inverse progression, which leads directly to the result already explained. I
shall, therefore, now direct attention to the third case, in which 7 and s have
been rendered equal to each other.

The equation now becomes

dS if
di,/(29) = Cee > secS.ds,

which has for its integral
i S
bay2g) = : log tan iG + 5)

so that if a heavy body be projected from the nadir-point of a circumference,
with a velocity just due to a free descent along the diameter, the time in which
it describes a given are is proportional to the meridional part on Mercator’s
projection of the sphere corresponding to a latitude homologous with the half
of that arc.

VOL. XXVI. PART TI. 6B

454 MR EDWARD SANG ON THE MOTION OF A HEAVY BODY
Thus, when we have continued the progression so far as to make GI = IH,
the time in which the angle GFE has been generated can readily be found.

In order to obtain a perfectly clear idea of this scheme of approximation,
let us draw through c, A, C, E, G, &c., lines perpendicular to CG, and inflect
to these Ad’ = Ad ; CB’= CB, ED’= ED, &c. ; then the angles d@’, B’, D’, F’ at
the extremities of these are evidently the maximum values of d, B, D, F; and
approach, toward the end of the series, more and more rapidly to a right
angle.

Suppose that we wish to compute the time in which an arc of 6° is described,
when the entire are of oscillation is 7° to each side of the vertical line, the
diameter of the circle being unit. Having made the angle B’Ad’ = 3°.. 35’,
since it is the angle at the circumference subtended by the arc of oscillation,

and measured Ad’ = a (<) in this case wit, we draw d’c perpendicular to

the horizontal line AB’; and then construct on the other side the trigon cAd,
having Ad = Ad’, and cdA = 3°; after which the formation of the series pro-
ceeds as already described.

The partial motion of the body through 6° of a total arc of 7° is now
synchronous with another motion through an are 2B of a whole arc 2B, but in
a different circle ; and lastly, it is synchronous with a motion through 2H", when
the body would just reach the zenith point of its circle. Hence the time would

be expressed by 7,/2g = = log tan (45°+ 4H). Hence the following scheme
of calculation :—
cA = -061 04854 8-785 6753 8-785 6753 33000 = I’
Ad = 1:000 00000 0-000 0000
1:061 04854 8:°785 6753
AC = :247 08000 9°392 8376 9-668 1324 27 00 26 = B’
CB = 53052427 9-724 7052 |
‘777 60427 9-117 5428
CE = +362 05235 9°5587714 9-969 0427 68 3721 =D’ |
ED = -388 80213 9:°589 7287
*750 85448 9:148 5001 /
EG = °375 18888 9:574 2500 9:999 7242 8757 30 = F” |
GF = -°375 42724 9-574 5258
‘750 61612 9:148 7758 |
GI = -375 30806 9:574 3879 0:000 0000 90 00 00 = H’
IH = :375 30806 9:574 3879
A = 7-099 44 0-851 2242

Here the data are Ad = 1:000, d’ = 3° 30’; having written these in their
places, and also the logarithm of Ad in the second column, the log sine of @

ALONG THE CIRCUMFERENCE OF A CIRCLE. 455

in the third column, we take the sum of these, which is the logarithm of cA,
and place it in the second column. The sums cA + Ad and log cA + log Ad
are next obtained. The half of the latter sum gives log AC, whence AC ; the
half of the former sum is CB, whence its logarithm. This very simple compu-
tation is repeated until we obtain GI and IH alike as far as our tables enable us
to proceed. Log AC — log CB is written in the third column, it is log sin B’,
whence we obtain B’, the maximum value of the angle B, and similarly for the
others. The maximum value of H is 90°, that is to say, since this is the angle
at the zenith point, the moving body would just describe the whole semicircum-
ference from N to Z. We have thus computed all the dimensions of the
diagram, fig. 10, on the left hand side of the line cl.

In order now to find the time in which the arc of 6° from the bottom of the
circle would be described, we make cdA = 3°. Adding to the log sine of this

the logarithm of = we obtain log sm Acd, whence Acd = 59° 00’ 46°.9
Half the sum of these angles is CAB; by adding to log sin CAB, log or

log sin B’, we obtain log sin B, whence B. And thus we proceed until we
Some to H = 21°37’ 38".
In the following scheme the calculations are given for the arc 6°, and also

for the whole are 7° :—

d= 3000000 8-718 8002 3 30 00:00 8-785 6753
1-214 3247 1:214 3247
c = 59 0046-90 9-933 1249 90 00 00:00 0-000 0000
A = 31002345 9-711 9215 46 45 00-00 9-862 3526
9-668 1324 9-668 1324
B = 13525307 9-380 0539 19 49 46°30 9-530 4850
CG = 22 26 3826 9-581 8127 33.17 53:15 9-739 5685
| 9-969 0427 9-969 0427
| D = 20492960 9°550 8554 30 44 43-40 9-708 6112
E = 21 3803-93 9-566 6532 32 0118-27 9-724 4733
9-999 7242 9-999 7242
F = 2137 11:98 9-566 3774 3159 56.40 9-724 1975

G = 21 37 37-96 32 00 37°34
90 90 ;

111 37 37-96 122 00 37°34
L. tan 55 48 48-98 = 0-167 9689 61 00 18-67 = 0-256 3407
Log log tan (4 + =) = 9-225 2289 9-408 8176
Log nep tan 10 = 0°362 2157 0-362 2157
Colog IH — 0-425 6121 0-425 6121
1-030 520 0-013 0567 1572 698 0-196 6454

456 MR EDWARD SANG ON THE MOTION OF A HEAVY BODY

To obtain a clear view of the import of these results, let us describe a series
of circles all having the common nadir point N, and with diameters proportional
inversely to the squares of Ad, CB, ED, GF, IH, or, in the present example,
proportional directly to the numbers 1:000, 3°553, 6615, 7:095, and 7-099.
Along these let ares NU, NV, NX, NY, NZ, of 7°, 54° 01’, 187° 15’, 175° 557,
180°, that is homologous to the doubles of the maximum angles d’, B’, D, F,

N
Fig. 11.

and G’ be laid off. Lastly, lay off arcs N', N? on each of these circles, homolo-
gous to the doubles of the angles at d, B, D, F, and H, as obtained by com-
putation in the two cases. Im the diagram, jig. 11, the semicircles only are

drawn out ; for the sake of clearness, they are placed alternately to the right ~

and left of the common diameter NZ; also in the smallest circle the arc NU
of 7° is also N’, and N? is so close to it as not to be seen in the figure.

Let now a body be projected along the smallest circumference with a
velocity which would just carry it from N to U, another along the second cir-
cumference with a velocity sufficient to carry it to V, a third body along the
third circumference with speed enough to reach X, and so on. And imagine
that all these bodies set off simultaneously from N, then will they all reach the
points marked 1, 2 on the respective circumferences at the same instants. But
when the velocity is just sufficient to carry the body to the zenith point Z, we
can compute the time, and thus we are enabled to resolve our problem.

As there are no tables of neperian log tangents, we must convert the
usual denary log tangents into neperian ones, by multiplying them by 2°3025,
the neperian logarithm of ten. Hence the final computations shown in the
preceding scheme ; these give 1:030520 and 1:572698 for the values of ¢,/(29),
corresponding to ares of 6° and 7° respectively.

Thus, by a very easily understood and by no means an operose process, we

ALONG THE CIRCUMFERENCE OF A CIRCLE. 457

are enabled to compute the time at which a heavy body moving along the cir- -
cumference of a circle will reach a given point in it, when the velocity is
insufficient to carry the mass over the zenith point.

The other case, when the body passes the zenith point, is resolved by con-
sidering the angles at a, A, C, &c. Thus, if we describe a circle having its
diameter inversely proportional to the square of Ac, and imagine a body to fall
from a height greater than that diameter in the ratio of dA : Ac, and to be pro-
jected along the circumference with the velocity so acquired, its motion would
be represented by the variation of the angle Acd.

But this same investigation enables us also to resolve the converse problem:
“to find the position of the body at any given instant,” for since the time of
describing any portion of the arc NZ, fig. 11 is proportional to a logarithmic
tangent, we can easily compute the log tangent, and thence the arc corre-
sponding to a given time; and having thus obtained the angle IHG of jigure
10, we can deduce, by the operations of ordinary trigonometry, all the other
angles, and thus the positions of the various bodies at the proposed instant on
all the other circumferences can be found.

In retracing backwards the series of trigons, beginning with GIH, we must
make IGF double of IGH, GED double of GEF, and so on; hence we must
soon obtain an obtuse angle, the double of which would be a reverse angle,
and it would seem as if our process failed. But in reality this reverse angle
merely shows that the moving body has overpassed the zenith point, and has
begun to descend along the other circumference, and thus our construction
turns out to be absolutely general.

When one of the angles, as ECD of jigure 10, becomes right, its double
ECD becomes half a revolution, and so CAB and CBA become zeroes.
Wherefore when the motion represented by the trigon CED has performed half
of its period, the system typified by CAB has made a whole one. Thus at each
step downwards along the series the periodic time is doubled ; and if we wish
_ to keep the same periodic time throughout, we must halve the dimensions of
each successive trigon—an operation which brings us exactly to the conclusions
obtained in the original paper.

VOL. XXVI. PART IL. 6 ¢

———

( 459 )

XVIL—On the Homological Relations of the Coelenterata.

By Professor ALLMAN.
(Read 29th May 1871.)

Independently of the general agreement which necessitates the association of
the Hydra, Actinia, and other Ceelenterate animals into one primary group of the
animal kingdom, we must also expect a special morphological correspondence
between the various forms of animals thus associated. In other words, a homo-
logical agreement ought to be determinable between the parts of animals included
in any one subordinate section of the C@LENTERATA with the parts of animals
included in any other.

A comparison of the two primary sections of the C@LENTERATA (Actinozoa
and Hydrozoa), and of the various orders of these with one another, will show
that such an agreement really exists, and that it is possible, by easily under-
stood and thoroughly consistent modifications, to convert each special type into
any of the others.

With the view of rendering apparent these relations, we shall compare an
actinozoon (Actinia) with a hydrozoon (Hydra), and shall further compare with
one another the various orders of the HypRozoa.

Agassiz has compared the radiating chambers, which in an actinozoon inter-
vene between the stomach sac and the outer walls, with the radiating canals of a
medusa.* I believe that he has thus struck upon the true homologies of those
parts; but when he maintains further that the differentiated stomach of an
actinozoon is only the proboscis (hypostome) of a hydrozoon inverted into its
body cavity, he suggests a conception of actinozoal homology which is incon-
sistent with the actual structure.

In order to form a correct notion of the homological relations between an
Actinia and a Hydra, we have to imagine the tentacles of a Hydra (figs. 3, 4) for
a greater or less extent connate with the sides of the hypostome and with one
another. The hypostome of the Hydra, while retaining its normal position, will
thus become the stomach of the Actinia (figs. 1, 2, 6), and this will at the same
time become connected with the outer walls by a series of radiating lamellee—
the connate tentacle-walls,—separated from one another by radiating chambers
a—the cavities of the tentacles,—while such portions of the tentacles of the

* Contr. Nat. Hist. U.S. vol. iv. p. 377.
VOL. XXVI. PART. II. 6 D

460 PROFESSOR ALLMAN ON THE RELATIONS OF THE CHLENTERATA.

Hydra as still continue free will be represented by a single circle of the ten-
tacles aw of Actinia.

Fig. 1.—Diagramatic longitudinal section of Actinia. «a, Radiating interseptal space ; a’, tentacle ; b, differentiated
stomach-sac ; 0’, somatic cavity ; c, aperture in radiating septa; d, genitalia borne by radiating septa.

Fig. 2.—Diagramatic transverse section of Actinia. «a, a, Interseptal spaces ; b, differentiated stomach-sac.

Having thus established a fundamental identity between the regions of an
Actinia and of a Hydra, there will be no difficulty in recognising the relations
between an Actinia and a hydroid medusa ; for, as I have elsewhere* attempted
to prove, the tentacles of a Hydra are represented by the radiating canals (figs.

Fig. 3.

Fig. 4.

Fig. 3.—Diagramatic longitudinal section of Hydra. a, Tentacle; b, hypostome ; b', somatic cavity.
Fig. 4.—Diagramatic transverse section of Hydra through hypostome and tentacles. «, Tentacle ; b, hypostome.

», 6, a), and those extensions of them (fig. 5, a’) which form the primary mar-
ginal tentacles of the medusa.

The distal ends of the radiating lamellee in Actinia are perforated each by
an opening (fig. 1, ¢), through which the radiating chambers communicate
with one another. Agassiz has compared these openings to the circular canal

* Report on the Reproductive System of the Hydroida. Brit. Assoc. Report for 1863.

PROFESSOR ALLMAN ON THE RELATIONS OF THE CHHLENTERATA. 461

of a medusa, and I believe that in this view he has correctly expressed the
relations in question.
If we further add that the generative apparatus is borne by the radiating

Fig. 5.—Diagramatic longitudinal section of a Hydroid Medusa. w, Radiating canal ; a, marginal tentacle ; 4,
manubrium ; 0’, atrium; c, lumen of circular canal ; d, generative elements; 7, atrial region of umbrella ; 7’, manubrial
region of umbrella ; v, velum.

Fig. 6.—Diagramatic transverse section of Hydroid Medusa through the manubrial region of the umbrella. «a, Radi-
ating canal; 6, manubrium; d, generative elements; 7’, manubrial region of umbrella.

partitions, we shall have all the leading points in the morphology of an actino-
ZOOn.

A comparison of the various orders of the Hydrozoa with one another will
result in the detection of close homological correspondencies, and will throw
important light on the morphology of each.

Between a siphonophore and a hydroid the homology is so obvious as to be
instantly recognisable. The siphonophore (fig. 7), as well as the hydroid, pre-
sents us with a colony of zooids, kept in organic union with one another by
means of a common connecting basis or ccenosarce ; but this coenosare, instead of
being fixed, as in the Hyprorpa, is in the SrpHonopHora invariably free, and
provided with a special apparatus for natation.

In consequence of the great extent to which heteromorphism is carried
among the zooids composing a siphonophoral colony, we can scarcely institute
a satisfactory comparison between the two orders without determining the
homologies of each kind of zooid in the siphonophore. Beginning with the poly-
pites or alimentary zooids (¢) of the siphonophore, and comparing these with the
hydranths of a hydroid, we shall find the two forms to agree in almost every
point, except in the number and position of the tentacles, which in the siphono-
phore are reduced toa single one (/), springing, in all the typical SrpHonopHora,
from the base or proximal end of the polypite. The branched condition of the
tentacle in the siphonophore is in no respect inconsistent with this comparison ;
and even if it were necessary to find a parallel to it among the HyproIpa, we
should have this in the branching tentacles of Cladocoryne.

462 PROFESSOR ALLMAN ON THE RELATIONS OF THE CQILENTERATA.

The hydrocysts (g) of the siphonophore are plainly arrested polypites, in
which the mouth has never become developed.

Again, the generative zooids (2) are exactly paralleled by those of the Hy-
pDRoIDA, and are, like them, referable to two
types, expressed in the Hydroida by the phane-
rocodonic and the adelocodonic gonophores, the
situation of the generative elements being pre-
cisely similar in the two orders ; while the necto-
calices or locomotor zooids (/, 4) are essentially
hydroid medusa, with specially developed um-
brella, but with the manubrium suppressed, and
the somatic cavity reduced to the atrium, from
which spring radiating canals, which, exactly as
in the hydroid medusa, open round the margin
into a circular canal.

The bracts or hydrophyleia (/) of the siphono-
phore are essentially ceecal offsets from the com-
mon canal of the ccenosarc, but with the ecto-
derm greatly developed and modified, as in the
umbrella of a medusa, so as to fit them to become
organs of protection for the other zooids. They
have thus essentially the same morphological
Fig. 7.—Diagram of a Siphonophore. _ foundation as the nectocalices, but, with a dif-

e, Polypite; f, tentacle springing from

proximal end of polypite; f', branches ferent functional destination, diverge widely from
given off by the tentacle ; g, hydrocyst; h,

tentacle of hydrocyst ; ¢, generative zooid these, and constitute an apparatus of protection
Slee et ane 7, instead of locomotion.
bract 5 m, m, ccenosare ; n, pneumatocyst. : Z E 5

All these zooids are kept in union with one
another by a coenosare (m, m), which, in the typical S1pHonopHora, is fili-
form, with an axial canal in free communication with the cavity of each
of its appended zooids, thus corresponding essentially with the filiform
tubular ccenosarc of a hydroid colony; while in the somewhat aberrant
forms with fusiform or discoidal coenosare (Physalide, Velellide), an ob-
vious comparison is suggested with the appressed expanded ccenosarc of
Hydractinia.

From the hydroid ccenosarc, indeed, that of the SrpHonoPHORA mainly
differs in the absence of an external chitinous sheath, and in its free mode of
existence, the siphonophore dwelling at large in the open sea, through
which, in the great majority of the order, it is propelled by the contrac-
tions of the nectocalices. In the siphonophorous section, Physophoride,
the proximal extremity of the ccenosarc, instead of forming, as in the
Hydroida, a hydrorhiza for fixation, is modified by an inversion of its

Fig. 7.

PROFESSOR ALLMAN ON THE RELATIONS OF THE CQ@BLENTERATA. 463

walls, so as to constitute an air-filled chamber (pneumatocyst) (7), which acts
as a float.*

Continuing to take the Hyproipa as a standard of comparison, the other
hydrozoal orders may be now contrasted with them. If the atrium, or that
portion of the somatic cavity (fig. 5, 0°) which lies at the base of the manu-
brium in a hydroid medusa, be expanded laterally, and the ectoderm of its
floor be projected along four or eight symmetrically disposed radiating lines
into as many thick pillars (figs. 8 and 9, 0, 0), which converge towards the axis,
and there meet the manubrial extension of the cavity, while the thin interven-
ing portions of the floor between the pillars become developed into generative
pouches (d), and the velum or perforated diaphragm, which stretches across
the codonostome in the hydroid, disappears, we shall have the hydroid medusa
converted, in the more essential points of its structure, into a discophorous
medusa (figs. 8, 9).

Again, a Lucernaria (figs. 10, 11) may be conceived of by imagining a hydra

Fig. 8.

Fig. 8.—Diagramatic longitudinal section of a Discophorous Medusa. a, Radiating canal; 8, manubrium ; J,
somatic cavity ; d, generative pouches ; 9, 0, 0, pillar-like extensions of the oral side of the umbrella; z, tentacula-like
processes of the inner surface of the somatic cavity.

Fig. 9.—Diagramatic transverse section of a Discophorous Medusa. «a, a, a, Radiating canals; 6, manubrium
d, d, generative pouches ; 0, 0, umbrello-manubrial pillars.

with four tentacles to have these tentacles expanded laterally, until their sides
meet and coalesce, the hypostome still continuing free, and the proximal
portion of the body becoming extended into a solid peduncle of attachment,
containing a simple prolongation of the somatic cavity, or traversed longitudi-
nally by four narrow prolongations of this cavity, while generative sacs become
developed on each side of the partitions formed by the coalescent sides of the
tentacles.

* In the above comparison of the siphonophora with the hydroida, I have adopted for the sipho-
nophora the terminology proposed by Huxiry, whose views of the homological relations existing
between the two orders I have also generally followed. See his “ Oceanic Hydrozoa,” page 8, &c.

MOT XV, PART IT. 6E

464 PROFESSOR ALLMAN ON THE RELATIONS OF THE C@LENTERATA.

Lastly, the CTENOPHORA (figs. 12, 13) admit of an obvious comparison with a
hydroid medusa. In order to understand this, we must keep in mind the pre-
sence in the hydroid medusa of an atrial segment of the somatic cavity. This
is formed by that portion of the somatic cavity which is immersed in the sub-
stance of the umbrella at the base of the manubrium, and from which the
radiating canals proceed (fig. 5, 6’). The hydroid medusa thus admits of a
division, by a transverse plane, into two regions: an atrial region (7), which
corresponds to the solid summit of the umbrella with the parts therein con-
tained, and a manubrial region (7), which corresponds to the manubrium, with
that portion of the umbrella which with its associated structures is projected
round the manubrium in the form of a bell.

Now, in a Beroe (figs. 12, 13), the manubrial region is never developed, and
the body is represented by the atrial region alone. From the atrium (0 0/ ) con-

Fig. 10.

Fig. 10.—Diagramatic longitudinal section of Zucernaria. a, Cireum-oral dise ; a’, marginal tentacle ; b, hypos-
tome; 0’, somatic cavity; c, aperture by which the chambers of the circum-oral dise communicate with one another
aeross the distal end of the partition ; d, generative bands; p, peduncle ; z, tentacle-like processes of the inner surface
of the somatic cavity.

Fig. 11.—Diagramatic transverse section of Zwcernaria across the circum-oral dise and hypostome. a, a, Cham
bers of the dise ; 6, hypostome ; d, generative bands.
tained within this region two radiating canals (a, a) are given off. These imme-
diately divide and subdivide, so as to become ultimately eight, which are,
moreover, united at their distal extremities by a circular canal, which corre-
sponds to that of the medusa, though here thrown back by the non-develop-
ment of the manubrial region of the umbrella. Besides the eight longitudinal
canals (x, #) into which the two radiating canals ultimately subdivide, these two
canals give off, each immediately after its origin, an accessory canal (2 2’),
which runs without division close to the main body cavity towards the oral
orifice, and opens, like the others, into the circular canal.

The generative sacs (d’, d) are developed as diverticula along the course of
the radiating canals, whence they extend into the gelatinous substance of the body.

PROFESSOR ALLMAN ON THE RELATIONS OF THE C@LENTERATA. 465

LEUCKART insisted on the association of the CTENoPHORA with the AcTINo-
ZoA rather than with the Hyprozoa, and the same view of their affinities has
been advocated by Huxtey. According to this conception of ctenophoral
homologies, the ctenophore must be provided with a stomach-sac, differentiated,
as in the actinozoon, from the general body cavity. Now, though the somatic
cavity in Beroe suddenly diminishes towards the aboral end, and is there pro-
vided with a pair of valve-like folds (fig. 12, s), so that the entire tract admits
of being distinguished into two regions, it is nevertheless as continuous and
simple as in Hydra.

The advocates of the actinozoal nature of the Ctenophora see in the canal
system of a Beroe or a Cydippe the radiating chambers of an Actinia, separated
from one another by partitions of relatively enormous thickness. I do not
desire to dispute the correctness of this view. We have already compared a

Fig. 12.— Diagramatic longitudinal section of Beroe in a plane at right angles to that of the compressed somatic cavity.
In order to give a sufficiently comprehensive view of the structure, a few parts are here represented, which are in reality
somewhat removed from the plane of the section. a, a, Transverse portion of the radiating canal system, two of the
primary branches being shown as if cut off close to their origin ; x, x, meridional portion of this system ; z'a/, deep or
accessory canals, their distal ends cut off ; 0’ b', somatic cavity ; c, lumen of circular canal ; ¢, one of the aboral outlets
of the somatic cavity ; b'b’, somatic cavity; ¢, external opening of one of the aboral canals ; s, valve-like processes of
the inner surface of the somatic cavity ; d, d', generative sacs, male and female.

Fig. 13.—Diagramatic transverse section of Beroe. 0’, Somatic cavity ; 7, z, meridional portion of the radiating
canal system ; 2’, 2’, deep or accessory canals ; d, d’, generative sacs, male and female ; y, vibratile lamelle.

hydroid with an actinozoon, and have seen in the radiating canals of a hydroid
medusa the homologues of the radiating chambers of an actinia ; so that, even
though the CrENopHora be truly Hyprozoa, we must expect to find in them the
same points of agreement with the AcTINozoa which we have endeavoured to
demonstrate for the other hydrozoal orders.

Now, the fact of the radiating canals being widely separated from the axial
cavity instead of being adnate to it, is exactly the point which essentially dis-
tinguishes a hydrozoon from an actinozoon ; and the fact of the intervening
space being in the ctenophore obliterated by the interposition of a voluminous
gelatiniform mass does not alter this relation, for it is exactly what we find in

466 PROFESSOR ALLMAN ON THE RELATIONS OF THE CCQRLENTERATA,

the atrial region of an ordinary hydroid medusa, while it is distinctly expressed
in the gonophore of clavatella, where the free or manubrial region of the
umbrella is rudimental, and the whole gonophore, apart from the marginal
tentacles, becomes comparable to the atrial region of an ordinary hydroid
medusa. :

The accessory canals of Beroe run, it is true, close upon the walls of the
axial cavity until they leave these to throw themselves into the circular canal ;
but this fact cannot, in opposition to the greatly preponderating hydrozoal
features of Beroe be used as an argument for the actinozoal nature of the
CTENOPHORA.

The accessory canals are not represented in the hydroid, while the Beroe
further differs from the hydroid in the presence of the two short aboral canals,
by which the aboral end of its somatic cavity communicates with the outer
world (fig. 12, ¢), as well as in the disposition of its so-called nervous system
and sense organs, and in its characteristic bands of vibratile lamellee (fig. 13, y);
all which features are among the special characteristics of the order, and in no
way justify the absorption of the CTENoPHORA into the ACTINOZOA.

In this attempt to determine the true affinities of the CreNopHora, I have
taken Beroe, instead of Cydippe or other ctenophorous genus, as the subject of
comparison, not only because Beroe is a typical ctenophorous form, and of com-
paratively simple structure, but because I have myself made its anatomy and
development a subject of special study.*

* Proc. Roy. Soc. Edinb., 1862,

G0r4G7 ei)

XVIIL.—On the Gravid Uterus and on the Arrangement of the Foetal Membranes
in the Cetacea. Plates XVII. and XVIII. By Professor TURNER.

(Received 20th March 1871.—-Read 8d April),

CONTENTS.
PAGE | 3 PAGE
Introduction, ; ‘ ‘ : . 467 Comparison of Placentation with that of
Uterus and Appendages : : . 470 other Mammals, agian ¢ . 486
Foetal Membranes, ; > 5 . 478 Physiological Conclusions, . : . 498
Position and General Form of Foetus, . 484

The distinguished French naturalist, Professor H. Mitne Epwarps, in the
ninth volume of his valuable Lectures on Comparative Anatomy and Physiology,
published only last year, when referring to the foetal membranes in the Cetacea,
states, that much information is still required to complete our knowledge of
that subject.*

It may perhaps be advisable, before I commence to describe the results
arrived at by my recent dissections, to give a brief account of the observations
made by previous inquirers into this department of anatomy, so that we may
more clearly recognise wherein our deficiencies le, and the direction in which
our researches ought to be conducted, in order to render our information as
complete as possible.

Karu Ernst von Baer in his celebrated memoir, “ Ueber Entwicklungs-
geschichte der Thiere,”+ says, “ I know nothing of the ovum of the cetacea from
my own observations ; the scanty notices which we find on this subject in
anatomical literature at least show that there is no definite placenta, and lead
us therefore to suppose that the ovum is similar to that of the pachydermata.”

D. F. Escuricut in an academic dissertation, published in the same year as
Von Barr’s memoir, recorded the dissection ‘‘ Delphini phoceene gravidi.”t He
described the free surface of the uterine mucous membrane as rugose, cellular,
and cribriform. The surface of the chorion was strongly marked by ruge,
which could not be obliterated. Almost the whole surface was covered by*
villi, which were separated from each other by intervals of nearly half a line.
| The villi possessed narrow stalks, and their free ends expanded into a globular
| branching crown, like the head of a cauliflower. In the hollows between the
Tugee very small villi were found. A beautiful capillary network was situated
within the crowns of the villi. The villi were adapted to the little recesses or

* Lecons sur l’anatomie comparée, vol. ix. note, p. 563. Paris, 1870.
+ Second part, p. 257. Kénigsberg, 1837.
De organis que respirationi et nutritioni foetus mammalium inserviunt, Hafnie, 1837.

VOL. XXVI. PART II. 6F

468 PROFESSOR TURNER ON THE GRAVID UTERUS AND

cells seen on the surface of the uterine mucous membrane, which surface was
very vascular. He also observed a layer of branching uterine glands, which
were so numerous and so closely set together that it was difficult to trace
out single specimens. The mouths of these glands opened into areole on the
uterine mucous surface, and he believes that their secretion is taken up by the
veins of the villi of the chorion, which are in apposition with these mouths, and
that the nutrition of the foetus is sufficiently provided for by the absorption of
this secretion.

Professor OWEN, in a note to JoHN HuNTER’s description of the parts of
generation of the cetacea,* states that the allantois is co-extensive in its
development with the chorion, and that both extend into the horns of the
uterus. The foetus has neither placenta nor cotyledons ; but, as in the hog and
camel, the general vascularity of the chorion is subservient to its nutrition and
respiration. In a foot-note on a previous page (70), he had characterised the
placenta in the sow and the mare as diffused over nearly the whole surface of
‘the chorion. In the catalogue of the comparative anatomy specimens in the
Museum of the College of Surgeons,t he mentions the presence of peduncu-
lated corpuscles of the amnios on the umbilical cord of a foetal dolphin (D.
delphis).

Dr C. D. Metes, in the course of some observations on the reproductive
organs and the foetus of Delphinus nesermak,t observed the plicated arrange-
ment of the uterine mucous membrane, and of the corresponding surface
of the chorion, the projections and sulci of the one being accurately adapted to
the sulci and projections of the other, so that the real surface of contact very
much exceeded the apparent surface. The foetus was developed in the left
uterine cornu, which was larger than the right, though the latter was partially
developed by the intrusion into its cavity of the chorion and allantois. The
amniotic outgrowths are figured on the umbilical cord, but are not described.

Professor RotiEsTon also directed attention to the prolongation of the
membranes of a solitary cetacean embryo,§ which he had examined, “from one
cornu round into the other, and projecting by a ccecal extremity into a
short corpus uteri.” He observed and described filiform outgrowths of
the amnion, where it invested the umbilical cord, and pointed out that the
cornual ends of the cetacean membranes were bare and glabrous as compared
with the villous character of the rest of the chorion.

* Collected works, Palmer’s Edition, vol. iv. p. 390. 1837.

t+ Vol. v. p. 200. 1840. Comp. Anat. of Vertebrates, vol. iii. p. 732.

t Journal Academy Natural Sciences of Philadelphia, 1849, vol. i. p. 267. I have not seen
this paper, and am indebted to my friend Dr Rotiaston for the above abstract of its contents. In
a paper in the Proceedings of the same Academy, vol. iv., 7th August 1849, Dr Mates related some
experiments made to ascertain the effects of deep-sea pressure on the uterus of the cetacea.

§ Trans. Zool. Soc. 1866, v. p. 307. The species was not determined.

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 469

In the account, necessarily very imperfect, owing to the greatly injured
condition of the membranes, which I gave of the chorion in the Longniddry
whale (Balenoptera Sibbaldii *), I described the folds and ridges of the chorion,
the villous character of its surface, and the presence of at least one well-defined
spot free from villi Like my predecessors, I was unable to determine the
relations of the amnios and allantois to each other and to the chorion. In the
present communication I hope to be able to supply not only these, but other
important blanks in our knowledge of the arrangement of the foetal membranes
in this interesting group of mammals.

In the early part of February of the present year, a shoal of whales entered
Bressay Sound, Shetland, on the cessation of a heavy storm, which had raged
for many hours. The fishermen at once started in pursuit, and succeeded in
driving the shoal ashore in a bay to the north of Lerwick, with the exception
of one specimen, which sank before it reached the beach. The animals stranded
were eighteen in number.

Through the great courtesy of Mr James GATHERER, collector of customs in
Lerwick, a gentleman well known to many naturalists as a careful and zealous
observer, I have not only learned the following interesting particulars respecting
these animals, but have also had the good fortune to receive the gravid uterus
of a pregnant female. Mr GaTuerer writes, “‘ The animals ranged in length from
17 feet to 24 feet. The Shetlanders call them the ‘ spotted caaing whale,’ or the
‘fleckit whale,’ or the ‘ pict whale,’ and in one locality the ‘ Lupster,’ a term which
has a Norse sound, and is possibly of Norse origin. Though ‘ caaing’ (driving)
whales, yet they are not the common ‘caaing’ whales of the Shetlanders (Go-
biocephalus deductor). The more prominent dorsal and the shorter pectoral
fins, the less rounded head and muzzle, and the piebald colour, show a marked
difference between them and the ‘ caaing’ whales, so frequently and in such
_ large numbers driven ashore on the Shetland coast. I suppose it will be
found that they were the Grampus (Phocena orca). The Shetlanders know
very little about them, although small herds have on several occasions been
driven ashore in different voes throughout the islands. It is seldom so many
or so large specimens are driven. The natives consider them far more active,
wary, and dangerous than the ‘caaing’ whale. They tell me a few are some-
times seen mingled with a herd of the ‘ caaing’ whales, on which occasions they
fail to drive the latter. They attribute their escape to the superior retreating

tactics of their more wide-awake congeners, who take the lead.

“TJ laid open the stomachs of two of the animals with the hope of finding
some evidence of the nature of their food, but with the exception of a large
number of worms, and some green frothy matter, the stomachs were empty.
We found a foetus lying on the beach which some of the flensers had extracted
* Proc. Roy. Soc. Edinburgh, 20th December 1869, and Transactions for 1870.

470 PROFESSOR TURNER ON THE GRAVID UTERUS AND

from the womb. Observing another female in an apparently ‘interesting con-
dition,’ we got one of the men to make a longitudinal opening in the abdomen,
which exposed a gravid uterus of large dimensions. To render the transmis-
sion of the parts safer, I ran off the uterine liquor before packing the uterus in
a barrel. I found the number of teeth on each side of each jaw to be eleven,
or forty-four in all. The teeth of many, I suppose the adults, were quite flat,
or entirely worn down until flush with the gums. When this was not the case,
they were of considerable length, and the teeth of one jaw fitted into the
intervals between the teeth of the jaw opposed to it.”

From the examination of the foetus, and of the skull of one of the adult animals,
I can substantiate the supposition of Mr GATHERER, that these animals were
Orcas. By recent systematic writers the Orca, or Killer Whale, is no longer
included within either the genus Grampus, or Delphinus, or Phocena, with
one or other of which it had been associated by many naturalists. . The special
characters which it exhibits are now considered to have a generic value, and
the name Orca gladiator is applied to this creature.

I shall now pass to the description of the specimen, and shall consider—
lst, the uterus and appendages; 2d, the foetal membranes ; 3d, the position
and general form of the foetus; 4th, a comparison of the cetacean form of
placentation with that of other mammals.

The Uterus and Appendages.—The uterus consisted of a cervix, a corpus,
and of two cornua (Plate X VII. fig. 1). The various subdivisions of the organ
were invested by a continuous layer of strong peritoneal membrane, which
extended in a broad double fold from the concave border of each cornu to
the side and surfaces of the cervix uteri. The cervix was 7 inches long, and
the corpus uteri 14 inches. The two cornua curved outwards from the body,
which seemed indeed to bifurcate at its anterior border into the two horns.
The left horn, about twice the size of the right, measured, along the convexity
of the curve, 6 feet 7 inches from the angle of bifurcation to the junction of
the tip of the horn with the Fallopian tube. The right cornu, along the cor-
responding border, measured 3 feet 6 inches. The greatest breadth of the left
cornu was 19 inches, of the right 9 inches.

A strongly muscular “ ligamentum rotundum,” flattened at the sides, was
attached to each horn, 3 inches from its free end, and lay in relation to the
inferior part of the broad ligament. Each horn terminated in a well-defined
Fallopian tube, upwards of one foot in length, which lay in the free margin of ~
the broad ligament, and terminated externally in a widely dilated trumpet-
shaped mouth, the wall of which was formed of a duplication of the peritoneum.
The dilatation was so wide that the entire ovary could be included within it.
Immediately on the uterine side of this mouth was an elongated, deep, pouch-
like recess, formed by a folding of that part of the broad ligament which

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 471

extended between the Fallopian tube and the root of the ovary. Numerous
tortuous blood-vessels accompanied the Fallopian tube, branches from which
ramified between the layers of peritoneum which surrounded the mouth of the
tube. When these vessels were turgid with blood, the otherwise lax membrane
would doubtless have become tense, and capable of being accurately adapted to
the surface of the ovary. The lumen of the tube was throughout so large that a
full-sized catheter could be passed along it. Its mucous lining was elevated in
longitudinal folds, continuous at the uterine end with folds to be afterwards
described in the mucous membrane of the cornu. At the opposite end, where
the mucous lining of the tube became continuous with the serous membrane
surrounding the mouth, the folds diverged from each other, and then passed
outwards as foldings of the serous membrane, along the inner surface of the
trumpet-shaped mouth, as far as its free edge.

The left ovary had been removed before the specimen came into my posses-
sion. The right ovary, about the size of a duck’s egg, was attached to the
uterine cornu by its proper ligament. It lay in relation to the upper surface
of the broad ligament, with which it was connected by a mes-oarium 3 inches
in depth by 33 inches in breadth, so that it could be freely removed to and fro.
Between the folds of the mes-oarium numerous blood-vessels passed to and from
the gland, and close to the hilum was a flattened body 23 inches long by 14 inch
in its greatest transverse diameter, the relation of which to the ovary reminded
one of that of the epididymis to the testicle. The ovary was somewhat flattened
at the sides, and presented near the free convex surface a linear, slightly puck-
ered depression, which was in all probability a cicatrix. In other respects the
outer surface of the ovary was smooth, and its investing membrane was con-
tinuous with the mes-oarium. Beneath this membrane, an abundant venous
plexus, which was readily injected, and through which the injection passed
freely into the veins within the ovary and the flattened body at its hilum. A
vertical mesial section was then made through the ovary and the flattened
body. The latter was found to be composed of a close plexus of dilated and
tortuous veins and arteries imbedded in a dense connective tissue. The ovary
itself appeared to be completely occupied with a large corpus luteum, which was
3 inches long by 2 inches broad. It possessed a strongly marked central cica-
trix, much broader at one end than the other, which measured 1°8 inch in length
(fig. 2). From this cicatrix numerous slender bands radiated into the corpus
luteum, which possessed the characteristic yellow colour. Owing to the great
size of the corpus luteum, the proper ovarian substance was not at first recog-
nised, and it was only after a number of thin sections had been made and
examined under the microscope, that the ovarian stroma, pushed entirely to the
periphery of the ovary, and forming a sort of capsule to the yellow body, was
detected. In many parts of the periphery the stroma formed little more than

VOL, XXVI. PART II. 6G

472 PROFESSOR TURNER ON THE GRAVID UTERUS AND

a connective tissue envelope for the larger superficial arteries and veins, but
elsewhere it was not so compressed, so that a distinct layer of stroma substance
was more readily recognised, from which processes passed between the sub-
divisions of the corpus luteum. In these processes small arteries and veins
were situated, which entered the yellow body, and formed in it a capillary
plexus. This plexus extended as far as the margin of the central cicatrix,
where it was much less abundant than in the peripheral portions of the
corpus luteum. It formed a beautiful polygonal network, the meshes of
which were occupied by the characteristic fusiform cells of the corpus.
Some of these cells were unicaudate, but others were split into several
processes at their opposite ends (fig. 3). The corpus luteum was much more
vascular than the ovarian stroma, in which latter numerous dumb-bell shaped
bodies were seen.

The vagina was 16 inches in length, and 8 inches in breadth. It possessed a
thick muscular coat, and its mucous membrane was elevated into the powerful
folds, corrugated on the surface, which are so well-known in the interior of
this tube in the cetacea. The bladder was closely attached to the anterior
wall of the vagina, about 4 inches behind the cervix. . It was pyriform, and re-
ceived on each side a large ureter, which ran obliquely through the muscular
coat, before it opened into the bladder. A well-defined urethra, partially im-
bedded in the inferior wall of the vagina, ran backwards, to open immediately in
front of the vaginal orifice. The posterior surface, summit, and sides of the
bladder were covered by peritoneum, which was prolonged on to the cervix
uteri, whilst it left the bladder at the summit, along the line of the slender
obliterated urachus.

At least twenty arteries, about the size of the human brachial and ulnar, but
the coats of which were relatively thicker, lay between the two layers of each broad
ligament, close to the side of the cervix uteri. They ran forward, diverging
somewhat from each other, to the cornu, and in their course did not present
the tortuous arrangement found in the arteries of the human gravid uterus. A
few small collateral branches arose from them, which passed to the tissue of
the broad ligament, and here and there an obliquely-extending anastomosing
branch united adjacent arteries. At some part of its course each artery bifur-
cated, and where these branches reached the uterine horn, some extended for-
wards on one surface, others on the opposite. The branches now became more
frequent, and on the convex border of the horn, the branches for the opposite
surfaces freely anastomosed with each other. Those arteries which lay nearest
to the cervix and corpus uteri entered their substance, and, like the arteries of
the cornua, subdivided in the muscular wall. Numerous veins were seen to
accompany the arteries, and these again were neither tortuous nor dilated into
venous sinuses, such as one sees in the pregnant human uterus. Nerves, also,

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 473

some of which were larger than the great splanchnic nerve in man, accompa-
nied the arteries in their distribution.

When the uterine cornua were opened into, their walls were seen to be not
more than from + to 4 an inch in thickness ; their cavities contained the bag of
foetal membranes, and in that part of the bag which lay in the left cornu a good
sized foetus had formed. The foetal membranes had become detached from the
uterine mucous surface, and could be readily drawn out of the uterine cavity.
The two cornua did not communicate with each other so freely in the corpus
uteri as might have been supposed from the external examination of that
part ; for a vertical fold of mucous membrane, 8 inches in length, the posterior
border of which was sickle-shaped and free, formed a mesial septum between
the two horns at the anterior part of the corpus ; and the orifice of communi-
cation between them, which was situated in immediate relation to the os uteri
internum, was not more than about 5 inches in diameter.

The mucous lining of the cornua had a reddish-brown colour. Its general
characters did not quite correspond on the two sides. In the right smaller
cornu the mucous membrane, near the funnel-shaped passage into the Fallopian
tube was elevated into strong succulent folds, which projected from half-an-inch
to an inch beyond the general plane of the mucous membrane. These folds
starting from the oviduct, as from a centre, slightly diverged from each other, as
they passed parallel to the long axis of the horn towards the corpus uteri, and
at the same time gradually subsided, so as almost to have disappeared where
the cornu joined the body of the uterus (fig. 1). On closer examination each
fold was seen to be subdivided into multitudes of ridgelets, with narrow sepa-
rating furrows, which lay almost parallel to each other, and in the direction of
the main fold.

In the large impregnated cornu, the folds were only distinctly visible at the
free narrow end of the horn, for they soon subsided to the common plane of
the uterine mucous membrane, in all probability owing to the great distension
of this cornu. But their original position and direction were marked on the
surface of the membrane by parallel lines, which obviously represented the
ridgelets previously referred to.

The vertical fold of mucous membrane, already described as forming an im-
perfect mesial septum between the two cornua, was continued along the inferior

| and superior walls of the corpus uteri, close down to the os internum, and a
_ number of folds of mucous membrane from the inner end of each horn
| converged to the same orifice. The mucous membrane of the os itself was
arranged in distinct and almost parallel lamine, which projected into the cervix
uteri. The orifice was filled up with a plug of very viscid and strongly smelling
mucus. The lining membrane of the cervix uteri did not show an arbor vite, but
simple longitudinal parallel folds, not so prominent as those of the os internum.

474 PROFESSOR TURNER ON THE GRAVID UTERUS AND

These folds reached as far as the os externum, the orifice of which was suffi-
ciently large to admit a good-sized orange. The wall of the cervix was half an
inch thick.

When the free surface of the mucous membrane of the uterine cornua was
examined with the naked eye in a good light, under either water or spirit, it
was seen to possess a delicate reticulated character. The strands of the net-
work were formed of slender bands of the mucous membrane, many of which
ran parallel, being connected at intervals by shorter transverse or oblique bars ;
whilst others, again, had a much more irregular arrangement. Small recesses,
or pits, or furrows opened on the surface of the membrane, between these
bands or bars, and sank some depth into its substance. By the use of low

hi |

7 i iN
i ik Gl ‘
it ! mt ily
a | am i

Surface view, under a low power of the microscope, of a portion of the uninjected uterine mucous membrane.
The recesses, furrows, and pits, into which the pockets or crypts open, are darkly shaded in the figure.

magnifying powers these recesses could be more accurately studied. Some-
times they formed elongated furrows, which were again subdivided by more
delicate bands of the mucous membrane into smaller crypt-like compartments,
which opened freely into the furrow. In other localities the recesses were
irregular polygonal pits, and sometimes even ovoid or circular in form. These
also, like the furrows, were subdivided into a variable number of crypt-like
compartments. In some places the recesses were so closely crowded together
that the surface of the mucous membrane had a honeycomb appearance, but
in others comparatively broad patches of membrane separated a cluster of
recesses and crypts from those which lay around. As a rule, the body of each
crypt was more dilated than its mouth, and the crypts formed little pockets or
pouches for the reception of the club-shaped processes of the chorionic villi.

‘,

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 475

The complete absence of any putridity led me to hope that a minute injec-
tion of the vessels of the mucous membrane might be obtained, and that the
relation of the capillaries to the crypts and to the uterine glands might be
studied. Injecting pipes were accordingly inserted into some of the uterine
arteries, and, with the skilled aid of my assistant, Mr Srrriine, the uterine
system of capillaries was successfully filled with a gelatine and carmine injec-
tion. But before I enter on an account of the distribution of the blood-vessels,
it will be necessary to point out the characters of the mucous membrane itself,
as displayed both in vertical sections, and in horizontal sections made parallel to
the plane of the free surface.

Vertical sections through the membrane confirmed the description just
given of the arrangement of the pits, recesses, and crypts already recognised by
the inspection of the free surface. The depth of the crypts was variable, but in
no instance did they seem to occupy more than about the superficial third of
the thickness of the mucous membrane, and those which passed deepest into
its substance had usually several shallower crypts grouped around them. The
form of the deeper crypts resembled more a funnel, that of the shallower a cup.
It may be convenient to call this part of the membrane the crypt-layer (q, fig. 4),
or perhaps, from its numerous capillaries, the vascular crypt-layer.

The deeper two-thirds contained numerous elongated tubular glands—the
proper utricular glands—and may appropriately be called the glandular-layer
(6, fig. 4). It lay in contact with the muscular coat. From their tortuous
course and direction, the individual glands could only be followed for a com-
paratively short distance, and they presented different appearances in the vertical
and horizontal sections. As their long axes lay mostly in a direction parallel to
the free surface, good views of their arrangement were obtained in the horizontal
sections, in which the glands were seen not only to be convoluted, but to branch
(fig. 5). Sometimes they bifurcated, at others three branches arose close
together, and the branches could be traced sometimes for a considerable dis-
tance, but at others they formed short diverticula closed at their free ends.
Each branch possessed throughout an almost uniform diameter, but the portion
of the gland-tube situated immediately beneath the crypt layer, which may be
called the stem of the gland, had a wider calibre than its various branches,
which lay in the deeper portion of the mucous membrane. In the vertical
sections, again, as a rule, only short lengths of any given gland could be traced,
_ for the tubes were divided, sometimes longitudinally, but not unfrequently
obliquely or transversely (fig. 4, 0). As it was of importance to ascertain the
relations of the glands to the crypts, I examined many vertical sections to see
if I could follow the stems of the gland tubes, through the thickness of the
mucous membrane, to their openings or mouths on the free inner surface of the
uterus. And in carrying on these observations I encountered considerable

VOL. XXVI. PART II. 6H

476 PROFESSOR TURNER ON THE GRAVID UTERUS AND

difficulty, for the stems of most of the gland-tubes, as they lay immediately
below the vascular crypt-layer, were, in many of these vertical sections, obliquely
or transversely divided, and consequently their precise mode of termination and
the position of their mouths could not be followed out (¢, fig. 4). But in other
sections, where the stem of the gland-tube lay perpendicularly to the plane
of the surface, and where the knife had passed through its long axis, a short
length could be seen going to the deeper surface of the crypt-layer, and inclin-
ing indeed directly to the bottom of one of the deeper funnel-shaped crypts,
with the cavity of which its lumen was continuous (d, fig. 4).

From the examination of surface views of the mucous membrane, more
especially when the vessels were injected, I was able to obtain also satisfactory
evidence that the glands opened into the deeper part of the funnel-shaped crypts.
For, on looking into these crypts through a binocular microscope, I not unfre-
quently saw that the deeper end possessed an opening which communicated
with the stem of a tubular gland. The direction of this opening was in most
cases oblique, so that the tube of the gland, immediately prior to its termina-
tion, lay with its long axis oblique or almost parallel to the bottom of the crypt,
and consequently was transversely or obliquely divided in many of the vertical
sections (fig. 4, c, d). The relation of the orifice of the gland to the bottom of
the crypt closely resembled the appearance figured many years ago by Dr
SHARPEY in the pregnant uterus of the bitch.* Additional evidence of the
communication of the glands with these deeper crypts was obtained in some of
the specimens by observing a little plug, formed in all probability either of
epithelial cells or of the coagulated secretion of the gland, projecting from the
mouth of the gland into the bottom of the cavity of the crypt (fig. 10, @).

Owing to the great complexity of the free surface of the uterine mucous
membrane from the multitude of crypts, it was not possible to say how many
gland tubes opened in a given area. It was evident, however, that they were
not so closely set together, but that several smaller cup-shaped crypts (fig. 4, @),
which did not receive glands, intervened between the deeper funnel-shaped
crypts with which the glands communicated. And it was also clear that the
number of what I have termed the stems of the gland tubes, which reached the
erypt layer, was very much smaller than that of the tubes in the deeper por-
tions of the gland-layer, so that the number of branches springing from each
‘stem must have been considerable.

The glands were lined by a very distinct cylindrical epithelium, which was

* Dr Baty’s Translation of Mtiuer’s Physiology, p. 1576, figure 212.

Dr Suarpey, to whom I showed my preparations during the meeting of the British Association in
Edinburgh in August of the present year, told me that in the uterus of a pregnant Manis, which he
had examined some years ago, he found an arrangement of the uterine glands almost identical with
that seen in this Orca, and that, like myself, he had experienced a difficulty in tracing the glands into
the erypts.

ON THE ARRANGEMENT OF THE F@®TAL MEMBRANES IN THE CETACEA. 477

closely arranged around the wall of each tube, but leaving a distinct lumen in
the axis of the gland. The epithelial cells exhibited no appearance of degene-
ration, and the glands had the aspect of secreting organs in a state of complete
functional activity. I may mention here that in the pregnant uterus of a pig,
the foetus in which weighed only 12 grains, which was examined at the same
time, the glands had only half the diameter of those observed in this Orca.

The mucous membrane, which formed the walls of the crypts, and in which
the glands were imbedded, consisted of a delicate connective tissue containing
numerous nucleated corpuscles (fig. 4,7). These corpuscles were in part the
spindle-shaped nucleated corpuscles of the wall of the capillaries, but more fre-
quently were proper to the tissue itself. In the connective tissue of the gland-
layer these corpuscles mostly had the fusiform shape, but in the walls of the
erypts a distinct layer of globular or ellipsoidal nucleated corpuscles was seen
immediately within the boundary line of the mucous membrane, which was not
unfrequently elevated in a gently wavy line immediately superficial to the cor-
puscles. From their position these cells may be called the sub-epithelial cells of
the mucous membrane of the crypts (fig. 4,7). In thin sections, more especially
where the capillaries were partially filled with the red injection, they could be
readily distinguished from the spindle-shaped corpuscles of the capillary wall
by the difference in shape, and by the same test there was no fear of confounding
them with the epithelial lining of the crypts to be next described.

In many of the sections I was able to trace without difficulty the epithelial
lining of the crypts, and obtained the clearest views of the cells in the injected
portions of the mucous membrane (fig. 11, a). This lining, where it had not been
disturbed, was in contact with, and closely followed, the various irregularities
of, the mucous surface ; but in many places it had been either partially or alto-
gether removed, so that it is obviously readily shed from the membrane. The
cells of which it was composed had the appearance of a pavement epithelium,
though they were not larger than the broad, free ends of the cylindrical epi-
thelium lining the glands, with which, indeed, the epithelial lining of the crypts
was anatomically continuous. I examined, but failed to detect any difference
in shape between the cells lining the gland-crypts and those which lined the
crypts into which the utricular glands did not open.

The close relation which exists between the uterine glands and the deeper
funnel-shaped crypts, and the manifest continuity of the epithelial lining of the
one with that of the other, seems naturally to justify the inference, that in the
pregnant cetacean, all the crypts into which the glands open are merely the
mouths of the glands “ somewhat enlarged and widened,” and thus to establish
a correspondence with the arrangement which Dr SHarpery first pointed out in
the pregnant bitch. The extent to which, if we assume the accuracy of this
inference, the dilatation of their mouths may have proceeded, I am not as yet

478 PROFESSOR TURNER ON THE GRAVID UTERUS AND

in a position to say, neither do I know if a general enlargement of each gland
occurs, as I have had no opportunity of comparing them with the glands in the
unimpregnated uterus of the same species of whale; but it is probable that, n
the cetacea, as in the other mammals in which they have been seen, the glands
undergo a marked increase in size during gestation.

The uterine mucous membrane was very vascular. Numerous small arteries
ran through it, either obliquely or in a slightly tortuous manner, to the super-
ficial crypt layer ; immediately beneath which they subdivided into terminal
branches, which ended in a close compact capillary network, distributed on the
sides and at the bottom of the crypts, and immediately beneath the free surface
of those bands of mucous membrane which separated the trenches or pits into
which the crypts opened from each other (fig. 10). In the elongated bands
which lay between the trench-like recesses, the capillaries formed an elongated
network. I frequently saw a terminal artery pass up one side of a crypt and
give origin to capillaries which arched in a series of festoons around the walls of
the crypts; in many cases a distinct capillary ring surrounded the somewhat
constricted mouth of a crypt (fig. 12, a). The capillaries in the walls of all the
crypts belonging to the same group formed a continuous network, which freely
anastomosed across the intermediate portions of mucous membrane with the
capillaries in the walls of adjacent groups of crypts, so that a continuous
capillary plexus was produced, which gave to the free surface of the injected
portions of mucous membrane a bright carmine-tinted appearance. In the
relative number of the vessels, and the closeness of the network, this plexus
may fitly be compared with the capillary plexus of the lungs.

As the small arteries passed through the gland-layer, they gave off branches —
which ended in a capillary network situated in the connective tissue between
and surrounding the tubular glands. Where the stems of the gland tubes
reached the funnel-shaped crypts, there the capillaries of the crypts became
continuous with those which surrounded the glands. Owing to the open
character of the capillary plexus in relation to the glands, there was a strongly
marked difference between the vascularity of the crypt-layer and the gland-
layer ; for the vascularity of the latter was not greater than, and may fairly be
compared with, the capillary plexus surrounding the tubular glands of the
human stomach. The principal vessels of the gland-layer, like the glands them-
selves, lay parallel to the general plane of the mucous surface.

The Foetal Membranes.—The chorion was prolonged from the left into the
right horn across the corpus uteri, and extended on each side as far as the
opening of the Fallopian tube. The left subdivision, like the corresponding
uterine horn, was about twice the size of the right, and contained the embryo.
Near the tip of the right horn the chorion was thrown into permanent ruge,
which corresponded in direction to the folds of the uterine mucous membrane,

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 479

and fitted into the intervals between those folds. They gradually subsided
towards the corpus uteri. In the left horn the ruge of the chorion were much
less strongly marked in correspondence with the feebler folds of the mucous mem-
brane. Except in three limited areas, the entire outer surface of the chorion was
thickly studded with villi. These spots, bare of villi, corresponded to the three
openings into the uterus, viz., the os uteri internum, and the mouths of the two
Fallopian tubes (fig. 13, a, 6, 6). The spot opposite the os internum had a stellate

Stellate non-villous portion of the chorion of Orca opposite the os uteri. About half the size of nature.

form ; the central space of which was nearly the size of a crown piece, and from
it about twelve radii extended for a short but varying distance. The radii were
separated from each other by folds of the chorion, thickly studded with villi, which
fitted between the folds of mucous membrane, already described as converging
to the os uteri internum.* The spots opposite the mouths of the Fallopian tubes
formed the poles of the chorion, and were not larger than kidney beans. They
were not very readily recognised, on account of the puckered rugose condition
of the polar portions of the chorion, and it was not until after the blood-vessels
of the villi had been injected with coloured gelatine that their form and appear-
ance were satisfactorily determined. The absence of villi on those parts of the
chorion, which corresponded to the uterine openings, bears obviously a special
relation to the absence of a mucous surface at those spots into the depressions
in which the villi could be received ; for though the chorion was lying loose in
the uterine cavity when I opened into it, there can be no doubt that, before the
uterine liquor had been evacuated by Mr Gatherer, the chorionic villi had been
lodged within the uterine crypts, as the hand and fingers fit into a glove. «

* When I described and figured (Transactions of this Society, vol. xxvi., fig. 17) the only bare spot
which I had recognised in the chorion of the foetus of the Longniddry Balwnoptera, I regarded it, in
all probability, as one of the poles of the chorion, as the non-villous spot opposite the os internum
was not then known. The further knowledge which I have gained from the examination of this
Orea leads me now to think, from its size and the projection of the marginal fold, that it was a
portion of the bare spot opposite the os uteri internum.

VOL. XXVI. PART TI. 61

480 PROFESSOR TURNER ON THE GRAVID UTERUS AND

The villi could be seen with the naked eye, but their form and general
arrangement were more distinctly recognised when examined with low magni-
fying powers. Considerable variety was displayed in the arrangement of the
villi, in their length, and in the number present in a given area. In many places
they were set in rows, so as to form parallel series of ridgelets. In other places
they were collected into little tufts, irregular in form and size, which sometimes
consisted of two, three, or four villi, but frequently of a much larger number.
Solitary villi were also met with, and in the irregular intervals of comparatively
smooth membrane, which lay between the bases of the tufts or ridgelets, it was
not uncommon, as Escuricut had also observed in his Phocana, to see shorter
stunted simple villi projecting from the general plane of the chorion. It is
evident that the crypts on the uterine surface, into which the stunted simple
villi had been inserted, must have opened directly on its free mucous surface,
and not into a trench or pit. Asa rule the villi were compound in form, and
subdivided into three or more secondary club-shaped villi. The compound
villus not unfrequently swelled out at the free summit into a branching crown,
which, to adopt Escuricut’s expression, formed a miniature representation
of the head of a cauliflower.

The chorionic villi were composed of a delicate connective tissue, in which
numerous spheroidal and fusiform nucleated corpuscles were imbedded. Some
of these corpuscles were situated in the walls of the finer blood-vessels, but
others were proper to the tissue itself. A layer of spherical or ovoid corpuscles
was situated immediately within the free surface both of the simple villi and of the
secondary portions of each compound villus, and not unfrequently the limitary
membrane of the villus was slightly elevated immediately above the individual
corpuscles, so that the outline of the villus had a gently undulating appearance.
From their position these cells may conveniently be termed the sub-epithelial
corpuscles of the villus (fig. 6, @). The chorionic membrane between the bases ot
the villi consisted also of a delicate connective tissue, containing both spheroidal
and fusiform nucleated corpuscles. The fusiform cells possessed in many cases
very elongated poles, and had distinctly granular protoplasmic contents. Besides
those connected with the coats of the finer blood-vessels, others were situated
in the membrane itself. The spheroidal corpuscles were proportionally fewer
than in the tissue of the villi. No epithelial covering was recognised on the
chorionic villi, though it was carefully looked for, but it is very probable that
the epithelium had been shed, or rubbed off from their surfaces, before 1
reached this stage of the examination. For not only had several days elapsed
after the death of the animal, but the chorion had soaked for some time in
warm water during the process of injection.

When the chorion was cut into, along that surface which was adapted to the
convex aspect of the left uterine horn, the umbilical cord, allantois, amnion,

ON THE ARRANGEMENT OF THE F@®TAL MEMBRANES IN THE CETACEA. 481

and contained foetus could be examined. The chorion itself was seen to be
distinctly divided into two layers; the outer villous, already described, and an
inner thin translucent membrane. These layers were attached to each other
by very delicate connective tissue, and between them, the chorionic arteries
and veins, passing to and from the villi, ramified.

As the chorion showed no trace of putrefaction, I decided to make an
injection of the umbilical vessels and, with the skilful co-operation of my
assistant, Mr Srrruine, have succeeded in obtaining some beautiful prepara-
tions in illustration of the vascularity of the foetal membranes.*

The umbilical cord, 15 inches long, consisted of two arteries, two veins, and
the pedicle of the allantois (urachus), which were held together by an areolated
connective tissue, and were invested by the amnion (Plate XVII. fig. 7, a, and
Plate XVIII. fig. 15). A careful search was made in the substance of the cord
for the umbilical vesicle, or its pedicle, and for the vessels of the vitellus, but
without any positive result ; for although some elongated threads, which could
be isolated from the surrounding tissue, were met with, yet their impervious
condition prevented one from concluding with any certainty that they were the
remains either of the vitelline duct or of its accompanying blood-vessels.

The two umbilical veins resulted from the bifurcation at the umbilical
aperture of the single intra-abdominal vein. They were of large size, and
placed at the sides of the cord ; and the urachus, with the two umbilical arteries,
was situated between them. Sixteen inches from the abdomen of the foetus
the cord bifurcated into a right and left branch, an artery and vein passing off on
each side, conducted by the allantois, and situated between that membrane and
the amnion, to the chorion. In their course the artery wound around the vein
in a spiral manner from left to right on the one side, from right to left on the
other. Ten inches from the angle of divergence the vessels reached the chorion,
along the line of attachment of the allantois to that membrane, and in their
course gave off several branches. As soon as they reached the chorion, the -
branches of the vein diverged from the corresponding arteries, each pursuing
an independent course, and ramifying in an arborescent manner between the
two layers of the chorion. The general mode of branching was dichotomous,
but occasionally, and this more especially with the arteries, collateral branches
arose, in which case two not unfrequently came off close together, and extended
for some distance side by side before they proceeded to their respective areas
of distribution. Several anastomoses were observed between the branches of
the larger veins, and the finer branches of the arteries were occasionally
observed to inosculate with each other.

The mode of origin of the terminal arteries varied with the arrangement of

* These and the other preparations obtained from this uterus are preserved in the Anatomical
Museum of the University of Edinburgh.

482 PROFESSOR TURNER ON THE GRAVID UTERUS AND

the villi. When these structures were in rows, or scattered, the terminal
branches which entered the villi arose independently ; but, when the villi were
in clusters, several terminal twigs arose close together as from an axis. The
arteries did not, like the branches from which they arose, le parallel to the
surface of the chorion, but immediately after their origin entered the villi at
the base of attachment, and in the case of a compound villus divided, within
its substance, into twigs for the secondary villi. These twigs ascended towards
the tip of the villus, and ended in a very compact capillary plexus, situated
beneath the free surface of the villus, and in close relation to the sub-epithelial
layer of spheroidal cells already described. This plexus may conveniently be
termed the intra-villous capillary network (fig. 14, a). From the intra-villous
plexus capillaries proceeded, which passed out of the villus at its base, and at
once joined a capillary network situated immediately beneath the general plane of
the chorion. This plexus may be termed, from its position, the sub-chorionic or
extra-villous capillary network (fig. 14, 6), The extra-villous plexus was much
less compact than that within the villi; its meshes were, as a rule, elongated.
but occasionally more irregular in form. Numerous minute veins arose directly
from it, which joined together to form the rootlets of the umbilical vein.

Occasionally I saw a vessel, which was somewhat larger than the capillaries
in its immediate neighbourhood, and might therefore be regarded as a direct
rootlet of an umbilical vein arisg within, and leaving*a villus at its base;
but this was quite exceptional, for repeated examinations have satisfied me that
the arrangement which most commonly prevailed was for the capillaries of the
intra-villous plexus to be directly continuous with those of the sub-chorionic net-
work, and for the latter to give origin to an umbilical vein. Hence the blood
in its passage from the terminal twigs of the umbilical artery to the rootlets of
the umbilical vein has to flow through a complicated capillary system, one
subdivision of which les within the villi, the other beneath the chorionic
membrane which connects them at their bases. When the chorion lay in its
proper position within the uterus, the first subdivision would have been in
relation to the maternal system of capillaries lining the walls of the crypts, the —
other to the capillaries situated immediately beneath the free surface of those
parts of the mucous membrane which separated the trenches or pits into
which the crypts opened from each other. This greatly diffused capillary area
has undoubtedly a special relation to the comparatively simple mode of union
between the maternal and foetal surfaces in the diffused form of placenta.

The three bare spots on the chorion were almost entirely destitute of blood- —
vessels, and, after the chorion was injected, contrasted strongly with the sur-
rounding highly vascular villous portion.*

* It may not be out of place to state that some portions of the chorion were injected with a blue-
coloured gelatine from branches of the umbilical artery only, when the intza- villous plexus was readily

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 483

The urachus freely communicated at its abdominal end with the bladder.
At the angle, where the umbilical vessels diverged from each other, the urachus
rapidly expanded into a huge funnel-shaped sac-like allantois, capable of hold-
ing several pints of fluid. When theallantois reached the chorion, it became
prolonged both to the right and left in the form of a comparatively wide, almost
cylindrical sac-like horn (fig. 7, 6, 6). The left horn-like prolongation extended
to 7 inches from the end of the left horn of the chorion, where it formed a
cul-de-sac. The right horn-like prolongation entered the right cornu, but did
not reach to within 21 inches of the tip of the right horn of the chorion. By far
the greater part of the attached surface of the allantois was in contact with the
amnion. But, at the points where the umbilical vessels reached the chorion,
opposite to the abdominal aspect of the foetus, the allantois came in contact
with the deeper layer of that membrane, to which it continued to be attached
by a limited portion of its surface, as far as the ends of its horn-like prolonga-
tions. The free surface of the allantois was perfectly smooth, and was bathed
by the allantoic fluid.

EXPLANATION OF D1AGRAMS.—Outline diagrams to show the arrangement of the membranes at the stage of develop-
ment described in the text. A, Longitudinal section. B, Transverse section. H, Embryo. ch, Chorion. al, Allantois.
am, Amnion represented in a dotted line. The umbilical vesicle is not shown.

The amnion formed a continuous bag from one horn of the chorion to the
other, but was not co-equal with it in extent. For though in the left horn it
reached to 2 inches from the pole of the chorion, in the right the chorion
extended 9 inches beyond the closed end of the amnion (fig. 13 ¢, c). Where
the amnion was in relation to the dorsal aspect of the foetus, it was connected
by a delicate filamentous tissue to the inner layer of the chorion ; but, where
the allantois was attached to the chorion, 7.¢., opposite the abdominal aspect,
the amnion was reflected on to the outer surface of the cylindrical horns
and funnel-shaped sac of the allantois, and was conducted by that membrane
to the umbilical cord, which it invested. Beyond the closed ends of the
filled ; that others were injected with a carmine-coloured gelatine from branches of the umbilical vein
only, when both the extra and intra-villous capillaries were readily filled ; and that others again were

injected both from artery and vein, until the coloured gelatines intermingled in the capillaries, and
produced there a purple tint.

VOL. XXVI. PART II. ; 6K

484 PROFESSOR TURNER ON THE GRAVID UTERUS AND

amnion the chorion formed the sole constituent of the foetal membranes.
Not only was the amnion surrounding the cord abundantly studded with
yellowish brown or olive-tinted bodies, smaller even than mustard seeds, but
they were also thickly congregated on the amnion where it covered the
funnel-shaped sac and horns of the allantois, though they were much more
sparingly distributed on that part of the amnion which was in contact with the
chorion (fig. 7,d@). In the right cornu they were absent on that portion of the
amnion which extended beyond the horn of the allantois, but on the left side
they extended further, and one was seen almost at the closed end of the
amnion (fig. 7, d'). By the presence of these corpuscles in connection with
the amnion, and their absence on the free surface of the allantois, these two
membranes were at once readily distinguished from each other. Some of these
corpuscles were pedunculated, others sessile, and they had obviously been
developed in relation to the attached, and not to the free surface of the amnion,
for each was invested by a prolongation of that membrane (as the spleen is
invested by the peritoneum), and, where the corpuscles were pedunculated, the
pedicle of attachment, which sometimes was ith of an inch long, was formed
by a slender filamentous process of the amnion. Thin sections of the corpuscles,
examined with a magnifying power of 480 diameters, were found to be composed
of cells, closely packed together, some of which were oval, others somewhat
elongated, others somewhat polygonal in shape. In many of the cells clusters
of brown pigment granules were contained (fig. 8).

The amnion and allantois were connected together by a very delicate fila-
mentous tissue, in which a few tortuous slender branches of the umbilical
arteries and veins were distributed, and frequently formed well-marked anasto-
mosing loops. When the amnion and allantois were separated from each other,
these vessels remained in contact with the deep surface of the former membrane.
By dissecting off the amnion from the cord, numerous vasa vasorum could be
seen distributed to the coats of the umbilical vessels and the urachus.

T also found between the amnion and allantois, close to the trunks of the
umbilical vessels which passed to the left horn, three peculiar-looking bodies,
the blood-vessels of which were injected from the above-mentioned amniotic
arteries. The largest, tri-radiate in form, was ? inch long by 4 inch in greatest
breadth, the others were much smaller, and of an ovoid form. They lay in
linear series, the highest and largest was 11 from the second, and that again 4
an inch from the third. They were connected together by intermediate vessels,
and resembled in appearance a chain of small lymphatic glands. When the
allantois and amnion were separated from each other, these bodies remained
attached to the amnion. When a section was made into one of these bodies, a
brown pultaceous mass, contained within a cavity, the wall of which was formed
of a fibrous capsule, was exposed. The brown mass, examined microscopically,

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 485

was seen to consist of cells, the great majority of which were circular in form,
and had a yellowish colour. The smallest of these cells were of the size of lymph-
corpuscles, but as a rule they were twice as large. Most of the cells contained
a single nucleus, but some possessed two or more, and in a few instances large
brood cells were observed, packed full of nuclei, or young cells. Some patches
of small hexagonal cells, fitted together by their margins, like the cells of the
choroid coat of the eyeball, were also seen ; and it is possible that these cells
formed a pavement epithelium for the cavity in which the brown pultaceous
mass was contained. (Fig. 9, Plate XVII.) These bodies, like the amniotic cor-
puscles already described, were developed in relation to the attached surface
of the amnion, but, unlike the majority of the latter, had not projected so as
to assume a pedunculated appearance. Inall probability they and the amniotic
corpuscles have common morphological relations, though what their function
may be it is not easy to determine.

Position and Form of the Foetus.—The foetus, a male, enveloped in its
membranes, occupied the left uterine cornu. It lay with its beak directed
towards the corpus uteri and os internum. The caudal end was curved forward
under the abdomen, so that the flanges of the tail were in close relation to the
penis (Plate X VIII. fig. 16). The curve of the back of the foetus corresponded
with the convex anterior surface of the uterine cornu. In its position in utero,
therefore, it closely corresponded with that of a foetus of Globiceps, figured by
VAN BENEDEN,* and it bears out the opinion entertained by that naturalist, that
the foetus in the cetacea is born with the beak foremost. The foetus possessed
the external characters of the genus Orca. The beak was not so pointed as in
| Delphinus, or so truncated as in Globio-cephalus; the falciform dorsal fin
lay in the same transverse plane as the umbilical cord; the pectoral fins
were broadly ovoid, flattened on the surfaces, with a rounded anterior and
sharp convex posterior border. The colour was dark-slate on the dorsum
of the head, back, tail, and upper part of the sides of the body. A
distinct pale patch extended horizontally backwards immediately above and
beyond the eye slit. The under surface of the lower jaw, throat, and the
entire ventral surface to a little beyond the anus, were also pale, and a
patch continuous with the ventral surface commenced in front of the penis and
extended upwards and backwards along the side of the body. The under
surface of the tail was also pale. In the adult these pale patches and surfaces
are white, which, contrasting strongly with the otherwise dark colour of the skin,
give to the animal a characteristic black and white piebald appearance. But in
the fcetus, from the comparative thinness of the cuticle over those surfaces where
it was devoid of pigment, the pale patches, instead of being white, were salmon-
tinted, owing to the colour of the vascular cutis being transmitted through the
* Bulletins de Acad. Royale de Belgique, 2d Series, xx. No. 12.

486 PROFESSOR TURNER ON THE GRAVID UTERUS AND

comparatively thin cuticle. Parallel to and immediately below the base of the
dorsal fin, the dark-slate colour was modified by a lighter pinkish-tinted patch,
which corresponded with the purplish spot represented by ScHLEGEL in the
adult Orca which he has figured.* The foetus measured from the tip of the
upper jaw along the middle of the back to the mesial notch of the tail 36
inches. Its greatest girth midway between the root of the flipper and the
attachment of the umbilical cord was 184 inches. From the attachment of
the cord to the root of the penis, 3°5 inches; from the latter to the anus, 2°0
inches ; and from the anus to the mesial notch of the tail, 10°3 inches. From the
tip of the lower jaw to the umbilical cord, 15°2inches. Length of pectoral fin, 4°8
inches; length of base of falciform fatty fin, 3-2; height measured along
anterior border, 4°2 inches. From posterior rise of dorsal fin to mesial notch
of tail, 14:2 inches. Between tips of tail lobes along their posterior concave
borders, 7°5 inches.

Comparison of Placentation with that of other Mammals.t—t shall endea-
vour, in making this comparison, to show, as far as the materials at my disposal
will permit, the features of resemblance and dissimilarity, not only as regards
the general arrangement of the foetal membranes and uterine mucous surface,
but their more minute structure.

The dissection of this Orca confirms the results previously arrived at from
the dissection of various specimens of the genus De/phinus, by the anatomists
quoted in my introductory observations, that in the cetacea the chorion extends
into both uterine horns, and its surface is so studded with villi as to forma ~
placenta of the “ diffused” type, in which, from the absence, so far as was
ascertained, of a uterine decidua intermingled with the chorionic villi, the fetal
and maternal surfaces readily separate from each other. It also agrees with
the specimen examined by Professor RoLLEsTon, in the presence of a bare spot,
free from villi, at each of the poles of the chorion. But I have also demon-
strated, what had previously been overlooked, that a third, larger, non-villous
area is situated opposite the os uteri internum,

Of the mammals, the placentation of which most commonly comes under
observation, the sow and the mare also offer well-known examples of the dif-
fused form of placenta. Of these the uniparous mare presents more points of —
resemblance to the uniparous cetacean than does the pluriparous sow. For in
the mare not only does the chorion of the solitary embryo extend from one uterine
cornu to the other, and possess small non-villous spots at the poles, but an
even larger, stellate, bare spot also exists in relation to the os internum.}

* Abhandlungen aus dem Gebiete der Zoologie und vergleichenden Anatomie. Part ii. fig. 7.
} October 1871. This section and the final one, entitled “Physiological Conclusions,” have
been re-written since the Memoir was read.

{ In one specimen, I observed that the non-villous pole of the horn of the chorion which contained
a foetal foal about 2 feet long, was somewhat smaller than that of the Orca, but the bare spot in the

ON THE ARRANGEMENT OF THE F@®TAL MEMBRANES IN THE CETACEA. 487

In the pluriparous sow, again, though the poles of the ovum are smooth and
almost non-vascular, the third bare spot is not present, for the membranes
surrounding each embryo are confined to a single horn, and do not pass across
the corpus uteri.* In the sow also the folds of the chorion and uterine mucous
membrane do not, as in the whale and mare, lie in the direction of the long axis
of the uterine cornu, but at right angles to it.

The bare spots at the poles of the greatly elongated chorion of the cetacean
are undoubtedly homologous with the smooth ends of the shorter ellipsoidal
chorion of the carnivora, but the non-villous portion is absolutely and relatively
much smaller in the former than in the latter. As the carnivora are, as a rule,
pluriparous, the conditions necessary for the formation of a third bare spot are
non-existent in them, just as is the case in the pig.

But the closer affinity of the mare than of the sow to the cetacean, in the
characters of the chorion, is shown also in the arrangement of the villi, and in
the distribution of the capillary blood-vessels. The villi of the chorion of the
mare are thickly distributed over its surface. They are for the most part com-
pound and arranged in tufts; though isolated simple villi are scattered in the
intervals between the tufts. The tufts are, as a rule, somewhat larger than
those in the cetacean, and they have more of a brush-like form than of that of
the head of a cauliflower, a difference which I find to be due to the secondary
villi being elongated and filamentous rather than club-shaped. Not only is a
rich capillary network situated within the villi, but this plexus, as I have
satisfied myself, from the examination of an injected chorion, freely communi-
cates, as in the whale, with an abundant extra-villous sub-chorionic plexus, from
which the rootlets of the umbilical vein arise, so that in both animals a diffused
capillary area is produced by similar modes of distribution.

Von Barr and Escuricut long ago pointed out that in the pig the trans-
_ verse folds of the chorion are notched at the free margin, and the teeth, by grow-
ing, become converted into villi. These villi are smaller than in the mare and
whale, and at an early period of development can scarcely be said to be present.
In a gravid uterus which I examined, where the embryos were 1:3 inch long,

opposite horn was considerably larger, being 24 inches long by from 4 to ? inch broad, and with radiated
bare processes passing off from its two ends. The bare spot opposite the os internum was twice as large
as in the cetacean, and had several strongly-marked, radiating, non-villous processes. The presence of
these bare spaces, or at least their exact position and signification in the chorion of the mare, seems to
have escaped the notice of veterinary anatomists. Neither CHavveau nor Gurit make any mention of
them in their well-known treatises, and Franck (“Handbuch der Anatomie der Hausthiere,” Stutt-
gart, 1871), whose work is the most recent and fullest in detail of any which I have been able to con-
sult, merely says, “in one or other horn roundish spaces are found, where the villi are sparse and
feeble, and here the chorion has a semi-transparent appearance.”

* In one pig’s uterus, I found that the membranes belonging to the embryo, situated lowest down
in the left horn, actually did pass across the corpus uteri into the right horn, but this of course was
not the case with the other ova.

MOM XOGVA. PART: Li, 6 L

488 PROFESSOR TURNER ON THE GRAVID UTERUS AND

multitudes of elongated feeble ridgelets traversed the surface of the chorion,
which fitted into corresponding shallow elongated depressions in the uterine
mucous membrane, but no true villi were recognised, and the smooth and feebly
vascular poles were absolutely and relatively larger than in the cetacean. A
distinct and compact capillary plexus, which was injected with carmine and
gelatine, was seen both in the ridgelets and in the intermediate portions of the
chorion. This plexus was elongated within the ridgelets, and had the same
direction, but in the intermediate membrane it formed a polygonal network, so
that the entire surface of the chorion, except at the poles, possessed a diffused
capillary vascularity, without any differentiation into intra- and extra-villous
areas of distribution.

In possessing a basis substance of delicate connective tissue, in which nume-
rous nucleated corpuscles lie, the chorionic villi in the cetacean agreed with the
structure of villi generally.. The layer of corpuscles situated immediately within
the periphery of the villus, which I have described as its sub-epithelial corpuscles
(p. 479), obviously corresponds in position, arrangement, and shape, to the
corpuscles figured and described as the internal cells of the villus,* by Pro-
fessor GoopsiR in the villi of the human chorion. In a recently published and
very elaborate memoir on the “Structure and Function of the Placenta,”
Professor ErcoLani of Bologna has also figured,t in connection with the human
villi, a layer of spheroidal corpuscles, which he terms the cells of the internal
epithelial layer, situated to all appearance within the villus close to the fcetal
vessel, for he represents a layer of membrane immediately outside the cells. His
drawing certainly gives one the impression that these cells corresponded in
position with the sub-epithelial corpuscles above referred to, but in his text he
speaks of them as not included in the thickness of the membrane, but as cells
of the decidua serotina which enter into the formation of a new glandular
organ which envelopes the villus. Moreover, he regards them as identical with
the single layer of flattened spheroidal cells which Dr Farre{ observed and
described as forming the sheath or outer case of the villus, and which undoubt-
edly belonged to the decidua serotina, and not to the chorionic villi. Although,
for the reasons already stated, I did not detect an epithelial layer on the free
surface of the villi, yet it is probable that not only they, but the whole outer
surface of the chorion, possessed an epithelial covering, as is the case with the
chorion in other mammals.

Passing now to the consideration of the characters of the uterme mucous

* Anatomical and Pathological Observations, p. 54, plate 2, fig. 19, f. 1845, reproduced in Ana-
tomical Memoirs, vol. ii. p. 18. Edinburgh, 1868.

+ Mémoire sur les glandes utriculaires de l’uterus, et sur l‘organe glandulaire de neo-formation, &e.
I know this work, which was published at Bologna in 1868, only in the French Translation by BrucH
and AnpREINI. Algier, 1869. Plate X. fig. 2, d.

+ Cyclop. of Anat. and Phys., article Uterus, p. 718, fig. 485.

ON THE ARRANGEMENT OF THE F@®TAL MEMBRANES IN THE CETACEA. 489

membrane, our attention should especially be directed to the appearance pre-
sented by its free surface, to the arrangement of the proper utricular glands,
and to the discussion of the question, whether the villi of the chorion in the
placental area do, or do not, enter within the mouths of the glands.

Escuricut had recognised that the free surface of the uterine mucous mem-
brane in the gravid porpoise, which he dissected, appeared “ cellulosa vel
cribrosa.” The form of the “cells” was “valde irregularis. Interdum quadrate
vel triangulares sunt, interdum rotundate, sepius oblonge unum punctum
elevatum irregulariter radiatim circumstantes’”—a description which closely
applies to the depressions termed recesses, trenches, and pits, which I have
figured on p. 474 in the gravid Orca. It does not very clearly appear, from
his description, whether he had recognised the small cup-shaped pouches
or pockets opening into or leading out of these larger depressions, which I
have named the crypts; so that I am inclined to think that Escuricut’s
term “cells” must be taken as equivalent to my pits, recesses, or trenches,
rather than to my crypts. His account of the arrangement of the closely-
packed, branching, uterine glands, applies, in most particulars, equally well
to what I have seen in the Orca; but from the statement, “ ostia areolis
seu maculis levibus insunt, quibus nullze omnino insident cellule,” he seems
to think that the glands open on the surface of the mucous membrane,
not in the “cells” in which the chorionic villi are lodged, but in separate
shallow areole ; and he goes on to say, that for so great a multitude of gland
ramifications, there are not more openings on the surface of the mucous
_ coat than in the pig, and the openings are separated by intervals of one or
two lines.

In the gravid uterus of a mare, where the foetus was about two feet long,
which I examined several years ago, I observed that the inner surface of its
mucous membrane was pitted with minute depressions, for the reception of the
villi of the chorion ; but unfortunately I omitted to notice the relation which
they bore to the uterine glands. In another uterus, in the sixth month of
gestation, the free surface of the membrane was crowded with multitudes of
crypts, not unlike those described in Orca. Opening sometimes within these
crypts, but at others on the ridges which separated different groups of crypts
from each other, were oval, or almost circular, orifices surrounded by a capillary
ring, which were evidently the mouths of utricular glands. In vertical sections
the glands were distinctly seen. In the deeper part, branched, tortuous, and with
diverticular off-shoots, but in their course to the crypt-layer they ascended almost
straight and vertically, so that they could be followed without difficulty, and
contrasted strongly, therefore, in this respect with the arrangement seen in
Orca. The crypt-layer was, as in the cetacean, much more highly vascular than
the gland-layer of the mucous membrane, but the vessels ascended to the crypt-

490 PROFESSOR TURNER ON THE GRAVID UTERUS AND

layer parallel to the tubes of the gland.* Both Fapricrus and Von Bakr had
previously recognised small openings on the surface of the uterine mucous
membrane of the mare into which the villi of the chorion entered, but Professor
GurR.ttTt was the first to affirm that the villi passed in this animal into the mouths
of the glands. Ercozani describes numerous simple gland follicles, lined by a
pavement epithelium, as receiving in the mare the chorionic villi. But he
considers these follicles to be newly formed during pregnancy, and to be entirely
independent of the uterine glands, which, lined by their cylindrical epithelium,
and of uniform diameter, open by separate orifices on the uterine mucous
surface, and do not receive the villi of the chorion.

In the pregnant sow the surface of the uterme mucous membrane is modi-
fied in accordance with the conformation of the corresponding aspect of the
choricn, as already referred to. Instead of presenting multitudes of crypts,
such as have been described in the whale, the surface is traversed by transverse
folds separated by intermediate furrows and fosse, so as to present an undu-
lating appearance. Shallow depressions, or areole, which are to be regarded
either as the dilated mouths of the glands, or, as depressions into which the
glands open, are scattered over the surface. In an uterus which I examined,
where the foetal pig weighed 12 0z., about twelve of these areolze, correspond-
ing to the mouths of an equal number of glands, were situated on a portion of
mucous membrane 74;ths of an inch square. The vessels of this specimen had
been injected with vermilion, and the preparation corresponded in appearance
to the specimens described by Escuricut, that in injected uteri the areole are
easily observed, as they are much less vascular than the surrounding portions
of mucous membrane.{ According to Von Barr and Escuricat, little circular
or star-like vascular elevations of the surface of the chorion are attached to
these dilated orifices of the glands.

In another pig’s uterus, where the foetus weighed only 12 grains, the mouths
of the glands could be distinctly seen opening sometimes directly, at others
obliquely, and these openings were either on the plane surface of the mucous
membrane, or in shallow depressions, and not unfrequently plugs of epithelium
were seen projecting through the orifices, exactly in the manner I have described ~
in the pregnant Orca; and the gland tube, as in that animal, did not pass verti-
cally to the plane of the mucous surface, but lay obliquely or parallel to it. The
intervals between the mouths of adjacent glands were so great that, examined
with a 4-inch objective, one, or at the most two, could only be seen at the same
time in the field of the microscope ; and the intermediate portions of the mucous

* This specimen was injected in 1853 by that excellent anatomist, the late Mr Joun Bartow of
Edinburgh.

+t Handb. der vergleich. Anat. der Haus Saugethiere, p. 431. Berlin, 1860.

+ This diminished vascularity, as it seems to be, is probably due merely to the vessels being less
perfectly filled with the vermilion injection.

ON THE ARRANGEMENT OF THE FETAL MEMBRANES IN THE CETACEA. 491

membrane between the gland mouths presented an undulating appearance, from
the fosse and furrows, with their intervening ridgelets, which it possessed.
The vessels in this specimen were very perfectly filled with a carmine and gela-
tine injection, and the capillaries were seen to form a close polygonal network,
which was quite as abundant around the mouths of the glands as in the inter-
mediate parts of the membrane.

The study of this specimen has been of great service as a guide in deter-
mining the signification of the appearances presented by the free surface of the
more complicated utermme mucous membrane in the gravid Orca. In both
animals it was clear that the utricular glands opened on the free surface of the
mucous membrane; only, in the Orca they opened at the bottom of deep funnel-
shaped crypts, but in the pig, either on the plane surface or in shallow fosse.
In both, the gland mouths were separated by intermediate portions of mucous
membrane, which in the Orca was folded with so much complexity as to form
numerous cup-shaped crypts, but in the pig to produce only shallow fossz and
furrows. In both animals the mucous membrane was highly vascular, not only
where the glands opened, but in the intermediate portions. In Orca the villi
of the chorion were lodged both in the crypts into which the glands opened and in
the intermediate crypts ; in the pig the fossee and furrows received the highly
vascular transverse folds of the chorion, which represented and performed the
functions of the villi, and would in course of time have become villous, whilst it
_ is probable that, if the circular or star-like elevations of the chorion had been
developed, they would, as Von Barr has shown, have been in relation to the
gland orifices. It is clear, therefore, that in the pregnant uteri of these ani-
mals, not only the uterine glands, but the intermediate portions of mucous
membrane, bear important relations to the chorion.

[have already stated, p. 477, that my observations on the mucous membrane
_ of Orca seem to justify the inference, that the deeper funnel-shaped crypts may
be regarded as the dilated mouths of the utricular glands which open into them.
But, from what I have observed in the mare, it does not appear that in this
animal all these glands do necessarily open, or dilate, into crypts at their mouths,
for some without doubt presented no such arrangement. I have said nothing

| as yet, however, of the probable mode of formation of the cup-shaped non-

glandular crypts, of which two hypotheses may be advanced in explanation, one
to be considered here, but the other to be more appropriately discussed when
comparing the cetacean and carnivorous modes of placentation.

It is well known that during pregnancy the uterine mucous membrane not
only becomes very vascular, but increases both in superficial area and thick-
ness. Many years ago, Joun Goopsir pointed out* that the swollen state of

* Op. cit. p. 57. 1845.
VOL, XVI. PART: II. 6M

492 PROFESSOR TURNER ON THE GRAVID UTERUS AND

the membrane in the human gravid uterus is due, not only, as SHARPEY had
shown, to changes in the uterine glands, but to an increase in the amount of
the inter-glandular tissue, by the formation of ‘‘a texture which consisted
entirely of nucleated particles.” ERcoLant had also observed, in the various
uteri which he had examined,* this increased growth of the uterine tissue, and
in accordance with our present modes of expression, described it as formed
by the proliferation of the connective tissue. But he also affirmed that, in
all the placental mammals, through the transformation and folding of the
uterine mucous membrane in the interspaces between the utricular glands, a
new glandular organ is formed, which, however flexuous the mucous membrane
may become in different animals, never loses the type of a simple glandular
follicle. And further, that it is into the follicles of this new gland-organ that
the villi of the chorion penetrate, and not into the utricular glands.

The examination which I have made into the minute structure of the uterine
mucous membrane of Ovca has satisfied me that in it also a great growth of the
inter-glandular tissue had taken place, for not only was the connective tissue with
its fusiform nucleated corpuscles largely developed, but I distinctly recognised
also a layer of sub-epithelial corpuscles, situated close to the surface of the
mucous membrane. Epithelial-lined, cup-shaped crypts, for the reception of
villi, had formed in great numbers, which had added largely to the superficial area
of the membrane. Now, there can be no doubt but that these crypts, which —
obviously correspond to the simple gland follicles of ERcoLANI, could be formed
by a flexuous growth of the mucous membrane, and that the difference presented
by this surface in Orca and in the pig is simply due to the foldings being
more complicated in the former than in the latter—a difference which is in
accordance with the greater size and numbers of the villi in the cetacean than
in the pachyderm.

In Ruminants, which furnish such characteristic examples of the Coty-
ledonary placenta, utricular glands can be readily recognised in the uterine
mucous membrane. In the gravid uterus of the sheep the mouths of these
glands may be seen with the naked eye opening on the plane surface of the
membrane in the intervals between the cotyledons, and, when vertical sections
are made, there is no difficulty in following them as comparatively straight tubes
through its thickness. In the deeper layer of the membrane they branch and
bear a strong resemblance to the utricular glands figured by Franck in the cow
and goat. They ascend also for some distance on the sides of the cotyledons,
but their relations to the centre and summits of each of these bodies, and to
the openings which admit the chorionic villi, are difficult to determine. Pro-
fessor SPIEGELBERG, however, maintains that the tubes which open on the sur-

* He gives no description of the cetacean placenta.

ON THE ARRANGEMENT OF THE F@®TAL MEMBRANES IN THE CETACEA. 493

face of a cotyledon for the reception of the chorionic villi are only remarkable
dilatations of the uterine glands.*

In the zone-like placenta of the Carnivora, the observations of Dr SHARPEY,*t
which WeEBER and Biscuorrt{ have confirmed, have shown that two kinds of
glands, simple and compound, open on the mucous surface of the uterus of
the bitch.§ After impregnation both kinds of glands dilate into pits, which
receive the foetal villii The simple undergo a uniform enlargement, whilst
only the mouths and adjacent part of the ducts of the tubular glands are dilated
into pits. JASSINSKY even states, || that in the bitch all the chorionic villi, without
- exception, pass into the uterine glands, so that a double membrana propria and
a double epithelial coverig may be recognised in connection with each villus,
one membrane and one epithelial layer belonging to the villus itself, the others
to the gland in which it is included.

And here I may refer to the other hypothesis, to which I alluded on p. 491,
in explanation of the mode of production of the crypts? in the cetacean.
It may be that in the Orca also both kinds of glands exist, and undergo dila-
tation during pregnancy, so that the crypts of my vascular crypt layer may
include both the uniformly enlarged, short, simple glands and the dilated mouths
of the utricular glands. In the absence, however, of any knowledge of the
existence of the simple glands in the unimpregnated uterus of the cetacean, the
hypothesis previously advanced is to be preferred ; as we do know that a great
growth of the inter-glandular connective tissue and increase in the superficial
_ area of the uterus take place, which, if thrown into folds, would produce such
a crypt-like structure as has been described, without the necessity of assuming
the pre-existence and subsequent enlargement of the short simple glands.

In the mammals which possess the Discoid form of placenta Leypic** has
observed the existence of utricular glands in the mole. Of the Rodents which

* HENLE and Preurer’s Zeitschrift, vol. xxi.

+ Op. cit., p. 1576. {t Hunde Ei, plate xiv.

§ Weper, Rotizuston, and Ercouani have pointed out the presence of utricular glands in the cat.
T have also seen them in the badger, in which animal they closely resemble the figure and description
given by Bisonorr of these glands in the bitch. Erconanr denies the existence of two kinds of glands
in the bitch’s uterus, and states that only the utricular glands are present. Cari FrrepLANpER has,
however, recently made some observations (“ Untersuchungen iiber den Uterus,” Leipzig, 1870), which
reconcile the opposite statements of SHarpry and Ercouani. For he points out that, whilst in the
quiescent condition of the uterus of this animal only the utricular glands are present, in the period of
heat, when the mucous membrane is swollen, and its vessels turgid with blood, simple glands are also
met with.

|| VircHow’s Archiv, xl. p. 350. 1867.

{| To prevent misunderstanding I may state that BiscHorr specially designates the short simple
glands of SHarpxy as the mucous crypts,whilst the longer, branching, convoluted glands are the proper
utricular glands. In my description of the uterine mucous membrane of Orca, I have employed the
term crypts to designate all the pouches or pockets which receive the chorionic villi, whether accord-

“ing to the above hypothesis they are the simple glands uniformly enlarged, or the dilated mouths of
the utricular glands.

** Histologie des Menschen und der Thiere, p. 517. 1857.

494 PROFESSOR TURNER ON THE GRAVID UTERUS AND

have been more especially examined, viz., the rat, rabbit, hare, and guinea pig,
there is a general agreement among observers that proper utricular glands in
the form of elongated tubes do not exist, but the mucous membrane is thrown
into complex foldings, which possess an appearance long since compared by
ReEIcHERT to the convolutions of the brain.* Leypic indeed is disposed to
regard the spaces between the folds, with their epithelial lining, as equivalent
to colossal glands, although they want the tubular form.

In the Human Subject, where the placenta possesses its most concentrated
character, utricular glands are present in the uterme mucous membrane. As is
well-known, however, these glands are difficult to demonstrate in the quiescent
uterus, and only acquire well-marked characters after conception has taken
place or during menstruation. They form important constituents of the decidua
vera or uterina on the free surface of which their mouths may readily be seen.
As the ovum enters the uterus it becomes enclosed within a chamber formed
of the decidua ovuli, the inner surface of which chamber is pitted with shallow
depressions, which receive the chorionic villi, but there is no satisfactory evidence
to show that these pits in the decidua ovuli are the dilated mouths of the uterine
glands.

In the area of the decidua serotina where the placenta is developed, various
observers have shown that a great production of globular and spindle-like cells
takes place, which are intimately intermingled with the chorionic villi. Cart
FRIEDLANDER has recently pointed out that both the decidua vera and serotina
may be divided into two principal layers, an inner cell-layer intermingled with
the chorionic villi, and an outer glandular, which latter lies next the muscular
coat. The cells of the inner layer are elongated or rounded colossal cells,
frequently with many nuclei, and correspond in appearance to those cells
which histologists now term giant cells (Aiesenzellen). Beautiful representations
_ of these cells have been given by ErcoLani in Plate X. of his memoir. The
glandular layer contains, amidst its corpusculated connective tissue, hollow
spaces clothed with an epithelium, the cells of which are partly flattened, partly
cylindrical. These spaces FREIDLANDER regards as the modified utricular
glands ; but he considers that we still need satisfactory proof of the penetration —
of the villi into these glands.

As yet only one anatomist has recorded, in a sufficiently precise form, distinct
observations which seem to show that some at least of the villi of the human
chorion enter the utricular glands in the placental area. JaAssinsky describest
thick villi (dicke Zotten) in the placental region, which differ from the ordinary
chorionic villi in having very few or even no lateral branches, and in possessing
knob-like dilatations at their free ends. In each thick villus, he says, two
structureless membranes and two epithelial layers may be recognised, the inner

* Miiller’s Archiv, 1848, p. 79. t Op. cit. p. 346.

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 495

epithelium is flattened, and with the structureless membrane with which it is in
contact belongs to the villus, whilst the outer epithelium is cylindrical, and with
its structureless membrane constitutes the utricular gland into which an ordinary
vascular villus has entered. Further, he states that all the uterine glands do
not contain villi, but that some remain free. The appearance described by
JASSINSKY is obviously the same as that represented by Professor Goopsir in
his well-known figure (Plate II. fig. 19). And although that anatomist did not
definitely describe the external cells of the villi as the epithelial lining of
the utricular glands, yet he stated that they belonged to the decidua, were
the remains of the secreting mucous membrane of the uterus, and played the
part of secreting cells by separating from the blood of the mother the matter
destined for the blood of the foetus. As regards therefore the morphological
position and the function of these cells, JASsINSky’s observations correspond
closely with those of Goopsir, though he employs a somewhat different mode
of expression in his description. Similarly the layer of flattened spheroidal
cells, which Dr Farre described as forming the sheath or outer case of the
villus, obviously corresponds with the external cells of Goopstr and the outer
epithelium of Jassinsky, and belongs, therefore, to the decidua serotina,
although Farre does not definitely state that it forms a part of that structure.

ERCOLANI again considers that the cells of the serotina, or cells of the internal
epithelial layer, which penetrate between and surround the chorionic villi in the
human placenta, constitute a new-formed glandular organ intervening between
the villi and the maternal blood spaces or lacune, and the cells of which
separate from the maternal blood nutritive material to be absorbed by the villi.
These cells he considers to be derived from cells, which multiply in great
numbers, furnished by the submucous connective tissue of the uterus. Whilst
agreeing, therefore, with Goopsir and JASSINSKY, in considering the cells of the
serotina, which surround a villus, to be concerned in an important way in
foetal nutrition, yet he does not, with the one, regard them as the remains of
the uterine mucous membrane, or, with the other, as the epithelium of the .
utricular glands, but as a new production formed after the period of conception.

Hence there is a common understanding not only amongst these, but other
investigators, as VAN DER Ko1k and Priestiey,* that the villi of the human
chorion are invested by cells, which intervene between the vessels within the
villi and the maternal blood-vascular system, and which play an important part
in foetal nutrition, though opinions differ as to their mode of origin.

But the development of a decidua serotina in the placental area, and the
intimate intermingling of certain of its anatomical constituents, which are
shed at the time of separation of the placenta with the chorionic villi, are not
confined to man, but are found in all animals which possess either the discoid

* Lectures on Gravid Uterus, p. 83.
VOL, XXVI. PART I. 6N

496 PROFESSOR TURNER ON THE GRAVID UTERUS AND

or zone-like form of placenta. It is quite unnecessary for me to go into the
proofs on which this statement rests, as ample evidence of its accuracy has been
already advanced by Professors Escuricut and RoLLEsTon in their valuable
memoirs already so frequently referred to. The shedding of the placental portion
of the serotina at the time of parturition, has led zoologists to class all those
mammals together in which it occurs as caducous or deciduate mammals.

We. may now inquire if there are any structures in the Orca, and inferen-
tially in the other mammals which possess a “diffused” placenta, at all
comparable to the decidua serotina, although it may not be actually shed along
with the foetal membranes.

I have already stated that the cells of the serotina intervene between the
villi and the maternal blood-vessels. In the Orca we have also seen that
although the vessels of the uterine mucous surface are not dilated into sinuses,
but preserve the form of ordinary capillaries, yet that they are separated from
the epithelial investment of the villi, not only by the epithelial cells lining the
crypts, but by the sub-epithelial corpuscles of the mucous membrane. In their
anatomical position these layers of cells correspond therefore to the cells of the
serotina. We have no evidence at present that these layers are separated at
the time when the membranes come away, though I think it very probable if
the chorion of a whale or of a mare, investing a foetus born at the full period of
gestation, were carefully examined, that the epithelial lining of the crypts might
to some extent at least be found to be shed along with it. But during the period
of involution which follows parturition, it is obvious that great changes, either
from actual shedding of portions of its substance, or from degeneration and
interstitial absorption, must take place in these and the other constituents of
the crypt layer before it.can be restored to its proper non-gravid condition.
The difference between the diffused placenta of a whale and the concentrated —
placenta in the human subject appears therefore to be not an essential difference
in the presence or absence of certain anatomical structures, but rather a
difference in arrangement. In the whale the cell structures developed in con-
nection with the uterme mucous membrane, and which occupy the position of
the decidua serotina, are diffused over an extensive surface, and possess a simple
laminated arrangement, and the maternal blood-vessels do not lose their capillary
characters. The chorionic villi, also, are lodged in comparatively shallow crypts,
out of which they can be easily enucleated, either with or without the simul-
taneous shedding of the cell-layers of the crypts. In the human subject, on the
other hand, the cell structures of the serotina, developed in connection with the
uterine mucous membrane, are concentrated in a limited area, and so inter-
mingled with the chorionic villi, that, when these are separated, the cells of the
decidua must necessarily be shed at the same time. The maternal blood-
vessels also assume the form and size of sinuses. Further, I believe that these

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 497

two forms of placentation, which at the first glance seem, and are indeed
usually regarded as, so widely separated, blend with each other through
gradational forms which occur in the other mammalia.

Hence, whilst it may be convenient to employ the terms caducous or non-
caducous, deciduate or non-deciduate, in zoological classification, as expressing
the shedding or non-shedding of distinctly recognisable portions of the uterine
surface, along with the foetal envelopes, they ought not to be regarded as signi-
fying absolute differences in anatomical structure in the two groups of placental
mammals.*

Turning now to a comparison of the arrangement of the membranes which
lie within the chorion, we find that, although the allantois in the cetacea is
elongated towards the two poles of the chorion, yet that it does not reach their
extremities as in the mare and ruminants; still less does it pass through and
beyond to form those pouch-like sacs, which Von Barr many years ago
described in the pig as the diverticula allantoidis. Neither do we find that it
lines either the whole, or even the larger part of the inner surface of the
chorion, as in one or other of these animals, or as in the carnivora; but its
chorionic attachments are limited to that aspect of the latter membrane which
is opposite the abdominal aspect of the foetus. In the persistent condition of
its allantois it differs moreover from the human subject and the other mammalia
in which that membrane disappears at a comparatively early period of gestation.

_ The amnion in Ova, though it does not reach the poles of the chorion, yet
preponderates over the allantois, which is just the opposite condition to the
arrangement met with in the solipeds, ruminants, and pachyderms. Projecting
into the sac of the amnion, though invested by that membrane, are the small
corpuscles, “filiform outgrowths, which are undoubtedly homologous with the
similarly placed growths in the early ruminant, and in the. soliped embryo, as
well as with those on the amnion of the Tenrec,” as RoLieston has already
pointed out. Dr SHArpey also informs me that he has met with a similar set
of bodies in connection with the amnion in Manis. It is to be observed that
these structures are not limited to the part of the amnion which invests the
cord, but are distributed irregularly in connection with its entire surface.

The umbilical vesicle, again, disappears in Orca some time before birth, as
in the mare, pig, and ruminants, and does not persist in the form of a consider-
able sac, as in carnivora, rodents, bats, and insectivora; or as a rudiment, as is
sometimes seen in the human subject.

In its placental affinities the Orca, as will be recognised from the statements
made in the comparison I have just instituted, approaches more closely to the

* It is right to state that Professor Huxtey, by whom the terms deciduate and non-deciduate
were introduced, “ by no means intended to suggest that the homologue of the decidua does not exist
in the non-deciduate mammals.” —-Elements of Comparative Anatomy. p. 108.

498 PROFESSOR TURNER ON THE GRAVID UTERUS AND

mare than to any other mammal, the placentation of which has been accurately
studied. These affinities may be briefly stated as follows :—Both animals are
uniparous and possess an elongated chorion, over the entire surface of which,
with the exception of three limited areas (two polar and one intermediate), well
defined villi are “ diffused.” In both, the amnion is studded with small cor-
puscles, and the umbilical vesicle disappears some time before birth. In both,
the allantois persists as a large sac, but whilst it preponderates over the amnion
in the soliped, it possesses a relatively smaller area in the cetacean. In both,
the highly vascular free surface of the uterine mucous membrane is crowded
with crypts for the reception of the villi of the chorion, and in both the utricular
glands are well developed; but in the mare the glands ascend with a com-
paratively straight stem almost vertically to the crypt-layer, whilst in the
cetacean they are so tortuous as to be followed with considerable difficulty to
their termination.*

Physiological Conclusions.—Finally, I may say a few words as to the physio-
logical conclusions to be drawn from the study of the arrangement of the

placental structures found in this Orca. And as it may help to give one a
clearer insight into this important subject, I shall, in the first instance, briefly
summarise and illustrate, by means of the accompanying diagram, the relative
position of the constituent elements of the placenta in this animal.

* Although the consideration of the placental affinities of the whale shows it to be more closely
allied to the mare than to any other mammal, yet I by no means wish it to be understood that im the
other organic systems a correspondence occurs between the cetacean and the soliped closer than can be
seen between them and any other class of the mammalia. For in their osteological characters, as
Professor Huxtey has pointed out, the cetacea are allied to the true carnivora through the extinct
Zeuglodon and the Seals; in the possession of a compound stomach and of a third bronchus, they
resemble the Ruminants; in the “diffused” character of the chorion, in the presence of a vena azygos
(RoLLEsTON ), and in the remarkable modifications of the cerebral and-intestinal arterial systems, for an
account of which I must refer to my memoir on the Longniddry Balenoptera, they are allied to the
Pachydermata. A full discussion, however, of the relative value of these characters, as determining
the zoological position of the cetacea, would be out of place on this occasion.

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 499

The placenta is distinctly subdivided into two portions, a maternal and a
foetal. The maternal consists of the specially modified uterine mucous mem-
brane, in which, from the attached to the free surface, the following parts may
be recognised :—The utricular glands c, imbedded in the connective tissue, and
surrounded by an open capillary plexus; and the crypt layer, consisting of a,
cup-shaped crypts, and 6, funnel-shaped crypts, into the latter of which the
utricular glands ¢c. open. In this crypt layer are found the connective tissue
(including the fusiform corpuscles d, and the layer of spheroidal sub-epithelial
corpuscles ¢ e) and the close capillary plexus g g; the crypts are lined by
the epithelial layer ff The fcetal portion consists of the chorion, from
which the villi 2 2 project and fit into the crypts of the maternal part of the
placenta. These villi are invested by the epithelial layer 27, and they consist
of connective tissue which contains a layer of sub-epithelial corpuscles 4, of
fusiform corpuscles /, and a close capillary plexus m m, derived from the
umbilical arteries, which plexus is continued into an extra-villous chorionic
network 2n, from which the umbilical vein arises. The foetal and maternal
vessels are not in contact, still less continuous, with each other, but are
separated by the layer of sub-epithelial corpuscles of the villus, the epithelial
investment of the villus, the epithelial lining of the crypt, and the layer of sub-
epithelial corpuscles of the crypt. The space between the two epithelial surfaces
is intended to show the interval between the foetal and maternal portions into
which the secretion of the uterine glands is poured.

It may be useful to compare the above diagram with the well-known diagrams
with which Professors JoHn Retp* and Goopsir have illustrated their views of
the structure of the human placenta. By both these anatomists the tufts of
foetal villi were regarded as intimately related to the dilated uterine sinuses,
the inner coat of which, or a structure continuous with that mner coat, being
reflected on to each tuft. But Goopsrr also represented two systems of cells :—
The external cells of the villus, belonging to the decidua, which lay in imme-
diate relation to the foetal aspect of the ling membrane of the maternal sinus;
the internal cells which lay within the villus between its vascular loop and its
_ external membrane; and he separated the two systems of cells from each other
by a space which he regarded as the cavity of a secreting follicle, of which the
external cells were the secreting epithelia.t The human arrangement, as repre-

* Edinburgh Medical and Surgical Journal, January 1841; and Physiological and Anatomical
_ Researches, p. 325. Edinburgh, 1848.

+ Although from the description which Professor Goopsmr gave of these external cells, he
undoubtedly considered them to be of the nature of secreting epithelium, yet he did not represent them
in his diagram as situated on the free surface of the maternal placenta, but as separated from the space
_ between the maternal and fcetal portions by a sharp line, as if a membrane intervened. How far he
intended this line to represent a definite structure [am unable to say. If such were his intention, then
the external cells would rather correspond in position to my layer of sub-epithelial corpuscles of the mucous
membrane than to a free epithelium. Similarly his layer of internal cells of the villus corresponds in
position, not to the epithelial investment, but to my layer of sub-epithelial corpuscles of the villus.

VOL. XXVI. PART II. 6 O

500 PROFESSOR TURNER ON THE GRAVID UTERUS AND

sented by those anatomists, differs from that in the whale in the sinus-like
dilatation of the maternal blood-vessels, and in the much more intimate relation
of the tufts of foetal villi to those sinuses; the closer approximation of the
foetal and maternal vascular systems being effected not only by the projection
of the tufts into the uterine sinuses, but by the absence, or at least the non-
recognition, of the epithelial investment of the villus. In the whale again, the
utricular glands have not, except perhaps at their mouths, lost their tubular
character, whilst the space between the two systems of cells in the human
placenta, which Goopsir regards as the cavity of a secreting follicle, has, if we
conceive it to be formed by a dilated uterine gland, altogether lost its tubular
form.

It is generally admitted by physiologists that the placenta is an organ in
which a double function is performed—unutritive and respiratory. But there is
a difference of opinion as to how far these different functions are carried on by
the same structures in this single organ, or how far there may not be separate
structures set aside for the performance of each function.

In the early stages of development of the ovum it has indeed been generally
admitted that the uterie glands, and the cells of the decidua reflexa and
serotina, where such exist, elaborate a material which, when absorbed by the
chorionic villi, is engaged in the nutrition of the embryo. But the idea has
been also entertained, that, after the new blood-vessels have been developed,
both in the villi of the chorion, and in the maternal portion of the placenta,
the uterine glands cease to perform their office, and the nutrition of the foetus
is effected by the passage of materials from the blood in the maternal
to that in the foetal blood-vessels. ‘Some have held that the mode of passage —
consists in the simple transudation of these materials through the walls of
the vessels from the maternal to the foetal vascular systems. Others again
have contended,* that two sets of cells intervene between the two systems of
vessels, a uterine set (external cells of villus of Goopstr), which selects from
the maternal blood, and transmits the selected material into the villus; and a
chorionic set (internal cells of villus of Goopsir), which absorbs the material
transmitted prior to its passage into the foetal vascular system.

ERCOLANI again has advocated a doctrine which, in some important parti-
culars, differs from those of his predecessors. He admits that the utricular
glands do furnish materials for the nutrition of the embryo, but only in the
early period of its development. And he strives to prove that, from a transfor-
mation and greatly increased growth of the uterine mucous membrane, and of
the sub-epithelial connective tissue, a new maternal glandular organ is formed,
which in its simplest form consists of secreting follicles, arranged side by side
and opening on the surface of the mucous membrane. In the human subject,

* See the Memoirs of Professor Goopsir, and Drs ArTHUR Farre and W. O. PriestLey.

ON THE ARRANGEMENT OF THE FGITAL MEMBRANES IN THE CETACEA. 501

he says, the typical form of the glandular organ is wanting, but the cells of the
serotina, which invest the chorionic villi, represent the fundamental portions of
the gland organ. Into these new-formed secreting follicles, and not into the
utricular glands, the villi of the chorion penetrate, and are bathed by the fluid
secretion, which they absorb for the nourishment of the foetus. He believes
that these observations completely overthrow the view so frequently enter-
tained that the nutrition of the foetus is due to an exchange of materials by
processes of exosmosis and endosmosis between the two vascular systems.

There can be no doubt that the structures which I have described, both in
Orca and in the mare, as the crypts of the mucous membrane, are the same as
those to which ERcoLANI gives the name of follicles. I have already discussed
the probable mode of formation of these structures, and whilst considering that
the deeper, funnel-shaped crypts* may, from their relation to the glands, be
regarded as their dilated mouths, yet I have inclined to the view that the cup-
shaped crypts are pouches formed during gestation by the folding of the surface
of the greatly hypertrophied mucous membrane. Hence they may be looked
upon as newly-formed follicles ; and, so far, I agree with Erco.ant.

Are they, however, to be regarding as secreting? Here I experience
greater difficulty in accepting ErcoLant’s theory; for although their epithelial
lining is anatomically continuous with that of the utricular glands, yet it is
not of the same character. Both in Orca and in the mare, whilst the glan-
dular epithelium is cylindrical, that lining the crypts is pavement. Now, we
are not in the habit of regarding the pavement epithelium as secreting in its
functions, for in the localities where it is customarily found, it seems to fulfil
merely a protective office, whilst the function of secretion is reserved for cylin-
drical, spheroidal, or polygonal epithelial cells.

Again, I am not disposed to consider that the utricular glands cease to

* perform their functions at an early period of embryo-life. In this Orca,

although the foetus had reached an advanced stage of development, the vascularity
of the glands, their epithelial contents, even the presence of plugs of epithelium
or inspissated secretion projecting through their orifices, all gave one the impres-
sion of structures in a state of active employment. If this be the case, then
the secretion would be poured out into the crypts, and brought in contact
with the villi of the chorion. In the mare, however (and it is quite possible
also in Orca, although I have as yet no positive observations), some of the

* It may be said, as an objection to the inference that the funnel-shaped crypts are the dilated
mouths of glands, that in the mare some of the utricular glands open on the surface by circular
orifices without exhibiting any dilatation, and that therefore the crypts into which the glands open are,
like the other crypts, merely due to a folding of the mucous membrane at that spot. The difference
in the character of the epithelial lining of the glands and crypts, and the similarity in the epithelial
lining of all the crypts, may also be advanced as additional reasons why all the crypts should be
regarded as formed after the same plan, and not by the dilatation of gland orifices.

902 PROFESSOR TURNER ON THE GRAVID UTERUS AND

glands do not open into crypts, and their secretion, consequently, would be
brought in contact not with villi, but with the plane surface of the chorion
between the bases of the villi.

Now in the diffused form of placenta, and it is probable also in some of the
other forms, the villi are not the only absorbing surfaces of the chorion. For
whether we regard the process of absorption as conducted through the agency
of special cells, or of capillary blood-vessels, both these structures are met with,
not only in connection with the villi, but with the plane surface of the chorion
between their bases. Hence, it seems to matter little whether the secretion
is poured into a crypt or not, as in either case it will meet with a chorionic
surface capable of absorption. I am disposed, therefore, to conclude, that in all
those forms of placentation in which the utricular glands preserve their structural
characters within the placental area they play an important, if not the whole,
part in the nutrition of the foetus, not merely in the early, but throughout
the whole period of intra-uterine life.

What office is to be ascribed to the cells which, from their position, I have
called the sub-epithelial corpuscles? I am not disposed to think that these
corpuscles, either in the chorionic or uterine surfaces, are to be regarded as
secreting cells. For in neither case do they lie on the free surface; they are
deeper than the epithelium in position, and are located within the delicate con-
nective tissue itself. In form and appearance they resemble those colourless
corpuscles to which, from their resemblance to the white corpuscles of the blood
or of lymph, the term “lymphoid” is now not unfrequently applied. And it is
quite possible that they may have “ wandered” out of the adjacent capillaries
into the connective tissue. It is probable, therefore, that they may be concerned
in the nutrition and growth of this texture, which processes undoubtedly go
on with great activity during the period of gestation.

I may now say a few words on the respiratory function of the placenta.
No one, I feel sure, could examine the injected preparations of the chorion,
and the uterine mucous membrane in Orca, without coming to the conclusion ~
that the great vascularity of these structures must have a definite relation to
the special functions of the organ. In the mucous membrane a striking con-
trast was exhibited between the vascularity of the glandular and crypt layers.
The former had no greater proportion of vessels than may be found in any
gland of the same type, enough to provide for its nutrition and special secre-
tion. The crypt layer, however, had a remarkable vascularity, very much
greater indeed than we see in connection with secreting follicles, and the
vessels were so arranged as to lie immediately beneath the free surface. The
capillary network closely followed the flexuosities of the membrane, and was
brought into close relation not only with the villi of the chorion, but with the
intermediate portions of that structure. The vascularity of the chorion similarly —

ON THE ARRANGEMENT OF THE F@TAL MEMBRANES IN THE CETACEA. 505

was very great, and the capillary network was distributed not only within the
villi, but beneath the intermediate portions of the membrane. Hence; two

_ extensively diffused highly vascular surfaces, a maternal and a foetal, were

brought into close apposition with each other, and it is to this arrangement,
I believe, that we are to look for the structures concerned in placental
respiration.

The conditions, however, under which the interchange of gases takes place,
differ very materially in the pulmonic and placental respiratory organs. In
the lungs the gases dissolved in the blood have to be interchanged with the
air in the air-cells. In the placenta the gases are in a state of solution on the one
side, in the foetal, on the other, in the maternal blood, and the transmission of
the dissolved gases would take place through the thin layer of fluid secreted by
the utricular glands, which bathes the surface of the chorion. The physical
conditions approach therefore more closely to what we find in aquatic rather
than in ordinary atmospheric respiration.

EXPLANATION OF PLATES.

Figures 1, 2, 7, 13, and 16, were drawn from nature, under my superintendence, by Mr J. B.
ABERCROMBIE ; figures 10, 11, 12, 14, and 15, by Minien Covcurrey, M.B.; and figures
3, 4, 5, 6, 8, and 9, by myself.

Puate XVII.

Figure 1. Gravid uterus of Orca gladiator, with the cavities opened into, reduced jth. The single
arrow is passed through the corpus uteri from one cornu to the other. The double-
headed arrow is passed into the canal of the cervix; a, the ovary; b 0, the Fallopian
tubes, of which the right is cut through; ¢ c, the round ligaments; d, the bladder with
the urethra leading from it; e, the mouth of the vagina.

Figure 2. A vertical section through the ovary reduced one-half. a, the large corpus luteum with its
central cicatrix ; b, the highly vascular body at the hilum.

Figure 3. Cells of the corpus luteum. x 320.

Figure 4. A magnified vertical section through the uterine mucous membrane. a, crypt layer; 8,
glandular layer ; ¢, transversely divided gland-tubes beneath the crypt-layer; d@, a funnel-
shaped crypt with a gland opening into it, the free ends, and not the sides of the
cylindrical epithelium of the glands, were seen in this specimen; e, a cup-shaped crypt;
g, sub-epithelial corpuscles of the crypt.

Figure 5. A magnified view of the utricular glands, with the intermediate connective tissue, as seen in
a horizontal section through the glandular layer of the mucous membrane.

Figure 6. A highly magnified view of an uninjected compound villus of the chorion. The secondary
club-shaped villi are shown with the corpuscles of their connective tissue. The sub-
epithelial corpuscles are indicated at a.

Figure 7. A view of the allantois and amnion displayed by everting these membranes, reduced sth ;
a, the funis; 6b, the horns of the allantois; ¢c, the horns of the amnion; d, the
amniotic corpuscles, of which d’ represents the corpuscle situated near the tip of the left
horn of the amnion.

Figure 8. Cell structures from an amniotic corpuscle. x 480 diameters.

VOL. XXVI. PART II. 6P

504 PROFESSOR TURNER ON THE GRAVID UTERUS, ETC., IN THE CETACEA.

Figure 9. Cell structures from one of the peculiar-looking bodies between the allantois and amnion ;
a, cluster of hexagonal cells; 0, large brood-cell. The cells to the left are sketches of the
more usual forms. x 480 diameters.

Pratt XVIII.

Figure 10. A magnified surface view of the injected uterine mucous membrane at a part where the
furrowed arrangement was well seen. Around the mouths of many of the crypts a
vascular ring may be seen, and at the bottom of more than one of the funnel-shaped
crypts a plug of epithelium projects, as at a, from the mouth of a gland.

Figure 11. A highly magnified view of a uterine crypt obtained in a vertical section through the
crypt-layer. The general arrangement of the capillaries is well shown, and the epithelial
lining, as at aa. An utricular gland, b, may be seen passing to the bottom of a erypt.

Figure 12. A magnified vertical section through the wall of the injected uterus ; a, the erypt-layer ;
b, the gland-layer ; c, the muscular coat. The relative vascularity of the different Pate
is lowe in the figure.

Figure 13. Chorion reduced aii a, the large intermediate non-villous spot opposite the os uteri ;
b b, the polar non-villous spots ; ¢ ¢, the poles of the right and left horns of the amnion.

Figure 14. A napatted surface view of the injected chorion; a a, ‘the intra-villous capillary plexus ;
b b, the extra-villous capillary plexus ; ¢, branch of the umbilical artery ; d, a rootlet of
the umbilical vein. In this figure the umbilical arteries and intva-villous capillaries are
coloured red, the extra-villous capillaries and umbilical vein black.

Figure 15. A transverse section through the cord reduced 4; a, the umbilical arteries; 6 b, the
umbilical veins ; «, the urachus.

Figure 16. Foetus of Orca gladiator reduced 4th.

a a

EXPLANATION OF WOODCUTS.

Page 474. Surface view, under a low power of the microscope, of a portion of the uninjected uterine
mucous membrane. The recesses, furrows, and pits into which the pockets or crypts open
are darkly shaded in the figure. J
Page 479. Stellate non-villous portion of the chorion opposite the os uteri.
Page 483. Outline diagrams to show the arrangement of the membranes at the stage of development
described in the text; A, longitudinal section; B, transverse section; E, embryo ; ch.
chorion ; am, amnion, represented by dotted line ; a/. allantois. .
Page 498. Diagram of placenta of Orca—
a. Cup-shaped crypt.
6. Funnel-shaped crypt, with
c. Gland opening into it.
d. Fusiform connective tissue corpuscles of crypt.
ee. Large sub-epithelial corpuscles of same.
jf f. Epithelial lining of crypts.
g g. Blood vessels of crypt-layer.
hh. Chorionic villi, with
ii. Their epithelial investment.
k. Sub-epithelial corpuscles of villus, and
i. Fusiform connective tissue corpuscles.
m m. Intra-villous capillaries.
nn. Extra-villous capillaries.
The interspace between the foetal and maternal portions, in which the letters a b f and 7
are placed, is for the sake of distinctness made comparatively wide in this figure.

‘. eae teat etree

B74.

*
os
t
‘

ns. Roy. Soc. Edin® Vol. XXVI, Plate XVII.

M‘¥arlane & Erskine, Lith*® Edin*

Roy. Soc. Edin*® Vol. XXVI, Plate XVIII.

M‘Farlane & Erskine’ bith™® Edin™

‘S}

(505)

XIX.—On some Abnormal Cones of Pinus Pinaster. By ALEXANDER Dickson,
M.D., Edin. & Dublin. ; Regius Professor of Botany in the University
of Glasgow. (Plates XIX.—XXIL.)

(Paper read 1st May 1871. Given in for publication 23d October 1871.)

It is well known that although the overwhelming majority of specimens of fir

cones exhibit one or other of the simple spiral arrangements represented by the

é alperlor Lard To) eB . :
terms of the ordinary series =, =, =, 5 &c., whose generating and successive

secondary spirals are indicated by the numbers 1, 2, 3, 5, 8, 13, &., yet excep-
tional cases occur now and again, where we find either conjugate spirals of the
ordinary system, or arrangements (usually simple, but sometimes conjugate)
belonging to other systems of spirals. Ofthese exceptional arrangements, perhaps

the most common are bijugates of the ordinary system, giving the numbers 2, 4,

6, 10, 16, 26, &c., and simple spirals belonging to the system = Hata yi &e.,

giving the numbers 1,3, 4,7, 11, 18, &c. Morerarely, trijugates of the ordinary

system occur, giving the numbers 3, 6, 9, 15, 24, 39, &c.; or spirals of the

system meee £80 1D? 8 &c., giving the numbers 1, 4, 5, 9, 14, 23, 37, &c.;

not to speak of various other arrangements, some of which will fall to be
considered in the special cases which form the subject of the present communi-
cation.

_ The occurrence of exceptional arrangements in structures exhibiting so much
general uniformity as fir cones, naturally raises the question as to whether these
deviations may not be deduced from the simple spiral of the ordinary system, or
from some fundamental form, affording a common basis for both.

In his great work on the Arrangement of the Scales on Fir Cones, ALEx-
ANDER Braun, in speaking of the occasional occurrence of cones where the
most apparent secondary spirals, instead of being, as usual, 5 and 8 in number,
are 4 and 7, refers to the temptation to explain away the anomaly by assum-
ing the abortion of the fifth and eighth secondary spirals respectively, only,
however, to reject such an assumption as an absurdity of the same kind, as
if 2-nary, 4-nary, and 7-nary flowers were considered respectively as 3-nary,
5-nary, and 8-nary ones reduced by abortion ; adding, that “it will be better
VOL. XXVI. PART IL. 6 Q

506 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

calmly to pursue the investigation, rejecting all premature hypotheses, which
can never be more than substitutes for science.”

In their essay on the arrangement of curviserial leaves, the brothers Bravats
approach this question somewhat more closely. In dismissing the idea of
abortion in such cases, Braun occupied his position mainly, it would appear, in
consequence of the absence of any supporting evidence. The celebrated
French authors, on the other hand, actually observed, in capitula of Dipsacus
and in fir cones, certain cases where, in the same inflorescence, there occurred
an actual change from one spiral system to another, accompanied by a diminu-
tion in the number of secondary spirals. They do not appear, however, to have
had a very definite idea as to the exact manner in which this diminution in
the number of secondary spirals is effected. Although on the whole they
seem inclined to treat the phenomenon in question as the result of abortion of
secondary spirals,—as, for example, when referring to certain capitula of
Dipsacus with 16 and 26 as the numbers of the secondary spirals at the base,
suddenly changing to 15 and 26, or 15 and 25, or 16 and 24, they state that
here “there is no doubt as to the abortion of the missing spirals, since they
are as evident as their fellows at the base of the capitulum,”’t—yet, in the
résumé at the end of their essay, considerable uncertainty is indicated in the
sentence, that “the convergence of two spirals into one is to be explained by
partial abortion of one of the spirals, or, if Dee by the coalescence of two
spirals into one.”{

Of cases of this kind occurring in fir cones, MM. Bravats describe two
cones of Pinus Pinaster (Pin maritime), one where the lower part of the cone
123 D8
2. DF 12 Gs

the apex the arrangement changed to 7 8S, 11 D (series 72? Tp ie &e.);§

and a second, in which the ee four-fifths of the cone exhibited secondary

1 2
spirals 9 S, 18 D (series — 5 = os

ment changed. by suppression of one of the spirals by 9, to 8 S, 13 D (ordinary

exhibited secondary spirals 7 S, 12 D (series , &c.), while towards

&c.), while at the upper fifth the arrange-

series oy = ae as » &c.)||. Such cases, along with some others, chiefly in capitula

of Dipsacus sylvestris, lead MM. Bravais into a discussion of the general
question of the transition from one arrangement to another, involving a change
in the number of secondary spirals. As regards their “curviserial” forms,

* Braun, Vergleichende Untersuchungen iiber die Ordnung der Schuppen an den Tannenzapfen ; :
Nova Acta Acad. cm xv. 1, p.d16,
+ L. et A. Bravats, Shi la disposition des feuilles curviseriées. Ann, des Sc. Nat, 2™° sér. vil. —
p. 100.
alc. Ppl OortOe 62.) p. 9s: || Zee. p. 103. =

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 507

however (under which they include such arrangements as those in fir cones),
they are disposed to admit the occurrence only of such transitions as take
place by way of “convergence” of secondary spirals, resulting in diminu-
tion of number. For example, after referring to the Lanes derivation of an

arrangement with 5 and 7 secondary spirals (series = ae &c.) from an

Oo as
ordinary one with 5 and 8, by abortion of one of the spirals by 8, and adding
se : i 1 2 3.5
mau the series 1,4,5,9 ... . « PB 9 12 8 , &e. |, does not admit of

explanation by the way of abortion, and that one can deduce it from the
ordinary series only by supposing a supen/etation, or addition of a new spiral,
among the secondary spirals by 8,” they continue, “This hypothesis appears
to us altogether improbable, since, in the face of an immense number of
instances where two spirals converge into one, we cannot, on the other hand,
cite one (apart from pocliscrial stems), where one spiral diverges into two
similar and parallel ones.’
Not having any strong haliet in the fundamental re enero between the
“ curviserial” and “rectiserial” forms of these authors, and knowing that
“ divergence” or bifurcation of vertical rows (which are, in one sense, to be
regarded only as the steepest secondary spirals) is not very rare in “ recti-
serial” Cacti and succulent Huphorbie, I have not been surprised to encounter,
as I have done, cases of “ divergence ” of secondary spirals in fir cones. In
the following remarks, however, I shall treat chiefly of the “ convergence” of
secondary spirals, a phenomenon which, I think, I shall be able to explain
more definitely than has hitherto been done ; reserving to a future occasion
more extended reference to the phenomenon of “ divergence ” of spirals.
For some of the cones to be described, I am indebted to the kindness of R.
Smytu, Esq., Emyvale, county Monaghan, Ireland; others I obtained in the
woods at Muirhouse (seat of H. Davinson, Esq.), near Edinburgh ; further, I
am indebted to Professor BALFour for permission to examine the collection of
cones in the Museum at the Royal Botanic Garden, Edinburgh, where I found
the remarkable cone of P. lambertiana, which was lately exhibited before the
Society,t and to which I pea recall attention in the following remarks, as also

a cone of P. Pinaster, with 2 =, 5 Spiral, which was exhibited on the same occa-

sion, and of which I give an ene figure in Plate DON fig. 8

I shall, in the first place, give a brief description of the cones forming the
subject of the present paper ; after which we shall be in a position to judge of
and discuss the bearings of the abnormal phenomena.

I. Cone of P. Pinaster—Mr Smytu—(Plate XX. figs. land 2; Plate X XI.

| fig. 1). Length of cone 54 inches. At the base there is a right-handed a

* L.é. pp: LO4, 105. + See Proceedings R. 8S. Edin., vii. pp. 398, 399.

208 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

spiral (series 4 sige lashetiny = &c.), with secondary spirals 9 S, 14 D, 23 8.

A. little above the base, however, two of the nine spirals to the left “ converge ”
into one, leaving, from that pomt up to about the middle of the cone, an
arrangement with secondary spirals 8 S, 14 D, 22S, = a left-handed bijugate
of the series = 5, : = = &c., whose two fundamental spirals have each the
i 2 5 About the middle of the cone, two of the fourteen spirals
to the right converge into one, leaving from thence to the top of the cone an

arrangement of secondary spirals 8 S, 13 D, 21 S, = a left-handed a spiral of

divergence

- i Lo OO is :
2 ss i CIC, MI Ego t
the ordinary series 5 S19 OT 3a &c. The following table (where the
three regions are roughly termed “top,” “middle,” and “bottom ”) will
render the arrangement intelligible :—

Taste A.*—Cone of Pinus Pinaster.

S D iS) D S D iS) 13
2 = —
Top, 1 3 5 8 13 21 34 34
5
Tide —- — 22 -36 =
Middle, 2 6 8 14 36 isxa
Bottom, — 1 4 5 9 14 23 i =

II. P. Pinaster—Mr Smytu—(Plate XIX. fig. 1; Plate XX. fig. 3; Plate
XXI. fig. 2). Length of cone 53 inches. From the base to near the top there
PedineBesSened
3 47 tgs
4D,758,11 D. Near the top of the cone, however, two consecutive scales in
one of the spirals by 4 to the right (adjacent scales of two of the spirals by 7 to
the left) have partially coalesced, giving beyond that point an arrangement of
secondary spirals 4 D, 6S, 10D, = a left-handed bijugate of the ordinary series,

whose two fundamental spirals have each the divergence oxo The following

is a right-handed x spiral (series &c.), with secondary spirals,

table represents the arrangement :—

Taste B.—Cone of P. Pinaster.
D Ss) D Ss D

3
T — =
op, 2 4 6 10 16 8x2
5

* In this and the following tables, under S, are indicated the numbers of spirals, generating as
well as secondary, running to the Jeff; under D, the numbers of those running to the right ; while

under V, are indicated the numbers of vertical rows.

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 509

Ill. P. Pinaster—Muirhouse—(Plate XX. figs. 4 and 5; Plate XXII.
fig. 1). Length of cone 34 inches. Some scales on one side are somewhat
damaged, by having been bitten, apparently before the cone was mature. The
lower third of this cone exhibits a left-handed trijugate of the ordinary sys-
tem, with secondary spirals 6 D, 9S, 15 D, 24 S, and whose three fundamental

oe Two of six spirals to the right
“ converge ” into one, producing in the middle third of the cone the arrangement
me, 9S, 14 D,23S,=a right-handed spiral (series = = = = se = &c.).
In the upper third of the cone, we have two further changes: in the first place,
two of the nine spirals to the left “converge” into one, giving the arrange-
ment 5 D, 8 S, 13 D, = a left-handed spiral of the ordinary series, probably
po
34’
by 5 to the right (adjacent scales of two of the spirals by 8 to the left) have
partially coalesced, almost precisely in the same way as the two scales near
the top of the last cone, giving beyond that, up to the top of the cone,
the arrangement 5 D, 7 S, 12 D, =a right-handed spiral of the series
m23 5 8 13

>» 5 7 19’ 19" &c., probably 31°

The changes in this very complicated cone are shown in the following
table :—

spirals have each the divergence

; and then, a little higher up, two consecutive scales of one of the spirals

Taste C.—Cone of P. Pinaster.

S D iS) D S D S V “8
= : a 2 9 a
Top, 1 2 5 7 1 i 3]
13
Below Top, 1 2 3 5 8 13 21 34 = 34
8
Middle. — 1 4 5 9 14-98 T=
)
— — : ) 15 24 39 =
Bottom, 3 6 9 a,

IV. P. Pinaster—Muirhouse—(Plate XX. figs. 6 and 7; Plate XXII. fig.
2). Length 44 inches. In this cone, nine secondary spirals to the left run
continuously from near the bottom to the top. At the very bottom, there is a
considerable amount of irregularity, some of the scales being of exceptional
size and shape. It is probable, however, that were it not for the mal-develop-

ment, the arrangement here would be a —, 7 spiral, similar to what we meet

16
further up on the same cone. However this may be, a “ divergence ” distinctly

takes place of one secondary spiral into two, which form members of a set of
eight running to the right. Thus the first arrangement which we can legitimately
| VOL. XXVI. PART IL. 6R

510 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

determine in this ae 24 one with secondary spirals 8 D, 9 S, = a left-handed

spiral (series 4 &c.). This spiral is continued through between 40

ily 8’ 5 A ‘
and 50 scales of the cone, when two of the eight secondary spirals to the right
converge into one, giving us the arrangement 7 D, 9 S, = a right-handed

4
279 16
a and 30 scales, gives place to a left-handed trijugate of the ordinary series,
with divergence

A g Spiral (series —, : iS &c.), which, after being continued through between

= 3 by two of the spirals by 7 becoming replaced by one
of a set by 6. Neglecting the very bottom of the cone, we have the changes
represented in the following table :—

TasLe D.—Cone of P. Pinaster.

D S) D Vv .

Top, — 3 6 9 15e— 5K 3
Middle, Ha Fid “ye nen
16
Bottom (a little above the) — 1 8 9 17 = 2

V. P. Pinaster—Muirhouse—(Plate XTX. fig. 3). Length of cone 33 inches.
i eae bans 8 ae i :
The upper part of this cone exhibits a left-handed Tl (or possibly 7 spiral.

Carrying the eye downwards, however, we begin to observe, a little below the
middle, rudimentary scales of small size and somewhat peculiar shape, inter-
calated with considerable regularity among the others, so as to appear as_pro-
jections placed between the angles of the larger scales. As the base is
approached, the scales in the downward continuation of the larger series become
gradually reduced in size, till they are practically indistinguishable from the
smaller ones; the general arrangement becoming, at the same time, so
crowded and confused as to render precise determination of the spiral ~
impossible.

In addition to the above, I would recall attention to the abnormal cone of Pinus
lambertiana which I exhibited to the Society on a former occasion. This cone is
noteworthy, not om as Showing a transition by convergence from a bijugate of the

system See m7 - &c., giving the numbers 4, 10, 14, &., to a simple spiral of
Y ¢ a
the system |, == 5° = a &c., giving the numbers 1, 4, 5, 9, 14, &c., but also

as showing a transition, conversely, by divergence, from the latter to the
former.

The following table represents the arrangement :—

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 511

Taste E.—Cone of Pinus Lambertiana, in Museum, Edinburgh Botanic Garden.

Ss bs KS D Ss Vv i
o
Top, il 4 5 9 14 Bh) = 33
: 5
Middle, — 2 4 10 14 4= iax2
5
Bottom, 1 4 5 9 14 23 = 53

Having submitted the foregoing facts, I shall now proceed to consider :—

1st, in what the so-called convergence of secondary spirals really consists ;

2d, what constitutes affinity of different spiral systems as regards their
possible or actual derivation one from another ; and

3d, whether it is possible to conceive of the varying spirals in fir cones, or

in other plants, being mediately or immediately derived from some one funda-
mental arrangement in a given set of cases.

1st, As to the nature of ‘‘convergence” of secondary spirals. It cannot
fail to strike the attentive observer that a certain absurdity is involved
in the idea of coalescence, or fusion of two secondary spirals into one.
Secondary spirals, it is to be remembered, have only a relative existence. For

Sapa :
example, in a cone with an — aI spiral the very same scales which constitute the

eight secondary spirals running to the one hand, make up the thirteen
running to the other. The same objection applies to “ convergence ” considered
as the result of an abortion or suppression of a secondary spiral; for it is as
difficult to conceive of the abortion of one spiral, which has only a relative
existence, as of the fusion of two similarly circumstanced. ‘To take a special
case. About the middle of Cone er described above, an ordinary trijugate

_ changes by “ convergence” into an = spiral; but it would be quite as correct

to say that coalescence (or abortion) occurred among the spirals by 15, or
by 24, as among those by 6, &c. (see Plate XXII. fig. 1). MM. Bravais
appear to have felt this difficulty, but content themselves with arguing for the
probability of the abortion occurring among the more apparent secondary
spirals, where the successive insertions are in contact, as contrasted with the
improbability of its occurrence among the less apparent secondary spirals
whose insertions are not contiguous.

Having disposed of the hypotheses of abortion and fusion of secondary
spirals, respectively,—hypotheses which are, in fact, little more than different
attempts to express what is simply one of the results of a certain disturbance,
viz., diminution in the number of secondary spirals,—we proceed to inquire if
there are any facts to guide us in ascertaining the proximate cause of the disturb-

912 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

ance itself. To do this we have only to look at the upper portion of Cones IT.
and III. In each of these a change is ushered in by the partial fusion of two
adjacent scales. These scales may be viewed either as consecutive members of
one secondary spiral, or as adjacent members of two parallel ones ; as, however,
their relative position is strictly defined by reference to the spiral in which they
are consecutive members, which it cannot be by simple reference to any two
parallel spirals in which they occur, the former description is that which must
be adopted. In these cases it is impossible to doubt that it is the coalescence or
Susion of two consecutive scales in one of the secondary spirals which leads to or
causes the general disturbance of the arrangement. When such a disturbance
takes place, a convergence of certain secondary spirals becomes prominently
visible ; but this convergence is not at all more real, although more apparent,
than the convergence of other secondary spirals whose component members do
not happen to be in contact. It will be noted, moreover, that this disturbance
affects the numbers of all the secondary spirals, excepting only those among
which the fusion of consecutive scales occurs ; for example, in Cone II. consecu-
tive scales in one of the spirals by 4 have coalesced, and it is these spirals by 4
alone which run continuously throughout the two systems without diminution in
number. That a similar explanation legitimately applies to all cases of ‘ con-
vergence,” even where the duplex nature of what I would term the scale of
convergence is not demonstrable, I think few will be inclined to doubt.

At this point, I may now conveniently refer to the method to be adopted in
numbering the scales on the cones exhibiting ‘‘ convergence.” From the
circumstance of the “scale of convergence” resulting from the fusion of two
adjacent scales which are usually at a considerable interval from each other on
the generating or fundamental spiral, it is evident that the disturbance conse-
quent thereon must, at the very least, extend to all the scales which would
have been included between the numbers of the two scales which have
coalesced, if, indeed, it does not involve a region of the cone extending both
above and below the level of the scale of convergence. At first sight it might
appear to be the simplest method to consider the scale of convergence as the
point of passage from the one system to that succeeding it—as at once the last
term of the lower system, and the first of the upper. Here we might reckon
the scales in the lower system either up to the lower, or up to the higher, of
the two components of the scale of convergence. In either way, however, we
should encounter a difficulty; as in the former case a considerable number of —
scales would escape the reckoning, while in the latter the double enumeration
of a considerable number of scales would be involved. The disadvantage of
double enumeration also attaches to a method suggested to me by a friend,
viz., to number the scales from below the level of the convergence, according to
the spiral of the lower system, as far up as one can go above it, and from above —

Py *

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 5138

that level, according to the spiral of the upper system, as far down as
one can go below it. The method now to be described, and which I have
employed in numbering the outline figures of the cones in Plate XX. figs. 1-7,
and in the construction of the plans or diagrams of the same in Plates X XI.
and XXII., although to a certain extent artificial and arbitrary, yet has
the advantage of reducing to a minimum the number of doubtful or ambiguous
scales. My procedure is as follows :—The arrangements above and below a
given convergence are noted; then, in order to ascertain how the secondary
spirals of two such arrangements would most naturally fit or run into one
another, constructions of the two arrangements are made in such a way that
the last term of the lower system coincides with the first (¢.e., No. 0) of the
upper; or if either or both of the systems happen to be conjugate, one of the
last terms, or of the first, or of each, as the case may be, is to be placed at the
common point of the two systems. Further, the constructions are made so
that those secondary spirals which correspond in number and direction
in the two systems shall be continuous ; these spirals being, as above indi-
cated, those among which the fusion of consecutive scales has, or is
presumed to have, occurred. To take an example. Supposing we have

to join or fit together a spiral below, and a 16 spiral above, as in Plate

2
ivy
XXII. fig. 2; a construction is made of the lower spiral, and its last term is
taken as the starting-point of the one above, which is thence constructed in
such a way that the lines of its secondary spirals by 9 are continuous with the
9 in the arrangement below. The lines of the secondary spirals by 8 in the
lower, and of those by 7 in the upper system, are now drawn (these being the
lines among which the convergence is most apparent, and which, of course,
have approximately the same direction), when the next proceeding is to join the
8 below to the 7 above, which is effected as follows :—the lower extremities of
the seven upper lines are joined to the upper extremities of those seven of the
eight below, which lie nearest them in the same direction. The upper extremity
of the eighth or remaining lower line is then joined to the nearest (in the same
direction) of the lower extremities of the seven above, when it becomes
apparent that, in the construction, it is this last point which is the point of
convergence, and, moreover, that this point corresponds to No. 6 of the upper
system. The cone under examination is now taken, and the “scale of
convergence” marked as No. 6 of the upper spiral. This upper spiral is now
reckoned back to No. 0, up to which point, on the other hand, the scales of the
lower spiral are continuously reckoned. When this has been done, it will be
found, in the case before us, that one scale has been excluded from the
enumeration; and, on turning to the construction, it will be seen that this
scale occupies a position corresponding to a point where one of the junction
VOL. XXVI. PART II. 6S

514 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

lines between the upper seven and lower eight spirals happens to intersect one
of the lines by 9 running continuously through the two systems in the other
direction. In the case before us, there is only one such intersection ; but, as
we shall presently see, there may be two or more such points in similar
associations of other spiral systems, which, in the same way, will be found to
correspond in position to scales excluded from the enumeration ; or, again,
there may be no such intersections, in which case every scale in the cone can
be included in the continuous enumerations. Again, it will be found that the
different transitions differ in the value in the upper system to be attached to
the scale of convergence. To illustrate the above, I may refer to the diagrams
in Plates X XI. and XXII., with which the outline figures of the corresponding
cones on Plate XX. may be compared. In doing so, it will only be necessary
for me to indicate the spirals by their systems,—thus, 1, 2, 3, 5, 8, &c., or 1, 3, 4,
7, 11, &c., and so on,—the particular term of its series to which a given spiral
belongs being quite immaterial as regards the present question. The unnumbered
scales on the outline figures of the cones, and the intersections corresponding
thereto in the diagrams, I have marked with asterisks. In Plate XXI. fig. 1,
we have the system 1, 4, 5, 9, 14, &c., passing into the bijugate 2, 6, 8, 14, &c.,
with three intersections corresponding to three unnumbered scales on the cone,
and No. 2’ of the upper system as the point or scale of convergence ; also, this
. same bijugate 2, 4, 6, 8, 14, &c., passing into 1, 2, 3, 5, 8,18, &c., with one
unnumbered scale, and No. 6 of the new system as the scale of convergence. In
Plate XXJ. fig. 2, the system 1, 3, 4, 7, 11, &c., passes into the bijugate
2, 4, 6, 10, &c., with no unnumbered scales, and No. 1’ of the new system as
the scale of convergence. In Plate XXII. fig. 1, we have the trijugate 3, 6, 9,
&c., passing into 1, 4, 5, 9, &c., with two unnumbered scales, and No. 4 of the
new system as the scale of convergence; then 1, 4, 5, 9, &c., passing into
1, 2, 3, 5, 8, &c., with no unnumbered scales, and No. 4 as the scale of
convergence ; and, lastly, 1, 2, 3, 5, 8, &c., passing into 1, 2,5, 7, &c., with one
unnumbered scale, and No. 3 as the scale of convergence. In Plate XXII.
fig. 2, we have 1, 8, 9,17, &c., passing, as above mentioned, into 1, 2, 7, 9,
16, &c., with one unnumbered scale, and No. 6 as the scale of convergence ;
and 1, 2, 7, 9, 16, &c., passing into the trijugate 3, 6, 9, 15, &c., with two
unnumbered scales, and No. 1 as the scale of convergence. *

With regard to the second point that I proposed to consider,—viz., as to what
constitutes afinity of different spiral systems as regards their possible or actual
derivation one from another; in other words, upon what the aptitude of
different spirals to pass into one another depends,—it might, @ priori, have been

* Tt might, perhaps, be possible for a mathematician to furnish a formula, whereby, from two
spiral systems given, to deduce the number of ambiguous scales, and the value in the upper system oF
the scale of convergence, thus saving the trouble of a preliminary geometrical construction.

PROFESSOR: DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 515

imagined that resemblance in the divergence of the generating spiral would be
the bond of union in cases of transition. It is manifest, however, from the
instances I have given above, that although in some of the cases we find a close

resemblance between the generating divergences of the two spirals between

which a passage occurs,—as, for example, between = and == —yet much more

frequently do we find the transition occurring between spirals with widely dif-
ferent generating divergences,—for example, between a and o , or between

17
= and = Such facts, coupled with the circumstance that in these transi-

tions ee a of the generating spiral is very frequently reversed, and that,
not unfrequently, we have a transition from a simple spiral to a conjugate or
vice versd, are sufficient to show that the aptitude of spirals of different systems
to pass into each other is quite independent of what is ordinarily supposed to
be the most essential element of a spiral arrangement, viz., the divergence of
the generating spiral. On the other hand, it is to be noted that the corre-
spondence in the numbers of the secondary spirals and verticals is always very
close between the spirals which pass into one another; indeed, so much so,
that the conclusion seems forced upon us, that here we have the essence of what
may be called the genetic afinity in such cases, which may be expressed in
the following terms :—that, as regards their production or origination, spirals
of different systems are to be considered as allied in proportion to the numerical
correspondence of their secondary spirals and verticals.

I shall now turn to the third and last point to be considered, viz., whether
itis possible to conceive of the varying spirals in fir cones, or in other plants,
being mediately or immediately derived from some one fundamental arrangement
ina given set of cases. This question opens up an interesting, but I am afraid

very dangerous, field of speculation. To simplify matters we may confine our
~ attention to a few of the commoner arrangements—arrangements which may be
found to prevail over an entire cone. Among such (as I have already indicated
at the commencement of this paper) are the simple spirals of the ordinary
system ; after which the bijugate of the ordinary system and spirals of the

system ee? TP ie &c., are conspicuous ; and after these the trijugate of the
1 ga

ordinary system and spirals of the system — 2B 9 14’

Have these different forms a common origin? If they have, it is in the highest
degree probable that their descent from that common origin is by way of
“convergence.” Although admitting the occasional occurrence of divergence of
secondary spirals, yet I am ready to agree with MM. Bravats in considering
it as an improbable element in the production or derivation of the commoner
forms.

&c., may be mentioned.

516 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

The idea which, perhaps, most naturally occurs is that the simple spiral
of the ordinary system is the fundamental form; we may therefore, in the
first place, look how the other forms may be derived therefrom.

(a.) The ordinary bijugate is derivable from the ordinary simpie spiral
through the intervention of the system 1, 3, 4, 7, 11, &c., thus,—

ie 2 Ono ate
eo ee a ee
2 4 6 10

(.) The system 3, 4, 7, 11, &c., is directly derivable from the ordinary

system, thus,—
eo oe Oe

eo Se ele

(c.) The ordinary trijugate appears to be derivable from the simple spira
of the ordinary system only by way of the bijugate, and hence thus,—

£2 13. oD aoe las
L in Dial Atlee ptt
Dye et. HAD
Dip ree Fo

(d.) The system 1, 4, 5, 9, 14, is similarly derivable from the ordinary
system by way of the bijugate, thus,—
bo Drs SVS rhs Qt
AOE SRS aS
2 4 6 10 16
fi C4 OIE

To the foregoing derivations the most serious objection is, that it seems
improbable that the bijugate, which is the commonest of the anomalous forms,
should be the result of two convergences ; and still more so, that the not very _
rare trijugate of the ordinary system, and the also not very rare simple system
1, 4, 5, 9, 14, &c., should each be the result of three such transitions. |

In the next place, let us take the ordinary bijugate and see how the other
systems may be derived from it.

(a.) The ordinary simple system is derivable by convergence from the
bijugate only, it would seem, by way of the system 1, 4, 5, 9, &c., thus,—

2 4 6 10
eae Seg
Ly. Soe te

* It will be remembered that in Cone III. the system 1, 2, 5, 7, 12, &c., is derived from the
ordinary system by coalescence of consecutive scales in one of the secondary spirals by 5. Here, the
system 1, 3, 4, 7, 11, &c., would be derived from the same by coalescence of consecutive scales in on
of the secondary spirals by 3. r

ihe

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 517

This derivation is highly improbable, and, indeed, quite imadmissible ;
although it is interesting to note that in Cone III. (see diagram in Plate X XII.

fig. 1), a spiral [a] of the ordinary system is actually derived from the system

1, 4, 5, 9, &c. Failing this derivation, we may ask if it is possible for the
ordinary simple system to be derived from the bijugate by abortion of one-half
of the scales? At first sight this derivation seems, and perhaps is, much more
improbable than the first one. In the very remarkable cone, however, described
above, and figured in Plate XIX. fig. 3, a process of abortion of this kind, arrested
half way, seems to have occurred. In looking at the lower part of this cone,
it is quite apparent that had the large and small scales been developed equally
we should have had a bijugate arrangement. It is just conceivable, however,
that these small scales may be the result of a “ superfcetation ” (to borrow MM.
‘Bravais’ very obstetrical term) ; but this does not seem at all probable. The
derivation by abortion would, of course, be represented thus,—

2 4 6 10 16
Ree 3 os 98

(.) The system 1, 3, 4, 7, 11, &c., is derivable from the ordinary bijugate
only by way of the simple spiral of the ordinary system, and this may be done
by either of the methods indicated under the last head.

(c.) The ordinary trijugate, as already indicated, is directly derivable from
the bijugate, thus,— vs

2 4
5) 9

6
6
(d.) The system 1, 4, 5, 9, &c., is also directly derivable from the bijugate
of the ordinary system, thus,—
a a
4 5 29
In looking at the two foregoing schemes of derivations, it is evident that the
great difficulty lies in the absence of any probable derivation, either of the
ordinary bijugate from the ordinary simple spiral, or conversely, of the ordinary
simple spiral from the bijugate. As to the other systems, it is evident that 1,
3, 4, 7, 11, comes most naturally from the simple ordinary spiral; while the
ordinary trijugate and the system 1, 4, 5, 9, 14, are most readily derived from

the ordinary bijugate. At one time I was inclined to accept the bijugate as the
fundamental form ;* but the more I reflect on the matter, the more I am con-

* This view I published (under reserve) in the abstract of this communication in the Society's
Proceedings, vol. vii. p. 453.

VOL. XXVI. PART II, OT

518 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

vinced of the futility of the endeavour to derive the different forms from one
origin. Indeed, we seem almost compelled to recognise both the ordinary simple
spiral and the ordinary bijugate as fundamental forms, i.e., forms with either
of which a cone may commence without the intervention of another; and if this
be done, the derivation of the various systems from the one or from the other
would be a very simple matter.

Before concluding, I would submit a tabular analysis of the five flower-
spikes of Banksia occidentalis, already brought under the notice of the Society,
but which may very profitably be reconsidered in connection with the above
communication.

Tabular Analysis of five flower-spikes of Banksia occidentalis in Museum of Economie Botany,
Royal Botanic Garden, Edinburgh.

One, with — — — 7 7 14 a
Two _ — 1 6 v6 13 = a
ey 13
One — ] Z 5 7 i a
; A 12
One 1 2 3 4) 8 Wy = 8
+ 13

I shall not stop to inquire from what one or more fundamental forms these
may be derived, but it is very interesting to note how, among these forms, we have
spirals of the most widely different fundamental divergences, closely resembling
each other in the number of their secondary spirals and verticals, just as we
have seen above in the case of the fir cones.

It is possible that, in ignorance, I have gone over ground which has been —
trodden before; for example, I have not had access to any of ScHIMPER’s
works on phyllotaxis. However, as it is in the highest degree improbable that
the abnormal forms above described should repeat themselves, I rest assured
in the belief that I have contributed at least new, if not valuable matter. }

In conclusion, I would specially thank my friend Professor Tair for the —
many valuable suggestions I have received from him in the course of this
work, and for his patience and readiness in assisting me when in any
difficulty.

PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER. 519

EXPLANATION OF PLATES XIX., XX., XXI., XXII

[In Plate XTX. the figures are drawn on stone from photographs.

In Plate XX. the figures are photo-lithographic reductions from outlines made in the following
manner :—Photographs of the cones were obtained (the cones having been previously painted of a
uniform grey colour), and on these the outlines were carefully gone over with a steel “ crow-quill” and
Indian ink. This done, the photographs were then washed out with a solution of Cyanide of Potassium
(about 5 grains to the ounce of water); the outlines drawn remaining, but now, of course, on white
paper. These outlines, after being retouched and intensified with Lamp-black, and receiving the
addition of the numbers, were then photo-lithographed to the scale required, and colour added by
chromo-lithography. I have thought it worth while to record the above process, as it may be found to
be very useful in many cases where an accurate outline of a given object is required.

The diagrams in Plates XXI. and XXII. are photo-lithographic reductions from drawings on a
larger scale. |

Puate XIX.

Figure 1. Cone II.; about natural size. The two secondary spirals by 7, which converge into one of a
double set by 3 at the top, are distinguished by being shaded of a lighter tint than the
others. The scale of convergence is obviously double.

_ Figure 2. Cone III.; somewhat magnified. Here, as in the last case, the scale of convergence near the
top of the cone is obviously double.

Figure 3, Cone V.; somewhat magnified. Showing the small scales intercalated among the larger ones
towards the lower part of the cone.

Puate XX.

Figures 1 and 2 represent different aspects of Cone I.; considerably reduced. In fig. 1, two secondary
spirals by 9, coloured red, are seen to converge into one by 8, which is continued to the
top of the cone. In fig. 2, two secondary spirals, coloured blue, in a double set by 7, are
seen to converge into one by 13, which is continued to the top of the cone. See diagram
in Plate XXI. fig. 1.

Figure 3. Cone II. ; considerably reduced. The same view as that in Plate XIX. fig. 1. Two secondary
spirals by 7, coloured red, are seen to converge into one of a double set by 3. The scale
of convergence is obviously double. See diagram in Plate XXI. fig. 2.

Figures 4 and 5. Different aspects of Cone III.; about naturalsize. Fig. 4 is the same view as that
in Plate XIX. fig. 2. Two secondary spirals, coloured blue, in a triple set by 2, converge
into one by 5, which is continued to the top of the cone. Of the three secondary spirals
by 9, coloured red, the two uppermost converge into one by 8; which last, and the lower-
most of the aforesaid three, converge in their turn into one by 7. The last scale of
convergence (coloured purple from the blue and red spirals happening to cross) is, like
that near the top of Cone II., obviously double. See diagram in Plate XXII. fig. 1.

Figures 6 and 7. Different aspects of Cone IV.; considerably reduced. In fig. 6, two spirals by 8,
coloured red, about the middle of the cone, are seen to “ diverge” from a single one at
the base. In fig. 7, two secondary spirals by 8, coloured red, converge into one by 7;
while, higher up, two of the spirals by 7, coloured blue, converge into one of a triple set
by 2. See diagram in Plate XXII. fig. 2.

Figure 8. Cone, from Museum of Economic Botany, Royal Botanic Garden, Edinburgh; considerably
reduced (natural size 44 inches). The arrangement in the lower part of this cone is
somewhat confused, and has not been determined. From about the middle, however, up
to the top, a regular 2% spiral (series 4, 3, 735, 2%, &c.) is exhibited. As has been pointed
out by MM. Bravais, this system is readily derivable by convergence from the system 1,
4, 5, 9, 14, &e.

520 PROFESSOR DICKSON ON SOME ABNORMAL CONES OF PINUS PINASTER.

Figure 9. Cone, from Mr Suyru, Emyvale ; consieomly reduced (natural size 44 inches). This cone
exhibits a very regular spiral of the series }, 4, 2, 3%, &c. It will be seen that the lines
by 37 are scarcely vertical, so that the spiral has probably the divergence 33.

[Puares XXI. and XXII. The diagrams or plans here hardly require explanation. The regions
exhibiting the different spiral arrangements are marked off from each other by horizontal lines projecting
laterally at the level where each new system commences; the arrangement in each region being
indicated by a fraction alongside of it. I have marked the “scales of convergence” with their number —
in the upper of the two systems in the respective cases ; adding, within brackets, the numbers of the
actual or presumed components of these scales according to Hee spiral of the lower system,—the actual —
without, the presumed with, a mark of query. ]

Pruate XXI.

Figure 1. Diagram of Cone I, It will be noted that here, in order to save room, the u
; ppermost spiral
is carried up only to No, 36 of the 106 or 107 scales in the sei es of the
cone.

Figure 2. Diagram of Cone I.
Prats XXII.

Figure 1. Diagram of Cone ITI.
Figure 2. Diagram of Cone IV.

UP OUT aunsay g BURLAP SPT

Roy. Soc. Edin*

Vol. XXVI, Plate KX.

Py Gren ~~. Q hg

- a . jet vio ee
b Om 7% O78 <
> 0 : za

f
af i A Dickson, M.D. M‘Farlane & Erskine, Lith"® Edin*

Trans Roy. Soc. Edin”

Vol. XXVI, Plate XXI.

as ee, |
| ee cS
LIPRARSCS

NRPY SD
NON = = PY
Sho / Oa
K ea Ca

(Bijug, axe )

SO TS, Ge <—= :
2a Ga

Qh.5 20h)

XX.— Account of the New Table of Logarithms to 200 000.
By Epwarp Sane, Esq.

(Read 20th February 1871.)

b
The essential character of all tabular aids to computation is, that the results

of many operations are recorded in some systematic way for easy reference,
and that thereby the computer is spared the toil of obtaining these results for
himself.

o In many cases this constitutes almost the whole advantage of the table.
Thus when, instead of extracting the cube root of some number, we take it
from a printed book, we are merely using another's labour. The gain to the
t calculating community is, that the oft-repeated extraction of the same root is
avoided. We also gain by the facility of systematic calculation ; the labour of
computing a series of successive results being in general only a small fraction
of that which would have attended the same work performed in a desultory
manner.

The possession of a table of the corresponding values of two connected
magnitudes enables us to perform the inverse operation, that of finding the
argument from its function, an operation generally much more difficult than
that of finding the function from the argument. Tables special to his own
pursuits are thus indispensible to every investigator.

- But tables of logarithms possess advantages of a peculiar nature. Except
in a few rare speculations, no one desires to know a logarithm for its own sake ;
no , im general, would that knowledge be of any use to us. The ability to
mpute the logarithm belonging to a number, or the number belonging to a
arithm would, of itself, be almost barren of useful result. If, in order to
ly the neperian process to ordinary multiplication, we had to compute the
ngarithm of each factor, add these together, and thence compute the corre-
ponding number, we should have expended a hundred times the labour of the
rdi nary process. Viewed abstractly, Napier’s process is ridiculously cir-
uitous ; its whole advantage is and was intended to be derived from tabula-
‘ion ; so much so, that the mechanical operations of paper-making and printing
nter among its constituent parts almost as essentially as the arithmetical
somputation itself. Napier’s original conception was of a table to subserve
bg ain ends, and his efforts were directed not to the discovery of a single
ogarithm, but to the construction of a logarithmic system. Here the table is
everything,
_ VOL, XXVI. PART III. 6U

524 MR SANG’S NEW TABLE OF LOGARITHMS TO 200 000.

From this example, the general principles which regulated the actual course
of proceeding may be understood. Two things have to be kept in view when
seeking for a convenient way of getting at the logarithm of a proposed prime
number, one to get an easy divisor, the other to obtain by a change of sign the
logarithm of some other number not previously found, preferably a prime
number.

For example, we have filled the list of prime numbers up to 29, the logarithm
of which has now to be found. Our first business is to search for some
multiple of 29 which ends in 0001, or in 9999, in order that the divisor terminate
in zeroes, the more the better; 29 ends in 9, and therefore we may use the divisor
30, which would also give us the logarithm of 31; this divisor, however, is too
slow, so we carry on our search thus :—

ee oa 29
30 '* 129 =O 870
ol x. 29°" 2809

900 x 29 = 26 100

931 x 29/=526,999

There the divisor 900 would have done, provided the logarithm of 31 had
been known ; wherefore we proceed another step, which brings us to the divisor
27 000; this divisor is available if the logarithm of 931 be known. On turning
to the filled-in table of natural numbers, we find the logarithm of 931 there; it
had come from the product 7°7:19. From 27 000 we also get 27 001, and there-
fore inquire whether this be a prime or a composite number. This research in ~
itself would have been enormously tedious, so much so that any saving from the
discovery of the factors would have been but a small set-off against the labour
expended. That most admirable table, however, of the Divisors of Numbers —
constructed by BurKHARDT makes the matter easy;* it shows us that 27 001 is
the product of 13°31°67; so that once the logarithm of 31 is found, that of 67

also may be obtained. Wherefore, making «= = a0? We obtain log 29 and log
27 001.
Subsequently making — we have log 31, log 901, and log 53; and

thence again log 97.

Here it may be proper for me to bear testimony to the great value of
BuRKHARDT’S work, which contains the divisors of all numbers up to three
millions. The prodigious amount of labour, in the face of an expected small

*In this particular instance we might have done without Burkuarpt’s help, because 27 001
= 30%4-18 and so is divisible by 3041.

MR SANG’S NEW TABLE OF LOGARITHMS TO 200 000. 525

return, is only equalled by the scrupulous carefulness of the execution. For
many years, and in various branches of research, I have habitually used the
Table des Diviseurs, and only in one instance have found a fault,—that fault
having been caused by a displacement of the types in the process of printing.

So long as the primes were under 1000, their logarithms were compared
with those given by CALLeT in his Tables Portatives to 60 places; and the
coincidence was held as a sufficient check. In no one instance was an error
found in Cater. Afterwards, however, each logarithm was computed twice,
generally once from a multiple ending in 01, and once from another ending in
99. At times the divisors were enormously large ; thus, for the prime 653 the
divisor 249 000 000 was used. In such cases the advantage of the second result
was lost, since it would have been a matter of great labour to have found the
divisors of 248999999. It would be still more laborious in the case of
7 580 000 001 attending the computation of the logarithm of 1277.

The computation of the logarithms of all the primes in succession to above
2000 was thus carried on, and those of many other primes incidentally found ;
these and the logarithms of their multiples up to 10 000 to twenty-eight places,

having been written in their places, a sufficient groundwork was obtained for a

table to fifteen places, beginning at 100 000, since the differences of the third
order there count only in the sixteenth place.
Paper having been ruled, and the lines numbered from 100 000 to 150 000,

_ the logarithms, but only to fifteen places, of the products of the numbers already

found, were inscribed in their proper places ; the first and second differences of
these were taken wherever they happened to be sufficiently grouped together,
and the gaps were then filled up by means of second differences.

These interpolations were easily accomplished, because, since the third
differences are less than units in the fifteenth place, the progress of the second
differences could be estimated. It was enough, then, to make trials with the
last three figures of each order of differences ; and after a little practice these
trials were quickly made. The final figures of the second differences having
thus been found, the others were obtained, and the first differences with the

logarithms themselves were found by subtraction and addition.

In this way a table of fifteen-place logarithms for all numbers from 100 000

~ to 150 000 was formed.

To all who are familiar with extensive tabular work, it must be apparent
that these results, though trustworthy on the whole, could not be depended
upon as accurate in each item. A wrong figure may have been written, and
yet, on account of the consecutiveness of the differences, may not have been
detected by the subtraction. It became necessary, therefore, to revise the
whole in such a way as to preclude this source of error.

For this purpose a new set of ruled pages were consecutively numbered, and
VOL. XXVI. PART. III. 6 x

526 MR SANG’S NEW TABLE OF LOGARITHMS TO 200 000.

the last two figures only of the second differences were copied into their places ;
the first line of the first page, that is, the logarithm of 100 000, with the first
and second difference complete, was also filled up by copying. The whole table
was then re computed by continued summation, the results bemg compared, at
each fifth step, with the previous work; but as this comparison was not made
until the result was actually written down, the possibility of the transference of
an error was avoided. In this way the new computation may be held as, in all
probability, quite free from error, except, indeed, the minute errors inseparable
from interpolation, and not exceeding one or two units in the fifteenth place.

This new computation was to serve at the compositor’s desk ; wherefore, for
the purpose of keeping it clean and free from injury, the pages were transferred
by the copying press, and the copy was made to serve both for the type-setting
and for farther calculation. The original sheets were bound up for preservation,
in volumes containing each 10 000 numbers.

The logarithms for the latter half of the new table were obtained from those
of the first half by adding to each alternate logarithm the logarithm of 15. In
this way each third logarithm was found ; the intermediate ones being obtained
by interpolation. To have constructed the table directly in this way would
have left us liable to unchecked individual errors. In order to avoid these, or
rather to convert them into running errors, which cannot fail to be detected,
the following plan was followed :—

The last two figures only of the alternate logarithms were considered, and
the last two figures of the second differences for interpolation were adjusted by
trial on the slate, and, after being tested, were written in their places on the
prepared paper. These trials are easily made, because, at this part of the table
the third difference only amounts to ‘25 of the fifteenth place, and, towards the
end of the work, comes down to ‘10 of the same. From these terminal figures
the table was constructed by continuous summation as before, and each third
result was checked by addition after having been written. This check also
afforded a test of the accuracy of the preceding part, and, in point of fact, one
solitary error was detected by it.

Thus, the whole manuscript table of fifteen-place logarithms, with their first
and second differences, was constructed by continuous summation from 100 000
to 200 000, and may almost be held as free from any but last-place errors. It
is contained in ten quarto volumes, which form the first ninth part of the manu-
script table of all numbers up to one million.

The importance of having the printed table absolutely free from error,
naturally brought up the question of the use of calculating machinery; and
that question had to be very seriously considered. All the calculating machines
hitherto contrived are capable merely of addition or subtraction. These opera-

MR SANG’S NEW TABLE OF LOGARITHMS TO 200 000. 527

tions are sufficient for our purpose. The only intrinsic difficulty is this, that
the final differences change irregularly in the last place; wherefore, in using
any machine for computing logarithms, the operator must set the final difference
by hand preparatory to each step. The machine thus cannot properly be called
self-acting ; it is liable to errors caused by mistakes of the operator, who is

_ under the necessity of examining each result. He dare not venture to overpass

several steps, because one error may have balanced another in the intermediate
work. If the instrument go only to the extent of the printed work the last
digit would thus be insecure.

Tf, on the other hand, the instrumental work be carried to more places, in
order that minute errors may not tell, the second differences would be brought
into account, the machine would become enormously complex, and the expense
of it would exceed many times what, in another way, may produce as good a
result. Besides, the amount of attention, that is, of mental fatigue, would be

~ much greater, and would be accompanied by a loss of time.

Again, the machine must not merely compute, it must record the results in

some solid form capable of transferring impressions to paper. Only two ways

have been proposed for this. One to cause the figures to be punched in a plate
of soft metal from which electrotype casts may be taken; the other, to arrange
moveable types by the machine. In the first way it is difficult to correct an
error otherwise than by going over the whole page. In the second plan a
wrong type may get into the group from which the figures are taken. In either
way the results have to be carefully watched. Thus, do what we will, the last
resort is to careful reading, and careful reading will accomplish the whole with-
out any machine. The determination, therefore, was to proceed by the ordinary
method of hand-setting.

_ The course actually followed did not differ essentially from the usual process.
The types were set up in pages, proofs were taken, corrections made, and the
pages were then subjected to the process of electrotyping. Jn two respects
only did the course actually followed differ from the usual one. After the

electrotype has been taken, the page is unlocked, and the types are distributed
into their proper boxes to be ready for future use. The usual way is to make
use of these types, which lie in disorder in the case. Instead of using the types

directly after distribution, these types were set up in regular packets, so as to
permit of the examination of their faces. In this way the errors of distribution
were completely obviated. Moreover, the types were now placed in the case
all ready arranged and in the position most convenient for the compositor, who
was no longer obliged to examine the manner in which each individual type
happened to be lying in its box, and who could now take up the types without
almost looking off his paper. By this arrangement the errors in composition

528 MR SANG’S NEW TABLE OF LOGARITHMS TO 200 000.

were themselves much lessened, while those of distribution, twenty or thirty —
times as numerous, were entirely removed.

The other change was in taking two electrotypes of each page, one set to be
used for printing, the other set to be reserved for the sole purpose of repro-
ducing working plates. In this way, as any errors which may be detected are ©
to be corrected in the reserved as well as in the working plates, the table will
eventually come to be entirely freed from faults.

It is now two centuries and a half since the logarithmic table was completed
to 100 000. The want of a greater extension has been felt by the few com-
puters engaged in researches requiring great precision; and in 1819 a proposal
was made in the House of Commons, that our Government should join with
that of France in the publication of new Logarithmic and Trigonometric tables.
No actual progress, however, was made.

It is then worth while to record the extension of the seven-place tables to —
twice the usual length; particularly when that is accompanied by the finished
calculations for one-ninth part of the million table. Which ninth part really
implies more than one half of the calculations needed to reach the next step in
the denary scale.

PHYSOSTIGMA,

# times the Minimum -lahal Dose
Fe - . .
ad . . - .
<
=
- 9) 2%. . . .
5
2
°
°
2s = . o
a
S
ta - . . .
o
w
”
o
o
The Miunum-lenal Dose
12 gr 0 Ercract of Fhywart 7
(? J pounds oF TabOI) 1 eiod
Bin = - z

Se tones the Mintriune-lethad Dose \—

DOSES

Ue» -» 5 5 5

peg or gan Netliad, Dose.
ot Sud ; 7 !
Hag AAAI POISE i

te - . .

DIAGRAM J. ILLUSTRATING THE 157 SERIES OF EXPERIMENTS, IN WHICH ATROPIA WAS ADMINISTERED EIVE MINUTES BEFORE PHYSOSTIGMA.

anu

tore

2igrs.

Sgr

yr.

gre,

S5YTF.

PHYSOSTIGMA

oF
te

DOSES

buts

DOSES OF

SULPHATE

ATROPIA

(PER “THREE POUNDS

WEIGHT

ANIMAL.) _

DIAGRAM 8. ILLUSTRATING THE 28? SERTES OF EXPERIMENTS, IN WHICH ATROPIA WAS ADMINISTERED FIVE MINU1 AFTER PHYSOSTIGMA.
14 a | i—} I
|
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DOSES OF SULPHATE OF ATROPIA (PER “THREE POUNDS WEIGHT OF ANIMAL) Wh Ad tdemes Leer

Uh»

The Minn -lechal Porte

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VT PL NATL

Trans Roy Soa dine Vo

DIAGRAM 2. ILLUSTRATING THE DF SERIES OF EXPERIMENTS.

Wie ve 1h 7 70 —b97,

‘f - TEL a
DOSES OF SULPHATE OF ATROPIA (PEM THREE POUNDS WEIGHT OF ANIMAL)

DIAGRAM 4 ILLUSTRATING TIE 28 SERIES OF EXPE RUMEN TS.

DOSES OF SULPHATE OF ATROPIA (PER THREE POUNDS WEIGHT OF ANIMAL)

a a ren

Trans Roy Soo Edin? Vol XNVE PLANNIN

DIAGRAM 5. ILLUSTRATING THE [St SERIES OF EXPERIMENTS. IN WHICH ATROPIA WAS ADMINISTERED FIVE MINUTES BEFORE PHYSOSTIGMA.
(THE ENTIRE REGION OF RECOVERY WITH NON-LETHAL AS WELL AS WITH LETHAL DOSES OF PHYSOSTIGMA IS REPRES iD.)

aa ! Jue 1 Hee 10 ul tT | iB Li ft HH +}
i! A i 7 N il tet 1 || ++ {it a ciciaistaislalalats tt

The Poems Lechol Bere
(ae Zao of Phyzostigmas

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COPE Tyre, Dips, Dore Tages) 1908 T5ghe eyrs Tare Dsgrs, Tagrs ers.

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—
i

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Pea Minna ark De

DOSES OF SULPHATE OF ATROPIA (PER THREE POUNDS WEIGHT OF ANIMAL.)

MINUTES

REPRESENTED IN

TIME

INTERVALS

SIMULTANEOUS
ADMINISTRATIO

“-

ATROPIA ADMINISTERED
TER PHYSOSTIGMA

PHYSOSTIGMA

ADMINISTERED BEFORE

ATROPIA

Trans Rev: Soc Ridin: Vol XXVT. PUXRV.
DIAGRAM 6. ILLUSTRATING THE 3° SERIES OF EXPERIMENTS, IN WHICH THE DOSE OF PHYSOSTIGMA WAS CONSTANT
(ONE AND A HALF TIMES THE MINIMUM-LETHAL DOSE), WHILE THE DOSE OFATROPIA AND THE INTERVAL OF TIME
BETWEEN THE ADMINISTRATION OF THR TWO SUBSTANCES VARIED. :

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Sara (Saat SL ee nr tee { ~ BOF A iM, j

XXI.—An Experimental Research on the Antagonism between the Actions of
Physostigma and Atropia. By Tuomas R. Fraser, M.D., Lecturer on
Materia Medica and Therapeutics at Surgeon’s Hall, Edinburgh. (Plates
POX to X XV.)

(Read 29th May 1871.)

INTRODUCTION.

It is natural to suppose that soon after it became known that injurious
effects follow the introduction of certain substances into the system, attempts
were made to remedy these effects, and also to discover counteragents, or
antidotes, to the hurtful substances. The success attending these attempts must,
of necessity, have been closely related to the existing state of knowledge regard-
ing the actions of active substances. When the effects of poisons were referred
to supernatural manifestations, it was chiefly charms and superstitious rites that
were trusted to as protectives and remedies. At a somewhat more advanced
period in the progress of human knowledge, vague notions of physiological laws

and processes supplied the indications of curative treatment. Alexipharmics,

_ Mithridates, and theriacee were compounded of substances possessing elimina-

tive and so-called “general stimulant” properties, and bezoars of such as
enjoyed a reputation as specifics against poisonous influences ; and these were
employed, almost indiscriminately, as universal antidotes. Still later, chemistry
suggested that, as the physical properties of poisons may be modified by various
_re-agents, so may their effects be prevented by the administration of suitable

- substances.

The recommendations derived from chemistry were at first only of the crudest

_ description ; but as the science advanced, many valuable hints were obtained,

and now the class of the chemical antidotes probably includes the largest
number of efficient counteragents to poisons. Alkalies and acids are employed
to neutralise each other, tannin to render insoluble tartar emetic and many yege-
table alkaloids, hydrated sesquioxide of iron to precipitate arsenious acid, and
soluble and inert sulphates to decompose lead salts, and render them unabsorb-
able. In these examples, as well as in the many others belonging to this class,
the operation of the antidote is limited to the chemical change it produces on
the poison while it remains in the alimentary canal. As soon as the poison
becomes absorbed into the blood, it appears to pass beyond the antidotal
influence of the chemical counterpoison, for no example exists of a chemical
antidote neutralismg a poison after absorption. Thus it is that the value of
such antidotes is considerably restricted.
VOL. XXVI. PART III. 6 Y

530 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

PHYSIOLOGICAL ANTAGONISM.

Localised Antagonism.—In order perfectly to neutralise the effects that
‘follow the introduction of a poison into the living economy, it would appear to
be necessary that the physiological functions of the affected organism should be
modified. The early, though, undoubtedly, crude notions that originated the
employment of alexipharmics, Mithridates, and theriace, to a certain extent
recognised this principle. The more perfect knowledge acquired within recent
times regarding the functions of structures and organs, has led to the discovery
that various substances are able to modify them in a definite and constant
manner, and that the modifications produced by certain substances are of a
nature contrary or opposite to that of those produced by others. By such obser-
vations, the existence of physiological antagonism between certain of the effects
of different active substances has been demonstrated. Several apparently well-
authenticated examples have been made known: among which may be instanced
the antagonism between the actions on the iris and on the minute blood-vessels,
of opium or morphia on the one hand, and belladonna, hyoscyamus, and
stramonium on the other; between the actions on the capillary circulation of
morphia and quinia; between the actions on the vagi nerves of physostigma
and atropia, hydrocyanic acid and atropia, and muscaria and atropia; and
between the actions on the iris and on visual accommodation of physostigma
and atropia.

General and Lethal Antagonism.—In some instances, the existence of such
limited counteractions has led to the supposition that the general, or, at least, —
the primary lethal action of one of the substances concerned is capable of —
being antagonised by the physiological action of the other. A notable instance
of this is to be found in the revival, by the late Dr THomas ANDERSON, in 1854,
of the old, but, at that time, almost forgotten doctrine, that belladonna is a
physiological antidote to the poisonous action of opium.* ANDERSON was led to
this idea from the fact that these two substances produced contrary effects on
the iris. The occurrence of an antagonism limited to a smgle organ in no
important degree related to the continuance of life is, however, an insufficient
reason for supposing that the general actions of any two substances are of an
antagonistic nature. In order legitimately to infer whether one substance is
capable of acting as a physiological antidote to another, it is necessary to acquire —
a definite knowledge of the exact nature of the general physiological action
exerted by each of them. As yet the action of only a few substances has been
ascertained with the completeness that is required ; and hence it is that the —
examples that have been advanced of general antagonism between the actions

* Edinburgh Medical and Surgical Journal, vol. xviii, 1854, p. 377.

as
a

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 531

of active substances are but few in number, while the evidence on which these
examples have been founded is generally imperfect.

Between Opium and Belladonna, Hyoscyamus or Stramonium.—Among the
_ various instances in which a general antagonism has been stated to exist between
the actions of active substances, in the sense that the lethal effect of the one
substance is capable of being prevented by the physiological action of the other,
the most familiarly known is that where the substances are, on the one hand,
opium, and, on the other, belladonna, hyoscyamus, or stramonium. The existence
of a belief in the power of belladonna to counteract the general physiological
action of opium; may be referred to so early a date as the year 1570, when it
was recorded by Perro Pena and Marui DE Lopet that certain Italian peddlers
gained much notoriety by employing the root of the belladonna plant to quench
thirst, and by administering opiates to remedy the evil effects that were occa-
sionally produced thereby.* In 1661, Horsrius reported a case in which the
injurious effects of a large dose of the inspissated juice of belladonna were
apparently removed by the use of opium.t Soon afterwards, FaBer related a
somewhat similar experience ;{ and, in 1766, Boucuesr, of Lille, published five
cases of poisoning by belladonna berries, in two of which opium was administered
as an antidote.§ At the commencement of the present century, JosEPH LIpPI
wrote an inaugural dissertation, “ De veneficio baccis belladonne producto atque
Opii in eo usu,” in which were recorded, according to GracominI, “ pleusieurs
guérisons 4 laide de laudanum de SYDENHAM.” || GIACoMINI himself expresses a
favourable opinion regarding the beneficial effects of opium in poisoning by bella-
donna; and mentions, further, that the Italians were accustomed to administer
opium to remove the stupor and convulsions that follow excessive doses of
hyoscyamus and stramonium. Within more recent times, many modern authors,
as ANGELO Poma,1 ANDERSON,** Cazin,tt Benzamin BELL, {{ Béuter,§§ LEE, |\||
Norris,11 and Constantin Pavt,*** have published evidence that appears to
favour a belief in the existence of this antagonism. This evidence has been
derived from cases of poisoning in man by opium, in which belladonna, hyoscyamus,

* Stirpium Adversaria Nova, authoribus Perro Pena et Maruta pe Losen, Medicis. Londini,
1570, p. 103. (Quoted by Dr Norris, The American Journal of the Medical Sciences, vol. xliv.
1862, p. 399.)

t+ Opera Medica. { Strychnomania, 1677.

§ Journal de Médecine, Chirurgie et Pharmacie, etc., tome xxiv. 1776, pp. 310-332.

|| Traité philosophique et expérimental de Matiere Médicale et Thérapeutique, traduit par Mason
et Roenerta, 1839, p. 537.

@ Gazette Hebdomadaire, 10 Avril 1863. 4 Loe, ett.

t+ Traité des Plantes Médicinales Indigenes, 1855.

{+ The Edinburgh Medical Journal, vol. iv. 1859, pp. 1-7.

$$ L’Union Médicale, Juillet 1859.

|||| The American Journal of the Medical Sciences, vol. xlii. January 1862, p. 54.

G4 Ibid., vol. xliv. October 1862, p. 395.

*** De L’Antagonisme en Pathologie et en Thérapeutique, 1866, pp. 92-115.

532 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

or stramonium was used as a physiological antidote; and, conversely, of poisoning
with one or other of the latter substances, in which opium was used as an antidote.
In presence of the numerous important fallacies that are inseparably connected
with such evidence, it would be vain to expect that from it alone an absolute
demonstration could be obtained of the existence of a general antagonism so
perfect as to constitute any one active substance the physiological antidote of —
another. This evidence must, therefore, be regarded as unsatisfactory, more
especially as several observers of recognised ability, as Dr Joun Hariey* and ~
L. OrrFita,t have pronounced it insufficient, after a careful examination of the
record of each case.

The general result of the investigations that have been made to decide this
question by experiments on the lower animals, is also of an inconclusive
character. No doubt, the experiments of Bois,{ Camus,§ Onsvum, || and
Brown-SEQUARD 1 appear to show that the lethal action of opium cannot be pre-_
vented by the physiological influence of belladonna, hyoscyamus, or stramonium,
nor that of the latter substances by opium ; but these expériments are open to
the objection, that the doses of the substances used as antidotes do not seem
to have been sufficiently varied.

At the same time, there can be little doubt that the evidence derived from
both clinical observation and experimental research is sufficient to show that
several of the actions of opium are of a contrary nature to those of belladonna,
hyoscyamus, and stramonium.** It is, however, equally undoubted that, in the
meantime, this evidence is insufficient to prove the existence of a general anta-
gonism; or of one between actions of sufficient importance to constitute opium
a physiological antidote to belladonna, hyoscyamus, or stramonium, or these
latter substances physiological antidotes to opium. The question still remains
an open one; but such knowledge as is already possessed renders it pro-
bable that a general antagonism does really exist, to the extent, at any rate,
of the primary lethal action of opium or morphia being preventable by the
physiological action of belladonna, hyoscyamus, or stramonium. A properly
devised series of experiments would in all likelihood justify the opinion of
those who, with no little courage, have practically affirmed their belief in the
existence of this antagonism.

* The Old Vegetable Neurotics, 1869.

+ Dictionnaire Encyclopédique des Sciences Médicales (Atropine), tome vii. 1867, p. 215.

{ Gazette des Hopitaux, 1864.

§ Etude sur l’antagonisme de lopium et de la belladonne. Thése de Paris, 1865.

|| Schmidt’s Jahrbucher, 1865, Bd. 128, p. 288.

{ Journal de la Physiologie de Vhomme et des animaux, tome 3™°, 1860, p. 726.

** Tnteresting accounts of several of these contrary actions, founded on careful clinical observa-
tion, have been published by Drs Mircnett, Kren, and Morenovuss (see their paper “ On the Antagonism
of Atropia and Morphia,” in the American Jour. of the Med. Sciences, Vol. L. 1865, p. 67; and also by
Dr Ertenmeyer (for an abstract of whose paper, see “ L’antagonisme de opium et de la belladonne,”
by Dr Raynavp, Paris, 1866, p. 40).

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 533

The recent development of the study of Pharmacology has led not only to
the acquisition of knowledge regarding the exact manner in which many active
substances influence the physiological conditions of vital structure, but also to
the differentiation of the special structures, by the modification of whose physio-
logical conditions the lethal action of these substances is produced. In a few
instances it has been shown that the nature of the modification produced in the
physiological condition of the structure or structures involved in the lethal action
of the substance is apparently contrary to that produced on the same structure
or structures by the physiological action of another substance. The estab-
lishment of such facts has led, within the last few years, to the suggestion of
two instances of antagonism,—the first being between the lethal action of prussic
acid and the physiological action of atropia, and the second between the lethal
action of muscaria and the physiological action of atropia.

Between Atropia and Prussic Acid—For the first of these instances Phar-
macological science is indebted to Professor Preyer of Jena. In the course of
an elaborate research* on the action of prussic acid,—a research that may
fairly be characterised as the most important that has yet been made on the
action of this substance,—PRreEYER established that the primary lethal action is
due to embarrassment of the respiratory and cardiac functions. He further
showed that the embarrassment of the former function is caused by stimulation
of the terminations of the vagi nerves in the lungs, and by impairment of the
activity of the respiratory nerve-centre, while the embarrassment of the latter

function is caused by excessive stimulation of the inhibitory cardiac fibres of

the vagi nerves. Previous investigators—more especially Von Brzotp and
Biespaumt—had already shown that atropia produces effects that are in a
remarkable manner contrary to these ; for, in certain doses, it accelerates both
the respiratory and the cardiac movements,—the former, by paralysing the
terminations of the vagi nerves in the lungs, and by stimulating the respiratory
nerve-centre, and the latter, by paralysing the inhibitory cardiac fibres of the

_ Yagi nerves. Guided by these facts, PREYER made a few experiments which
strongly support the opinion he has arrived at, that atropia is a physiological

antagonist to prussic acid, even to the extent of being able to prevent the primary
lethal action of that poison. It is, however, to be regretted that no attempt
was made absolutely to demonstrate that the dose of prussic acid used in each
experiment was a lethal one, more especially as the subsequently performed
experiments of Professor BarrHoLtow—limited, no doubt, in their scope—do
not seem to confirm PREYER’s opinion.t

* Die Blausiure. Physiologisch Untersucht. Von W. Pruyer, Dr. Med. et Phil. Bonn, 1870.
T Ueber die physiologischen Wirkungen des schwefelsauren Atropins. Von A. vy. Brzonp und Dr
Frrepr. Buasaum. (Untersuchungen aus dem physiologischen Laboratorium in Wiirzburg. 1867.)
t The Physiological Effects and Therapeutical Uses of Atropia and its Salts, 1869, p. 25.

MOL, XXVI. PART III. 6 Z

534 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Between Atropia and Muscaria.—The second of the instances mentioned
was first made known by SCHMIEDEBERG and Kopps, ina very interesting memoir
on Muscaria, published in 1869.* This active principle was separated by them
from Agaricus muscarius, L., and found to possess an action in many respects
contrary to that of atropia. The general nature of its lethal action was observed
to be similar to that of prussic acid; and, accordingly, the reasons which
induced SCHMIEDEBERG and Koprer to examine as to an antagonism between it
and atropia were analogous to those by which PREYER was led to investigate the
influence of atropia in counteracting the primary lethal action of prussic acid.
In this instance, likewise, only a very few experiments were made. Their
results, however, are strongly in support of the existence of a more or less
general physiological antagonism between the two substances.

Various other instances of General and Lethal Antagonism.—tIn addition to
these, many other examples of general or of lethal antagonism have been ad-
vanced. Their existence, however, has rarely been inferred from a knowledge
that the substances concerned influence the same structures in contrary modes,
but has been conjectured from a knowledge merely of the general pheno-
mena which are produced by these substances. The conspicuous spasmodic
effects by which the action of strychnia is characterised, appear to have sug-
gested the employment, as physiological counteragents, of various substances
whose general action includes the production of paralysis; and, accordingly, the
list of proposed antagonists to this alkaloid includes opium,t curara,} aconite,§
nicotia,|| bromide of potassium,‘| chloroform,** chloral,tt and nitrite of amyl.{{
Opium and quinia have been proposed as antidotes to each other, because the
former exalts several of the organic functions, whilst the latter depresses them.{§
General antagonism has been inferred between chloroform and sulphuric ether,
solely on the ground that the anzsthetic action of the former is supposed to be
accompanied with depression, and that of the latter with excitement ;|||| and the

* Das Muscarin. Das Giftige alkaloid des Flegenpilzes. Von Dr Oswatp ScumiepreBere und Dr ~

Ricuarp Koprr. Leipzig, 1869.

+ Pexretier et Caventou. See Dictionnaire Encyclopédique des Sciences Médicales (Antidote),
tome 5™°, 1866, p. 322.

+ L. Venta. Comptes Rendus des Séances de | Académie des Sciences, xlix. 1859, p. 330, and
li. 1860, p. 353.

§ E. Woaxes. The British Medical Journal, October 26, 1861, p. 440.

|| S. Haueutoy. Dublin Quarterly Journal of Medical Science, August 1862.

q F. A. Saison. Du Bromure de Potassium et de son Antagonism avec la Strychnine. Paris, 1868.

** T. Gatnarp. Annales d’Hygitne publique et de Médicine Légale, t. xxiv. 1865, pp. 182-184.

tt Oscar LiepreicH, Comptes Rendus des Séances de ]’Académie des Sciences, Ixx. 1870, p. 403;

Bennett, Edinburgh Medical Journal, 1870, v. 16, part 1, p. 262; Groves, Medical Press and

Circular, 1870, p. 398.
tt J. Sr Crain Gray. Glasgow Medical Journal, February 1871, p. 188.
§$ Guster. Société Médicale des hopitanx, 10 Février, 1858; and Commentaires Thérapeu-
tiques du Codex Medicamentarius, 1868, p. 591.
|||| Fann. Thése, 1860. (Quoted by Camus, op. cit. p. 122.)

_—

—_

—~

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. D900

physiological actions of iodine and bromine are said to neutralise each other
because the former produces sedation, and the latter excitation of certain
general functions.*

Among these examples there are several worthy of further examination,
and it is not impossible that their existence may thereby be established. Mean-
while, the criticism of the Professor of Therapeutics at Paris, in reference to the
majority of recorded examples of antagonism, appears to be a just one, that “la
précision fait souvent défaut dans lanalyse des faits, les inductions manquent
de rigueur, et la pratique attend de nouvelles lumieres de la part de la physio-
logie expérimentale et de la thérapeutique rationnelle.”t

Between Physostigma and Strychnia.—This criticism is also applicable to

much that has been advanced regarding antagonism between physostigma:and

certain other substances. The first instance that has been suggested of an
antagonism in which physostigma is concerned, is that between it and strychnia.
The spinal excitant action of the latter substance was naturally looked upon as
more or less contrary to the paralysing influence exerted by physostigma on the
spinal cord. Ina paper published by me in 1862,{ an experiment is described
which lent some countenance to this surmise. Since that time experiments
have been made by NunneLey,§ V&z,|| and Espen Wartson,1 which, on the
whole, support the opinion that the spasmodic effects of strychnia may be
diminished by the paralysing action of physostigma. They are, however, insuffi-
cient to decide whether the lethal action of the one substance can be prevented

by the physiological action of the other.

Between Physostigma and Chloral.—In the remaining instance, the power of
chloral to counteract the lethal action of physostigma has been experimentally

tested by Professor Benner. It is, however, impossible to decide how far the

Opinion expressed by this observer, that chloral has a most marked influence in

counteracting the lethal action of physostigma, is justified by the results of his
experiments, as only a very brief account of them has as yet been published.**

* Guster. Bulletin Général de Thérapeutique, tome Ixvii. 1864, p. 9.

t Guster. Dictionnaire Encyclopédique des Sciences Médicales (Antidote), tome 5™°, 1866,
p. 322.

¢ “On the Characters, Actions, and Therapeutic Uses of the Ordeal Bean of Old Calabar.”
Edinburgh Medical Journal, vol. ix. 1863, p. 245; and reprint, p. 19. See also, “On the Physio-
logical Action of the Calabar Bean.” ‘Transactions of the Royal Society of Edinburgh, vol. xxiv.
‘part ii. 1866-7, p. 740.

: § “On the Calabar Bean : its Action, Preparations, and Use.” Lancet, 1863; and pamphlet, pp.
2-15.

|| Recherches Chimiques et Physiologiques sur la Féve du Calabar (These), Parle Dr Am#pie
Vis. Paris, 1865, pp. 28-30.

{| “On the Physiological Actions of the Ordeal Bean of Calabar, and on its Antagonism to
Tetanus and Strychnia Poisoning.” Edinburgh Medical Journal, vol. xii. May 1867, p. 999; and
Teprint, pp. 17-25.

** “ On Chloral in Phthisis, and its Antagonism to the poisonous effects of Calabar Bean.” The
Practitioner, vol. iv. 1870, p. 262.

536 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

This account, however, does not very obviously support Professor BENNETT’s
opinion ; for, of the eight experiments mentioned, in which rabbits were sub-
jected to the influence of the two substances, seven terminated in death, and
only one in recovery. Further, there is no evidence to show conclusively _
that the rabbit that formed the subject of the apparently favourable experi-
ment had received a dose of physostigma sufficient to have caused its death
had no chloral been administered.

In the preceding historical sketch every important alleged example of anta-
gonism has been referred to. It has been shown that although in many cases
the @ priori reasons in favour of the existence of a lethal or of a more or less
general antagonism are extremely plausible, the experimental data by means of
which it has been attempted to establish the reality of the antagonism are
probably, without exception, imperfect, and therefore insufficient to do so, I
trust, however, that the description of the research forming the subject of the
present communication will render it obvious that the reality of a lethal anta-
gonism may be readily and certainly established by experiment.

ANTAGONISM BETWEEN THE ACTIONS OF PHYSOSTIGMA AND ATROPIA.

This research on the antagonism between the actions of physostigma and
atropia was commenced in April 1868, and the results of some of the earlier
experiments were published in a preliminary note read before this Society, on
the 30th of May 1869,* In this note were described a number of experiments,
which prove that the lethal action of physostigma may be prevented by the
physiological action of atropia.

Previous to this time, however, the attention of more than one observer had
been attracted to the subject. In 1864, KiernwAcuTeEr narrated an interesting
case of poisoning with an unknown quantity of atropia, in which the internal
administration of physostigma produced a marked amelioration of the symp- —
toms.t Three years subsequently, BouRNEVILLE, of Paris, in a paper on the
treatment of tetanus by physostigma,{ described an experiment in which he, in
the first place, introduced into the stomach of a cabiai a quantity of powdered
kernel of physostigma, sufficient, in his opinion, to cause death, and then, while
severe symptoms were present, injected subcutaneously a small quantity of
atropia, with the result that the symptoms quickly diminished in severity, and
the cabiai ultimately reassumed a normal condition. At the time when my
preliminary note was published, BouRNEVILLE’s experiment was quite unknown
to me, and it is with much satisfaction that I now draw attention to it as an
independent observation harmonising with the results I had obtained when my

* Proceedings of the Royal Society of Edinburgh, 1868-69, pp. 587-590.
t Berliner Klinische Wochenschrift, No. 38, 1864, p. 369.
$~ De ’Emploi de la Féve de Calabar dans le Traitment du Tetanus Paris, 1867.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 537

preliminary note was published, and have since greatly extended.* The obser-
vations of another experimenter, Professor Bartuotow, of Cincinnati, have
likewise only recently come to my knowledge. The publication, in the “ Prac-
titioner” of February 1870, of a paper by me on “Atropia as a Physiological Anti-
dote to the Poisonous Action of Physostigma,” directed Dr BARTHotow’s atten-
tion to my researches, and by his courtesy and kindness I have been favoured
with a copy of an essay on “ Atropia,” which he had published in 1869. I am
thereby enabled to supply an omission that would otherwise have occurred in
this account of the literature of the subject, for the essay contains not only an
interesting theoretical discussion on the antagonism between atropia and
physostigma, but also several experiments bearing on its existence. The
experiments were performed on frogs and cats, and a description is given of
two experiments on each of these species of animal. One experiment on a frog
and one on a cat terminated in recovery, while the two others terminated in
death. From these experiments Dr BartHotow deduces a number of general
conclusions regarding the mutual counteraction of the two substances on several
of the structures and functions modified by them. The following quotation
‘contains an epitome of his views :—“ Atropia is not a physiological antagonist to
physostigma, except in regard to their action on the organic nervous system. It
would be improper, then, to use atropia against poisoning by Calabar bean. . .” +
The second of these propositions seems to imply that the existence of a lethal
antagonism was not favoured by the results of the experiments. The account
given of the experiments, however, does not justify any opinion as to how far the
non-existence of a lethal antagonism is supported by them, for, unfortunately, the
obviously necessary information is omitted by which to judge if a lethal dose of one
or other substance had been administered to either of the animals that recovered.

Preparations used in the Research.—In this research physostigma was admin-
istered in the form either of an alcoholic extract, or of the sulphate of the
active principle.

The alcoholic extract was prepared by placing a moderately fine powder of
the kernel in a percolator, acting upon it with alcohol (84 per cent.) until the
powder was exhausted, and then concentrating the tincture by distillation and
by evaporation on a water bath, until an extract of ordinary consistence was

* Since this sentence was written, I have received a more recent paper by M. Bournevitye, which
contains evidence of an absolutely satisfactory nature regarding the power of atropia to counteract the
lethal action of physostigma. It is entitled, ‘‘ De l Antagonisme de la Féve de Calabar et de l Atropine,”
and appears to be a reprint from the “ Revue Photographique des Hépitaux,” of June 1870. <A de-
seription is given of five experiments on guinea pigs, in which non-lethal doses of atropia were adminis-
tered a few minutes after lethal doses of extract of physostigma, with the result that recovery took
place in all of the experiments. The great value of the evidence contained in this paper depends on
the fact that the doses of physostigma given were proved to be at least equal to the minimum lethal.
This was accomplished in much the same way as has been described in my preliminary note in the
Proceedings of this Society, and in my communication to the “ Practitioner” of February 1870.

+ Loc. cit. p. 46.

VOL. XXVI. PART III. 7A

998 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

obtained. This preparation contains a considerable quantity of fatty matter,
which prevents its complete solution in water ; and as the division into separate
doses of a mere watery suspension would lead to many inaccuracies, it was
found necessary to weigh the requisite quantity separately for each experiment.
It is also hygroscopic, which further required that it should be dried and kept
in an exsiccator, in order to ensure an unvarying preparation. Nearly all the
experiments in which an extract was used were made with one prepared in this
manner, and a sufficient quantity was obtained by one process to serve for the
entire research. A few experiments, however, were made with an extract for
which I am indebted to Dr Cook, of the firm of Messrs T. and H. Surru, of Edin-
burgh. It will be seen, from the description of these experiments, that Dr
Coox’s extract is more powerful than that prepared by myself, and this may be
accounted for by the fact that absolute alcohol was employed in its preparation.
The active principle, physostigmia,* whose sulphate was also used in this re-
search, was obtained by the following process. Alcoholic extract of physostigma
was mixed with distilled water, and the fatty matters were completely removed
by agitation with successive portions of sulphuric ether. An excess of bicar-
bonate of sodium was then added to the watery solution, and the resulting alka-
line liquor was shaken with successive portions of ether. The decanted etherial
solutions were washed with water, concentrated by distillation, and then evapo-
rated spontaneously, by which means a residue consisting of an impure physo-
stigmia was obtained. This was dried over sulphuric acid, and treated with
anhydrous ether, and on evaporating the etherial solution, a less impure physos-
tigmia was obtained in the form of a pale brown extract-like substance. From it
the sulphate was prepared by neutralising a solution in rectified spirit with very
dilute sulphuric acid, and evaporating at a low temperature. This sulphate is a
pale brown amorphous substance, readily soluble in distilled water. As watery
solutions of the vegetable alkaloids gradually undergo decomposition, it was
considered advisable to weigh separately the dose required for each experiment.
Physostigmia, in common with the extract, possesses the inconvenient property
of absorbing moisture from the atmosphere, and for this reason, the obviously
necessary precaution was adopted of keeping the sulphate in an exsiccator. }
The atropia was administered in the form of sulphate, which salt was pur-
chased from Messrs T. and H. Smirx of this city. The doses were generally

* This alkaloid was first separated by me in 1863 ; and in a paper published in 1864 (“On the
Moth of the Eseré, or Ordeal Bean of Old Calabar,’ Annals and Magazine of Natural History, May,
1864), I named it Hserinia, from Eseré, the usual name of the ordeal poison at Calabar. Since then
I have, in various publications, adhered to this name, and it has been almost invariably adopted by
French physiologists and chemists. The reasons in favour of designating an active principle, derived
from the vegetable kingdom, by a modification of the generic name of its botanical source are, however,
so numerous and weighty, that I have thought it right in the present communication to follow the
usual practice. This I have the more readily done, as the name physostigmia (or physostigmin) is now
commonly to be met with in the writings of German physiologists.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 539

weighed separately for each experiment, but in several instances it was found
necessary to subdivide a recently prepared solution, as such minute doses were
required that it would have been impossible to weigh them accurately.

Subjects of Experiment.—With a few exceptions, wherein dogs were used,
the experiments were performed on rabbits. The animals were invariably
in a state of perfect health, and in full digestion. The latter is a condition
of great importance, the plan of research requiring a strict attention to the
relation between the weight of the animal and the doses of the substances.
The amount of food contained in the stomach appreciably modifies the weight
both of dogs and rabbits, but it does so to a very marked extent in rabbits, for
on several occasions I have found that an increase of three or four ounces oc-
curred after food had been taken. As, generally, the rabbits employed were
about three pounds in weight, the difference represented by such an increase
is obviously of importance in estimating the doses of the substances.

Plan of Experiments.—The following plan was adopted for the experiments,
as it appeared to be the one by which the most conclusive results were to be
obtained :—In the first place, the minimum fatal dose for rabbits of the extract
of physostigma and of the sulphate of physostigmia employed was determined
by a number of preliminary experiments, so that, on the weight of the animal
being ascertained it was an easy matter to be certain of the dose that could kill
it. Then, in those experiments in which recovery followed the administration
of a dose of atropia given in combination with a dose of physostigma equal to
or in excess of the minimum fatal, the animal used was killed many days after-
wards, and when the effects of the two substances had completely disappeared, by
a dose of physostigma less than or only equal to that from which it had pre-
viously recovered. Therefore, when the administration of atropia prevented an
otherwise fatal dose of physostigma from causing death, a perfect demonstration
was obtained of the power of atropia to produce some physiological action or actions
that counteracted some otherwise lethal action or actions of physostigma.

The administration of the substances was effected by subcutaneous injection.
There is an abundance of evidence to show that, when exhibited by subcutaneous
injection, the activity of a substance, relatively to its dose, is considerably greater
than when it is exhibited by introduction into the stomach. By adopting this
method, therefore, the existence of a lethal antagonism was subjected to a more
severe test than if the substances had been introduced into the stomach ; for, not
only are the general physiological effects produced with greater rapidity and
certainty, but also the lethal action of a minimum fatal dose is induced in a

shorter time when the substances are injected under the skin than when they

are introduced into the stomach. This method of administration has, besides,
the great recommendation of being followed by results constant as to both
character and time of occurrence ; for not only is the total quantity, within cer-

540 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

tain limits, of the substance absorbed into the blood, but also the process of
absorption is commenced directly after the injection is effected. Further, it
has the great advantage of convenience, wherein it is greatly superior to the
method by introduction into the stomach.

Chief Objects of the Research.—As evidence was obtained at an early period
in this research of the existence of an antagonism between the general actions
of physostigma and atropia, a wide field for further investigation was thereby
opened up. In the experiments that will be described in the first portion of
this communication, I shall endeavour to show, as clearly as possible, that
atropia possesses, in a remarkable degree, the power of counteracting the lethal
action of physostigma. In the subsequent portion of the communication the
extent of this power will be examined and its limits defined.

Section AA—EXAMINATION OF THE INFLUENCE OF ATROPIA UPON THE
LETHAL ACTIVITY OF PHYSOSTIGMA.

DETERMINATION OF THE MINIMUM LETHAL DOSES OF THE PREPARATIONS.

In accordance with the plan that has been adopted for this research, several
preliminary experiments were made in order to determine the minimum lethal
dose for rabbits of each of the preparations employed. For the present pur-
pose it is sufficient to mention only the leading results of these experiments.

Minimum Lethal Dose of Sulphate of Atropia.—tn the following table a
summary is given of experiments undertaken to determine the minimum lethal
dose for rabbits of sulphate of atropia.

Number : Dose per
| of Experi- ae ‘| Actual Dose. | 3 Ibs. Weight Result. Notes.
ment. pe of Animal.

1. 4 Ibs. 13 oz. 4 ors. 2°49 ors, Recovery. The only effects were dila-
tation of the pupils, slight
restlessness, and accelera-
tion of the cardiac contrac-
tions and of the respira-
tions.
2: 4 Ibs. 10 oz. 5 grs. 3°24 grs. Recovery. Do.
3. 4 lbs. 8 oz. 9 gers. 6 grs. Recovery. Do. Also some obvious
symptoms of visual de-|
rangement.
4. 3 lbs. 6:5 prs, | 6°5 grs. Recovery. Distinct, though unim-
portant, paralytic symp-
toms were produced, the
respirations were reduced
in frequency, the cardiac
action was accelerated,
and the pupils were
dilated.
5. 3 Ibs. 5 oz. 79 grs. | 7°5 gts. Recovery. Do.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 54]

; Dose per
Weight of | Actual Dose. | 3 Ibs. Weight Result. Notes.
Rab tte of Animal. .

2 Ibs. 15 oz. 7°34 ers.| 7°5 grs. | Recovery. Do.; excepting reduction
in the frequency of the
respirations.
2 Ibs. 4 02. 6 grs, 8 grs. —_| Recovery. Do. do.; and production
of hypnosis.
3 lbs. 9 grs. 9 grs. Recovery. Dilatation of the pupils,
increase in the frequency
of the cardiac and respira-
tory movements, and
slight paralysis were pro-
duced.
3 lbs. 2 oz. 15°6 grs. 15 grs. Recovery. Dilatation of the pupils,
acceleration of the heart’s
action, increase followed
by reduction in the rate
of the respirations, dis-
tinct paralysis, and tre-
mors and starts, were
produced.
2 Ibs. 12 oz. | 16°5 grs. 18 grs. ~ | Recovery. Do.
3 lbs. 19°5 ers. 19°5 grs. | Recovery. . The chief effects were
dilatation (not extreme)
of the pupils, acceleration
followed by slowing and
weakening of the heart’s
action ; reduction in the
rate of the respirations ;
hypnosis; and well-
marked paralysis.
2 lbs. 13: 0z. | 19:9 srs. 21 ers. Death, in more | Do.
than 1 hour, and
less than 5 hours

30 minutes.

2 Ibs. 134 oz.| 19°9 grs. | 21 grs. Recovery. Do.

3 Ibs. 2 oz. | 23°43 grs.| 22°5 grs. | Recovery. Do.

3 Ibs. 14 oz. | 22°9 grs. | 22°5 grs. | Recovery. Do.

albs., 6 0z. | 27 gers. 24 ers. Death in 35|The chief effects were
minutes. dilatation (not extreme)

of the pupils; accelera-
tion soon followed by
slowing and great weak-
ening of the heart's
action ; reduction in the
rate and strength of the
respirations ; paralysis ;
and feeble tremors and
spasms.

In each of these experiments every precaution was taken to prevent fallacy
It will, however, be observed, that while, in one case, a dose of 21 grains, and
n another one of 24 grains per three pounds weight, was found sufficient to cause
VOL. XXVI. PART III. 7B

542 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

death, in two other cases recovery followed the administration of 22°5 and of
21 grains respectively. It must, therefore, be allowed that the minimum fatal —
dose has only approximatively been determined. A more accurate determina-
tion could not be effected without greatly increasing the number of experiments.
Fortunately, however, it was unnecessary to incur the trouble and expense*
that would thereby have been entailed, as an approximative determination of
the minimum fatal dose of atropia was all that was needed for the purpose of
this research.t

Minimum Lethal Dose of Extract of Physostigma.—The experiments which
are mentioned in the next table were undertaken to determine the minimum
lethal dose for rabbits of extract of physostigma.

dixperiment, | Raghit:, | Aetual Dose. hyesene of animal Result.
5 be pe 3 Ibs. 3 oz. 0°8 er. 0°7 gr. Recovery.
18. 3 lbs. 0°9 gr. 0°9 er. Recovery.
19. 3 lbs. 2 02. 0°93 gr. 0°9 gr. Recovery.
20. 3 lbs. 8 oz. 1°2 gr. 1°02 er. Recovery.
21, 3 lbs. 6 oz. TS pr. 1:05 gr. Recovery.
22: 2 Ibs. 14 oz. 1 or. 1:05 gr. Recovery.
93. 3 lbs. 1 oz. 1-2 or, 1:2 gr. Death, in about 27 minutes.
94, 3 Ibs. 6 oz. 1°35 gr. 1:2 gr. Death, in about 23 minutes.
25. 3 Ibs. 5 oz. 1°32 gr. 1-2 gr. Death, in about 33 minutes.
26. 3 Ibs. 2 oz. 1°87 gr. 1°8 gr. Death, in about 16 minutes.

The results of these experiments indicate that the minimum lethal dose for
rabbits of extract of physostigma is 1:2 grain for every three pounds weight of
animal, or 0°4 grain for every pound. :

Minimum Lethal Dose of Sulphate of Physostigmia.—The minimum lethal
dose for rabbits of sulphate of physostigmia was discovered by the experiments —
that are epitomised in the next table.

* The price of sulphate of atropia. being a little more than fifteen shillings for sixty grains, the ques-
tion of expense becomes worthy of consideration. )

+ The minimum lethal dose of sulphate of atropia, administered subcutaneously, appears to be
smaller for dogs than for rabbits. Among other experiments, I have performed the following :—A
dog, weighing seven pounds and fifteen ounces, received twenty grains, and recovery followed ; but
when a dose of twenty-five grains was given to the same dog, eight days subsequently, death occurred
in twenty-three minutes. Another dog, weighing sixteen pounds, which, seven days previously, had
recovered after the administration of ten grains, died on the fourth day after it had received fifteen
grains. -

i
i

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 545

Taporimest. | Rathi, | Actual Dose. |yratehe of Animal Result.

: 97. 3 lbs. 8 oz. 0:035 er. 0:03 or. Recovery.

98. 3 lbs. 1 oz. 0076 er. 0'075 gr. Recovery.
29. 3 Ibs. 11 oz. 0-1 gr. 0081 er. Death, in about 33 minutes.

30. 3 lbs. 2 oz. O'l er. 0096 er. Recovery.
31. 3 Ibs. 1 oz. 0°12 er. 0-117 gr. Death, in about 37 minutes.
32. 3 lbs. 5 oz. 0°13 er, 0-117 er. Death, in about 44 minutes.
30. 3 lbs. 4 oz. 0°13 er. 0°12 er. Death, in about 34 minutes.
34. 2 Ibs. 14 oz. 0°13 gr. 0°13 er. Death, in about 22 minutes.
35. 3 lbs. 6 oz. 0°15 gr. 0:13 gr. Death, in about 16 minutes.
36. 3 Ibs. 4 02. 0°16 er. 0°147 gr. Death, in about 25 minutes.
of. 3 Ibs. 0°15 gr. 0°15 er. Death, in about 28 minutes.
38. 3 lbs. 3 oz. 0°16 gr. 0°15 er. Death, in about 21 minutes.
39. 3 lbs. 2 oz. 0:19. er. 0°18 gr. Death, in about 16 minutes.
40. 3 lbs. 1 02. 0°18 er. 0°18 gr. Death, in about 19 minutes.

From these experiments, it would appear that in rabbits the minimum lethal

ounds weight of animal, or 0°04 grain for every pound. The experiment in which
death occurred after the administration of 0-08 grain per three pounds weight
( ixpt. 29), must be regarded as an exceptional one, seeing that during it
th e rabbit was in a.violently excited state; and the constant energetic move-
m ents that were made placed the animal in an unfavourable condition to resist
the toxic influence of a poison that materially embarrasses both the cardiac and
ie respiratory functions. Still, even after excepting this experiment, the table

ows that 0°12 grain per three pounds is a dose rather in excess of the minimum

one to ten that is thereby obtained between corresponding lethal doses of
sulphate of physostigmia and extract of physostigma is a very convenient one,

It may not be altogether unnecessary to point out that the results of these
leterminations are applicable only to the special preparations with which they
have been obtained ; for the composition, and therefore the lethal activity, of
zach of them varies somewhat in accordance with the processes followed in its
manufacture.

INFLUENCE OF ATROPIA ON THE LETHAL ACTION OF PHYSOSTIGMA.

_ The minimum fatal dose for rabbits of the extract of physostigma and of
the sulphate of physostigmia having thus, with considerable accuracy, been

544 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Atropia administered before Physostigma.—The nature of this influence when
atropia is administered before physostigma is shown by the following experi-
ments :—

EXPERIMENT 41-a,—In a rabbit weighing two pounds and fifteen and a half
ounces, it was found that the number of the cardiac impulses was 40 in ten
seconds, and of the respirations 12 in ten seconds, and that the pupils measured
14ths x 28ths of an inch.

Three-tenths of a grain of sulphate of atropia, dissolved in 30 minims of dis-
tilled water, was injected under the skin of the left flank. Intwo minutes and
thirty seconds thereafter, the pupils measured 4$ths x 48ths of an inch ; and in
four minutes, the cardiac impulse occurred 54 times in ten seconds.

Five minutes after the injection of sulphate of atropia, one grain and a fifth
of extract of physostigma, suspended in 25 minims of distilled water, was injected
under the skin at the right flank ; and, immediately afterwards, the syringe was
washed out with 15 minims of distilled water, and this solution injected under
the skin at the right hip—the whole operation lasting thirty seconds.

In two minutes after i total dose of physostigma had been injected, the
pupils measured 2%ths x 4%ths of an inch, and infrequent fibrillary twitches were
occurring at the right flank and hip. In nine minutes, the rabbit became rest-
less, having been perfectly quiet until now, and the pupils measured }8ths by
t&ths of an inch. Soon afterwards, fibrillary twitches were occurring generally
over the surface of the rabbit, some unsteadiness was apparent in the move-
ments, and often slight tremblings took place, especially marked in the head. In
fifteen minutes, the fibrillary twitchings were more frequent and more strongly
marked, so that it was difficult to distinguish the cardiac impulse, but it ap-
peared to occur about 46 times in the ten seconds. In twenty-six minutes, the
general symptoms had become slightly aggravated, as a normal posture was
maintained only with difficulty; the arching of the back becoming gradually
less prominent, and the head drooping a little. At the same time the fibrillary
twitches had become more marked, so that the skin of the whole surface of
the animal was in constant movement, and occasionally a weak spasmodic
start occurred. In thirty-seven minutes, the head had so far subsided as to
permit the chin to rest on the floor, but this latter posture was maintained for
only a few minutes, and was succeeded by a more natural one in which the head
was raised. In fifty-seven minutes, the rabbit was in a normal sitting attitude,
and the chief symptom was well marked universal fibrillary twitching. The
pupils measured 2Sths x 48ths of an inch, the cardiac impulse was at the rate of
41 in the ten seconds, and the respirations 16 in the ten seconds. In one hour
some urine was voided, and three minutes afterwards a considerable quantity
of pultaceous and wet fseces was passed. In one hour and sixteen minutes,
slight mucous sounds, apparently originating in the larynx, were heard during”
the respirations, and fseces having the unnatural appearance above described

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 045

were again passed. In one hour and thirty minutes, the rabbit was still in the
normal sitting posture it had for some time assumed, and when it was obliged to
move about no obvious difficulty could be detected. The fibrillary twitches had
decreased considerably; the rate of the cardiac impulse was 40 in ten seconds ;
and the pupils measured 46ths x 4§ths of an inch.

On the following day, twenty-three hours after the commencement of the
experiment, the rabbit appeared to be perfectly well, for it went about actively
and fed well. The rate of the cardiac impulse was 52, and that of the respira-
tions 12 in ten seconds, and the pupils measured 2&ths x 48ths of an inch.
From this time, a gradual diminution went on in the rapidity of the heart’s action,
and in the size of the pupil; until, on the fifth day, the former had assumed
the normal rate of 41 in ten seconds, and the latter measured exactly the same
as before the experiment was commenced, namely, 44ths x 18ths of an inch.

On the eleventh day this rabbit was subjected to the influence of a
- minimum lethal dose of extract of physostigma, and the result is described in the
next experiment. It is of importance to note, that during all this time food
had been supplied to the rabbit ad libitum, as this is of importance in the
maintenance of a state of absolute health, and that the same was also done in
all the similar experiments of this research.

EXPERIMENT 41-b.—This rabbit now weighed three pounds, and it was ascer-
tained that in ten seconds the cardiac impulse occurred 41 times, and the respira-
tory movements 17 times, and that the pupils measured }4ths x 43ths of an inch.

One grain and a fifth of extract of physostigma, suspended in 25 minims of
distilled water, was injected under the skin at the right flank, and the syringe
washings under that at the right hip. The first effect observed was the occur-
rence of infrequent and slight twitchings of the panniculus carnosus muscle
in the neighbourhood of the regions where the injections had been made, and
this effect commenced in about one minute and thirty seconds after the first of
_ the two injections. Beyond this, there was no obvious symptom until six
_ minutes, when some slightly restless general movements were made, and at the

“same time movements of the mouth and lips occurred, as if an accumulation of
Saliva were being removed. Soon afterwards, there was evident difficulty in
going about ; gradually slight stiffness showed itself in the anterior, and then in

the posterior extremities, which by-and-by became extended, and thereafter the
| rabbit stumbled about, or stood shaking with the body elevated on the extended
limbs. In eight minutes, the above condition was present, and besides, the
fibrillary twitchings had become more general and frequent, and the pupils
slightly larger, having increased from 1éths x 1%ths to i3ths x Léths of an inch.
In ten minutes, the extended state of the limbs disappeared, and was succeeded
by partial paralysis, so that the rabbit now sank down on the abdomen and chest.
In thirteen minutes, great general weakness, accompanied with constant
VOL. XXV] PART III. ne

546 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

tremblings, was present, now and then somewhat severe tremors occurred,
fluid (salivary) was escaping from the mouth, soft and pultaceous feces, wet
on the surface, but preserving the pellet shape, were being passed, and the pupils
measured 43ths x 42ths of aninch. In fourteen minutes, the respiratory move-
ments were somewhat embarrassed, and accompanied with moist sounds, while
their frequency was diminished to about 10 in ten seconds. The head of the
animal was now lying on the table, the back was scarcely at all curved,
but the general tremors had almost disappeared, although the fibrillary
twitchings had rather imcreased in frequency. In seventeen minutes,
the rabbit fell over on the side. Only slight fibrillary twitchings were now
present ; the respirations were laboured, greatly impeded by mucus accumulated
in the mouth and larynx, and accompanied with struggling movements of the body
and limbs ; the pupils measured 443ths x 44ths of an inch; the cardiac impulse
was weak and infrequent ; frothy saliva was escaping from the mouth, and liquid
feeces were being passed at intervals. Very soon afterwards, the respiratory move-
ments became mere laboured gasps, the pupils still further diminished in size,
and general weak tremors succeeded each other. By-and-by it was a matter
of difficulty to distinguish any respiratory movement or cardiac impulse, and then,
at twenty-two minutes after the administration of the poison, death occurred.

After death, fibrillary twitchings continued for more than twenty minutes,
and the first appearance of rigor was seen in thirty minutes, the extremities
having then become slightly stiff (temperature of laboratory, 63° F.). The post
mortem changes in the condition of the pupils were as follows :—at the moment
of death, they dilated to 42ths x 4%ths of an inch; in one minute, they had con-
tracted to }2ths x 42ths; in two minutes, to ths x 18ths; in three minutes,
to ths x ;4ths; in four minutes, to ths x >ths; in six minutes, to 5,ths x
Asths; and they continued at the last size until twenty-four minutes after
death, when they became dilated to ths x ~ths. On the following day and
while strong general rigor was present, the pupils measured }8ths x 18ths of
an inch.

The influence of atropia on the lethal action of a much larger dose of the
extract was tested in the next experiment.

EXPERIMENT 42-a.—In a rabbit, weighing three pounds and four ounces,
preliminary observations showed that the average rapidity of the heart’s action
was 42 in ten seconds; and of the respiratory movements, 26 in ten seconds ;
and that the pupils measured 44ths x 4%ths of an inch.

A seventeen-hundredth of a grain of sulphate of atropia, dissolved in 30
minims of distilled water, was injected under the skin at the left flank. Two
minutes thereafter, the rate of the heart’s action was 50 in ten seconds. In
four minutes, it had still further increased, having attained a rate of 54 in ten”
seconds, while now the respiratory movements occurred 18 times in ten

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 547

seconds, and the pupils measured i$ths x 48ths of an inch. With these
exceptions, there were no appreciable symptoms present.

Five minutes after the administration of atropia, I injected under the skin
at the right flank a mixture of three grains and nine-tenths of a grain of extract
of physostigma with 40 minims of distilled water ; and, afterwards, under the
skin at the right hip, the few drops of distilled water with which the syringe
was subsequently washed. In three minutes after the injection of the extract of
physostigma, the cardiac impulse occurred 58 times in ten seconds; the pupils
measured 48ths x 28ths of an inch, and infrequent and slight twitches were
present at the right flank and hip. In five minutes, the animal was some-
what restless, and the heart’s rate was now 60 in ten seconds. In seven
minutes, the restlessness was accompanied by slight involuntary shaking of
the head; and, soon after, a great increase took place in the frequency of
the fibrillary twitchings of the panniculus carnosus muscle over the whole
surface of the body. In ten minutes, some weakness was present in the
anterior extremities, and gentle tremors, brief in their continuance, occurred
whenever movements were made, or the animal was startled by any cause.
The weakness of the anterior extremities soon became so great that they were
unable to support the fore part of the body, and then the animal sank down
on the abdomen and chest. Several series of tremors followed this change of
posture ; and, on their termination, the head drooped until the lower jaw was
rested on the table. In fifteen minutes, this posture was still unchanged,
except that the arching of the back had disappeared. The cardiac impulse
occurred 61 times, and the respiratory movements 19 times, in ten seconds,
and the pupils measured 28ths x 48ths of an inch. This general condition
was maintained unchanged for about fifteen minutes, with the exception of a
marked decrease in the frequency of the fibrillary twitchings, and an unim-
portant diminution in the rate of the respiratory movements. Soon afterwards
the symptoms became more serious ; for in forty minutes the respiratory move-
ments occurred only thirteen times in ten seconds, and their character was
somewhat abnormal ; for not only were they weak, and almost entirely confined
to the diaphragm and the abdominal muscles, but the expiratory movements
were abrupt and slightly spasmodic. This depreciation in the character of the
respiratory movements appeared to cause considerable distress, as the animal
every now and then raised the head in an uneasy manner, notwithstanding that
there seemed to be great difficulty in doing so. At this time the heart’s action
was at the rate of 57 in ten seconds. These symptoms continued for about one
hour and ten minutes, but at the end of this time a slight improvement was
manifested ; for, in two hours after the injection of physostigma, the respirations
had increased in rate to 12 in ten seconds, and had become almost normal in their
character. In two hours and thirty minutes, the improved state of the animal

548 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

was still further indicated by the head -being often kept up, without any
trembling, for several seconds, and by the back being again arched ; but the
limbs were still sprawling helplessly, and no general movement could be accom-
plished. The cardiac impulse was now found to recur 47 times in ten seconds,
and the pupils to measure 1%ths x 44ths of an inch, while only rarely a weak
twitch in some portion of the panniculus carnosus muscle could be detected.
The observations were now interrupted until six hours and thirty minutes after
‘the injection of physostigma, by which time a very great improvement had taken
place in the condition of the animal. A normal sitting posture had been
resumed ; paralytic symptoms had almost disappeared, and the rabbit was able
to go about without much difficulty ; and neither general tremors nor fibrillary
twitchings occurred. The rate of the heart’s action was 31 in ten seconds ; the
respirations were irregular, being 20 in one period of ten seconds, and 27 in
another ; and the pupils measured 14ths x t7ths of aninch. It was seen that in
the interval during which no observations were made a large quantity of feces, —
having normal characters, had been passed ; but no urine had yet been voided.

On the following day, the rabbit was found going about actively, and freely
consumed the food that was given to it. The cardiac impulse was at the rate
of 27, and the respirations were at that of about 12, in ten seconds; but the
latter were very irregular. The pupils measured 33ths x 44ths of an inch.

On the third day, the cardiac impulse was at the rate of 37, and the respira-
tions (now pretty regular), were at that of 24 in ten seconds; and the pupils
measured 13ths x 43ths of an inch.

On the fifth day, the’cardiac impulse was at the rate of 40, and the respira-
tions were at that of 23, in ten seconds; and the pupils measured +4ths x
13ths of an inch. By this time, therefore, every appreciable effect of the ex-
periment had disappeared.

On the ninth day, a dose of extract, weighing only one-third of that which —
had been given in this experiment, was administered to the same rabbit ; and
the results of this administration will now be described.

EXPERIMENT 42-b-—The rabbit now weighed three pounds and five ounces;
and immediately before the administration, the rate of the heart’s impulse was 42,
and that of the respirations 21, in ten seconds. One grain and three-tenths
of extract of physostigma was mixed with 20 minims of distilled water, and
the mixture injected under the skin at the right flank. The syringe was then
washed out with a few drops of distilled water, and the washing in its turn
injected under the skin at the right hip. Within one minute and thirty
seconds thereafter, faint fibrillary twitchings occurred, at rare intervals, at the
right flank.. These gradually increased in frequency, until they became a pro-
minent symptom, within four minutes from the commencement of the injection. —
At this time, the heart’s rate had diminished to 33 in ten seconds; but the

¥ ;

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 549

respirations still retained their previous frequency. In six minutes, the rate of
the heart’s impulse was 28, and that of the respirations 21, in ten seconds ; the
fibrillary twitches had become rather more frequent and marked ; and move-
ments of the mouth and lips occurred, which were of such a kind as to suggest
that some substance was being moved from the anterior part of the mouth and
swallowed. There was no other symptom present, and the rabbit sat quietly
on the elevated table on which it had been placed. In eight minutes, however,
uneasiness was manifested by some restless movements, which at first were
somewhat unsteadily performed, and by-and-by were attended with stumblings
and occasional slight tremors. The latter symptoms appeared to be caused by
an undue extension rather than by flaccidity of the limbs. In ten minutes, the
four limbs were in almost complete extension, and the rabbit either stood
unsteadily, or went about stiffly and with stumblings on the limbs thus extended.
_ The pupils measured 33ths x 43ths of an inch; and moist sounds frequently
accompanied the slightly accelerated respiratory movements. No marked
change occurred in the condition of the rabbit for several minutes; but at
fourteen minutes after the injection, the extended state of the anterior extre-
mities had almost entirely disappeared, and the thorax not infrequently rested
on the table, while the pelvis and posterior parts of the body were elevated on
the still extended posterior extremities. The pupils had now contracted to
iiths x 43ths of an inch, and the heart’s rate had decreased "to 22 beats in ten
seconds. In eighteen minutes, the rabbit lay on the abdomen and chest, with
the head drooping, and occasionally resting on the table; the respirations
occurred 25 times in ten seconds, and were accompanied with noisy bubbling
sounds ; frothy saliva was escaping from the mouth; and feces, of a green
colour and semi-liquid consistence, were being passed. Soon, the respiratory
movements became laboured, less frequent, and often greatly obstructed by
accumulations of frothy fluid in the mouth and air-passages ; and the rabbit
was extended on the abdomen, with the head resting on the table, from which
it was raised, though with difficulty, whenever the respiratory movements were
much impeded. In twenty minutes, some general struggling movements
occurred, obviously due to obstructed respiration, and the rabbit fell over on
the side. The cardiac impulses were now at the rate of 18 in ten seconds ;
the pupils measured -6,ths x ;5,ths of an inch ; and the fibrillary twitchings were
very frequent, and affected the whole surface of the animal. The difficulty in
the performance of the respiratory movements gradually became greater, until,
in twenty-nine minutes, only one very laboured, gasping respiration occurred
every ten seconds. Soon afterwards, two or three series of weak tremors affected
the animal, and at the termination of the last of these the respirations alto-
gether ceased, and death took place—thirty-one minutes after the commence-
ment of the first injection.
VOL. XXVI. PART III. a)

550 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

At the moment of death, the pupils measured ths x ths of an inch;
three minutes afterwards, their size had increased to ;4ths x 35ths of an inch ;
and this increase gradually became greater until one hour and thirty minutes,
when they measured 43ths x 4}ths of an inch.

The first appearance of post mortem rigidity was observed at thirty-two
minutes after death, and it consisted of a very slight degree of stiffness re-
stricted to the posterior extremities. The rigidity next appeared in the anterior
extremities and the neck, and finally it became universal, but not until one hour
and fourteen minutes after death. (Temperature of laboratory, 56° F.)

A considerable quantity of opalescent urime was removed from the bladder, ©
and when tested it was found that the opalescence was due to suspended
phosphates, and that the urine was perfectly free from albumen.

In the next experiment, in place of extract of physostigma, the sulphate of
the active principle was administered. .

EXPERIMENT 48-a.—In a rabbit weighing three pounds, it was found that
the rate of the heart’s impulse was 44, and that of the respirations 15, in ten
seconds, and that the pupils measured 14ths x 18ths of an inch.

A solution containing half a grain of sulphate of atropia in 15 minims of
distilled water was injected under the skin at the right flank. In one minute
afterwards, the rate of the heart’s impulse was 47 in ten seconds; in one minute
and thirty seconds, the respirations occurred 22 times in ten seconds ; in two
minutes, the pupils measured 43ths x 48ths of an inch; in three minutes, the
rate of the heart’s impulse was 53 in ten seconds; in three minutes and thirty .
seconds, the respirations occurred 21 times in ten seconds ; and in four minutes,
the rate of the heart’s impulse was 55 in ten seconds, while the pupils measured
46ths x 4&ths of an inch.

Five minutes after the sulphate of atropia had been injected, a solution
. containing six twenty-fifths of a grain of sulphate of physostigmia in 25
minims of distilled water was injected under the skin at the right flank, and
then the syringe was washed out with a few minims of distilled water, which
was injected under the skin at the right hip—the entire operation occupying
thirty seconds. The first symptom that followed was the occurrence of in-
frequent and slight twitches of small portions of the panniculus carnosus
muscle, in the neighbourhood of the regions where the two last injections were
made. These twitches were first observed one minute and twenty seconds after
the commencement of these injections of physostigmia, and they gradually ex-
tended over the surface of the animal, until in three minutes they had become
general. In four minutes, the rate of the heart’s impulse was 49, and that of the
respirations 21, in ten seconds ; and the pupils now measured 48ths x }8ths of
aninch. At this time, also, the respiratory movements were often accompanied
with a hiccup-like start. In six minutes, the rate of the heart’s impulse had

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 551

decreased to 42 in ten seconds; and now the rabbit was affected with occasional
tremblings, restlessness was present, and the movements were somewhat impeded
by a slight degree of extension of the anterior extremities. In eight minutes,
urine and feces were voided, the latter having a perfectly normal appearance,
tremors were of frequent occurrence, and the fibrillary twitchings had become
greatly exaggerated, the entire surface of the animal being in constant move-
ment. In ten minutes, the extension of the limbs had given place to undue
flaccidity, so that they could scarcely support the body; weak general
tremors succeeded each other at intervals; the muscles of the neck seemed unable

_ properly to support the head, which often subsided until the lower jaw nearly
rested on the table ; the respirations occurred 30 times in ten seconds ; and the
pupils measured 4ths x +8ths of an inch. In seventeen minutes,the animal
fell on the abdomen and chest, and remained in this position. Tremors still
occurred, though weaker and less frequent than before, and the fibrillary
twitching of the panniculus carnosus muscle had rather diminished ; but it was
apparent that similar twitchings were occurring in the deeper muscles. In twenty-
two minutes, the lower jaw was rested on the table, and the arching of the back
had almost disappeared. Attempts were made to count the heart’s impulse,
but when the hand was placed on the animal, tremors so severe and continuous
were excited that it was impossible to ascertain the rate with accuracy. In
twenty-five minutes, the general weakness had still further increased, so that the
limbs were extended helplessly at right angles to the body, and the side of the
head was resting on the table. The respirations were now 20 in ten seconds, the
pupils measured i8ths x 48ths of an inch, and the fibrillary twitches had
become less prominently marked. In thirty minutes, a slight improvement was
manifested in the condition of the animal, for spontaneous tremors but rarely
occurred, nor were they excited in their former severity when the hand was

_ placed on the body. It was therefore possible to count the heart’s impulses,
which were ascertained to occur 41 times in ten seconds. A general improve-
ment was still more distinctly perceived at forty minutes after the injection of
physostigmia, when the head was now and then quietly elevated, and attempts
were made to raise the body from the table. The latter were at first unsuccess-
ful, but at forty-nine minutes the rabbit succeeded in rising, and at once assumed
a perfectly normal posture. In fifty-two minutes, several feecal pellets of natural
appearance were passed ; the heart’s impulse was at the rate of 36, and the re-
Spirations were at that of 22, in ten seconds; the fibrillary twitchings were
pretty well marked ; and the rabbit was able to go about, though with consider-
able difficulty. After this, the animal usually sat quiet in a normal attitude, and
in a short time it was able to go about without any perceptible difficulty. In
one hour and thirty minutes, a great number of large feecal pellets were passed,
which were of a somewhat pultaceous consistence. At this time, the rate of

552 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

the heart’s impulse was 36, and that of the respirations 19, inten seconds ; the
pupils measured 4%ths x 4%ths of an inch, and distinct fibrillary twitchings
were still present ; with these exceptions, the animal was in a perfectly normal
state.

On the following day—at twenty-six hours after the injection of physostigmia
—the rabbit was active and well; and it was observed that, since the last note,
a considerable quantity of pultaceous feeces had been passed. The rate of the
heart’s impulse was 32, and that of the respirations 15, in ten seconds ; and the
pupils measured 44ths x 3ths of an inch.

By the fourth day, a normal rate of the cardiac contractions and respiratory
movements, and a normal condition of the pupils, had been reassumed.

On the tenth day, the rabbit was found to weigh three pounds and half an
ounce ; and it was then made the subject of the following experiment :—

EXPERIMENT 43-b.—Having dissolved six twenty-fifths of a grain of sul-
phate of physostigmia in 25 minims of distilled water, I injected the solu-
tion under the skin at the right flank, and then washed the syringe with a few
drops of distilled water, and injected this water under the skin at the right hip.
Before this experiment was commenced, the rate of the cardiac impulse was 42,
and that of the respirations 19, in ten seconds ; and the pupils measured 14ths

x }3ths of an inch. .

In one minute and thirty seconds after the commencement of the admini-

stration, rare fibrillary twitches occurred near the regions of injection; but no
marked general symptoms appeared until four minutes and forty seconds, when
the limbs, especially the two anterior, became extended. The animal then
went about unsteadily, and with considerable difficulty ; and the rate of the
cardiac impulses was 37 in ten seconds. In six minutes, some fecal pellets
were passed ; tremors occurred almost without intermission ; stumbling and
somewhat excited movements were made; and the extended state of the limbs
disappeared, and the rabbit subsided on the abdomen and chest. These symp-
toms rapidly became more and more serious; the pupils contracted to =>ths
x ;8ths of aninch ; general paralysis became well marked ; frequently-recurring
tremors, weaker now than before, impeded the respiratory movements, and
saliva escaped from the mouth. In eight minutes, the respirations consisted
of mere gasps, laboured in their character, and greatly obstructed by mucus,
while the rate of the cardiac impulses had become diminished to 13 in ten
seconds. Soon afterwards, only rarely-occurring gasps were observed, and it was
impossible to detect any cardiac impulse. The former ceased on the occur-
rence of death, nine minutes and fifty seconds after the commencement of the
injection.

At the moment of death, the pupils measured ,5,ths x “ths of an inch, and
they slowly increased in size until, at forty-one minutes after death, they

cf
5

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 553

measured }%ths x ths of an inch; and now, for the first time, post mortem
' rigidity had been initiated, slight stiffness being present in the posterior ex-

tremities (temperature of laboratory, 58° F.)

It is very obviously shown by these experiments that the fatal action of
certain lethal doses of extract of physostigma, and sulphate of physostigmia, may
be prevented in rabbits by the previous administration of atropia.

Atropia and Physostigma simultaneously administered.—In the two following
experiments, extract of physostigma and sulphate of atropia were administered
simultaneously, or nearly so, only an unavoidable interval of a few seconds in-
tervening between the administration of the two substances.

EXPERIMENT 44-a.—In a rabbit that weighed three pounds and twelve
ounces, I injected under the skin of the left flank half a grain of sulphate of
atropia, dissolved in 15 minims of distilled water, and immediately after-
wards three grains of extract of physostigma, suspended in 28 minims of dis-
tilled water. Without any loss of time, the two syringes employed in these
injections were washed out with a few drops of distilled water, and the wash-
ings were separately injected under the skin at different regions.

Except dilatation of the pupils and fibrillary twitches of the muscular struc-
ture beneath the skin, obvious symptoms were but slowly produced, and it
was not until eleven minutes after the injections had been finished that para-
lytic effects were produced. These effects, however, increased in severity some-
what quickly, and in fifteen minutes the rabbit fell over on the side, though it soon
turned again, and lay on the abdomen and chest with the back well arched. At
this time, the pupils were in full dilatation, fibrillary twitches occurred over the
whole surface of the animal, and feces of normal consistence and colour were
passed, while now and again a spasmodic contraction of the abdominal muscles
accompanied the inspiratory movements. Unsuccessful efforts were frequently

_ made to raise the body on the limbs, and often resulted in the production of
general tremors, during which the rabbit several times fell over on the side.
This state of great muscular weakness, attended with well marked fibrillary
twitches of the panniculus carnosus muscle, and apparently also of muscles
more deeply situated, continued, without any improvement, until one hour and
ten minutes after the commencement of the experiment, when further observa-
tions were interrupted.

On the following day the general state of the rabbit appeared a perfectly
normal one. The pupils were, however, in full dilatation, and it was observed
that a large quantity of semi-liquid feces had been passed.

On the third day the feeces that were passed were in every respect normal.
Dilatation of the pupils was still present, and this, the most persistent of the
symptoms, did not disappear until the sixth day.

On the tenth day, the following experiment was performed; the rabbit
VOL. XXVI. PART IIL. 7E

554 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

being perfectly well, and weighing three pounds and twelve ounces and a
half.

EXPERIMENT 44-b.—I injected one grain and a half of extract of physo-
stigma, suspended in 15 minims of distilled water, under the skin at the
right flank, and immediately afterwards washed out the syringe and injected
the washings under the skin at the right hip. Effects were produced in every
essential character analogous to those that have been described as occur-
ring in the preceding experiments with the extract, and in fifteen minutes —
the animal was lying on the abdomen and chest. At this time the only
noteworthy symptoms were an unusually abundant escape of saliva from the
mouth, and a remarkable frequency in the voiding of pultaceous feces. In
twenty-three minutes the rabbit fell over on the side, and while it remained
in this position the respirations were laboured and greatly obstructed by
mucus accumulated in the larynx and air passages, the pupils were con-
tracted, and the cardiac impulses of infrequent occurrence. In twenty-five
minutes a marked improvement occurred in the general condition of the rab-.
bit ; it turned so as to rest on the abdomen and chest, the head was frequently
raised, and the respirations became more frequent and almost free from
obstruction. This improvement was, however, of but short duration, for in
forty minutes the respirations again became embarrassed, tremors and irregular
and somewhat energetic general movements occurred, and the rabbit again
fell over on the side. Gradually the respiratory movements became less
frequent, frothy saliva escaped from the mouth and accumulated in the larynx,
the pupils diminished in size, and the heart’s impulses became feeble and in-
frequent. Soon afterwards the respirations assumed the character of laboured
gasps, greatly impeded by an abundant accumulation of frothy mucus, and they
finally ceased at fifty-four minutes after the injection of the extract.

EXPERIMENT 45-a.—In a young rabbit weighing two pounds and eight
ounces, I injected half a grain of sulphate of atropia, dissolved in 15 minims
of distilled water, under the skin at the right flank, and then one grain of ex-
tract of physostigma, suspended in 15 minims of water, under the skin at the
left flank. Immediately afterwards the water used in washing out each of the
syringes was injected under separate parts of the skin.

Before the experiment the pupils measured 12ths x 44ths of an inch, aa
in we minutes after the commencement of the a injection they had enlarged

+8ths x 23ths of an inch, while slight fibrillary twitches were observed at
ae right side in the immediate neighbourhood of the regions where physo-
stigma had been injected. In seven minutes the rabbit became restless ; in
thirteen minutes the pupils had still further enlarged to }$ths x 4&ths of an
inch; and in fourteen minutes several fecal pellets were passed, and the
fibrillary twitches were more marked, and occurred over the whole surface of

‘

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 555

the animal. With these exceptions, the rabbit appeared to be in a normal
state, until eighteen minutes after the commencement of the injections, when
symptoms of loss of power occurred in the thoracic extremities ; but these
symptoms did not become severe until twenty-five minutes, when stumbling
movements were made, and soon after the rabbit subsided on to the abdomen
and chest. It remained quietly in this attitude, with the head normally raised,
for about eight minutes, and then rose up and stood or went about somewhat
unsteadily. A large additional quantity of feces having a normal character
was passed, and some urine voided. Soon afterwards the partial paralysis
was recovered from, and at forty-five minutes there were no marked general
symptoms present, except a dilated state of the pupils and fibrillary twitches.
A perfect recovery ultimately occurred.

Thirteen days afterwards the following experiment was performed on this
rabbit, which then weighed two pounds and nine ounces.

EXPERIMENT 45-b.—One grain of extract of physostigma, mixed with 15
minims of distilled water, was injected under the skin at the right flank, and im-
mediately afterwards the syringe was washed out with a few drops of distilled
water, and this too was injected under the skin.

The phenomena usually produced by such a dose occurred in their ordinary

sequence : fibrillary twitches, stiff extension of the limbs succeeded by their
partial paralysis, slight tremors, unimportant and transient dilatation followed
by marked contraction of the pupils, defecation, excessive flow of saliva, and
then a state of general flaccidity, interrupted now and then by laboured and
_ gasping respirations. In eighteen minutes after the injection of the extract
the rabbit was dead.*

Physostigma administered before Atropia.—tlt is evident, therefore, that

-atropia is able to prevent the occurrence of death after lethal doses of physo-

| stigma, if the two substances be simultaneously administered. The influence

that it exerts on the lethal action of physostigma when administered after the

dose of this substance is shown by the following experiments. In the first of
these, the extract of physostigma was employed.

EXPERIMENT 46-a.—Having ascertained, in a rabbit weighing three pounds

and two ounces, that the average rate of the cardiac contractions was 40, and

* As has been frequently noticed in similar experiments, the bladder of this rabbit contained a
large quantity of urine. I endeavoured to ascertain whether physostigma is excreted by the kidneys,
by the following process :—About two ounces of this urine was evaporated at a low temperature on a
water bath, and the residue carefully mixed with a little rectified spirit, and the mixture was then
filtered, and in its turn evaporated to dryness at a low temperature. Then the extract thus obtained
was triturated with a very little distilled water, and a drop of the resulting fluid was applied to the
tight eyeball of a rabbit, it having been previously ascertained that both pupils measured }$ths x
$$ths of an inch. The pupils were carefully measured at frequent intervals during the following hour.
and twenty minutes, and it was found that no change occurred in the size of either. It is therefore
highly improbable that the physostigma is excreted by the kidneys.

556 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

of the respirations 27, per ten seconds, and that the pupils measured 19ths x
_ 49ths of an inch, I injected two grains and a half of extract of physostigma,
suspended in 30 minims of distilled water, under the skin of the right flank, and
immediately afterwards the washings of the syringe under the skin at the
right hip. ‘Two minutes thereafter, the respirations were at the rate of 28 per
ten seconds, and infrequent fibrillary twitches occurred at the right side. The
animal became slightly restless and appeared uncomfortable. In four minutes
the cardiac impulse was at the rate of 30 in ten seconds, but the pupils were
unchanged in size.

Five minutes after the commencement of the physostigma injection, half
a grain of sulphate of atropia, dissolved in 15 minims of distilled water,
was injected under the skin at the left flank, and the washings of the
syringe under the skin at the left hip. In one minute after the injection of
atropia, the pupils. had increased to the size of 12ths x 43ths of an inch, and
movements of the lips and mouth, symptomatic of the action of physostigma,
were being made. In two minutes, the rate of the cardiac action had increased
to 45 in ten seconds, and the fibrillary muscular twitches had become frequent
and general over the whole surface of the rabbit ; and in four minutes, the size
of the pupils had still further increased to 4$ths x 4%ths of an inch, while the
movements of the lips referred to still continued. In eight minutes, the rabbit
was sitting normally, though with some slight shaking, the heart’s rate was 51 in
ten seconds, and the fibrillary muscular twitches had become greatly ex-
aggerated. It was not until fifteen minutes after the administration of atropia,
and therefore twenty minutes after that of physostigma, that distinct symptoms
of paralysis manifested themselves, and they consisted of merely a slight yielding
of the forelimbs during the movements, and a little drooping of the head. At
this time the respirations were at the rate of 22 per ten seconds, and the heart’s
action was very frequent, though it was impossible to ascertain its rate with
accuracy, on account of the incessant recurrence of the fibrillary muscular
twitches. These various symptoms continued unchanged until twenty-five
minutes after the administration of atropia, when the paralytic symptoms became

more marked, for whenever the animal attempted to go about it stumbled,
and occasionally even fell on the abdomen. It was seen that the pupils had

now diminished in size to téths x 24ths of an inch. In forty-five minutes
a large quantity of urine was voided; in fifty minutes, the pupils measured
déths x 4%ths of an inch, and normal respiratory movements occurred at the

rate of 26 in ten seconds; and in one hour several large and somewhat soft

feecal pellets were passed. There was not, as yet, any decided change in the
general state of the animal ; stumbling occurred when movements were made,

and although a normal sitting posture could be assumed, there was distinct

drooping of the head while this posture was being maintained. Now, however,

. THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 557

_ the fibrillary muscular twitches had somewhat diminished, and accordingly it
was possible to ascertain with certainty that the rate of the cardiac contractions
was 44 per ten seconds. In one hour and ten minutes a little moisture was
seen at the mouth, and soon afterwards moist sounds occasionally accompanied
the respiratory movements. It was observed, at this time, that the two pupils
had become unequal in size, the right measuring, in full light, +2ths x +2ths, and
the left 44ths x téths, of an inch. In one hour and fifteen minutes the animal
went about quite steadily ; there was no drooping of the head ; the respirations
were frequent and no longer accompanied with moist sounds ; the cardiac con-
tractions were at the rate of 35 in ten seconds; and several soft and wet fecal
pellets were passed, and a little urine was voided. The accumulation of mucus in
the larynx had not, however, been entirely got rid of; for, every now and then,

a curious discordant sound, cough-like in its character, was heard, which was
unmistakably caused by an effort to get rid of some soft substance in the larynx.
In two hours these sounds had altogether ceased; the rate of the cardiac ©
impulses was 41, that of the respiratory movements 22, per ten seconds ; and the
size of the right pupil was ths x 4°ths, and of the left 43ths x 43ths of an inch.
But with the exception of infrequently occurring fibrillary twitches, there was
no obvious symptom present.

On the following day—twenty-seven hours after the commencement of the
experiment—the rabbit seemed to be perfectly well. The cardiac contractions
were occurring at the rate of 31, and the respiratory movements at that of 19,

_ inten seconds ; and the pupils were still unequal, the right measuring }%ths x
13ths, and the left t4ths x 4éths of an inch.

On the third day the most notable change that had occurred was in the
rate of the cardiac contractions, which had by that time reassumed a normal

rate of 41 in ten seconds. It was not, however, until the seventh day, that the
_ pupils had resumed their previous size of 4$ths x 48ths of an inch.

On the twelfth day the rabbit was in a state of vigorous health ; its weight
was three pounds and two ounces and three quarters ; the rate of the heart’s
contractions was 41, and that of the respirations 18, in ten seconds; and the
pupils measured 44ths x 4$ths of an inch.

_ Exprrmment 46-b.—Two minutes after the last observations had been made,

the rabbit received, by subcutaneous injection, two grains and a half of extract

of physostigma. In three minutes and thirty seconds the rate of the cardiac

impulses had fallen to 36 in ten seconds; the respirations were normal, and

there were no general symptoms except infrequent fibrillary twitches and

movements of the lips and mouth. In four minutes and forty seconds, how-

ever, the limbs became extended; and im seven minutes stumbling move-
ments were made, while a slight increase in the size of the pupils was
VOL. XXVI. PART III. 7 *F

558 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

observed. In eight minutes a series of tremors occurred, which were frequently —
repeated until the animal, in nine minutes, sank down on the abdomen and
chest. At this time, the rate of the heart’s contractions was only 19 in ten
seconds, and the respirations were considerably diminished in frequency, and
somewhat laboured, obstruction being apparently caused by mucus accumulated
in the mouth and throat. Very soon afterwards the embarrassment of the
respiratory movements became greater, to such an extent that each respiration
was accompanied by energetic struggling movements of the whole body ; and
in eleven minutes they assumed a gasping character. In eleven minutes and
thirty seconds the head was drawn back, and a few slight tremors occurred,
after which the rabbit was dead.

The first appearance of rigor, consisting of slight stiffness of the posterior
extremities, occurred twenty-four minutes after death (temperature of laboratory,
58° F.).

I shall now describe, but with less minuteness, three other experiments,
where the administration of atropia was preceded by the administration of a
lethal dose of extract of physostigma, and where the interval of time sepa-
rating the administration of the two substances was greater than in the last
experiment.

EXPERIMENT 47-a.—Two grains of extract of physostigma, previously sus-
pended in 20 minims of distilled water, was injected under the skin at the right ©
flank of a rabbit weighing three pounds and eleven ounces and a half. In eight
minutes, the rabbit was lying on the abdomen and chest, saliva was escaping
abundantly from the mouth, the pupils were somewhat contracted, the respira-
tions were noisy and laboured, and moist feeces were being copiously passed.

At eight minutes and thirty seconds, half a grain of sulphate of atropia, dis-
solved in 15 minims of distilled water, was injected under the skin at the left
flank. In four minutes afterwards the pupils were dilated and the flow of
saliva and passage of feeces had ceased. In six minutes vigorous efforts were
made to rise; but these were not successful until fifteen minutes. In about
one hour and twenty minutes, the rabbit was nearly well, though a slight degree
of paralysis was still present. In one hour and forty minutes, every obvious
symptom had disappeared, except dilatation of the pupils and fibrillary twitches
of the muscles.

EXPERIMENT 47-b.—Four days afterwards, this rabbit, while in a perfectly
normal condition, received, by subcutaneous injection, one grain and a half of
extract of physostigma, suspended in 15 minims of distilled water. Tremors,
paralysis, and great increase of the salivary and bronchial mucus secretions,
were quickly produced ; moist faeces were, by-and-by, evacuated in large quan-
tity ; the pupils became contracted ; and death occurred fifteen minutes and —
thirty seconds after the administration. ae

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 559

In the next experiment, an interval of ten minutes and thirty seconds was
allowed to intervene between the administration of the two substances.

EXPERIMENT 48-a—A young rabbit, weighing two pounds and fourteen
ounces, received, by subcutaneous injection, one grain and a half of extract of
physostigma, suspended in 15 minims of distilled water. Symptoms of phy-
sostigma action appeared in one minute and thirty seconds; but they did not
assume a serious aspect until six minutes after the administration, when the
rabbit had great difficulty in maintaining a sitting posture. In nine minutes, it
fell, and rested on the abdomen, chest, and lower jaw. In ten minutes, feces
were passed, and saliva escaped from the mouth ; while the animal lay flaccidly,
quite unable to move about, and, every now and then, was affected with tremors.

At ten minutes and thirty seconds, half a grain of sulphate of atropia, dis-
solved in 15 minims of distilled water, was injected under the skin at the left
flank. No obvious result occurred until four minutes and thirty seconds,
when the state of flaccidity somewhat lessened, the back becoming normally
curved. A few seconds afterwards, the head was again raised, the flow of

_ saliva was considerably diminished, and the pupils were slightly dilated. In
eight minutes the rabbit succeeded in rising, and it then sat im a natural pos-
ture. At this time, the exaggerated secretion of saliva had become completely
checked, and the pupils widely dilated.

EXPERIMENT 48-b-—Twelve days afterwards, one grain and a fifth of extract

of physostigma was suspended in 15 minims of distilled water, and injected
under the skin of this rabbit. Death occurred in thirty minutes.

In the following experiment, likewise, extract of physostigma was adminis-
tered ten minutes and thirty seconds before atropia.

EXPERIMENT 49-a.—Having suspended one grain and a half of extract of
physostigma in 20 minims of distilled water, I injected it under the skin at

the right flank of a strong white rabbit, whose weight was three pounds and

three ounces. Ten minutes thereafter the rabbit was suffering from an
advanced stage of physostigma poisoning. It was lying, unable to make any
movements, except irregular struggles; saliva was freely escaping from the
mouth ; feeces were being passed ; the entire surface of the animal was affected
with fibrillary twitches ; and the pupils were contracted to -ths x ths of an

‘inch, their size before the injection having been 43ths x 44ths of an inch.

At ten minutes and thirty seconds, I injected half a grain of sulphate of
atropia, dissolved in 15 minims of distilled water, under the skin at the left
flank. In three minutes after the administration of atropia, the pupils measured
#oths x 2,ths of an inch, and loud mucous sounds accompanied the respirations.
In six minutes, however, a decided improvement occurred, as the rabbit got
up and went about, though with considerable difficulty, and in a hurried and

excited manner. It was only at rare periods that mucous sounds were heard

560 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

with the respirations ; the exaggerated flow of saliva ceased; and the pupils
became enlarged to ths x =4ths of an inch. This improved state gradually
became more decided, until at thirty-seven minutes after the injection of
atropia, the rabbit seemed to have recovered from every abnormal symptom,
except that extremely well-marked fibrillary twitches and dilatation of the
pupils to the extent of {$ths x 4§ths of an inch were present. At about one
hour and thirty minutes, unmistakable symptoms of general physostigma poison-
ing somewhat unexpectedly again manifested themselves. Feces, normal
at this time in their characters, were passed; the pupils contracted to 42ths x
22ths of an inch, and saliva appeared at the mouth. In one hour and forty
minutes, the respirations were constantly accompanied with mucous sounds ;
soft and, now and again, semi-liquid feeces were passed, and urine was voided ;
the surface of the eye-balls became unnaturally moist ; the pupils measured
only }{ths xi8ths of an inch; and the animal lay on the abdomen and chest,
apparently fae to go abouin However, an improvement in the general
condition again occurred at two hours and twenty minutes; and from this
time the symptoms became less and less severe, until a perfectly normal con-
dition was established.

ExPERIMENT 49-b.—Nine days afterwards this rabbit, being in a state of
vigorous health, and having a weight of three pounds and five ounces, received,
by subcutaneous injection, one grain and three-tenths of extract of physostigma.
In thirty-four minutes thereafter it was extended on the side; infrequent,
laboured, and noisy respiratory gasps were occurring ; soft and almost liquid
feeces were being passed, along with which there were occasionally some small —
pieces of a clear jelly-like substance ; the cardiac contractions were occurring
at a very reduced rate of frequency ; and the pupils were in extreme contrac-
tion. In forty-six minutes and ten seconds after the administration of physo-
stigma, the rabbit was dead.

In two other experiments that will now be described, physostignia was.
administered in the form of sulphate of the active principle, and between the
administration of the two substances a period even longer than that in the last
experiment intervened.

EXPERIMENT 50-a.—In a rabbit, weighing three pounds and ten ounces, the
average rate of the cardiac contractions was found to be 38, and that of the re-
spiratory movements 22, in ten seconds; while the pupils measured 4iths x
48ths of an inch. A solution, containing nine-fifteenths of a grain of sulphadi
of physostigmia in 20 minims of distilled water, was injected under the skin at
the right flank, and, immediately afterwards, the syringe was washed with a
few drops of distilled water, and this water was injected under the skin at the
right hip. The following symptoms then occurred, the time being computed
from the moment when the first injection was commenced :—

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 561

Cardiac Respirations, | Size of Pupils, in
Contractions, per fractions of an General Symptoms, &Xc.
per 10seconds.| 10 seconds. inch.
_ | In 1 min. 30 sec. — 19 a Infrequent fibrillary twitches at right
i. side.
Ly : 32 14
Gar; 30). 29 — $2ths x }2ths | Movement of lips commenced.
(ae ; — — — Shght restlessness and some extension
of the limbs,
Bess : 25 16 34ths x }4ths | The extension of limbs more marked,
and some unsteadiness and shaking. |
HEL 5 ‘ — _— = Excited movements, and often
stumbles, while series of tremors
occur.
HS yy . — 18 — Head droops, the stiff extension of the

limbs has given place to some flac-
cidity and weakness, and the fore
limbs often give way. Somewhat
soft feeces are passed.

14 =, : 14 = iiths x }4ths | The respirations are noisy, impeded
by mucus, and laboured; and the
animal lies on the abdomen and
chest. Saliva escapes freely from
the mouth.

i PS. 5, , 9 — gaths x ;>ths | The animal is on the side. Infre-
A: quent, laboured, and noisy respira-
tions occur, which are accompanied
with general struggles. The cardiac
impulse is extremely weak.

‘sulphate of physostigmia, a solution containing seven-tenths of a grain of
phate of atropia in ten minims of distilled water was injected under the skin
the back ; and immediately afterwards, the syringe was washed and the few
ims of water employed was injected under the skin at the right shoulder.
r the commencement of these injections of sulphate of atropia, which were
made while the animal appeared to be at the point of death, the following
symptoms occurred :—

Cardiac Respirations, | Size of Pupils, in
Contractions, per fractions of an General Symptoms, &c.
per 10seconds.| 10 seconds. inch.
| In 1 min. 30 sec. == 7 goths x gaths
2: » . 50 18 8.ths x ;8;ths | The cardiac impulse is strong.

4 ,, 5 52 18 15ths x 15ths | Still on side. The respirations are
| almost normal in character, and
now unaccompanied with moist
sounds. Fibrillary muscular twitches
very frequent, and occurring over
the whole surface. Now and then
some spasmodic tremors.
Br, : 59 — — The tremors are less frequent, and the
head is occasionally raised.

VOL. XXVI. PART IL. 7G

562 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Cardiac Respirations, | Size of Pupils, in
Contractions, per fractions of an General Symptoms, &c.
per 10seconds.| 10 seconds. inch.
In 9 min. 30 sec. 61 19 4Ziths x iZths | Saliva has ceased to flow from the
mouth.
‘lpi Me GLO pe — — — Has succeeded in turning from side,

and is now lying on the abdomen

and chest, with the lower jaw rest-

ing on the table.

PAU es : — 20 — Moist sounds occasionally accompany
the respirations, Efforts are made
to arch the back. Fibrillary twitches
have become so frequent that it is

. impossible to count the cardiac im-
pulses, but they are ascertained to
be very frequent.

2a, : — 21 tiths x }Zths | The back is now arched, but the lower

jaw still rests on the table, and the

anterior extremities are extended
flaceidly at right angles to the body.

3055, co Ons; 61 16 = The fibrillary twitches are less marked.

DO) 55 ‘ — — 2iths x }8ths | The rabbit got up and walked a short

distance slowly and unsteadily. The

transverse diameter of the pupils is
greater than the perpendicular.

CO, , 50 US, iZths x }$ths | Now and then the rabbit goes about

with great difficulty, but usually

rests quietly on the abdomen and
chest. At rare intervals moist |
sounds accompany the respirations.

1200; : = — — There is only a little difficulty present

when the rabbit goes about. It

usually sits normally, with slight }
drooping of the head. The fibrillary
twitches are prominently marked.

CONN : 42 1G iSths x 1Zths | The general condition remains as last

' noted, except that the fibrillary
twitches are now only slightly
marked. Neither defecation nor |
urination have occurred since the
injection of the atropia.

_-= eo]

dade A Dictate A deeeenicet ae

sik

—

es SS i a ae

ba! se Se

On the following day—twenty-three hours after the commencement of the.
experiment—the rabbit was lively and well. It was ascertained that the cardiac
impulses occurred at the rate of 28, and the pele el movements at that of
9, in ten seconds. The pupils measured 13ths x 1$ths of an inch.

On the fourth day, the restoration of every ‘affected function to a norma
state appeared to have been perfected ; for now the cardiac contractions anf
the respiratory movements had returned to their usual rate of 39 and 22 in ten
seconds, while the pupils measured 43ths x 49ths of an inch, which exactly cor-
responded to their measurement Barre this pra was made. 4

ExpERIMENT 50-b.—On the tenth day this rabbit received, by subcutaneous
injection, nine-fiftieths of a grain of sulphate of physostigmia. It had previou

ie i, cee bie

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 563

been ascertained that the weight of the animal was now three pounds and ten
ounces and a half, that the pupils measured 38ths x ~%ths of an inch, and
that the cardiac contractions were at the rate of 38, and the respirations at
‘that of 15, in ten seconds.

The phenomena usually produced by such a dose quickly made their appear-
_ ance ; among which may be noted a reduction in the rate of the heart’s action
to 29 in ten seconds, five minutes after the commencement of the administration.
‘Stumbling and excited movements, accompanied with slight increase in the size
‘of the pupils, were, by-and-by, succeeded by partial paralysis, accompanied
with frequently recurring tremors, slight contraction of the pupils, noisy infre-
“quent respirations, a flow of saliva from the mouth, and the passage of feeces.
In thirteen minutes, the rabbit fell over on the side, the respirations were gasp-
ing and obstructed by mucus, the heart’s contractions were at the greatly reduced
ate of 14 in ten seconds, and the pupils measured only =5ths x ;§ths of an inch.
Sreasly incessantly the limbs were moving in a to-and-fro direction, and occa-
: sionally they were affected by more vigorous spasmodic movements. In fifteen
mi nutes and thirty seconds, it was impossible to discover any cardiac impulse,
the respirations consisted of rarely occurring gasping movements, the pupils
a ad contracted to ,ths x {ths of an inch, and the sensibility of the eyeballs
had entirely disappeared. Only a few more gasps occurred, and in sixteen
minutes after the commencement of the experiment the rabbit was dead.
After death, the first appearance of rigidity was detected in twenty-five
minutes, but decided general rigor did not occur until the end of fifty-five
minutes (temperature of laboratory 56° F.). At this latter period the pupils
measured 43ths x 29ths of an inch.

In the next experiment, also, it is conspicuously shown that atropia is able
to prevent the lethal action of physostigma even when its administration is
c deferred until death appears to be on the point of occurring.’

_ EXPERIMENT 51-a—In a rabbit, weighing three pounds and eight ounces, it
was ascertained that the rate, per ten seconds, of the heart’s contractions was 40,
and that of the respirations 20, and that the pupils measured 2éths x 18ths of an
inch. The rabbit then received, by subcutaneous injection, seventeen-hundredths
of a grain of sulphate of physostigmia. The following effects were noted :-—

Cardiac Respirations, | Size of Pupils, in
Contractions, per fractions of an General Symptoms, &c.
per10seconds.| 10 seconds. inch.
3 je a ERO RS | Pe MP
| In 1 min. 30 sec. — — — Slight fibrillary twitches at right side.
(a : — 16 — Twitches more frequent.
é 32 — L4ths x i3ths | A little restlessness.
As : 29 16 — Movements of lips and mouth.
a 15 18ths x }4ths | Limbs slightly extended. Shaking a

little.

564

In 10 min
JEON S tae
LA ys
17 red
2008;
23 ”
26° 5
28) ts
28

», 30 sec.

Cardiac
Contractions,
per 10 seconds.

bo
bo

Respirations,
per
10 seconds.

Size of Pupils, in
fractions of an
inch.

DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

General Symptoms, &c.

Stumbles frequently. Wet fecal
pellets passed.

Pultaceous feeces passed. The fibril-
lary twitches are well marked and
general. The rabbit is lying on the
abdomen and thorax.

The respirations are often accompanied
with mucous sounds. Saliva is
escaping from the mouth.

Tremors occurred, and the rabbit fell
on the side, but soon again lay on
the abdomen and thorax.

Tremors occurred. Semi-liquid feces
were passed.

The respirations are considerably im-
peded by saliva and mucus. There
is no arching of the back, and the
lower jaw rests on the table.

The respirations are laboured, and now
and again obstructed bymucus, which
is removed only after energetic
struggles. Rabbit has again fallen
over on side, and is unable to turn
itself. The fibrillary twitching is
now only slight.

Only infrequent laboured respirations
occurring at irregular intervals. The

rabbit is lying on the side, and =
movements of the limbs accompany |
The cardiac im- |

the respirations.
pulse is very weak.

zisths x 285ths | Infrequent and laboured gasps, greatly

obstructed by mucus.

At twenty-nine minutes after the commencement of the injection of physo- —
stigmia, half a grain of sulphate of atropia was administered to the rabbit by i
subcutaneous injection, when the symptoms became modified in the following

manner :—

In 1 min
sig ae
3 ”
6

Cardiac
Contractions,

per 10 seconds.

Respirations,
per
10 seconds.

S|
oS

Size of Pupils, in
fractions of an
inch.

goths x ¢

General Symptoms, &c.

gasps occurring.
Respirations are no longer gasping,

mucous sounds. ie
Respirations are less noisy. 7

Respirations are regular and full, and |

only occasionally ‘accompa with |

moist sounds.
Fibrillary muscular

again become prominent.

5ths | Rabbitisstillonside. Weak struggling | :

ie |

twitches ha ve

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 565

Cardiac Respirations, | Size of Pupils,

Contractions, per in fractions of an General Symptoms, &c.
per 10 seconds. | 10 seconds. inch,

Seep min. . — 15 aths x zths | Rabbit is still on the side, but usually
- quiet, except the shaking that is
caused by the incessant fibrillary

twitches.
) 61 _ ==

Unsuccessful attempts are made to
turn from the side.
he) é — 16 ;éths x 4$ths | The respirations are now quite free
from mucous sounds. The incessant
fibrillary contractions of the muscles
cause twitches not only of the skin,
but also of the toes, legs, ears, tail,
and even eyeballs.
fo” ', ; 62 18 iths x 4$ths | After many efforts, the rabbit suc-
ceeded in turning from the side.
aw, : — — — Some efforts were made to raise the
thorax on the anterior extremities,
but these efforts excited a series of
tremors, during which the rabbit
; fell on the side.
BZD 5, : 63 16 i€ths x {ths | The rabbit hasnowturned from the side.
: — A normal sitting posture was assumed
and maintained. There is, however,
a little drooping of the head.

a 62 — — The rabbit walked about with only a
little difficulty.
HO) +55 : — 14 +$ths x ¢§ths | Occasionally some moist sounds are

heard accompanying the respirations.
There is still a decided degree of
weakness present in the anterior
extremities. The fibrillary twitches
are still incessant in their occurrence.
HOO +s, : 40 14 The fibrillary twitches have diminished
in frequency, but not in the general-
ity of their occurrence, the eyeballs,
ears, feet, &c., being still sometimes
| twitched.
marzo |, : — — tSths x £$ths | The rabbit goes about without any
a unsteadiness. For the first time
since the injection of atropia, some
feecal pellets were now passed, which
are rather larger and slightly softer
than normal pellets.
nae 35 16 i5ths x +$ths | The general condition of the rabbit
seems to be a perfectly normal one.
It is only with difficulty that some
/ rarely occurring fibrillary twitches
can be detected. ‘A few more fecal
pellets have been passed, but no
urine has been voided since the
commencement of the experiment.

_ On the following day this rabbit seemed to be in an absolutely normal con_
dition. It fed largely, and went about actively and well. The pupils measured

+$ths x 18ths of an inch, and the rate per ten seconds of the cardiac contractions
VOL. XXVI. PART III. . iG HL

566 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

was 36, and that of the respiratory movements 18. No further observation was
made until the seventh day, when it was found that the heart was contracting —
41 times in ten seconds, that the respirations were occurring 20 times in ten
seconds, and that the pupils measured 22ths x ti ths of an inch.

The usual testing experiment to prove that the animal had received a lethal
dose of physostigma, was made on the ninth day; but in this instance a smaller
dose was administered than that from which the animal had already recovered.

EXPERIMENT 91-b,
subject of the preceding experiment now weighed three pounds five ounces
and three quarters, I administered to it, by subcutaneous injection, thirteen
one-hundredths of a grain of sulphate of physostigmia. In four minutes there-
after, the rate of the cardiac contractions had diminished to 34 per ten seconds,
and in six minutes to 29 per ten seconds. At the latter time, the limbs of
the animal were extended, and it stood or went about unsteadily with the body
abnormally elevated. Soon afterwards, it became excited, and went about
with hurried stumbling movements; and during these movements, it was found ~
that the heart’s action was accelerated to the rate of 44 in ten seconds. In ~
fourteen minutes, pultaceous freces were passed, moisture appeared at the
mouth, frequent fibrillary twitches were occurring, and occasionally moist
sounds accompanied the somewhat frequent respiratory movements. In
seventeen minutes, the pupils were markedly contracted, and the rabbit lay on
the abdomen and thorax. In twenty minutes, tremors frequently occurred, the
respirations were now laboured and greatly obstructed by mucus and saliva, and
the heart contracted only 16 times in ten seconds. The rabbit was dead in ~
twenty-four minutes.

Immediately before death occurred, the pupils became dilated to 48ths x 4
16ths of an inch; and at the moment of death they became contracted to
8 x Aoths of an inch. After this, their size diminished to jths x “ths of an
inch, at one minute and thirty seconds ; but soon afterwards, gradual dilatation
set in, until they measured -{ths x ths of an inch, twenty-four minutes after —
death. At this time, post mortem rigor had appeared in the posterior extre-
mities (temperature of laboratory, 58° F.)

In these various experiments, the influence exerted by atropia upon the
action of physostigma is shown to be a most remarkable and conspicuous
one, for it effectually counteracts the lethal activity of certain doses of
physostigma, whether it be given within a certain time before, simultaneously
with, or within a certain time after that substance. .

Experiments on Dogs.—The experiments I have described, whereby the cxeamy
tence of this counteraction is demonstrated, were performed on rabbits. In
the absence of proof to the contrary, and in the absence likewise of any reason-
able grounds for entertaining a different opinion, I feel entitled to assume that
this counteraction exists in all the species included in the higher subdivisio

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 567

of the animal kingdom. It was, therefore, with the greatest confidence as to
the result that the following experiments on dogs were performed.

EXPERIMENT 52-a,—I injected three-twentieths of a grain of sulphate of
_atropia, dissolved in 10 minims of distilled water, under the skin at the left
fiank of a cross-bred Spanish terrier, weighing eleven pounds; and the usual
plan was followed of injecting immediately afterwards the washings of the
syringe, so as to ensure that the whole of the dose mentioned should be
introduced.

At five minutes after the commencement of this injection, a dose of nine-
tenths of a grain of sulphate of physostigmia, dissolved in 30 minims of
distilled water, was injected under the skin at the right flank, the syringe was
washed out with a few drops of water, and this water was injected under the
skin at the right hip.

‘In five minutes after the injection of physostigmia, the dog was lying quietly,
apparently but little inconvenienced, and the pupils were dilated. Soon after
distinct fibrillary twitches were observed, a little discomfort was manifested,
and quite suddenly the dog fell over on the side. A normal crouching posture
was, however, soon assumed, but it was maintained for only a few minutes, and
in eleven minutes the dog again fell on the side. A few feeble and unsuccessful
efforts were made to turn, soon afterwards incessant tremors made their appear-
ance, and the fibrillary twitches became greatly increased in their frequency and

) It was not until fifty minutes after the administration of physo-

stigmia, that any decided evidence of an improvement in the general condition
of the dog was observed. It now, however, appeared to take some interest in
ie events that were occurring near it, and when spoken to, elevated the ears
nd even slightly raised the head. In fifty-nine minutes, it got up and walked
about the room slowly and unsteadily.
a On the morning of the second day, the dog eat a large meal with evident
Satisfaction, and with the exception of some languor and of a slowness in the
cardiac contractions, it appeared to be in a normal state. The pupils were now
somewhat contracted.
_ Experiment 52-b.—On the eleventh day—ten days after the performance of
the previous experiment,—the dog, being active and well, and weighing eleven
pounds and four ounces, received, by subcutaneous injection, three-tenths of a
grain of sulphate of physostigmia dissolved in 20 minims of distilled water.
Immediately before this injection, it was ascertained that the heart’s contrac-
tions occurred at the rate of 23 in ten seconds.

_ The first obvious effect occurred in five minutes, and consisted in the pro-
duction of fibrillary twitches. In seven minutes, feces were passed; and
soon afterwards, there was some unsteadiness in the movements, and gentle
tWemors and almost incessant movements of the lips and mouth took place.

568 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

In nine minutes, the dog lay down, but almost at once rose again, though with
great difficulty; and now frothy saliva escaped from the mouth. In ten
minutes, it was lying extended on the floor, with the head resting on the lower
jaw. In eleven minutes, the head fell on the side, starts frequently occurred,
the respirations were gasping, laboured, and obstructed by mucus, and well-
marked fibrillary twitchings were present, which involved the whole surface
of the animal, and seemingly the deeper muscles also. In thirteen minutes, the
animal was altogether on the side, in a flaccid state. In fifteen minutes, the
heart’s contractions occurred at the rate of only 4 in ten seconds, and so long |
were the intervals between the feeble respiratory gasps that more than once it
was thought to be dead. This event, however, did not occur until two minutes —
afterwards, or seventeen minutes after the commencement of the administration —
of physostigmia. 7

In the two next experiments, atropia and physostigma were injected nearly
simultaneously.

EXPERIMENT 53-a.—A vigorous English terrier dog, weighing ten pounds,
received, by subcutaneous injection, eight grains of sulphate of atropia, dissolved —
in 80 minims of distilled water, and immediately afterwards, three grains of —
extract of physostigma, suspended in forty minims of distilled water. These
injections, as well as those subsequently made, by which the washings of the
syringes were introduced under the skin, occupied altogether two minutes.

The chief symptoms that appeared were dilatation of the pupils, partial —
paralysis, frequent vomiting, and hypnotism. Of these, the first continued for
several days, and the two last for less than twenty-four hours. The partial
paralysis was nearly completely recovered from in forty minutes, after wae
the dog was in a perfectly normal condition, except that the pupils were in full
dilatation and that a tendency to sleep was manifested.

EXPERIMENT 53-b.—Three weeks afterwards, this dog being now ten pounds”
and two ounces in weight, received, by subcutaneous injection, eight grains of
sulphate of atropia, and immediately afterwards six grains of extract of

physostigma.

Dilatation of the pupils and considerable loss of motor power were again
produced, but no vomiting occurred. In addition to these symptoms, however,
certain others appeared that were undoubtedly due to physostigma poisoning,
such as tremors and exaggerated bronchial and salivary secretions. The
paralysis and tremors continued for more than three hours, and the dilatation
of the pupils for several days, after which the dog perfectly regained its former
condition. ie

EXPERIMENT 93-c.—Fifteen days after the second of these experiments, tne
dog, being in every respect in a normal condition, received, by subcutaneous i
jection, three grains of extract of physostigma—a dose equal to that from whie

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 569

j it recovered in the first experiment, and only one-half as large as that from
which it recovered in the second. The results were, that fibrillary twitches, par-
tial paralysis, and tremors were quickly produced; that the lachrymal, salivary,
_ and bronchial secretions were profusely increased; that the cardiac contractions
_ became gradually slower and slower; that defsecation and urination occurred ;
_ and that the respirations became more and more laboured and shallow, until they
_ ceased on the occurrence of death, at seventeen minutes after the administration.
It was ascertained after death, that the weight of the dog was ten pounds
and one ounce.

me In the experiment that will now be described, atropia was administered five
- minutes after a lethal dose of sulphate of physostigmia had been injected under
the skin.

EXPERIMENT 54-a.—An active young Scotch terrier dog, weighing ten
pounds and three ounces, received, by subcutaneous injection, three-fifths of a
grain of sulphate of physostigmia, dissolved in 25 minims of distilled water.
Before the injection, the rate per ten seconds of the cardiac impulses was
32, and that of the respirations 4, andthe size of the pupils, in a full light, was
ths x 22ths of an inch.

In two minutes after the commencement of the administration, symptoms of
discomfort were manifested, and the lips were moved and licked with the tongue,
as if an unusual quantity of fluid were present in the anterior part of the mouth.
In four minutes, slight tremors frequently occurred, and fibrillary twitches were
present.

In five minutes, a solution containing three-tenths of a grain of sulphate of
_atropia in 15 minims of distilled water, was injected under the skin at the right
- flank. In two minutes thereafter, the tremors already noted had become more
prominent and strong, the limbs were unable properly to support the body, urine
_ was voided, saliva escaped from the mouth, and the eyeballs were unnaturally
moist. The tremorsand weakness quickly increased, so that, on account of the
Tormer, it became impossible to determine the rate of the cardiac and respiratory
\ “movements; while, on account of the latter, stumbles occurred, and the head
began to droop, until often it touched the floor. In five minutes, the pupils
were greatly dilated, but now the secretions of the salivary and lachrymal
glands were diminished. In seven minutes, the dog lay quietly on the abdomen
| and thorax, and in thirteen minutes, it fell over on the side. An endeavour was
“made to count the cardiac impulses ; but when the hand was placed over the
heart, the tremors referred to became so greatly increased that it was im-
‘possible to distinguish the heart’s impulse. It was not until thirty-eight
minutes that an attempt to “ete the heart’s contractions was successful,

econds. At the same time, the respirations had a rate of 7 in ten seconds, and
VOL. XXVI. PART III. a1

570 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

the pupils measured 44ths x 24ths of aninch. In forty-eight minutes, the con-
dition of the dog had so far improved, that, after some efforts, it rose on the
limbs, and then lay down in a normal crouching attitude, with the head raised. _
In fifty-three minutes, the dog attempted to vomit, but it was not until one
hour and sixteen minutes that emesis occurred. Soon afterwards, it again got
up and walked about the room, with only a little unsteadiness. In one hour
and fifty-five minutes, the animal seemed to be perfectly well. The rate per
ten seconds of the cardiac contractions was 47, and that of the respirations 10,
and the pupils measured about 14ths x 4%ths of an inch. During all this time,
urine had been voided only os and no feeces had been passed.

On the following day, the dog was active and in a perfectly normal general
condition. The cardiac impulses occurred at the rate of 48, and the respiratory
movements at that of 5, in ten seconds, while the pupils measured 38ths x 18ths
of an inch.

It was not, however, until the sixth day that the heart’s action had become
reduced to the normal frequency of about 30 contractions in the ten seconds
and the pupils remained more or less dilated for other eight days, but on the
fifteenth day they had returned to the condition that existed previously to the
experiment.

This dog afterwards received without any atropia a dose of physostigmia
only one-half as large as that from which it recovered when atropia also was
given, and the following effects were produced :—

EXPERIMENT 54-b.—Nineteen days after the performance of the previous ex-
periment, the dog that had been used in it received, by subcutaneous injection,
three-tenths of a grain of sulphate of physostigmia, dissolved in a small quantity
of distilled water. In five minutes, symptoms of discomfort, slight unsteadiness —
of the limbs, and fibrillary twitches were observed; and soon after, struggling
and stumbling movements occurred, and the flow of tears and saliva became
increased. In eight minutes, decided paralysis of the posterior extremities was
present. In ten minutes, the dog lay down on the abdomen, and rested the
lower jaw on the floor. Series of gentle tremors succeeded each other in rapid
succession, and at the end of one of them the dog fell over on the side. Saliva
now escaped freely from the mouth, wet and soft feeces were passed, and the
respirations became rapid, noisy, and shallow. In fifteen minutes, the respira-
tions were very laboured and jerking, though still abnormally frequent, and the
tremors had somwhat increased in severity. In a short time, however, the
tremors became less severe and frequent, but at the same time the respiratory
movements became laboured, somewhat shallow, and greatly obstructed by
mucus accumulated in the mouth and larynx, and the cardiac impulse became
infrequent and weak. In ninteen minutes, the respirations consisted merely of
rarely occurring gasps, the pupils were contracted, and the sensibility of the

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 571

eyeballs had disappeared, while it was only with difficulty that now and again a
cardiac impulse could be detected. The gasping respiratory efforts became
gradually separated from each other by longer and longer intervals, until they
altogether ceased at twenty-two minutes after the commencement of the injec-
_ tion of physostigmia.
After death, it was ascertained that the weight of the dog was ten pounds
and four ounces.
It is shown by these experiments, that in dogs, as in rabbits, atropia exerts
a powerful counteracting influence to the lethal action of physostigma. It
_ would have been a matter of surprise had this result not been obtained, for there
was no reason to anticipate that either atropia or phyostigma would act other-
wise than in comformity with the general law, that every active substance
- influences the same histological structures in the same way in whatever animal
these structures are present. No doubt the prominence and importance of the
‘results that are produced by essentially the same action vary somewhat in
different animals; but in judging of the probable existence of an antagonism
between two substances, the prominence or importance of an effect resulting
from any primary action is of secondary moment to the fact of the existence of
; the action. Accordingly, if atropia be capable of producing upon one species
- of animal an influence of such a nature as to antagonise in it the lethal action of
_ physostigma, it is difficult to imagine why it should not produce the same
antagonising influence in all animals of equally high development. The mere
‘fact of there beimg a difference in the lethal activity of atropia in different
animals, is not sufficient to lead to the supposition that it will not in them
“successfully counteract the lethal action of physostigma; for the same primary
actions are produced, notwithstanding the differences that may exist in the
“lethal activity. Many circumstances of a more or less accidental nature may
‘modify the lethal activity of poisonous substances, and among these is the
m anner in which the substance is administered. In the case of atropia, its
lethal activity in rabbits may be enormously increased by introducing it directly
into a blood-vessal.
Experiments in which Atropia was injected into a vein and Physostigma
under the skin.—It seemed therefore of importance to administer atropia by
‘injection into a vein, in order to determine whether, when so administered, it
still, notwithstanding the great increase that is thus produced in its lethal
| activity, retains the power to counteract the lethal action of physostigma.
___ Experiment 55-a.—A rabbit, weighing four pounds, was secured by means
. of a CzErMax’s rabbit-holder, and one of the external facial veins was exposed,
and two ligatures were loosely applied to a small portion of it dissected from
| its connections. Two grains of extract of physostigma was then administered
_to the rabbit by subcutaneous injection.

572 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Five minutes afterwards, a thirtieth of a grain of sulphate of atropia, dissolved
in 10 minims of distilled water, was injected into the exposed facial vein. The
previously applied ligatures were then carefully secured, and the incision closed
by silk sutures. On the animal being set free, it was observed that movements
were performed with difficulty. In seven minutes after the injection of atropia,
the animal lay down on the abdomen and thorax, occasionally mucous sounds
were heard with the respirations, and it was observed that the entire surface of
the animal was affected with fibrillary twitches. In eight minutes, the pupils were
dilated (48ths x 33ths of an inch), the animal had assumed a normal sitting pos-
ture, and mucous sounds no longer accompanied the respirations. Indeed, with
the exception of great dilatation of the pupils and fibrillary twitches, the rabbit
seemed perfectly well. In seventeen minutes, however, symptoms of paralysis
again appeared, and wet feeces were passed; while in twenty-three minutes, the
respirations again became accompanied with mucous sounds, and the dilatation
of the pupils somewhat diminished. These symptoms continued, without any
improvement in the condition of the animal, for twenty-six minutes; but in
forty-six minutes after the injection of atropia, the rabbit raised itself with some
difficulty, and went about unsteadily. The pupils now measured 13ths x 44ths
of an inch, the respirations were laboured and noisy, and often the rabbit went
about ina very excited manner. In one hour and fifteen minutes, a large quan-
tity of urine was voided, and pultaceous feces were passed. It was not until -
three hours and ten minutes after the injection of atropia that a nearly normal
condition was assumed, and at this time no symptoms were present except
dilatation of the pupils, and rarely occurring mucous respiratory sounds. Ulti-
mately the rabbit recovered perfectly, and it was afterwards subjected to the
action of a lethal though smaller dose of physostigma than that from which it —
had thus recovered.

EXPERIMENT 53-b.—Seven days after the performance of the last experiment,
the rabbit which had been used in it received, by subcutaneous injection, one
grain and seven-tenths of extract of physostigma.

Previously to the performance of the injection, it was ascertained that the
weight of the rabbit was four pounds and three ounces; that the rate, per ten
seconds, of the cardiac contractions was 47, and that of the respirations 29;
and that the pupils measured 43ths x 34ths of an inch.

After the injection, symptoms of poisoning quickly manifested themselves,
and in seven minutes the rabbit was suffering from well-marked general para-
lysis, the rate per ten seconds of the cardiac contractions was 39, and that of
the respirations 28; while the pupils measured i$ths x 13ths of aninch. A short
period occurred during which the limbs were extended, and stumbling, excited
movements took place, and then the rabbit fell on the abdomen and thorax,
the respirations became noisy, wet and soft feeces were passed, and tremors —

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 573

succeeded each other at short intervals. In twelve minutes, the respirations
were laboured, and at the rate of 15 in ten seconds, the cardiac contractions
occurred 34 times in ten seconds, and the pupils measured only 19ths x ths
of an inch. The respiratory embarrassment soon became much greater, and
at the same time the general paralysis increased ; until, in twenty-one minutes,
only laboured, gasping and noisy respirations took place, the rabbit fell over
’ on the side, the pupils contracted to “jths x ths of an inch, and the cardiac
contractions were weak, and occurred only 12 times in ten seconds. At
twenty-three minutes after the administration of physostigma, the rabbit was
dead.

This method of administration was followed in the next experiment like-
wise.

EXPERIMENT 56-a.—Having exposed one of the external facial veins on
the left side of a rabbit, weighing three pounds and two ounces, I injected
under the skin, at the left flank, one grain and three-fifths of extract of
physostigma, mixed with 15 minims of distilled water.

Five minutes afterwards, I injected into the exposed and dissected vein a
forty-fifth of a grain of sulphate of atropia, dissolved in 8 minims of distilled
water. The vein was then ligatured, the wound closed with silk sutures, and
the rabbit set free from the CzERMAxk’s holder by which it had been secured.

In nine minutes after the injection of sulphate of atropia, the rabbit was
lying on the abdomen and chest, frequent fibrillary twitches were occurring over
the whole surface, the pupils were dilated, and the cardiac action was abnormally
frequent. In twenty-three minutes, it rose and went about, though somewhat
unsteadily. From this time, the general condition of the rabbit steadily im-
proved, until, in one hour and thirty minutes, there were no symptoms present
except pupillary dilatation, abnormal frequency of cardiac action, and slight
fibrillary twitches. During the experiment, there had not been any obvious
increase in the secretions of the salivary, bronchial, or lachrymal glands; nor
did defzecation or urination take place until more than two hours and fifteen

minutes after the experiment had been commenced.
_ Expertment 56-b.—The rabbit that had formed the subject of the preceding
experiment received, eight days afterwards, one grain and three-tenths of extract
of physostigma, suspended in a few minims of distilled water. Atthis time the
weight of the rabbit was three pounds and four ounces.

In nine minutes after the injection of the extract, the rabbit was lying on the
abdomen and chest, affected with pretty severe tremors, and breathing some-
what rapidly and noisily. In fourteen minutes, the pupils were contracted, and
the respirations were laboured and embarrassed by an accumulation in the
mouth of mucus and saliva, while the cardiac contractions were occurring
infrequently. By this time, = a considerable quantity of soft and pultaceous
VOL. XXVI. PART III. 7K

574 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

feeces had been passed. In fifteen minutes, the rabbit was lying on the side, and
laboured and infrequent gasping respirations were occurring. Soon afterwards,
the sensibility of the conjunctiva disappeared, the cardiac impulses became
extremely weak, and it was only at long intervals that a feeble gasp occurred.
Death occurred at nineteen minutes after the administration of physostigma.

It is shown by these two experiments that in rabbits atropia retains its
remarkable power of counteracting the lethal action of physostigma even when
its toxic activity in these animals is greatly increased.

Experiment with a Preparation of Physostigma different from that used in
all the other Experiments.—As the preceding experiments were, without excep-
tion, made with extract of physostigma and sulphate of physostigmia prepared
by myself, it seemed not altogether superfluous to check the results that were
obtained, by making some additional experiments with a preparation for whose
activity and properties I was not responsible. Accordingly, several experiments
were made with an extract prepared by Dr Cook, of the well-known firm of
Messrs T. and H. Smirn. With this extract essentially the same results were
obtained as with the preparations used in all the other experiments. It is,
therefore, unnecessary to give a description of more than one experiment in
which it was emplayed. .

EXPERIMENT 97-a.—A_ rabbit, weighing three pounds and eight ounces,
received, by subcutaneous injection, two grains of Dr Coox’s extract of physo-
stigma, suspended in 40 minims of distilled water. One minute and a half
afterwards, it received, also by subcutaneous injection, half a grain of sulphate
of atropia, dissolved in 10 minims of distilled water.

Tn three minutes after the injection of sulphate of atropia, the pupils measured
14ths x Léths of an inch, the measurement immediately before the experiment
having been }$ths x ths. In seven minutes, the pupils measured 1$ths x ths
of an inch, the rate of the heart’s contractions was considerably accelerated,
fibrillary twitches were occurring, and a little restlessness was present. In
thirteen minutes, this restlessness had become somewhat greater, and the animal
had decided difficulty in moving about. Soon afterwards the pupils became still
more dilated, and in eighteen minutes they measured 48ths x 44ths of an inch.
In twenty-five minutes, the difficulty in moving about had become greater—_
even to such an extent that often the anterior extremities yielded, and the rabbit
fell on the thorax. It appeared also to be in a somewhat excited state, as con-
fused and stumbling movements were frequently made. In fifty-two minutes,
the pupils measured 43ths xtéths of an inch, but no obvious change had
occurred in the general condition of the animal. In one hour and ten minutes,
however, evidences of recovery were manifested ; the rabbit went about with
but little difficulty, no restless excitement was present, and frequently a per-
fectly normal sitting posture was assumed. Indeed, the only symptom of an

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. D795

abnormal condition that was distinctly apparent consisted of frequently occur-
ring and well-marked fibrillary twitches. From this time the condition of the
animal steadily improved until perfect recovery took place.

EXPERIMENT 87-b.—Nine days afterwards, the rabbit which formed the sub-
ject of the previous part of this experiment received, by subcutaneous injection,
one grain of Dr Coox’s extract of physostigma, suspended in 20 minims of
distilled water. Before the injection was made, it was ascertained that the
weight of the rabbit was three pounds and eight ounces and a half, and that the
pupils measured i3ths x 49ths of an inch,

Symptoms of poisoning very quickly appeared. In six minutes, the posterior
extremities were trailing, and the anterior considerably extended, and stumbling
movements occurred, while well-marked fibrillary twitches were present. In
eight minutes, saliva was escaping from the mouth in drops, and tremors
frequently occurred. In eleven minutes, the rabbit was lying on the abdomen
and chest; several faecal pellets were passed; the pupils were contracted
(.ths x ths of an inch) ; and the respirations were rapid and accompanied with
mucous sounds. Soon afterwards, the -head subsided: until the lower jaw rested
on the table; the arching of the back disappeared ; the pupils became still
- further contracted (ths x 4;ths of an inch); the cardiac contractions greatly
diminished in frequency and strength; and the respiratory movements assumed
a gasping character. Feeble tremors then occurred, and the head was drawn
backwards ; after which a condition of general flaccidity set m. A few more
feeble gasps occurred, and then the respirations altogether ceased, at thirteen
minutes and thirty seconds after the commencement of the injection of
_ physostigma.

In this experiment the power of atropia to counteract the lethal action of
the extract of physostigma prepared by Dr Coox is displayed in a very remark-
able manner. '

The second portion of the experiment shows that the lethal activity of this
extract is considerably greater than that of the extract prepared by myself.
Nevertheless, atropia so completely and successfully antagonised the lethal
action, as to prevent the occurrence of any symptom of serious import after the
administration of a dose twice as large as that-by which death was afterwards
produced in about thirteen minutes.

Summary of the preceding Experiments.— Before passing to the second por-
tion of this research, it may be of advantage to give a brief summary of the
various facts that have been brought forward.

1. It has been shown by a statement of the result of several experiments,
that the minimum lethal dose for rabbits of extract of physostigma is 1:2 grain,
and that of sulphate of physostigmia 0°12 grain, for every three pounds weight
of animal.

576 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

2. The influence that is exerted by atropia upon the lethal action of extract
of physostigma and sulphate of physostigmia has been examined in rabbits, and
a description has been given of several experiments that were performed for
this purpose. The following facts have been stated among the conditions and
. results of these experiments :—

EXPERIMENT 41-a.—A_ rabbit, weighing 2 Ibs. 154 oz., received 0°3 grain of
sulphate of atropia; and, five minutes afterwards, 1:2
grain of extract of physostigma. Recovery took place.

41-b.—Ten days afterwards, the same rabbit, now weighing 3 lbs.,
received 1:2 grain of extract of physostigma. Death
occurred in twenty-two minutes.

EXPERIMENT 42-a.—A rabbit, weighing 3 lbs. 4 0z., received 0°17 grain of sul-
phate of atropia; and, five minutes afterwards, 3-9 grains
of extract of physostigma. Recovery took place. .

42-b.— Eight days afterwards, the same rabbit, now weighing 3 lbs.
5 oz., received 1°3 grain of extract of physostigma. Death
occurred in thirty-one minutes.

EXPERIMENT 43-a.—A rabbit, weighing 3 lbs., received 0°5 grain of sulphate of
atropia; and, five minutes afterwards, 0°24 grain of a
phate of physostigmia. Recovery took place.

43-b.—Nine days afterwards, the same rabbit, now weighing 3 Ibs.
4 oz., received 0:24 grain of sulphate of physostigmia.
Death occurred in nine minutes and fifty seconds.

EXPERIMENT 44-b.—A rabbit, weighing 3 lbs. 12 oz., received 0°5 grain of sul-
phate of atropia; and, nearly at the same time, 3 grains -
of extract of physostigma. Recovery took place.

44-b.—Nine days afterwards, the same rabbit, now weighing 3 lbs.
123 oz., received 1°5 grain of extract of physostigma.
Death occurred in fifty-four minutes.

EXPERIMENT 45-a.—A_ rabbit, weighing 2 lbs. 8 oz., received 0°5 grain of sul-
phate of atropia; and, nearly at the same time, 1 grain of
extract of physostigma. Recovery took place.

45-b.—Thirteen days afterwards, the same rabbit, now weighing
2 Ibs. 9 oz., received 1 grain of extract of physostigma.
« Death occurred in eighteen minutes.

EXPERIMENT 46-a.—A rabbit, weighing 3 lbs. 2 oz., received 2°5 grains of
extract of physostigma ; and, five minutes afterwards, 0°
erain of sulphate of atropia. Recovery took place.

46-b.— Eleven days afterwards, the same rabbit, now weighing 3 lbs.
22 oz., received 2°5 grains of extract of physostigma.
Death occurred in eleven minutes and thirty seconds.

;
z
a
»

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 577

Experiment 47-a.—A rabbit, weighing 3 Ibs. 114 0z., received 2 grains of
extract of physostigma; and, eight minutes and thirty
seconds afterwards, 0°5 grain of sulphate of atropia. Re-
covery took place.

47-b.—Four days afterwards, the same rabbit, now weighing 3 lbs.
8 oz., received 1°5 grain of extract of physostigma. Death

. occurred in fifteen minutes and thirty seconds.

ENT 48-a.—A rabbit, weighing 2 lbs. 14 0z., received 1°5 grain of
extract of physostigma; and, ten minutes and thirty
seconds afterwards, 0°5 grain of sulphate of atropia. Re-
covery took place.

48-b.—Twelve days afterwards, the same rabbit, now weighing
3 lbs. 1 0z., received 1:2 grain of extract of physostigma.
Death occurred in thirty minutes.

EXPERIMENT 49-a.—A rabbit, weighing 3 lbs. 3 0z., received 1°5 grain of extract
| of physostigma ; and, ten eannabos and thirty seconds after-
wards, 0°5 grain of sulphate of atropia. Recovery took
place. .
49-b.—Nine days afterwards, the same rabbit, now weighing 3 lbs.
5 oz., received 1°3 grain of extract of physostigma. Death
occurred in forty-six minutes and ten seconds.

EXPERIMENT 50-a.—A rabbit, weighing 3 lbs. 10 oz., received 0°18 grain of sul-
phate of physostigmia; and, fifteen minutes and ten
seconds afterwards, 0°7 grain of sulphate of atropia. Re-
covery took place.

50-b.-_Nine days afterwards, the same rabbit, now weighing 3 lbs.
104 oz., received 0°18 grain of sulphate of physostigmia.

Death occurred in sixteen minutes.

EXPERIMENT d1-a.—A rabbit, weighing 3 lbs. 8 oz., received 0°17 grain of sul-

phate of physostigmia; and, twenty-nine minutes after-

wards, 0°5 grain of She of atropia. Recovery took

place.
51-b. the same rabbit, now weighing 3 lbs.
52 oz., received 0°13 grain of sulphate of physostigmia.
x Death occurred in twenty-four minutes.

3. Several experiments have been described, in which the influence exerted
| by atropia upon the lethal action of extract of physostigma and sulphate of
physostigmia was examined in dogs also. The following facts were stated
among the conditions and results of these experiments :—

VOL. XXVI. PART III. 71

578 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

EXPERIMENT 52-a.—A dog, weighing 11 lbs., received 0°15 grain of sulphate of
atropia, and, five minutes afterwards, 0°9 grain of sulphate
of physostigmia. Recovery took place.

52-b.—Ten days afterwards, the same dog, now weighing 11 lbs.
4 oz., received 0°3 grain of sulphate of physostigmia.
Death occurred in seventeen minutes.

EXPERIMENT 93-a.—A dog, weighing 10 lbs., received 8 grains of sulphate of
atropia, and, immediately afterwards, 3 grains of extract
of physostigma. Recovery took place.

53-b.—Three weeks afterwards, the same dog, now weighing
10 lbs. 2 oz., received 8 grains of sulphate of atropia,
and, immediately afterwards, 6 grains of extract of physo-
stigma. Recovery took place.

53-c.— Fifteen days after the second experiment, the same dog,
now weighing 10 lbs. 1 oz., received 3 grains of extract of
physostigma. Death occurred in seventeen minutes.

EXPERIMENT 54-a.—A dog, weighing 10 lbs. 3 oz., received 0°6 grain of sul-
phate of physostigmia, and, five minutes afterwards, 0°3
grain of sulphate of atropia. Recovery took place.

54-b.—Nineteen days afterwards, the same dog, now weighing
10 Ibs. 4 0z., received 0°3 grain of sulphate of physo-
stigmia. Death occurred in twenty minutes. .

Although these experiments clearly demonstrate that atropia is able to —
counteract the lethal action of physostigma in rabbits and dogs, it is possible to
suppose that it will not do so m other animals of equally high development. —
Some support is given to this surmise by evidence tending to show that the
action of atropia in certain animals is different from its action in others. The
only difference, however, that is known to exist, is in the lethal activity of the
substance, relatively to certain animals; and in rabbits and dogs, this lethal
activity is less than in several other animals. Accordingly, if the lethal activity
of atropia for rabbits and dogs be increased, and if, notwithstanding this —
increase, successful antagonism be still produced in them, the only reason for
supposing that successful antagonism will not be produced in certain other
animals will be shown to be an insufficient one. For any given species of
animal, the lethal activity of atropia may be modified by the method of ad-
ministration, and it is very much greater when this substance is directly
introduced into the blood-stream, than when it is injected under the
skin. ;

4. The influence exerted on the lethal action of physostigma by atropia
injected directly into the blood-stream, was examined by experiments on rabbits.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 579

The following facts were stated among the conditions and results of these ex-
periments :—

_ EXPERIMENT 55-a.—A rabbit, weighing 4 lbs., received 2 grains of extract of
physostigma; and, five minutes afterwards, 0:03 grain of
sulphate of atropia by injection into a facial vein. Re-
covery took place.

55-b.—Seven days afterwards, the same rabbit, now weighing 4 lbs.

3 oz., received 1°7 grain of extract of physostigma. Death

occurred in twenty-three minutes. |

A rabbit, weighing 3 lbs. 2 0z., received 1°6 grain of ex-

tract of physostigma, and, five minutes afterwards, 0:02

grain of sulphate of atropia by injection into a facial vein.

Recovery took place.

56-b.—Eight days afterwards, the same rabbit, now weighing 8 lbs.
4 oz., received 1°3 grain of extract of physostigma. Death
occurred in nineteen minutes.

EXPERIMENT 956-a.

5. An experiment was described, which had been undertaken to determine -
whether a preparation of physostigma, different from that used in any of the other
experiments of this research, has its lethal action counteracted by sulphate of
atropia. The following facts were stated among the conditions and results of
this experiment :—

A rabbit, weighing 3 lbs. 8 oz., received 2 grains of ex-
tract of physostigma, prepared by Dr Cook; and, one
minute and thirty seconds afterwards, 0°5 grain of sul-
phate of atropia. Recovery took place.

57-b.—Nine days afterwards, the same rabbit, now weighing 3 lbs.
84 oz., received 1 grain of extract of physostigma, prepared
by Dr Coox. Death occurred in thirteen minutes and
thirty seconds.

EXPERIMENT 57-a.

_ These various experiments prove so clearly that atropia is able to counter-
act the lethal action of physostigma, as to be of themselves sufficient for the
purposes of this section. Their evidence may, however, be largely added to

580 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Section B.—DETERMINATION OF THE EXTENT OF THE COUNTERACTING
INFLUENCE OF ATROPIA UPON THE LETHAL ACTION OF PHYSOSTIGMA.

In the “ preliminary note” which I communicated to this Society, on the
antagonism between physostigma and atropia, the opinion was expressed, that
as this antagonism is “‘ concerned with two substances, each of which possesses
a number of actions, it is not unreasonable to anticipate that several of them are
not mutually antagonistic,” and that “above certain doses, a region may, there-
fore, be entered where the non-antagonised actions are present in sufficient
degrees to be themselves able to produce fatal results.” * Besides this con-
sideration, there are others derived from our knowledge of the physiological
action of physostigma, which render it probable that such a region exists.

Certain of the actions of the two substances are of a similar nature. When
a dose not much above the minimum-lethal of the one is counteracted by a
small dose of the other, the similar actions are not produced in sufficient
intensity to become, even in combination, important toxic actions. When,
however, a dose considerably above the minimum lethal of the one substance
is given along with a large dose of the other, the similar actions may be
produced in such intensity as to assume the importance of lethal actions.

Further, with regard to the counteracting actions themselves, it is to be
observed that various of the facts mentioned in the record of experiments
that is given in the preceding section tend to make mutual antagonism
probable, not only of one but of several of the actions of physostigma and
atropia ; and it is legitimate to suppose, that with a given dose of physostigma,
the counteraction produced by a certain amount of atropia will be more perfect —
in the case of one or more of the antagonistic actions than in that of others, and
that with certain doses of the two substances such incompleteness of counter-
action may exist as would, even without the occurrence of non-antagonised
action, suffice for the production of death.

Guided by these considerations, I anticipated that the counteracting influence
of atropia upon the lethal action of physostigma is successfully exerted only
within a definite range of doses, and that this range may be determined by —
experimental research. The somewhat laborious task of making this deter-
mination has been undertaken because it seemed likely that results would
thereby be obtained of the greatest interest and novelty, in connection not
only with this special instance of counteraction, but also with the general
subject of physiological antagonism and its important and direct bearing
on the principles of therapeutics.

* Loe, cit., p. 589.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 581

In order to define the limits of the counteracting influence of atropia upon
the lethal action of physostigma, three series of experiments were made.
The chief objects of the two first of these were to ascertain the maximum
_ dose of physostigma that can be successfully antagonised by atropia, and the
range of doses of atropia that can successfully antagonise lethal doses of
_ physostigma. In each series, a constant interval of time was maintained
_ between the administration of the two substances; but in the first, atropia was
administered five minutes before physostigma, while in the second, physostigma
_ was administered five minutes before atropia. These intervals of time were
selected in preference to simultaneous administration because, practically, it is
_ impossible for one experimenter to inject the two substances into different
regions exactly at the same moment, and further, because it seemed probable
that a difference would be found to exist in the counteracting power of atropia
according as it is given before or after physostigma. In both of these series,
experiments were made, in the first place, with the minimum-lethal dose of
_physostigma, and in combination with it, various doses of atropia were
administered, ranging from one that was too small to prevent the lethal action,
through a number that were able to prevent death, until a dose was .found
whose administration resulted in death. Similar experiments were made with
a dose of physostigma one and a half times as large as the minimum-lethal,
_ then with one twice as large as the minimum-lethal, and so on, at the same rate
of progression, until a dose of physostigma was reached that was too large to
be successfully antagonised by any dose of atropia.
The chief object of the third series of experiments was to ascertain within
what limits of time between the administration of the two substances successful
antagonism occurs. In the experiments of this series, a constant dose of physo-
‘stigma was given along with various doses of atropia, and with each dose of
3 atropia several experiments were made which differed from each other by a
} ifference in the interval of time between the administration of the two
substances. On this plan two sets of experiments were performed, in one of

given after; and subsequently these two sets of experiments were connected
together by a third, in which atropia, in various doses, was administered nearly
Simultaneously with the same dose of pePyResusnte as was given in the two
other sets of experiments.

| All the experiments of this portion of the research were performed on
| rabbits. In the great majority of the experiments, the weight of the animal
Was about three pounds, but when it was below or in excess of this, the doses
| of the substances administered were calculated for three pounds weight of
| animal.

In the description that will now be given of these experiments, the doses
VOL. XXVI. PART III. 7M

582 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

per three pounds weight of rabbit will alone be mentioned. For further details,
and more especially for the actual doses, the weight of the animals, and the
chief symptoms, I must refer to the Tabular Summary at the end of the

paper.

1. DETERMINATION OF THE LIMITS OF ANTAGONISM WHEN ATROPIA IS

ADMINISTERED FIVE MINUTES BEFORE PHYSOSTIGMA.

In this series of experiments, physostigma was administered in the form of
extract. It has already been shown that the minimum lethal dose of this
preparation is 1'2 grain per three pounds weight of rabbit.

Experiments with the Minimum-Lethal Dose of Physostigma.—tin accordance
with the plan which has been indicated, the first experiments of the series were
made to determine what doses of atropia can prevent the fatal action of the
minimum-lethal dose of physostigma. In the experiments performed for this
purpose,* the following results were obtained :— -

ExpermMent 71.+— With 0-005 srain of sulphate of atropia, death occurred.

EXPERIMENT 72-a. ,, 0°009 ms i recovery ,,
EXPERIMENT @73-a. ,, 0°015 : a recovery ,,
EXPERIMENT 74-a. ,, 0°02 an - recovery ,,
EXPERIMENT 73-a. ,, 0°025 3 F recovery ,,
EXPERIMENT 76-a. ,, 0°031 a 2 recovery ,,
EXPERIMENT @7-a. ,, 0°05 ne og recovery ,,
EXPERIMENT 41-a.t ,, 0°3 Fe recovery ,,
EXPERIMENT 78-a. ,, 0°92 a recovery ,,
EXPERIMENT 79-a, ,, 2° grains 2 recovery ,,
EXPERIMENT 80-a. ,, 3° , he recovery ,,
EXPERIMENT 81-a. ,, 4:3 sf Ps recovery ,,
EXPERIMENT 82-a. ,, 5° 4 " recovery ,,
EXPERIMENT 83-a, ,, 5:2 it 24 recovery ,,
EXPERIMENT 84, oe 3 » death P
EXPERIMENT 88, es: i ;. death _,,
EXPERIMENT 86. 5 De £ 3 death a
EXPERIMENT 87. ean 3) oN re death 5

‘a Tabular Summary, Series 1, Table 2.
+ Except in two cases, the numbers of these experiments have reference to the arrangement that —
has been followed in the Tabular Summary at the end of the paper, where the leading facts connected
with-all the experiments belonging to this Section ofthe research are mentioned. ;
‘t A full description of the éxperiment has already been given in Section A. (see p 544).

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 585

It is shown by this brief statement of the experiments with the minimum-
lethal dose of physostigma, that while one two-hundredth of a grain of sulphate
of atropia is a dose insufficient to prevent death, nine one-thousandths of a
grain is one sufficiently large to do so; that any dose of sulphate of atropia
ranging within the wide limits extending from the nine one-thousandths of a
grain to five grains and one fifth is able to prevent the fatal effect of this
_ dose of physostigma; and that if the dose of sulphate of atropia amount to
five grains and three-tenths, the region of successful antagonism is left, and
death occurrs.

Experiments with One and a half times the Minimum-Lethal Dose of Physo-
_ stigma.—In the next place, experiments were made to determine what doses of
atropia are able successfully to counteract a dose of physostigma one and a half
times as large as the minimum-lethal.* The following results were obtained :-—

EXPERIMENT. 88.— With 0-014 grain of sulphate of atropia, death occurred.

EXPERIMENT 89. » 0°015 a e death a
EXPERIMENT 90-a. ,, 0°02 a 3 recovery ,,
EXPERIMENT 91. 0302, rf . death ‘
EXPERIMENT 92-a. ,, 0°02 f. Ht recovery ,,
EXPERIMENT 93-a. ,, 0°03 - ere 55 recovery ,,
EXPERIMENT 94-a. ,, 0°05 . ,, 3; recovery ,,
EXPERIMENT 95-a. ,, 0°05 ‘5 Ms | SVECOVeLY.
EXPERIMENT 96-a. ,, 1°5 * is recovery ,,
EXPERIMENT 97-a. ,, 2° grains _ recovery ,,
EXPERIMENT 98-a. ,, 2°6 . P recovery ,,
EXPERIMENT 99-a. ,, 3°3 i sail, recovery ,,
EXPERIMENT 100-a. ,, 4:1 8 a recovery _,,
EXPERIMEN? 101. mon 2 ms 4 death ue

From these experiments, it appears that while three two-hundredths of a
grain of sulphate of atropia is a dose too small to prevent the occurrence of

In the presence of various causes of fallacy, which cannot alto-
gether be obviated, it is not surprising that results of an exceptional character
should occasionally be obtained. The occurrence of death in Experiment
| 91, where the dose of sulphate of atropia is one-fiftieth of a grain, may legiti-

* Tabular Summary, Series 1, Table 3.

584 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

mately be placed among these exceptional results. Several others will after-
wards be noted.

Experiments with Twice the Minimum-Lethal Dose of Physostigma.—When
atropia was administered five minutes before a dose of physostigma twice as

EXPERIMENT 102.— With 0:19 grain of sulphate of atropia, death occurred.

EXPERIMENT 108. » 0°02 ~ Li death Me
EXPERIMENT 104. ». 0°02 i He death is
EXPERIMENT 105. pe a2 A 5 death "
EXPERIMENT 106-a. ,, 0°025 pa Pa recovery ,,
EXPERIMENT 107, » 90:03 - " death ;
EXPERIMENT 108-a. ,, 0°04 a 2 recovery ,,
EXPERIMENT 109-a. ,, 0°05 % * recovery;
EXPERIMENT 110-a. ,, 0°3 3 recovery ,,
EXPERIMENT lll-a. _,,_ 1° c - recovery,
EXPERIMENT 112-a, ,, 2° grains - recovery ,,
EXPERIMENT 118-a. ,, 2°3 - a! recovery ,,
EXPERIMENT 114-a. ,, 3° a Behn recovery ,,
EXPERIMENT 115-2. ,, 3°2 se e recovery ,,
EXPERIMENT 116. ee Pe is death 5
EXPERIMENT 117. + ebeane. a ee death

One-fiftieth of a grain of sulphate of atropia is therefore too small a _
quantity to prevent death from following the administration of a dose of
physostigma twice as large as the minimum-lethal, but one-fortieth of a grain —
is sufficient to do so; and doses ranging from one-fortieth of a grain to three —
grains and a fifth can successfully antagonise this dose of physostigma. —
When, however, a dose larger than three grains and a fifth is administered, f
death occurs. Experiment 107 is another instance of an exceptional result
being produced. By referring to the description of this experiment in the —
Tabular Summary, it will be seen that soon after physostigma had been
administered, the rabbit became excited and_ran about and struggled ener-
getically. Such movements and struggles appear greatly to favour the toxic
‘action of physostigma ; and it has already been pointed out that when thea
occur, the minimum-lethal dose of physostigma is appreciably lessened.

Experiments with Two and a half times the Minimum-Lethal Dose of Phy- q
sostigma.— When two and a half times the minimum-lethal dose of physostigma —
was administered, the following results were obtained :—t :

* Tabular Summary, Series 1, Table 4. + Ibid., Table 5.

THE ACTIONS OF PHYSOSTIGMA AND ATROPTA. 585

EXPERIMENT 118,—With 0-011 grain of sulphate of atropia, death occurred.
EXPERIMENT 119. » 0°0187 i a death i
-__s-EXperrMenr 120. » 0°025 . P death: ©
EXPERIMENT 121-a. ,, 0:025 ss af recovery ,,
EXPERIMENT 122-a. ,, 0-027 x ‘ recovery ,,
EXPERIMENT 128-a.. ,, 0-028 mi 2 recovery __,,
EXPERIMENT 124-a. ,, 0-034 = f recovery ,,
EXPERIMENT 125-a. ,, 0-0375 EF i. recovery _,,

EXPERIMENT 126-a. ,, 0:05 ss $5 recovery

EXPERIMENT 127-a. ,, 0-088 a s recovery ,,
EXPERIMENT 128-a. ,, 0:43 ma A recovery _,,
EXPERIMENT 129-a, ,, 0:44 a me recovery ,,
EXPERIMENT 130-a. , 1- 2 a recovery ,,
EXPERIMENT 13l-a, , 1-2 sa recovery ,,
EXPERIMENT 132-a. | 1:25 Bs 3 recovery ,,
EXPERIMENT 13838-a, ,, 1:63 * 3 recovery _,,
EXPERIMENT 184-a, ,, 2: “3 : recovery ,,
EXPERIMENT 135. ,, 2° ‘ bs death i
EXPERIMENT 136-a. , 2:2 ie ‘ recovery ,,
EXPERIMENT 187, ,,_- 23 i: x death a
EXPERIMENT 138. ,, 2°6 i - death fi
EXPERIMENT 1389. pa22:66 3 x death s

The smallest dose of sulphate of atropia that can prevent the occurrence of
death after the administration of two and a half times the minimum-lethal dose
gi ee yeostigma is thus seen to be about one-fortieth of a grain; and it is

] iil ewise show that doses of aie of atropia ranging fort one-fortieth of a
ain to two grains and a fifth are able to prevent the fatal action of two and
a half times the minimum-lethal dose of physostigma; and that death occurs
if the dose of atropia amount to two grains and three-tenths. In Experiment

te look upon this result as exceptional; for in the previous experiment the
7 pene dose was followed by recovery, and in the subsequent eoeanr the

7 by which the general vigour was depreciated.

t Experiments with Three times the Minimum-Lethal Dose of Physostigma.—
| When atropia was administered five minutes before a dose of physostigma
--VOL. XXVI. PART IIL. te 7 N

586 DR. THOMAS R. FRASER ON THE ANGTAGONISM BETWEEN

three times as large as the minimum-lethal, the following results were
obtained :— *

EXPERIMENT 140.—With 0-043 grain of sulphate of atropia, death occurred.

EXPERIMENT 141. » 0:05 ie a death a
EXPERIMENT 142-a. ,, 0°06 8 Dh recovery .,,
EXPERIMENT 143-a. ,, 0:076 e ” recovery ,, 3
EXPERIMENT 144-a. , 0:088 2 a recovery ,, :
EXPERIMENT 42-a.t ,, 0°16 % mi recovery, \,,
EXPERIMENT 145-a, ,, 0°5 es z FECOVEEY,... »,
EXPERIMENT 146-a,_ ,, 1° %. & recovery _,,
EXPERIMENT 147-a. ,, 1:2 5s a recovery _,,
EXPERIMENT 148. + de - Hs death “3
EXPERIMENT 149. * alee be . death if
EXPERIMENT 190, Baas Fe) - ee death #

These experiments show that while one-twentieth of a grain of sulphate of
atropia is insufficient to prevent the occurrence of death after the administra-
tion of three times the minimum-lethal dose of physostigma, three-fiftieths of a
grain is sufficient to do so. They likewise show that the lethal action of this —
dose of physostigma may be prevented by any dose of sulphate of atropia —
from three-fiftieths of a grain to one grain and a fifth; but that if the latter —
dose be exceeded, the region of successful antagonism is left and death occurs. —

Experiments with Three and a half times the Minimum-Lethal Dose of Physo- —
stigma.—When the dose of physostigma was three and a half times the minimum _
lethal, the following results were obtained :—t

EXPERIMENT 161.— With 0-044 grain of sulphate of atropia, death occurred.

- ExperIMEenT 152. ,, 0°071 ‘ * death ¥
EXPERIMENT 153.8 ,, 0-1 vi 9 recovery ,, .
EXPERIMENT 154-a. ,, 0-2 . oA recovery ,, zp
EXPERIMENT 158. on O's ‘. % death i a
EXPERIMENT 156. sje AND *y 5 death ar y
EXPERIMENT 157. Per | . we death i“

Accordingly, when the dose of physostigma is so large as three and a half
times the minimum-lethal, the range of doses of atropia that can prevent death —
is a very limited one, extending only from one-tenth to one-fifth of a grain.

* Tabular Summary, Series 1, Table 6.

+ A full description of this experiment has already been given in Section A. (see p. 546).

t Tabular Summary, Series 1, Table 7.

§ Five days after this experiment, the rabbit became weak and languid ; from that time it~
gradually lost weight and condition ; and on the twentieth day, it died. The usual experiment with
a dose of physostigma alone could not, therefore, be made.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 587

It has occasionally happened, especially when the subject of the experiment
was a young animal, that the atropia effects were unusually slight. If the
description in the Tabular Summary of Experiment 152 be compared with that
of other experiments in which the same relative dose of atropia was ad-
ministered, it will be observed that the action of atropia was not developed

_ with its usual prominence, and that, consequently, the physostigma action was
; only feebly counteracted.
Experiments with Four times the Minimum-Lethal Dose of Physostigma.—
When atropia was administered five minutes before a dose of physostigma
equivalent to four times the minimum-lethal dose, the following results were
- obtained :—*

EXPERIMENT 158.—With 0:1 grain of sulphate of atropia, death occurred.
EXPERIMENT 189. - Oa - oF death i
EXPERIMENT 160. ay We. Fe death f
EXPERIMENT 161. oe will ee wr death 7.
EXPERIMENT 162. ide Wee res ~ death 4

It was unneeessary to proceed further with these experiments, as it had been
‘rendered obvious by those previously made, that the lethal action of this large
dose of physostigma cannot be prevented by atropia, if a dose of this substance
between one-tenth and one-fifth of a grain be unable to do so.

It has, accordingly, been shown that the maximum dose of physostigma
which can be rendered non-fatal by atropia administered five minutes pre-
viously is three and a half times the minimum-lethal dose. It has also been
shown that the range of doses of atropia capable of preventing death after the
administration of lethal doses of physostigma diminishes as the ee of
physostigma increases.

_ When these experiments are represented in a diagrammatic form, the results
that have been obtained may be clearly and readily appreciated. A simple
plan on which to construct a diagram is obviously suggested by the arrange-
ment that has been followed in the description of the experiments. By placing
symbolic representations of the results of the experiments performed with
ach lethal dose of physostigma in horizontal lines, and by arranging these
lines so that the doses of physostigma shall succeed each other in regular
_ sequence and at proper intervals, we may obtain a picture which graphically
! _ represents the various results that have been mentioned.

| , Diagram 1 (Plate XXIII.) illustrates the experiments described above, in
| which atropia was administered five minutes before lethal doses of physostigma.
| The experiments that terminated in death are represented by crosses, and those
1a

* Tabular Summary, Series 1, Table 8.

588 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

that terminated in recovery by dots, while the position assigned to each experi-
ment is determined by the doses of physostigma and atropia which were ad-
ministered. The doses of atropia are represented by the distance, in a
horizontal direction, from the perpendicular line forming the left margin of the
diagram, and increase at the rate of one-tenth of a grain for every two sub-
divisions of the horizontal. lines. The doses of physostigma increase from
below upwards,—the minimum-lethal dose being represented by the red hori-
zontal line, a dose one and a half times as large as the minimum-lethal by the
black horizontal line immediately above the red line, a dose twice as large as
the minimum-lethal by the second black horizontal line, and so on until a line
is reached at the top of the diagram, which represents a dose of physostigma
four times as large as the minimum-lethal. The curved line, abc, separates
the fatal experiments (crosses) from those which terminated in recovery (dots) ;
and, accordingly, the blue space on the one side of it represents a region in
which death always occurs, and the pink space on the other side a region in
which recovery occurs. The doses that were given in any experiment within
each of these regions are readily ascertained from the position of the experi-
ment; the dose of physostigma being determined by the horizontal line on
which the symbolic representation of the experiment is placed, and that of
atropia by the exact spot in the horizontal line which is occupied by the repre-
sentation.

With these explanations, the results of the experiments will be rendered
apparent by a mere glance at the diagram. It may again be pointed out that
the more obvious of these results are, that the maximum dose of physostigma
which can be rendered non-lethal by atropia administered five minutes pre-
viously is about three and a half times the minimum-lethal dose, and that the —
range of the doses of atropia which are able to render non-fatal various other- —
wise fatal doses of physostigma, diminishes as the dose of physostigma increases.
The general nature of these results is well illustrated in the diagram by the
triangular form of the pink region of recovery after lethal doses of physostigma
(abc), of which the apex indicates the maximum antagonisable dose of physo-
stigma, and the gradual increase in breadth from the apex to the red horizontal
line, the gradual increase in the range of doses of atropia that can prevent the
lethal effect of doses of physostigma diminishing from three and a half times”
the minimum-lethal to the minimum-lethal.

In this diagram, the pink region, and the curved line, adc, have been ex-
tended below the red line representing the minimum-lethal dose of physostigma,
and therefore into a space where the doses of physostigma are too small of —
themselves to cause death. The lateral extension of the diagram, however, is
insufficient to exhibit the chief interest of this space ; but it will be pointed out
in the description of the next and only remaining group of experiments con-

nected with the present series. |

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. . 589

Experiments with half the Minimum-Lethal Dose of Physostigma.—The con-
siderations which led me to anticipate that the counteracting influence of atropia
upon the lethal action of physostigma is successfully exerted only within a
definite range of doses, and that death may be produced when .a lethal dose
of physostigma, which is capable of being rendered non-lethal by atropia, is
given in combination with a somewhat large non-lethal dose of atropia, also
led me to anticipate that death may be produced by the combined adminis-
tration of non-lethal doses of the two substances. Experiments were accord-
ingly made,* in which half the minimum-lethal dose of physostigma was adminis-
tered five minutes after various doses of atropia with the following results :—

EXPERIMENT 58.—With 5°3 grains of sulphate of atropia, recovery occurred.
EXPERIMENT 39.

» 616 xs me recovery ms
EXPERIMENT 60. rea oy 59 of recovery s
EXPERIMENT 61. yo O's 3 we recovery ‘%
EXPERIMENT 62. ae Oro a bs recovery ie
EXPERIMENT 68. aan: ‘a 5 recovery i
EXPERIMENT 64. 4) oe . MS recovery oe
EXPERIMENT 685. 5 (S'S b: es recovery a
EXPERIMENT 66. Oto) “ ay recovery .
EXPERIMENT 67. pala 5) oe es recovery a
EXPERIMENT 68. 98 23 death Fr
EXPERIMENT 69. sr POs x Rs death 3
EXPERIMENT 70. ,, 10°5 “a & death ‘5

It is shown by these experiments that when sulphate of atropia is adminis-
tered five minutes before half the minimum-lethal dose of physostigma, death
occurs if the dose of the former substance be nine grains and four-fifths, or more.
This result appears a very remarkable one, when it is considered that a very
decided counteraction is exerted by much smaller doses of atropia against the
poisonous action of doses of physostigma greatly in excess of the minimum-
lethal, and that the minimum-lethal dose of sulphate of atropia is about twenty-
one grains. It, however, merely confirms a legitimate anticipation when certain
of the results of the experiments with lethal doses of physostigma are borne in
mind. Actions essentially the same as those that are produced in excessive
amount when the administration of five grains and three-tenths of sulphate of
atropia along with the minimum-lethal dose of physostigma is followed by death,
also become excessive after the administration of nine grains and four-fifths
of sulphate of atropia along with half the minimum-lethal dose of physostigma.
The result may be simply explained by supposing some action or actions of
both physostigma and atropia wherein there is no mutual counteraction.

* Tabular Summary, Series i. Table 1.
VOL. XXVI. PART III. 70

590 DR. THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

The effects that are produced by a combination of half the minimum-lethal
dose of physostigma with sufficiently varied doses of atropia being thus
determined, the entire regions of recovery and of death in the series of experi-
ments in which atropia was administered five minutes before physostigma may
now be considered.

This series is completely represented in Diagram 5 (Plate XXIV.), which
has been constructed on the same plan as Diagram 1, from which, however, it
differs in so far that the increase in the doses of atropia, represented by the
distance in a horizontal direction from the perpendicular line forming the left
margin of the diagram, proceeds at the rate of one-tenth of a grain for every
subdivision of the horizontal lines, in place of for every two subdivisions. This
modification was required to curtail the lateral extension of the diagram, so as
to retain it within convenient limits.

The area covered by the diagram includes every possible dose of physo-
stigma from the minutest fraction of the minimum-lethal dose to one four
times as large as the minimum-lethal, and every possible dose of atropia below
the minimum-lethal. In the previous section of this paper, a series of experi-
ments was described which rendered it probable that the minimum-lethal dose of
atropia for rabbits is about twenty-one grains for every three pounds weight
of animal; and I have specially indicated the position of this dose by a red
perpendicular line, which will be seen near the right margin of the diagram.

In the diagram; the region of recovery (pink) appears to be a very
restricted one when contrasted with the region of death (blue); and it is
almost unnecessary to point out that this relation may be greatly exaggerated
by enlarging the diagram so as to include within it a portion of the almost
unlimited area in which death occurs after combinations of physostigma and
atropia unrepresented in the present diagram. As the region of recovery after
lethal doses of physostigma occupies only that portion of the pmk space which
extends above the red horizontal line, the area that it occupies appears almost
insignificantly small. Seeing, however, that a dose of physostigma three and
a half times as large as the minimum-lethal is included within this region, its
insignificance in relation to the entire region of death becomes of but little
importance, when the interesting fact of the counteraction of so enormous a
dose is realised.

The diagram very clearly displays the singular result, that death may follow
the administration of physostigma and atropia in doses both below the mini-
mum-lethal. The combinations that are able to produce this result are included
within the blue space below the red horizontal line. The direction of the
line separating this space from the subjacent area of recovery (pink) is
much more horizontal than that of the line separating the region of death from
that of recovery after lethal doses of physostigma. The change of direction

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 591

occurs somewhat abruptly at the intersection of the red horizontal line, repre-
senting the minimum-lethal dose of physostigma; and it very graphically
represents some of the results to which attention has been drawn in the
description of the experiments. It was shown by these experiments, that when
physostigma is administered in lethal doses, the range of the doses of atropia that
are able to produce successful counteraction increases by about one grain for
each successive decrease by half the minimum-lethal dose in the dose of physo-
stigma. When, however, physostigma is administered in a dose equivalent only
to half the minimum-lethal, the range of doses of atropia that may be given with-
out the occurrence of death is increased beyond the range for the minimum-lethal
dose, not by one grain only, but by four grains and three-tenths. Accordingly,
while in the region where the doses of physostigma are lethal, the line separating
the area of death from that of recovery possesses a direction which indicates
an increase of one grain of sulphate of atropia for each decrease by half the
minimum-lethal dose in the quantity of physostigma, in the region where the
dose of physostigma is less than lethal, it possesses a direction which indicates
an increase of about four grains and a half of sulphate of atropia for each
decrease by half the minimum-lethal dose in the quantity of physostigma.

Il. DETERMINATION OF THE LIMITS OF ANTAGONISM WHEN ATROPIA IS
ADMINISTERED FIVE MINUTES AFTER PHYSOSTIGMA.

The second series of experiments was undertaken to determine the limits of
‘successful antagonism when atropia is administered five minutes after physo-
stigma. In the experiments of this series, physostigma was administered in
the form of sulphate of the active principle; of which preparation it has already
been shown that the minimum-lethal dose is about 0°12 grain per three pounds
weight of rabbit. As this dose is the one-tenth of the minimum-lethal dose of
extract of physostigma, the experiments of the first series may readily be com-
pared with those of the present.

Experiments with the Minimum-Lethal Dose of Physostigma.—When the
minimum-lethal dose of sulphate of physostigmia was administered five minutes
before various doses of sulphate of atropia, the results were as follows :—*

EXPERIMENT 168.—With 0-01 grain of sulphate of atropia, death occurred.

EXPERIMENT 169. » O015 oe . death i"
EXPERIMENT 170-a. ,, 0:02 Ps Fe recovery _,,
EXPERIMENT 171l-a. ,, 0°05 ¥, re recovery __,,
EXPERIMENT 172-a. ,, O°1 ie a recovery _,,
EXPERIMENT 173-a. ,, 0°2 + ” recovery .,,

* Tabular Summary, Series ii. Table 2.

592 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

EXPERIMENT 174-a.— With 0°5 grain of sulphate of atropia recovery occurred.
EXPERIMENT 175-a. winds ra recovery

EXPERIMENT 176-a. pitp2 Stalls! - 5 v recovery _,,
EXPERIMENT 177-a. so 53 ” recovery _,,
EXPERIMENT 178-a. guZD ) a recovery _,,
EXPERIMENT 179. yy 6 “ - death £
EXPERIMENT 180. Lee :, : death =.
EXPERIMENT 181. errn ne 56 Fs death

EXPERIMENT 182. warelie P ¥ death ts
EXPERIMENT 1838. ee 4 35 a death Z
EXPERIMENT 184. hae - i death

Accordingly, if sulphate of atropia be administered five minutes after the
minimum-lethal dose of physostigma, death occurs when the dose of sulphate
of atropia is not more than three two-hundredths of a grain, recovery when
the dose is from one fiftieth of a grain to two grains and a half, and death again
when the dose is larger than two grains and a half. The range of the doses of
atropia that can prevent the lethal effect of this quantity of physostigma is,
therefore, considerably less when physostigma is administered five minutes
before atropia, than when it is administered five minutes after it; and it will
be observed that there is a like difference between the results in all the other
corresponding groups of experiments in the two series.

Experiments with One and a half times the Minimum-Lethal Dose of Physo-
stigma.—In the next instance, the limits of successful antagonism were deter-
mined in the case of a dose of physostigma one and a half times as large as the
minimum-lethal :—*

EXPERIMENT 185.—With 0:03 grain of sulphate of atropia, death occurred.

EXPERIMENT 186-a. ,, 0°05 + Rs recovery _,,
EXPERIMENT 187-a. ,, 0:1 Pp 53 recovery _,,
EXPERIMENT 188-a. ,, 02 5 5 recovelyyy
EXPERIMENT 189-a. ,, 0°3 5 - recovery ,,
EXPERIMENT 190-a. ,, 0-4 Pe rr recovery _,,
EXPERIMENT 191-a. ,, 0°5 a x recovery
EXPERIMENT 192-a. ,, 0°7 gs > recovery sa
EXPERIMENT 198-a. ,, 1:2 = is reCOVery ae
EXPERIMENT 194-a. ,, 1°5 * recovery aa
EXPERIMENT 195-a. ,, 2:0 grains ,, _ recovery
EXPERIMENT 196-a. ,, 2:1 . 2s recovery as
* Tabular Summary, Series ii. Table 3. a

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 598

EXPERIMENT 197-a.— With2‘1 grains of sulphate of atropia,recovery occurred.
EXPERIMENT 198-a. §) 2D - - MECOVErY 5;
EXPERIMENT 199. Pe ye ie A death 43
EXPERIMENT 200. Hy 02S a rs death

EXPERIMENT 201. ap) i s death |
EXPERIMENT 202. ss Dre. a £ death Ki
EXPERIMNNT 203. a5) s 5 death 5
EXPERIMENT 204. 30 i o death 3

It is shown by these experiments, that when sulphate of atropia is adminis-
tered five minutes after this dose of physostigma, successful antagonism is
produced by any dose of sulphate of atropia ranging from one twentieth of a
grain to two grains and one tenth. In Experiment 198-a., successful antag-
onism likewise followed the administration of two grains and one fifth of
sulphate of atropia ; but as this result was not obtained in the next experi-
ment, in which the same dose was given, I have not included it within the
limits of success.

Eaperiments with Twice the Minimum-Lethal Dose of Physostigma.—The
following results were obtained when atropia was administered five minutes
after a dose of physostigma twice as large as the minimum-lethal :—*

EXPERIMENT 205.— With 0-05 grain of sulphate of atropia, death occurred.

EXPERIMENT 206. mr 0S + us death A
EXPERIMENT 207. » 0°08 ~ be death -
EXPERIMENT 208. » 0:09 Bs Hf death .
EXPERIMENT 209-a. ,, 0-1 33 3 recovery _,,
EXPERIMENT 210-a. ,, 02 * 5 recovery ,,
EXPERIMENT 2l1l-a. ,, 0°3 2 i recovery ,,
EXPERIMENT 212-a. ,, 0°4 a 4 recovery ,,
EXPERIMENT 213-a. ,, 0°5 3 ie recovery ,,
EXPERIMENT 214-a. ,, 0°5 :, recovery ,,
EXPERIMENT 215-a. ,, 0°8 es #3 LECOVELY, ae,
EXPERIMENT 216-a. ,, 0°9 5 i recovery _,,
EXPERIMENT 217-a. ,, 1:0 x as TECOveryy y.
EXPERIMENT 218-a. ,, 1:2 ss os recovery ,,
EXPERIMENT 219. th ie Ps FA death a
EXPERIMENT 220. ee a - a death i
EXPERIMENT 221, » Ls Fd . death |
EXPERIMENT 222. 2 eras |, * death 4
EXPERIMENT 228. aera . , death

* Tabular Summary, Series ii. Table 4.
VOL, XXVI. PART III. (on

594 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

These experiments prove that when sulphate of atropia is administered five
minutes subsequently to a dose of physostigma twice as large as the minimum-
lethal, nine one-hundredths of a grain of the former substance is too small a
dose to prevent death; that doses ranging from one-tenth of a grain to one
grain and a fifth are sufficient to do so; and that if the dose be larger than one
grain and a fifth, the higher limit of success is passed, and death occurs.

Experiments with Two and a half times the Minimum-Lethal Dose of Physo-
stigma.—The experiments of the next group were made with a dose of physo-
stigma two and a half times as large as the minimum-lethal.* The doses of
sulphate of atropia that were given with this dose of physostigma, and the
results of the experiments are as follows :—

EXPERIMENT 224.— With 0°05 grain of sulphate of atropia, death occurred.

EXPERIMENT 220. » 0°08 hs Me death .)
EXPERIMENT 226-a. ,, 0-1 - recovery” 3,
EXPERIMENT 227-a. ,, O15 . be recovery _,,
EXPERIMENT 228-a. ,, 02 ¥f As recovery __,,
EXPERIMENT 229-a. ,, 0°3 ‘A + recovery ,,
EXPERIMENT 2380-a. ,, 0°5 5 . recovery — ¥,
EXPERIMENT 281l-a. ,, 0°6 A sf recovery _,,
EXPERIMENT 282-a. ,, 0°7 Ae be recOVEnva
EXPERIMENT 288-a. ,, 0°8 i - recovery |
EXPERIMENT 234. af) ee) Ks death e
EXPERIMENT 295, es Way) > . death 3
EXPERIMENT 236. eels 2 s death F
EXPERIMENT 237. ea 5 - ene death 3

Accordingly, when two and a half times the minimum-lethal dose of physo-
stigma was given five minutes before sulphate of atropia, the range of doses of
the latter substance, capable of producing successful counteraction, extended
only from one tenth to about seven tenths of a grain.

Experiments with Three times the Minimum-Lethal Dose of Physostigma.—
When the dose of physostigma was three times as large as the minimum-
lethal,t the following results were obtained :—

EXPERIMENT 238.—With 0:1 grain of sulphate of atropia, death occurred.

EXPERIMENT 239. - O45 ie es death Ag

EXPERIMENT 240-a. ,, 0°16 es . recovery

EXPERIMENT 241. Prin (2 4 - death s

EXPERIMENT 242. » Os a FS death 3
* Tabular Summary, Series ii. Table 5. + Ibid., Table 6.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 595

EXPERIMENT 243.—With 0°3 grain of sulphate of atropia, death occurred.
EXPERIMENT 244. 5) 0:5 ys 5; death s,

Recovery, therefore, occurred in only one of the experiments in which atropia
was administered five minutes after a dose of physostigma three times as large
as the minimum-lethal. The administration of three twentieths and of one
fifth of a grain of sulphate of atropia resulted in death, but recovery took place
with the intermediate dose of four twenty-fifths of a grain.

Experiments with Three and a half times the Minimum-Lethal Dose of Physo-
stigma.—The results of the previous experiments having made it obvious that
the largest dose of physostigma that can be rendered non-lethal by atropia
administered five minutes subsequently, is one three times as large as the
minimum-lethal, it was evidently unnecessary to perform many experiments
with a larger dose. Accordingly, only two such experiments were made, with
a dose three and a half times as large as the minimum-lethal;* and the chief
purposes of these experiments were to complete this portion of the series, and
to ascertain the nature of the phenomena that are produced when this dose of
physostigma is given five minutes before atropia.

EXPERIMENT 248.— With 0°16 grain of sulphate of atropia, death occurred. .
EXPERIMENT 246. 3 Or? ¥ i death -

As both of these experiments terminated fatally, it was needless to continue
the series by making experiments with a larger dose of physostigma.

The result of the whole series of experiments is therefore to show that
when atropia is administered five minutes after physostigma, the largest quan-
tity of the latter substance that can be rendered non-lethal by the former is
three times the minimum-lethal dose, and that the range of the doses of atropia
that are capable of preventing the lethal action of physostigma diminishes
according as the dose of physostigma is increased.

- The results of this series of experiments are illustrated in Diagram 3 (Plate
XXIII.), which has been constructed on the same plan and scale as Diagram 1,
illustrating the first series of experiments, in order to facilitate a comparison
with it. It will be seen that the most prominent of the differences between
the two diagrams are, that the region of recovery after lethal doses of physo-
stigma (distinguished as a pink area enclosed within the curved line a b ¢, and
the segment a ¢ of the red horizontal line) is smaller both in its perpendicular
and in its horizontal extent, and that the curved line @6cis much more
irregular in the diagram of the second series (Diagram 3) than in that of the
first (Diagram 1).

The former of these differences very clearly illustrates the greater counter-

* Tabular Summary, Series ii. Table 7.

596 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

acting power of atropia when it is administered five minutes before, than when
it is administered five minutes after physostigma. It has been shown that in
the one case, three and a half times the minimum-lethal dose can be rendered
non-fatal, whereas in the other, only three times the minimum-lethal dose can
be successfully counteracted; and not only does this difference exist, but the
range of the doses of atropia that can prevent the lethal action of any given
dose of physostigma is also greater in the former case than in the latter.

The greater irregularity of the curved line abc in Diagram 3 than in
Diagram 1 renders very manifest certain other of the results, which also, it is
true, have already been mentioned in the description of the experiments, but
which cannot be so well appreciated from a mere verbal description as from
such a graphic representation as is afforded by the diagrams. It will be re-
membered that this lme separates the experiments that terminated in death
from those that terminated in recovery. For convenience of description, it may
be regarded as consisting of two portions, a 6 and 6b c,—the former separating
the experiments that terminated in recovery after the smallest successfully
counteracting doses of atropia from those that terminated in death after still
smaller doses of atropia, and the latter separating the experiments that ter-
minated in recovery after the /argest successfully counteracting doses of atropia
from those that terminated in death after still larger doses of atropia. In con-
nection with each of these portions of the line a 6 ¢, there are several points
to which attention may be directed. ;

In Diagram 1, the portion d ¢ is a straight line, because, when physostigma
is administered five minutes after atropia, the largest doses of atropia that
can produce successful counteraction differ from each other by one grain for
each difference by half the minimum-lethal dose in the quantity of physostigma.
In Diagram 8, however, 6c is a curved line, because when physostigma is
administered five minutes before atropia, the largest successfully counteracting
doses of atropia do not diminish regularly as the doses of physostigma are re-
gularly increased.

The greater irregularity of the curved line @0c¢ in Diagram 3 than in
Diagram 1 is apparent also in the portion a 6; and it will be seen that this
portion has a less perpendicular direction, as well as a less straight course, in
the former than in the latter diagram.

In both diagrams, the steep rise of @ 6 contrasts in a marked manner
with the gradual descent of 6c. This contrast brings into prominent relief
those already mentioned results that show the smallest of the various
doses of atropia capable of successfully counteracting different doses of
physostigma to differ from each other much less than the largest. It has been
ascertained by the first series of experiments, that when atropia is adminis-
tered five minutes before physostigma, the difference between the smallest

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 597

dose of atropia capable of preventing death after the minimum-lethal dose of
physostigma and the smallest capable of doing so after a dose three and a half
times as large is only nine one-hundredths of a grain, whereas the difference
between the largest doses is so great as five grains. When atropia is given
five minutes after physostigma, the difference between the smallest dose capable
of preventing death after the minimum-lethal dose of physostigma and after a
dose three times as large is only thirteen one-hundredths of a grain, whereas
the difference between the largest doses is so great as two grains and nine
twentieths.

In order more clearly to display the differences between a } in the two series
of experiments, I have drawn other two diagrams, in which the irregularities of
this line are more distinctly shown than in Diagrams 1 and 3. This has been
effected by simply diminishing the value of the subdivisions of the horizontal
lines, so that each tenth of a grain of sulphate of atropia is indicated by twenty
subdivisions in place of by two. By this modification, the direction of the line
a b has been rendered less perpendicular, and at the same time its course has
been more accurately defined. Diagram 2 represents the experiments of the
first series, and Diagram 4 those of the second, and only so much of each series
has been included as is required to exhibit the course of a 6. It will be seen
that in Diagram 4 the line @ 6 is more irregular in its course than in Diagram 2,
and that in Diagram 2 a number of irregularities are displayed in this line, which
are not apparent in Diagram 1, where the same series of experiments is repre-
sented in a more contracted form.

It will likewise be seen from an inspection of the diagrams illustrative of
these two series of experiments, that in rabbits a dose of sulphate of atropia
equivalent to four twenty-fifths of a grain per three pounds weight of animal is
able to prevent the fatal effect of any quantity of physostigma which can be
rendered non-fatal by atropia, and that even a very slight modification of this
dose suffices to curtail the extent of successful antagonism. There can be
little doubt that in every species of animal some dose of atropia occupies a
similarly important position, and bears a similar relation to the range of suc-
cessfully counteracting doses. A result of some practical value has probably
been obtained by the establishment of the fact that this dose is much nearer
the minimum than the maximum in the range of the doses of atropia capable
of preventing the lethal effect of physostigma.

Experiments with half the minimum-lethal dose of physostigma.—The next
experiments were made in order to determine the smallest dose of atropia that
in conjunction with half the minimum-lethal dose of physostigma administered
five minutes before it is sufficient for the production of death.* The following
results were obtained :—

* Tabular Summary, Series ii. Table 1.
VOL. XXVI. PART III. 7Q

598 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

EXPERIMENT 163.—With 5: grains of sulphate of atropia, recovery occurred.

EXPERIMENT 164." ,, 7° yt . recovery rE
EXPERIMENT 165. LD ey Fe recovery A
EXPERIMENT 166. it geh Mi 35 death Ke
EXPERIMENT 167. ornesk6 - 6 death A

The smallest quantity of atropia that in conjunction with half the minimum-
lethal dose of physostigma administered five minutes before it is sufficient for
the production of death is thus shown to be about eight grains per three pounds
weight of rabbit. In the analogous experiments of the first series a similar result
was obtained, for although there death did not occur unless the dose of atropia
was at least nine grains and four fifths, it is probable that this comparatively
slight difference may be due to the physostigma having been given in that series
in the form of extract. It will be remembered that the minimum-lethal dose
of sulphate of physostigmia is somewhat less than one-tenth of that of extract
of physostigma. For convenience of comparison, however, it has been assumed
in the second series of experiments, that sulphate of physostigmia is exactly ten
times as active as extract of physostigma.

With these experiments, the second series is completed. I have not con-
sidered it necessary to construct a diagram of the entire series, as all its special
characters are displayed in the diagrams representing the experiments in which
the doses of physostigma were lethal. With less than lethal doses, the results
are so similar to those of the first series of experiments that the region of
recovery would be of essentially the same form as that represented in |
Diagram 5.

III. DETERMINATION OF THE INFLUENCE OF THE INTERVAL OF TIME BETWEEN THE
ADMINISTRATION OF THE TWO SUBSTANCES, UPON THE DosE OF ATROPIA
REQUIRED TO COUNTERACT A GIVEN DOSE OF PHYSOSTIGMA.

In the two series of experiments that have already been described, the two
following series of dose-limits of successful antagonism have been determined,
namely, those limits where the atropia is given five minutes before, and those
where it is given five minutes after the physostigma. Further, it has been shown
that the limits in the one series differ somewhat from those in the other; and
when this result is taken in connection with several obvious considerations, it
is evident that the series of dose-limits of successful antagonism will be different
for every different time-relation in the administration of the two substances.
It is, however, evident that to make for each of any considerable number of
other time-relations in the administration of the two substances, a complete
series of experiments on the plan of the two series already described, would
entail an amount of labour quite out of keeping with the value of any resilag

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 599

that might fairly be looked for. The most interesting of such results would be

the determination for each case of given doses of the two substances compatible
with the production of successful antagonism of the maximum period separat-
ing the administration of the one substance from that of the other, both when
the atropia is administered before, and when it is administered after the physo-
stigma.

In the experiments of the present series (8d), I have contented myself with
determining this period in the case of one constant dose of physostigma with
doses of atropia ranging from the one-hundredth of a grain to five grains.
When the results derived from this series of experiments are considered along
with those of the first and second series, an indication will, I believe, be obtained
of the limits in the period separating the administration of the two substances
within which successful antagonism may occur, even for the cases where the
combination of doses of physostigma and atropia is different from any combina-
tion included in the present series.

The dose of physostigma I have selected for these experiments is one
equivalent to one and a half times the minimum-lethal dose; and it was
administered in the form of sulphate of the active principle, of which prepara-
tion this dose is represented by three twenty-fifths of a grain per three pounds
weight of rabbit. With each dose of atropia that was given in combination
with this dose of physostigma, several experiments were made, which differed
from each other by a difference in the interval of time separating the adminis-
tration of the two substances; this interval being in the first experiments
such as to permit of successful antagonism, and being in each subsequent ex-
periment altered until at length it became such as no longer to permit of suc-
cessful antagonism. ‘This, at least, was the general plan followed in this series,
but it was somewhat departed from on several occasions, when the circum-
stances of the experiments prevented or rendered inconvenient its adoption.
Briefly stated, the distinguishing characters of the series were that the dose of
physostigma was constant, while the dose of atropia and the interval of time
between the administration of the two substances varied.

In certain of the experiments atropia was administered before physostigma,
and in others physostigma before atropia; and in order to connect together
these two groups of experiments, a third group was undertaken in which atropia
and physostigma were administered as nearly simultaneously as possible. In
describing the chief results, I shall, as a matter of convenience, in the first place
consider (a) the experiments in which the two substances were simultane-
ously administered; then (4) the experiments in which atropia was administered
after physostigma; and finally (c), the experiments in which atropia was
administered before physostigma.

(2) Experiments in which Atropia and Physostigma were administered

600 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

Fas

simultaneously.—In the experiments of this group,* the administration of the
two substances was effected in exactly the same manner as in experiments
44-a@ and 45-a, Section A (pp. 553 and 555). Before mentioning the results
that were obtained, it is proper to point out that a brief interval necessarily
elapsed between the administration of the two substances. This interval,
however, was only one of a few seconds, and, practically, was of little moment,
especially as uniformity was obtained in all the experiments by care being taken
always to inject the dose of atropia before that of physcstigma. When, with
this precaution, a dose of physostigma one and a half times as large as the
minimum-lethal (0:12 grain of sulphate of physostigmia per three pounds weight
of rabbit), was administrated nearly at the same moment with various doses of
sulphate of atropia, the following results were obtained :—

EXPERIMENT 247.— With 0-02 grain of sulphate of atropia, death occurred.

EXPERIMENT 248, 5, 90:02 4s H death a
EXPERIMENT 249-a. ,, 0°05 i recoyery) 7);
EXPERIMENT 250-a. ,, 0°5 3 i. recovery.
EXPERIMENT 231l-a. ,, 1° a f PeCOVEEV =,
EXPERIMENT 252-a. ,, 1°5 3) z: recovery 5,
EXPERIMENT 253-a. ,, 2° grains ,, 1 recovery= ,,
EXPERIMENT 254-a. ,, 2°5 e ~ recovery ,,
EXPERIMENT 255-a, ,, 3° a 4 recoveryas,,
EXPERIMENT 256-a. ,, 3°3 S - recovery ,,
EXPERIMENT 2957. JUSS a %, death aa
EXPERIMENT 258. ens ¥ sf death a
EXPERIMENT 289. » 45 a) 3 death i
EXPERIMENT 260. ae Loy st M death 3

It is shown by these experiments that when a dose of physostigma one and
a half times as large as the minimum-lethal is administered simultaneously with
sulphate of atropia, one fiftieth of a grain of the latter substance is a dose
insufficient to prevent death, but that one twentieth of a grain is a dose
sufficiently large to do so. It is likewise shown that the fatal effect of
this dose of physostigma may be prevented when any dose of sulphate of —
atropia ranging from one twentieth of a grain to three grains and three tenths
is given simultaneously with it, and that death occurs when the dose of sulphate
of atropia is three grains and a half or greater than this.

() Experiments in which Atropia was administered after Physostigma.—A
considerable amount of interest is attached to the experiments of the next
eroup, in which the administration of physostigma preceded that of atropia.t
In briefly describing these experiments, I shall, in the first place, consider the —

* Tabular Summary, Series iii. Table 1. + Ibid. Table 2.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 601

results obtained when the administration of atropia was effected five minutes
after that of physostigma, then, those obtained when the administration of atropia
was effected ten minutes after that of physostigma, and so on until a period is
arrived at which is too prolonged to permit any dose of atropia to counteract
successfully the dose of physostigma invariably administered in this series.

The range of the doses of sulphate of atropia that are able successfully to
counteract one and a half times the minimum-lethal dose of sulphate of physo-
stigmia, when the administration of the former substance was effected jive
minutes after that of the latter, has already been ascertained by experiments
contained in the second series. It was shown by these experiments (p. 593) that
this range extends from the one-twentieth of a grain to two grains and one
tenth. Death was found to occur when the dose of sulphate of atropia was so
small as three one-hundredths of a grain, and also when it was so large as two
grains and three tenths. It will be observed that this range is smaller than
that which is obtained when the two substances are simultaneously administered,
for in the latter case it extends from the one twentieth of a grain to three
grains and three tenths.

In the experiments where the sulphate of atropia was administered ten
minutes after one and a half times the minimum-lethal dose of sulphate of
physostigmia, the following results were obtained :—

EXPERIMENT 261.— With 0°05 grain of sulphate of atropia, death occurred.

EXPERIMENT 262-a. ,, 0°3 Ph if recovery ,,
EXPERIMENT 263-a. ,, 0°5 is h recovery _,,
EXPERIMENT 264-a. ,, 1° os 33 TECOVELY 5,
EXPERIMENT 265-a. ,, 1°5 es i recovery ,,
: EXPERIMENT 266-a. ,, 2° grains ,, a eECOVELY
EXPERIMENT 267-a. ,, 2°3 . . recovery _,,
| EXPERIMENT 268-a. ,, 2°4 e bs recovery ,,
EXPERIMENT 269-a. ,, 2°5 a = recovery ,,
EXPERIMENT 270. Bes aad, 3 death .
EXPERIMENT 271. Nee be death i

From these experiments it is seen, that when sulphate of atropia is admin-
istered ten minutes after sulphate of physostigmia, any dose of the former
substance ranging from three tenths of a grain to two grains and a half, is able
to prevent the fatal effect of one and a half times the minimum-lethal dose of the
latter substance. The range is, again, a more limited one than that obtained by
simultaneous administration. It is, however, a somewhat more extended one
than that obtained where the atropia is administered five minutes after the physo-
stigma, and the greater extension is due to the maximum successfully antagonising
dose of sulphate of atropia being considerably greater when the administration
VOL. XXVI. PART Ili. 7B

602 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

of atropia succeeds that of physostigma by ten minutes, than when it succeeds
it by only five minutes. This difference is one, certainly, which I did not
anticipate. My expectation was rather that the maximum successfully
antagonising dose of sulphate of atropia would be greater when the interval
was one of five minutes, than when it was one of ten minutes. It is difficult
to account for the result that has been obtained. I cannot attribute it to any
known cause of fallacy in the circumstances of the experiments; and the expla-
nation that it is simply due to some of the causes of fallacy that are unavoidable
in such a research, seems to be opposed by its being derived, not from one or
two experiments only, but from seven, of which four belong to the interval of
five minutes, and three to that of ten minutes. Of the experiments belonging
to the former interval, death occurred in one where the dose of sulphate of
atropia was 2°3 grains, in two where it was 2°4 grains, and in one where it was
2°5 grains ; while of the experiments belonging to the latter interval, recovery
occurred in one with each of these doses. Further, of these experiments, two
differing in the interval but agreeing in the dose of sulphate of atropia (2°4
grains) were performed on the same day, on rabbits of nearly the same weight,
and as far as could be judged, of equally healthy condition, and yet, as has
already been stated, death occurred in the experiment with the former interval
(Experiment 202), and recovery in that with the latter interval (Experiment
268-a).

Still, notwithstanding these various ¢ircumstances, it may be that the result
is due to a mere accident. If, however, it be not so, its occurrence may possibly
be explained by supposing that the non-antagonised action or actions of
physostigma produce their maximum effect after a greater interval of time
from the administration than is the case with atropia. If this be assumed, it
is obvious that death will be most easily produced when the administration
of the two substances is so timed that the two maxima of effect may coincide.
These various suppositions being granted, the apparently anomalous result
of a larger dose of sulphate of atropia being within the range of successful
antagonism when the interval is one of ten minutes than when it is one of five
minutes, may be accounted for, by assuming that certain actions produced by
the two substances are not present in so great a degree of combined intensity
when atropia is given ten minutes after physostigma as when it is given only
five minutes after it.

Passing now to the experiments in which the interval of time was greater
than ten minutes, I find that only one experiment was made in which atropia
was administered fourteen minutes after physostigma (EXPERIMENT 272-a). In
this experiment, the dose of sulphate of atropia was three tenths of a grain,
and with it the fatal effect of one and a half times the minimum-lethal dose of
physostigma was successfully antagonised.

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 603

Several experiments, however, were made in which the administration of
the atropia was effected ji/teen minutes after that of the physostigma. Their
results are as follows :—

EXPERIMENT 273-a.— With 0°3 grain of sulphate of atropia, recovery occurred.

EXPERIMENT 274-a. peel ie) - . HUECONELY 5,
EXPERIMENT 275-a. aris |e 5 5 recovery ,,
EXPERIMENT 276. pete es) ap ve death
EXPERIMENT 277. a 2 Oras: 9 death i

With this interval, therefore, death is prevented from occurring by doses of
sulphate of atropia ranging from three tenths of a grain to one grain. It will
be observed that the range is a more limited one than that which was obtained
when the two substances were simultaneously administered, and also when the
interval was less than fifteen minutes. It is, however, very satisfactory to find, as
an indication of the remarkable efficacy of the antagonising influence of atropia,
that even when one and a half times the minimum-lethal dose of physostigma is
allowed to exert its toxic power without any interference for so long a period
as fifteen minutes, the administration of atropia is still able to prevent death.
The details of the experiments are of so great interest that I regret that it is
inadvisable to describe them fully,—since this could not be done without greatly
increasing the already formidable dimensions of this communication. I must
therefore content myself with referring to the abridged accounts contained in
the Tabular Summary. In all of the experiments, the animal was at the point
of death before the atropia was administered, and yet, in two or three minutes
threreafter, the gravity of the symptoms lessened with the most extraordinary
rapidity, not only in those experiments where perfect recovery was ultimately
effected, but even in those where the final result was death. On several
occasions also, an experiment that had been commenced could not be com-

_ pleted, because death occurred in less than fifteen minutes after the administra-

tion of physostigma, and, therefore, before the proper time had arrived for the
administration of atropia.

When, indeed, the interval was greater than fifteen minutes, some difficulty
was experienced in obtaining any evidence whatever of the influence that is
exerted by atropia upon the toxic effect of this dose of physostigma. It was
only after several attempts, that I succeeded in performing the two following
experiments, in which atropia was administered seventeen minutes after one and
a half times the minimum-lethal dose of physostigma.

EXPERIMENT 278.— With 0°3 grain of sulphate of atropia, death occurred.

EXPERIMENT 279. me AOD be ,, death

In both experiments, death occurred : in the one, after the administration

604 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

of three tenths of a grain of sulphate of atropia ; in the other, after the adminis-
tration of half a grain.

From the experiments of this group we learn that the fatal effect of one
and a half times the minimum-lethal dose of physostigma can be prevented by
any dose between one-twentieth of a grain and two grains and one tenth of
sulphate of atropia, if it be administered within five minutes afterwards ;
by any dose between three tenths of a grain and two grains and one tenth of
sulphate of atropia, if it be administered within ten minutes afterwards ; and by
any dose between three tenths of a grain and one grain of sulphate of atropia,
if it be administered within fifteen minutes afterwards.

(c) Experiments in which Atropia was administered before Physostigma.—
In the last group of experiments to be considered, the administration of atropia
preceded that of physostigma.* I shall, in the first place, describe those experi-
ments that were undertaken for the purpose of determining what range of
doses of atropia can successfully counteract one and a half times the minimum-
lethal dose of physostigma, when the former substance is given five minutes
before the latter. In Series i. of this section, this range has already been
determined in the case of the extract of physostigma; but it was considered
advisable also to perform with the sulphate of physostigmia a few experiments
in which this interval was observed, as it has been shown that the dose of this
substance adopted as the minimum-lethal is somewhat more powerful than the —
dose of extract of physostigma adopted as such (p. 543).

Experiments were accordingly performed, in which atropia in various doses
was administered jive minutes before one and a half times the minimum-lethal
dose of sulphate of physostigmia. The doses of atropia given in each experi-
ment, and the results obtained, were as follows :—

_ EXPERIMENT 280. — With 0°01 grain of sulphate of atropia, death occurred.

EXPERIMENT 281-a. ,, 0:02 ~ ae recovery |
EXPERIMENT 282-a. ,, 0°05 - recovery ,,
EXPERIMENT 288-a. ,, 3°5 grains ,, recovery ,,
EXPERIMENT 284-a. ,, 3°7 as recovery ,,
EXPERIMENT 2885. var 3HG ., rf death be
EXPERIMENT 286, » 40 be 7 death 3
EXPERIMENT 287, Maes Lo _ death __,,
EXPERIMENT 288, JOT eS - death __,,
EXPERIMENT 289. SB? - _ death -

It is shown by these experiments that when sulphate of atropia is adminis-
tered five minutes before one and a half times the minimum-lethal dose of
sulphate of physostigmia, any dose of the former substance ranging from one
fiftieth of a grain to three grains and seven tenths is able to produce success-

* Tabular Summary, Series iii. Table 3. ,

fon |
ar

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 605

- ful counteraction. The smallest of these doses of sulphate of atropia is exactly
the same as that which can successfully counteract one and a half times the
minimum-lethal dose of extract. The largest, however, is smaller by two fifths
of a grain than the largest dose that can successfully counteract one and a half
times the minimum-lethal dose of the extract ; and this difference, as tending
to show that the dose of sulphate of physostigmia adopted as the minimum-
lethal is somewhat more powerful than the dose of extract adopted as such,
confirms the result of the experiments by which the minimum-lethal dose of
these two preparations of physostigma was determined.

In the description of the other experiments of this group, those performed
with each of the doses of sulphate of atropia administered will be separately
considered, commencing with the smallest dose that was given. The experi-

ments made with each dose of sulphate of atropia will be briefly described in
an order proceeding from the shortest to the longest interval of time that
separated the administration of the two substances. In the account of these
experiments, the interval of time, the dose of sulphate of atropia, and the result
of the experiments, will alone be mentioned.

The smallest dose of sulphate of atropia that was given at various intervals
before one and a half times the minimum-lethal dose of sulphate of physo-
stigmia, was one twentieth of a grain (0°05 gr.) ; and with this dose the follow-
ing experiments were performed :—

EXPERIMENT 290-a.— With an interval of 10 minutes, recovery occurred.

EXPERIMENT 291-a. i 15 ‘ recovery A
EXPERIMENT 292-a. - 20 7 recovery ae
EXPERIMENT 298. - 25 . death i
EXPERIMENT 294. A 30 ‘3 death

These experiments show that the administration of one twentieth of a grain
of sulphate of atropia may precede that of one and a half times the minimum-
lethal dose of physostigma by an interval of twenty minutes, or less, and still
successful antagonism will occur. If, however, this interval be prolonged
_ beyond twenty minutes, as to twenty-five or thirty minutes, successful anta-
-gonism does not occur.

In the next experiments, the dose of sulphate of atropia was half a grain
(05 gr.) ; and the intervals that elapsed between its administration and the
subsequent administration of one and a half times the minimum-lethal dose of
physostigma, as well as the results of the experiments, were as follows :—

EXPERIMENT 295-a.— With an interval of 15 minutes, recovery occurred.
EXPERIMENT 296-a. sss D5 Y recovery
EXPERIMENT 297-a. x 30 s, recovery
VOL. XXVI. PART III. 78s

606 DR. THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

EXPERIMENT 298.—-With an interval of 35 minutes, death occurred.
EXPERIMENT 299. 3 40 i} death .

It appears, therefore, that successful antagonism occurs when half a grain
of sulphate of atropia is administered thirty minutes, or less, before one and
a half times the minimum-lethal dose of physostigma but not when the interval
is one of thirty-five minutes, or more.

The next dose of sulphate of atropia with which experiments of this kind
were made was one grain and a half (15 gr.). The intervals of time that
elapsed before the administration of physostigma and the results obtained
were the following :—

EXPERIMENT 300-a.— With an interval of 15 minutes, recovery occurred.

EXPERIMENT 801-a. F 30 » « recovery, _
EXPERIMENT 3802-a. % 40 3g TECOVERy, i
EXPERIMENT 308-a. _ 60 be recovery 2
EXPERIMENT 3804-a. = 65 » recovery 7
EXPERIMENT 905. 2 70 »» death e
EXPERIMENT 306, x 80 » death :

Accordingly, the longest interval compatible with the production of success-
ful antagonism that may elapse after the administration of one grain and a half
of sulphate of atropia, and before that of one and a half times the minimum-
lethal dose of physostigma, is one of about sixty-five minutes.

I have next to describe, in a similarly abridged manner, a number of experi-
ments in which the dose of sulphate of atropia was three grains.

EXPERIMENT 307-a.— With an interval of 40 minutes, recovery occurred.

EXPERIMENT 308-a. i 65 a recovery 3
EXPERIMENT 809-a. = 90 P recovery A:
EXPERIMENT 3810-a. i 95 » recovery -
EXPERIMENT 911. - 100 ie death ee
EXPERIMENT 312. ae 105 us death Fe

EXPERIMENT 3818. a 120 P death r

It is shown by these experiments that successful antagonism occurs when a
dose of three grains of sulphate of atropia is administered ninety-five minutes
(one hour and thirty-five minutes), or at any shorter period, before one and a
half times the minimum-lethal dose of physostigma ; but that it does not occur
when the period is prolonged to one hundred minutes, or still further.

The doses of sulphate of atropia that were given in the four sets of experi-
ments of this group that have last been described, namely, 0:05 gr.,.0°5 gr., 18
er., and 3 grs., are all included within the range of the doses of this substance
able to prevent the fatal effect of one and a half times the minimum-lethal dose of ' |

:
’

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 607

sulphate of physostigmia both when the atropia is administered five minutes
before the physostigma, and when the two substances are simultaneously
administered. The dose with which the last-mentioned experiments were
performed, namely, three grains, is, however, near the maximum limit of the

_ range in the case of simultaneous administration, and, accordingly, not far from

this limit in the case where atropia is administered five minutes before physo-
stigma.

I have in the next place to describe two sets of Soenimiont: made respec-
tively with one and the other of two doses of sulphate of atropia greater
not only than the maximum dose that produces successful antagonism when
given simultaneously with one and a half times the minimum-lethal dose of
physostigma, but also than the maximum that does so when given five minutes
before it.

The first of these doses of sulphate of atropia is four grains and a half. It
was administered before De osaane at the intervals and with the results that
will now be stated :—

_ EXPERIMENT 314,— With an interval of 10 minutes, death occurred.

EXPERIMENT 315-a. a 15 4 recovery ,,
EXPERIMENT 316-a. ss 15 . recovery ,,
EXPERIMENT 817-a. 20 Rs RECOVERY. 55

The very interesting and remarkable character of the results of these experi-
ments becomes apparent when they are considered along with those of two
experiments previously described, in which the same doses of sulphate of
atropia and sulphate of physostigmia respectively were administered. In the
first of these (Experiment 259), the administration of the two substances was
simultaneously effected, and in the second (Experiment 288), the atropia was

_ administered five minutes before the physostigma; and in both cases death

occurred. It has now been shown that when, with the same respective doses,
the atropia is given ten minutes before the physostigma, the result is stil/a
fatal one ; but that when the atropia is given fifteen or twenty minutes before
the physostigma, recovery, and not death, occurs. _

I have not made any experiments with this dose of sulphate of atropia for
the purpose of determining the maximum interval of time that may, without
hindering the production of successful antagonism, be allowed to intervene
between its administration and the subsequent administration of one and a half
times the minimum-lethal dose of sulphate of physostigmia.

Such a determination, however, was accomplished in the experiments where
the dose of sulphate of atropia was jive grains. The intervals of time separating
the administration of the two substances, and the results obtained in these
experiments, were as follows :—

608 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

EXPERIMENT 3818.—With an interval of 15 minutes, death occurred.

EXPERIMENT 3819. “s 20 » death
EXPERIMENT 320-a. i 25 oy peetovery 8
EXPERIMENT 821-a. es 30 so, reeoveryni ge
EXPERIMENT 822-a. e 65 ~ ecovery . 5,
EXPERIMENT 9828. @ 65 » death .
EXPERIMENT 3824. A 105 » death As
EXPERIMENT 8265-a. s 105 by ke OROCOVETI Ads
EXPERIMENT 826-a. " 140 »9 0 EOCOWMEFY iy,
EXPERIMENT 827-a. a; 170 ji ar ReCoveny:
EXPERIMENT 828-a. r 175 jy | | RECOVERYA ing
EXPERIMENT 829. e 180 » death P
EXPERIMENT 880. ‘3 185 » death FS
EXPERIMENT 3981. 55 200 » death 5

It appears from these experiments, that when a dose of five grains of sul-
phate of atropia is administered before one and a half times the minimum-
lethal dose of sulphate of physostigmia, death occurs if the interval of time be
one of fifteen minutes, or of twenty minutes ; that recovery generally occurs if
the interval be one included within the wide limits extending from twenty-five
minutes to one hundred and seventy-five minutes (two hours and fifty-five
minutes) ; and that death again occurs if the interval be one so great as a
hundred and eighty minutes (three hours). In connection with these results
also, it is of interest to point out that in two experiments previously described,
where the same respective doses of sulphate of atropia and sulphate of physo-
stigmia were given, death occurred both when the two substances were simul-
taneously administered (Experiment 260) and when the atropia was adminis-
tered five minutes before the physostigma (Experiment 289).

A very interesting and suggestive chain of events is therefore presented hy

the experiments in which five grains of sulphate of atropia was administered ~

in combination with one and a half times the minimum-lethal dose of sulphate
of physostigmia. For it is seen that certain actions produced with sufficient
intensity to cause death when the two substances are simultaneously adminis-
tered, lose the power of doing so when the atropia is administered at an interval
of twenty-five minutes before the physostigma; while the now unobscured

counteraction of the lethal effect of this dose of physostigma, which makes this —

loss perceptible, persists till the interval is increased to three hours.

These various changes are no doubt caused by there bemg a progressive
increase followed by a decrease in the intensity of certain of the actions —
that are produced by this dose of atropia. The progressive increase isa
probably influenced to some extent by the rate at which the atropia is—

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 609

absorbed. The decrease may be referred to various causes, such as the elimi-
nation of the atropia, or its destruction in the tissues, or it may be due merely
to a diminution in the degree of the actions produced by this substance,
altogether independent of either elimination or destruction.

To whatever cause we refer the decrease in intensity of the actions of atropia,
in some exceptional circumstances, connected in all probability with the condition
of the animal employed in the experiment, a delay seems to occur in the rapidity
with which those actions are decreased, that are accountable for the production
of death when the interval by which the administration of the atropia precedes
that of the physostigma is one of twenty minutes, or one of shorter duration.
The occurrence of this delay is well illustrated by the fatal termination of
Experiments 323 and 324 ; in the first of which the atropia was administered
sixty-five minutes, and in the other one hundred and five minutes before the
physostigma. In both experiments, the symptoms that were observed closely
resemble those of the experiments in which an interval too brief for the pro-
duction of successful antagonism had separated the administration of the two
substances. A reference to the description of these experiments in the Tabular
Summary will confirm this statement.

The various results of this, the third series of skporiinents, have been
graphically represented in Diagram 6 (Plate XXV.). This diagram agrees
with the diagrams already described in showing by distance from one and the
other of two straight lines placed at right angles to one another, what amount
of one and the other respectively of two variables is present in each of certain
combinations of them further diagrammatically distinguished by the character
of the mark indicating their diagrammatic position into fatal and non-fatal :
and differs from them in substituting for one of their variables, namely, the
dose of physostigma, a new variable, namely, a varying interval of time separat-
ing the administration of the one substance from that of the other ; the differ-
ence depending on the fact, that while in the two previous series of experiments
the dose of atropia and the dose of physostigma varied, and the interval of time
separating the administration of the one substance from that of the other was
constant, in the present series the dose of atropia and the interval of time
separating the administration of the one substance from that of the other varied,
and the dose of physostigma was constant.

The representation of the order in which the administration of the one sub-
stance stands to that of the other, is provided for by the arrangement that in
the representation of length of interval of time by distance from the line of zero
interval, that distance is taken on the upper or on the under side respectively
of that horizontal line, according as the administration of the atropia precedes
or follows that of the physostigma. Each of the equal subdivisions of distance
from the line of zero interval, which aremarked out by the lines drawn parallel

VOL. XXVI. PART IIL (ue

610 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

to that line, represents five minutes ; and each of those of distance to the right
from the perpendicular line forming the left margin of the diagram, which are
marked out by the lines drawn parallel to that line, represents (as in Diagrams
1 and 3) a twentieth of a grain of sulphate of atropia. The constant dose of
physostigma was one and a half times the minimum-lethal one.

The conditions of each experiment may, therefore, at once be apprehended
from the position occupied by the representation of the experiment in the
diagram : whether atropia was administered before or after physostigma is
seen from the representation of the experiment being placed above or below
the zero line; what the interval of time separating the administration of the
two substances was, from the distance of this representation in a perpendicular
direction from the zero line of time ; and what dose of sulphate of atropia was
administered, from the distance of this representation in a horizontal direction
from the left margin of the diagram.

In this diagram, as in the others as yet described, the experiments that ter-
minated in recovery (dots) have been separated from those that terminated in
death (crosses) by a black line ; and the regions of recovery and death that are
thereby mapped out have been coloured respectively pink and blue.

When the diagram is examined, the two points to which attention will pro-
bably be attracted first are the irregular form of the region of recovery (pink),
and the much greater extent, both horizontal and perpendicular, of the portion
of this region where atropia was administered before physostigma than of the
portion where atropia was administered after physostigma. The existence of
this difference illustrates very distinctly a general result of the experiments of
this series, namely, that successful antagonism occurs with a greater range of
doses of atropia and with a greater range of intervals of time between the two
administrations, when atropia is given before physostigma, than when it is given
after it. In the latter case, the length. of the intervals of time is obviously
limited by there being a limitation to the time within which this dose of physo-
stigma itself produces death. In the former case, the intervals are not subject
to a similar curtailment, seeing that the doses of atropia represented in the
diagram are all considerably below the minimum-lethal dose. |

In reference to the irregularity in the form of the region of recovery, the —
only special point to which attention need be drawn, is the existence of the —
curious anomaly in the portion where atropia was administered after physo-
stigma. ‘This anomaly brings into almost too great prominence the fact, already
at some length considered, that the maximum dose of atropia that produces
successful antagonism with the dose of physostigma employed in this series was
found to be greater when the latter substance is administered ten minutes
before the atropia than when it is administered only five minutes before it. It
has already been shown that the existence of this seemingly anomalous result is

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 611

founded on the evidence of a sufficiently large number of trustworthy experi-
ments to prevent its being regarded as certainly due to some of those unavoid-
able variations in the conditions of the experiments that we do not at present
know how to make allowance for, although this is the explanation that is most
‘naturally suggested.

Soon after presenting this anomaly, the line of demarcation between the
two regions crosses the line representing simultaneous administration, and
then continues its gradual ascent until it reaches the right margin of the
diagram. In this course, there are indicated certain points of interest relating
to the maximum dose of atropia that produces successful antagonism at different
intervals of time. It is seen that this maximum dose is considerably smaller
when atropia is administered after physostigma than when the two substances
are simultaneously administered; that is also smaller, though by a less difference
than in the previous instance, when the two substances are simultaneously
administered, than when atropia is administered five minutes before physo-
stigma; and finally, that it augments in size with each increase of the interval
of time separating the administration of atropia from the subsequent adminis-
tration of physostigma. So that, as I have already pointed out, when such
a dose of sulphate of atropia as five grains is used, death occurs when it is
administered at any interval after the physostigma, simultaneously with it, or at
any interval less than twenty minutes before it; but, on the other hand, recovery
generally occurs when it is given at any interval from twenty-five to one hundred
and seventy-five minutes before it.

The portion of the lme of demarcation that forms the upper boundary of
the region of recovery rises gradually and at a tolerably uniform rate from
where it cuts the perpendicular line indicating a twentieth of a grain of sulphate
of atropia to where it reaches that indicating five grains ; and as this rise implies
_ a corresponding increase in the interval of time by which the administration of
_atropia preceded that of physostigma, it displays very clearly another general
result of the experiments, namely, the establishment of the fact that the maxi-
mum interval of time by which the administration of atropia may compatibly
with the production of successful antagonism precede that of physostigma—in
other words, the length of time the antidotal influence produced by the adminis-
tration of a dose of atropia lasts—gradually and with tolerable regularity
increases as the dose of atropia is augmented from one-twentieth of a grain to
five grains. How far increase of that interval goes on in the case of further
increase of the dose of atropia, has not been tested by experiment ; but it seems
likely that were this done, it would be found to stop at a dose of atropia some-
where between the largest already tested (five grains) and the minimum-lethal
(about twenty-one grains). Near this point, the portion of the line of demarca-
tion forming the upper boundary of the region of recovery will reach its highest

612 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

elevation (an elevation above that represented in the diagram), and either there
or somewhere further in advance, it will meet the portion of the line forming the
lower boundary of this region, and in this manner the boundaries of the
region of recovery would be completed.

General Characters of the Symptoms produced by different Combinations of
Atropia and Physostigma.—In the account that has been given of the experi-
ments contained in this section, I have avoided all details of the nature of the
symptoms that were produced, believing that any special allusion to them would
probably have distracted the attention from the primary purpose of this portion
of the research. Further, the minute details of the kind that in the previous
section were in many instances given were not concerned with experiments per-
formed at any special portions of the region of recovery, nor in any instance
with experiments performed in the region of death. The experiments in each
of these regions may be divided into two great classes, in accordance with the
symptoms which they presented. In the one class, certain of the effects of
atropia were prominently developed and maintained for considerable intervals,
while the effects of physostigma were but slightly, or even not at all exhibited.
In the other class, several of the effects of physostigma were present in a decided
form, and masked either completely or in part the effects of atropia.

In the first and second series, the former class of symptoms characterised
the experiments where recovery followed the administration of a large dose of
atropia, and also those where death followed the administration of an excessive
dose of this substance. The latter class of symptoms were present in the experi-
ments where recovery followed the administration of a small dose of atropia,
and also in those where death followed the administration of a dose of atropia
insufficient to counteract successfully the lethal effect of the dose of physostigma
given in combination with it.

In the third series, after both substances had been administered, atropia
effects were most distinctly produced in the experiments where recovery
followed the administration of a large dose of sulphate of atropia simultane-
ously with, or five or ten minutes after the dose of physostigma there given ;
and also in those where recovery followed the administration of the larger of the
doses of sulphate of atropia that were given before this dose of physostigma,
provided the interval of time separating the administration of the two sub-
stances were not a very prolonged one. These effects were likewise promi- -
nently displayed in the experiments where death followed the administration
of an excessive dose of sulphate of atropia simultaneously with, or five or ten
minutes after, the physostigma; and also in those where death followed the
administration of from 3:9 to 5 grains of sulphate of atropia before the physo-
stigma at an interval of time too short to permit of successful antagonism. On
the other hand, the effects of physostigma were in this series most prominently

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 613

developed in the experiments where recovery followed the administration of the
smaller of the doses of sulphate of atropia that were given simultaneously with,
after, and before the dose of physostigma, or of a large dose of sulphate of
atropia at a long interval of time before the physostigma. These effects were
also produced in a marked form when death followed the administration of the
smaller of the doses of sulphate of atropia that were given simultaneously with,
and five minutes after and before the dose of physostigma, and when death
occurred where sulphate of atropia in somewhat large doses was given before
physostigma at too prolonged an interval of time to admit of successful anta-
gonism.

Such, in general terms, were the characters of the symptoms in different
portions of the regions of recovery and death. The data for amore complete
analysis of these symptoms are contained in the Tabular Summary: in this
very general analysis, I have, with regard to the region of recovery, contented
myself with showing that the symptoms produced when atropia successfully
counteracts the lethal effect of physostigma vary greatly, according to the con-
ditions of administration. Successful antagonism is not necessarily accom-
panied with any special class of symptoms. It may be attended by a greater
prominence of the effects of atropia, but the same is true also of those of physo-
stigma. And, further, it does not appear that any special action belonging to
one or other substance requires to be obviously or prominently produced, in
order that the antagonism shall be successful.

It is almost unnecessary to add, that the experiments in which recovery
occurred differed much from each other in the severity of their symptoms. In
many experiments the animal was only slightly affected, and there was no
reason at any time to anticipate a fatal result ; in others, however, symptoms
of a very serious character were developed, and in several cases it was for a
long time a matter of doubt whether the animal would recover.

Combined Representation of the Three Series of Eaperiments.—In the three
series of experiments that have now been described, I have demonstrated the
limits of antagonism between atropia and physostigma,—firstly, when atropia
is administered five minutes before physostigma; secondly, when atropia is
administered five minutes after physostigma; and thirdly, when atropia in
various doses is administered at various intervals of time before and after
one and a half times the minimum-lethal dose of physostigma.

In each series of experiments, of the three quantities (namely, dose of
physostigma, dose of atropia, and interval of time between the administration
of the substances) only two vary, and the results of any one series may there-
fore be represented by a diagram constructed on a plane. Such diagrams have
been constructed, and were described when the several series of experiments
_ were being separately considered.

VOL. XXVI. PART III. ou

614 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

A combined representation of the results of the three series, involving, as it
must, three variable quantities, will be best effected by means of a model in
three dimensions. Such a model may be constructed by bending wires into the
shape of the plane curves separating the pink and blue regions of Diagrams
1, 3, and 6, and fixing them in the manner to be described to two boards placed
at right angles to one another. The wire of Diagram 1 may be called a, that of
Diagram 3 6, and that of Diagram 6 ¢, and the boards may be distinguished as
A and B. Wire @ is to be so fixed to the boards that its plane shall be at
right angles to both of them, and intersect A in the position of the left-hand
margin, and B in that of the lower margin of Diagram 1. Wire d is to be so fixed
to the boards that its plane shall be parallel to, and at a distance corresponding
to an interval of ten minutes from that of a, and intersect A in the position of the
left-hand margin, and B in that of the lower margin of Diagram 3. Lastly, wire
cis to be so attached to wires a and 8, that its plane shall be parallel to, and at
a distance corresponding to one and a half times the minimum-lethal dose of
physostigma from B, and intersect A in the position of the left-hand margin of
Diagram 6 and a plane parallel to and half way between the planes of a and 8,
which may be called plane C, in the position of the line of simultaneous adminis-

tration.

The conditions represented by any point in the model may be found by draw-
ing from it perpendiculars to the planes A, B and C. The perpendicular upon A
represents the dose of atropia ; that upon B the dose of physostigma ; and that
upon C the interval of time between the administration of the two doses, —
atropia being administered first where the point is on the one side, and physo-
stigma first when it is on the other side of the plane C.

Diagram 7 is an orthogonal projection of such a model, in which the
three variables are represented on a scale somewhat different from that of
Diagrams 1, 3, and 6; but this difference does not cause any difficulty in the re-
cognition of the corresponding parts. The continuous line a @ represents the
boundary of the region of recovery in the experiments where atropia was
administered five minutes before physostigma (Series 1); the continuous line 0 0’
the boundary of this region where atropia was administered five minutes after
physostigma (Series 2); and the dotted line ¢ a’ l’ b ad the boundary of this
region where atropia was administered in various doses and at various intervals
of time before and after one and a half times the minimum-lethal dose of physo-
stigma (Series 3). It is obvious that these lines lie upon a curved surface, on
whose one side every point represents conditions leading to death, and on whose
other side every point represents conditions leading to recovery. The surface, —
of course, cannot be fully known from the three sections of it that have been
obtained by these experiments. It could be known only by greatly increasing
the number of the experiments, so as to obtain a number of other curves of

THE ACTIONS OF PHYSOSTIGMA AND ATROPTA. 615

perpendicular sections parallel to and on either side of 6 0’ and a a’, and of
horizontal sections parallel to and below and above ca’ b’/bad. To obtain a

Diagram 7.

This woodcut is an orthogonal projection of a model in which the curves separating the regions
_ of recovery and death in the three series of experiments are brought into proper relationship to each
other. The curve of the first series of experiments is represented by a a’, that of the second series by
6 ¥, and that of the third series by c a’ b'’ bad. Doses of physostigma are indicated by the distance
(parallel to the axis of z) from the plane Y O X ; doses of atropia, by the distance (parallel to the axis
of ) from the plane Z O Y ; and intervals of time between the administration of the two substances, by
the distance (parallel to the axis of y) from the plane Z O X, points on the Y side of this plane indi-
cating atropia administered before physostigma, and points on the Y’ side indicating atropia administered
after physostigma. The curve ca’ b’ b ad intersects the curve a a’ at a and a, and the curve b U’ at
and 6 ; the points a’ and a indicating the positions respectively of the largest and the smallest doses
of sulphate of atropia that produce successful antagonism when administered five minutes before one
and a-half times the minimum-lethal dose of physostigma, and the points 0’ d the positions respectively
of the largest and smallest doses of sulphate of atropia that produce successful antagonism when
administered five minutes after this dose of physostigma.

In this diagram, the line ¢ a’ b’ b ad has been drawn without taking into account the apparently ano-
Malous experiments already discussed in page 602. The interrupted line ¢ d occupies the supposed posi-
tion of a line that would represent the results of a series of experiments in which a fixed dose of
‘sulphate of atropia (5 grains per three pounds weight of animal), and varying doses of physostigma
| were administered at varying intervals of time. Such a series of experiments has not been made,
| but the points of intersection of this line with the lines bb’,aa’, and ca’ b'bad are fixed by the
| position of the latter lines.

___ Iam indebted to my friend Professor Crum Brown for the drawing from which this woodcut has
| been made, as well as for many valuable suggestions relative to the preparation of the other diagrams
in this paper.

f
oO

616 DR THOMAS R. FRASER ON THE ANTAGONISM BETWEEN

sufficient number of such curves, however, the labour and expenditure of time
would be very great, seeing that so large a number of experiments as two
hundred and seventy-six were made in order to obtain the curves represented
in the diagram. Besides, a tolerably accurate conception of the form of the
curved surface may be gained from the curves of the three series of experiments
that have been made.

In all probability the summit of this curved surface does not occupy an
elevation materially above that of the apex of the curve a a’; but if it reach a
higher elevation, the highest point will probably be situated at only a short
distance behind that apex. From the highest point the surface slopes gradually
to dc, somewhat steeply to @ b’, with decided steepness to 0’ 6, and with still
greater steepness to 6 a.

The region included within this curved surface represents every possible
variation in the doses of atropia and physostigma and im the intervals of time
separating the administration of the two substances that is compatible with the
production of successful antagonism between physostigma and atropia.

General Summary.—Although the above combined representation of the
three series of experiments in reality presents a complete summary of the more
important of the results that have been obtained, it may be convenient to briefly
recapitulate these results. At page 540, I have stated that the chief objects of
the research are to show that atropia possesses in a remarkable degree the
power of counteracting the lethal action of physostigma, and to examine the
extent of this power and define its limits.

The former object has been effected by a detailed account in Section A of
several experiments in which the fatal action of a dose of physostigma equal to
or greater than the minimum-lethal was prevented by the physiological action
of a non-lethal dose of atropia, as well as by a brief account in Section B, of
a larger number of similar experiments, which, however, are also described
with greater detail in the Tabular Summary. The total number of these ex-
periments is one hundred and sixty one; and in each of them the animal used
was killed many days afterwards, and when the effects of the two substances
had completely disappeared, by a dose of physostigma less than or only equal
to that from which it had previously recovered.

The examination of the extent of the counteracting influence of atropia upon
the lethal action of physostigma, as well as the defining of the limits of this
influence have been accomplished in the manner and with the results fully
described in Section B. By means of the three series of experiments contained
in this section, it has been ascertained what is the maximum dose of physo-
stigma that can be counteracted successfully by atropia, what are the doses of
atropia that can counteract any given dose of physostigma, and what relation-

THE ACTIONS OF PHYSOSTIGMA AND ATROPIA. 617

ship exists between the doses with which this mutual counteration occurs and
the length of the interval of time by which the administration. of atropia pre-
cedes or follows that of physostigma.

In presence of the many obvious proofs to the contrary contained in this
paper, I have considered it superfluous to enter into any discussion of the
possibility of this counteraction being the result either of some chemical
reaction between atropia and physostigma, or of an increased rapidity in the
elimination of the one substance produced by the action of the other. The
conditions of the experiments, and the symptoms that were observed, render it
certain that atropia prevents the fatal effect of a lethal dose of physostigma
by so influencing the functions of certain structures, as to prevent such modifi-
cations from being produced in them by physostigma as would result in death.
The one substance counteracts the action of the other; and the result is a
physiological antagonism so remarkable and decided, that the fatal effect even
of three and a half times the minimum-lethal dose of physostigma may be pre-
vented by atropia. The existence of such an antagonism encourages the hope
that the power of directly counteracting disease is far from unattainable, and it
supplies a strong incentive to efforts designed to determine the conditions of
disease and the actions of remedies with an exactitude sufficient to show how
the remedial action may be applied as a counteracting influence to the diseased
condition.

Explanation of Tabular Summary, &c.—In the Tabular Summary of Experi-
ments, with which this paper ends, I have included only those experiments that
are mentioned in Section B, and have endeavoured to state the leading condi-
tions and symptoms of each experiment in as brief a manner as possible. The
_ time of occurrence of each symptom is computed from the moment when
the administration of the last-mentioned substance was commenced. It is
proper to explain, that in the column of effects on secretion and excretion, the
phrase “ slight increase of secretion of certain buccal glands” implies merely
that such an increase was inferred from certain movements of the lips sug-
gestive of it; and that the phrase in the same column “with atropia
none” implies merely that there was no evidence of any obvious effect ; but it
does not imply that diminution of secretion or excretion did not occur—for in
such experiments the occurrence of this effect could not without great difficulty
be certainly established. It will be observed that the size of the pupils is
always indicated by two measurements : the first mentioned being the size in a
perpendicular direction, and the second that in a horizontal one.

I have not considered it necessary to mention the symptoms that were
observed in the } experiments (where a lethal dose of physostigma alone was
VOL, XXVI. PART III. hee

618 DR THOMAS R. FRASER ON PHYSOSTIGMA AND ATROPIA.

administered to an animal that had previously recovered from the combined
administration of atropia and physostigma) ; for the symptoms were always very
much the same, and a sufficient account of them has already been given in
Section A. The a and 6 portions of each experiment in which they occur were
performed on the same animal.

According to the system of enumeration that has been followed, the number
of the experiments contained in this paper appears to be 331. This number,
however, does not adequately represent the labour involved in the research,
for it includes 159 experiments that consist of two parts (@ and 0), and one that
consists of three parts (a, band c); and as each of these parts is in reality a
separate experiment, the total number is 492.

All these experiments were performed in the Materia Medica Laboratory of
the University of Edinburgh, and I cannot sufficiently express my gratitude to
Sir Ropert CuristTIson for having placed his laboratory at my disposal.

[This Paper was received for publication on Friday, November 10th, 1871.
Since that time, several additions have been made to it by the Author, the
most important of which is the insertion of Diagram 7 and its accompanying
description. —J. H. B. March 4th, 1872. |

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ELT

“OLT

VOL. XXVI. PART III.

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(6.6)

VOL. XXVI. PART III.

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DR THOMAS R. FRASER ON PHYSOSTIGMA AND ATROPIA. 713

EXPLANATION OF PLATES XXIII, XXIV., AND XXV.

In each of the diagrams represented in these plates, the experiments that terminated in recovery
are marked by dots, and those that terminated in death by crosses ; and a line (distinguished in several
of the diagrams as ac) has been drawn so as to separate the dots from the crosses. The area on
the side of the line where the dots occur has been coloured pink, while the area on the side where
the crosses occur has been coloured blue; and, accordingly, the pink area represents the region of
recovery, and the blue area the region of death. In the diagrams of Plates XXIII. and XXIV., the
red horizontal line indicates the position of the minimum-lethal dose of physostigma.

Puate XXIII.

Diagram 1 illustrates the first series of experiments, in which atropia in varying doses was administered
five minutes before varying doses of physostigma.

Diagram 2 illustrates the small portion of the first series that extends to °2 gr. of sulphate of atropia.
It is drawn on a different scale from Diagram 1, as each tenth of a grain of sulphate of
atropia is indicated by twenty in place of by two subdivisions of the horizontal lines.

Diagram 3 illustrates the second series of experiments, in which atropia in varying doses was adminis-
tered five minutes after varying doses of physostigma.

Diagram 4 illustrates the small portion of the second series of experiments that extends to 2 gr. of
sulphate of atropia; and the scale on which it has been drawn differs from that of
Diagram 3 to the same extent as the scale of Diagram 2 differs from that of Diagram 1. ~

Diagrams 1 and 3 are mainly designed to illustrate the experiments extending from the minimum-
lethal dose of physostigma to the largest dose that can be counteracted successfully by atropia. They
have been drawn on the same scale in order that the results of the two series of experiments represented
by them may be compared. Diagrams 2 and 4 exhibit the course of the line a bin the first and second
series of experiments respectively, with greater distinctness and accuracy than Diagrams 1 and 3.

Puate XXIV.

Diagram 5 illustrates the first series of experiments; but it differsfrom Diagrams 1 and 2 in so far that
the entire region of recovery (pink) is represented, and that each subdivision of the
horizontal lines indicates a tenth in place of a twentieth of a grain of sulphate of atropia,
The perpendicular red line marks the position of the minimum-lethal dose of sulphate of
atropia.

The main purpose of this diagram is to show what combinations of atropia with less than the
minimum-lethal dose of physostigma are able to produce death. These combinations are represented
in the blue region below the red horizontal line.

PuatTE XXV.

Diagram 6 illustrates the third series of experiments, in which the dose of physostigma was constant
(one and a half times the minimum-lethal dose), while the dose of atropia and the in-
terval of time varied. In this diagram, as in Diagrams | and 3, each subdivision of the
horizontal lines represents one-twentieth of a grain of sulphate of atropia. The intervals
of time are represented by distance in a perpendicular direction from the thick horizontal
line, which indicates the zero interval or simultaneous administration ; and points below
this line indicate atropia administered after physostigma, while points above it indicate
atropia administered before physostigma.

(zis

XXII.—On the Decomposition of Forces externally applied to an Elastic Solid.
By W. J. Macquorn Rankine, C.E., LL.D., F.R.SS. L. & E.

(Received, 5th January ; read, 15th January, 1872.)

Introductory Remarks.—The principles set forth in this paper, though now
(with the exception of the first theorem) published for the first time, were com-
municated to the French Academy of Sciences fifteen years ago, in a memoir
entitled “ De lEquilibre intérieur d’un Corps solide, élastique, et homogéne,”
and marked with the motto, “Obvia conspicimus, nubem pellente Mathesi,”
the receipt of which is acknowledged in the Comptes Rendus of the 6th April
1857.

(1.) Principle of Isorrhopic Azes.—The following theorem was first pub-
lished in the “ Philosophical Magazine” for December 1855.

Prop. I. “ Theorem. Every self-balanced system of forces applied to a con-
nected system of points is capable of resolution into three rectangular systems
of parallel self-balanced forces applied to the same points.

“« Demonstration.—Assume any set of rectangular axes, to which reduce the
forces and the positions of their points of application ; and let X, Y, Z be the
components of the force applied to any point (a y z).”

“Let

Then, in linear transformations of rectangular co-ordinates, A is covariant with
«*, D with yz, &e.
“Conceive the surface of the second order, whose equation is

Az? + By? + Cz” + 2Dyz + 2Ezx + 2F vy = constant d (1).

Then if the forces, and the positions of their points of application be reduced
anew to the principal axes of that surface, we shall have

0h 0h 0

and consequently, each of the three systems of component forces parallel to
VOL. XXVI. PART IV. 8Z

716 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

those three principal axes will be self-balanced, independently of the other two
systems. Q.#.D.”*

Remark.—The values of A, B, &c., obviously depend solely on the directions
of the axes, and not on their point of intersection.

(2.) Definitions.—The following terms will be employed in the sequel, rela-
tively to any given system of forces.

Rhopimetric Surface.—The surface (1).

Rhopimetric Co-efficients—The quantities A, B, C, D, E, F.

Isorrhopic Axes.—The principal axes of the surface (1).

Principal Rhopimetric Co-efficients.—The values of the co-efficients A, B, C,
for the isorrhopic axes.

Arrhopic System—A system of forces for which A=0, B=0, C=0,
D =0, E=0, F = 0; and for which, consequently, every direction is an isor-
rhopic axis. ,

(3.) Application to Elastic Solids.—The utility of the above principle of isor-
rhopic axes in the theory of the equilibrium of elastic solids arises from the
fact, that although, in treating of the equilibrium of a solid body as a whole
supposed to be perfectly rigid, it is allowable to suppose the point of appli-
cation of any force to be anywhere in the line of action of that force ; yet, when
the solid body is considered as being strained by the forces applied to it, no
such supposition is admissible ; and in every mathematical process for deter-
mining such straining effect the actual point of application of each force must
alone be considered. When the straining forces to which an elastic solid is
subjected are restricted within certain limits, the straining effect of any number
of self-balanced systems of forces combined is sensibly equal to the sum of the
effects which those systems respectively produce when acting separately.

Consequently, the principle of Isorrhopic Axes affords the means of re-
ducing the problem of finding the straining effect of any self-balanced system
of forces applied to an elastic solid to that of finding the separate straining
effects of three self-balanced systems of parallel forces.

Prop. IJ.—Prosiem. To jind the Rhopimetric Co-efficients for a system of
Forces applied over the surface and throughout the interior of a solid body.

Let X, Y, Z, denote the components of the attractive or repulsive accelera-
tive force applied to a molecule of the solid whose co-ordinates are 2, y, 2, its
volume dz dy dz, and its density p; let P, Q, R, denote the components of
the external stress (tension positive) per unit, of area, which acts on an element

* It is easy to see how this theorem may be extended to a system of moving masses, by putting

d?x
x—- ae for X, &e.

FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. 717

of the external surface of the solid whose co-ordinates are 2’, y/, z, and area
d*s; then we
A=S/ffaXp. dx dy dz + ffx’ Pd’s ;
D =f/ffzYp . dx dy dz + ff'Q.a@s (2),
=/{fyZp . dx dy dz + ffy'R. d’s

and the expressions for the other co-efficients will be similar, mutatis mutandis.
Q.£.1.

In the case of normal external stress, let ,, »,, n, be the direction-cosines
of the normal to the element d?s, and S the intensity of normal stress on
that element ; then pra

PSs; O=s2,> hi] sn

and in finding the values of the double integrals, we may put n, d’s = dy dz,
&c.; observing, that for each set of an even number of elements d?s, which
have a common projection such as dy dz, the quantity to be integrated is of a
form such as >Sz’, and contains as many terms as there are elements having a
common projection ; the sign of each term being positive or negative, accord- -
ing as the direction-cosine (as 7,) is positive or negative.

(4.) Summary of the Relations between Internal Stresses and Applied Forces
in an Elastic Solid in Equilibrio.

The following principles having been long known through the investigations
of various mathematicians, are here recapitulated for the sake of convenience.
In expressing internal stresses, the notation of M. Lame is adopted, viz., let
dx dy dz be a rectangular molecule, and let

ees Ne
x y 2

be the normal stresses per unit of area, on the pairs of faces normal respec-
tively to

X,Y, ~,;

such stresses being considered as positive when tensile, negative when compres-
sive. Also, let

Led gla

be the tangential stresses per unit of area on. the double pairs of faces parallel
respectively to

Ly UP

718 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

T,, being considered as positive when it tends to elongate the positive and
shorten the negative aa of the faces Lz, &c. In the transformation of
co-ordinates, N._, ,N, NZ pees are covariant respectively with 2”, y’, <’, yz,

zn, xy. Let n,, Ny, M,; + és ee of the external normal to any
point of the body’s surface.

Then the following are the conditions of equilibrium between the internal
stresses and the applied forces.

Conditions relative to each Internal Molecule :—

GN a idee aly

dl aN. valle

aT, dT, aN, i

Conditions relative to each Point of the Surface :—

P = aN, Had, Aa,
QOv=an,T, + a,N, + 21, ; : (4).
R=n,T, + 2,7, + 0,N,

(5.) Effect of Terrestrial Gravitation.—In a homogeneous heavy body near
the earth’s surface, the internally applied forces pX, pY, pZ, are constants,
being simply the components of the weight of unity of volume of the body
along the three axes of co-ordinates.

Prop. IJJ.—Prosiem. Yo Balance the Weight of a Homogenous Body by
Pressure applied to its Surface, so as to form an Arrhopic System of Forces ; and
to Determine the corresponding Internal Stresses—Assume a vertical direction
positive downwards, for the axis z, and let the plane of yz pass through the
centre of gravity of the body; then pY = = 0, pZ = 0; and pX = gp is the weight
of unity of volume of the body. For the applied external pressures, make
Q,= 6,8, =0; =

Pi == gpe nes , : a. ~~

Then the system of pressures P, balances the weight of the body ; for let the
horizontal projection of an element d’s of the body’s surface be dy dz, then d’s =

dy d Soi net teen : di : :
ne. and that projection is to be considered as positive or negative according

to the sign of m,; consequently

/{ Pd’s = — gp/fx' dy dz = — gpV.

FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. 719

where V is the entire volume of the body. Also the entire system of forces is
Arrhopic. For it is evident that ,

b= C=) = k= F =;

A= gp tf[fa da dy dx — ff x'*dy dz.
Now the first term of the above expression is well known to be null when the
plane yz traverses the centre of gravity ; and by attending to the rule, that
dydz is positive or negative according to the sign of 2,, it appears that
the second term is null also. Therefore, A = 0, and the system of forces is
Arrhopic.
Lastly, for the system of internal stresses, make

ON — O05 Os i 05 0 NG, = gph . - (6):

and

This system evidently satisfies equations 3, 4, and 5; that is to say, the re-
quired system of internal stresses consists in a vertical normal stress at each
molecule, proportional to its vertical distance from the horizontal plane of the
centre of gravity of the body, tensile above that plane, and compressive below.
Q.4.L.

DerFInitions.—Antibarytic Pressures: the externally applied pressures |
which (as in the above problem) form, with the gravitation of a body, an
Arrhopic system. <Antibarytic Stresses: the corresponding internal stresses.

Remark.—tt is characteristic of the Antibarytic pressures that their inten-
sity for each unit of area of the horizontal projection of the body’s surface is a
linear function of the vertical co-ordinate, viz.,

=I = — ¢ ’ ° ° ° °
7 gprs (6A).

Abarytic Pressures.*The system of pressures left after taking away the
Antibarytic Pressures from the Actual Pressures applied at the several elements
of the body’s surface.

Corottary.—The Abarytic Pressures are self-balanced ; their Rhopimetric
co-efficients are the same with those of the] entire system of Applied Forces ;
and in calculating their effects, the force of gravity is to be left out of consider-
ation. Hence the internal stresses corresponding to a system of Abarytic
Pressures must fulfil the following equations :—

die | aa adi

ae Page eae

Ry aN

Gee GE da (7).
at, | at, aN, _ 9.

dike t edypna day 4?

VOL. XXVI. PARTIV. | 9A

720 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

(6.) Decomposition of Abarytic Pressures.

Derinition.—Homalotatic System of Pressures: A system of Abarytic Pres-
sures applied to the surface of a body, and producing an uniform state of stress
throughout its internal molecules.

Prorv. [V.—Prosiem. To Decompose any Abarytic System of Pressures
applied to the Surface of a Solid into a Homalotatic System, and an pe rie
System.

Having computed the six Rhopimetric co-efficients for the given Abarytic
system of Pressures, as referred to any set of orthogonal axes, take the follow-
ing values for a set of six uniform internal stresses; V being (as in Proposi-
tion III.) the volume of the solid :—

These quantities being constant, fulfil equation (7).
The corresponding external pressures are as follows, according to equa-
tion (4) :—

= y [mat mF +E |
Op= {ak + mB + nD \ . «Neale
ih = vy {rE + nD + nC }

If the Homalotatic system of pressures given by these equations be taken from
the entire Abarytic system, an Arrhopic system will remain.

For the Rhopimetric co-efficients corresponding to the Homalotatic pres-
sures are as follows :—

Ay = [fe Pigs
ie | Affe dydz + Ff/fa'dzda + Effa'dady }

(and similar equations for the others, mutatis mutandis).
Now, observing as before, that

dydz, dzdx, dady
are to be considered positive or negative according to the sign of

Nez, Ny, Nz)
it appears that

‘FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. fon

Sfa dydz =N ; ffa' dzdx =0; ffa'dydz =0;
so that
A, =A;

and in like manner, each of the Rhopimetric co-efficients for the system of
Homalotatic Pressures is proved to be equal to the corresponding co-efficient
for the entire system of Abarytic Pressures; so that the system of residual
pressures, left after taking away the Homalotatic Pressure from the Abarytic
Pressures, is Arrhopic. Q.E.F.

Remark.—The Homalotatic system of six uniform stresses obviously repre-
sents the mean state of stress of the whole body.

Corottary.—The set of six uniform stresses given by equation (8) are
equivalent to three principal rectangular uniform normal stresses along the Isor-
rhopic Axes. For the three principal normal stresses of the system (8) are in
direction parallel to, and in magnitude represented by, the reciprocals of the
squares of the principal semi-axes of the surface,

Nee EON, oe Nee"
+ 2T,., 9% + 21,.,2@ + 27,.,2y =1; ; (10),

which is similar and parallel to the Rhopimetric surface.

(7.) Recapitulation, and Statement of the Advantages of the Method described.
—The following is a summary of the processes of the before-described method
of decomposing any self-balanced system of forces applied to an Elastic Solid
near the earth’s surface :—

First. By Proposition II. Equation 2, find the six Rhopimetric Co-eficients
of the system of applied forces, including gravitation.

Secondly. By Proposition III. Equations 5, 6, compute the system of verti-
cal Antibarytic Pressures, with their corresponding vertical internal stresses ;
which pressures at once balance the force of gravitation, and form with it an
Arrhopic system.

Thirdly. Take away from the entire pressure applied at each element of the
body’s surface the Antibarytic pressure at the same element, so as to leave a
system of Abarytic Pressures, which is self-balanced, independently of gravi-
tation, and whose Rhopimetric co-efficients are the same with those originally
computed.

Fourthly. By Proposition IV. Equations 8 and 9, compute from the given
set of Rhopimetric co-efficients the system of sex uniform mean internal stresses,
with the corresponding system of Homalotatic external pressures. ;

Fifihly. Take away from the Abarytic pressure on each element. of the

722 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

body’s surface, left by the third operation, the Homalotatic pressure found by
the fourth operation, so as to leave an Arrhopic system of externally applied
pressures ; whose effects in producing internal stress and strain remain to be
found.

The advantages of the method of decomposing the forces applied to an
Elastic Solid arise from the following circumstances :—

First. It is impossible to determine the effect of any system of forces applied
to an elastic solid, unless such system be self-balanced.

Secondly. It is, if not impossible, extremely difficult to determine directly
the effect upon an elastic solid of any self-balanced system of forces which are
not all parallel, unless they correspond to an uniform state of stress.

Thirdly. The difficulties of any problem respecting the stress of an elastic
solid are often much increased if the applied pressures are not parallel to an
axis of co-ordinates chosen with reference to the figure of the solid.

It is, therefore, desirable that all those pressures whose effects are not
capable of being expressed by a state of stress uniform at every molecule of the
solid (like that due to Homalotatic Pressure) should be reduced to a system or
systems whose components parallel to any axis whatsoever are self-balanced,
and may therefore have their effects separately computed—that is, to an
Arrhopic system, or systems; and this is what is accomplished by the pro-
cesses above described. To complete the solution, therefore, of the problem of
the internal equilibrium of any elastic solid near the earth’s surface, it is only
necessary to find the separate effects of three Residual Arrhopic self-balanced
systems of parallel pressures, parallel respectively to such axes as the figure of
the body may render most convenient.

(8.) Cases in which the Distribution of Internal Stress is Independent of the
Co-efficients of Elasticity of the Sold.

THEOREM.— When the molecular displacements are expressed by algebraical
functions of the co-ordinates not exceeding the second degree, and the stresses
(consequently) by constants and linear functions of the co-ordinates, the distri-
bution of internal stress is independent of the co-efficients of elasticity of the
solid.

Demonstration.—The cases in which the distribution of internal stress is
independent of the co-efficients of elasticity, are those in which the number of
arbitrary constants in the functions expressing the internal stresses is not greater
than the number of arbitrary constants in the functions expressing the mole-
cular displacements ; so that, consequently, the internal stresses can be deter-
mined from the external forces alone.

The internal stress at any point is expressed by six components, linear

FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. 2a

functions of the first differential co-efficients of the molecular displacements,
which last are three in number.

When these are expressed by transcendental or other irrational functions,
the former number of constants is double of the latter; hence this class of
cases is excluded.

When the molecular displacements are expressed by three homogeneous
rational and integral functions of the three co-ordinates of the n” degree, the

: : : athe
number of arbitrary constants contained in them is 5 (m + 1) (m + 2).

In the same case, the component stresses are expressed by six rational and
integral homogeneous functions of the degree » —1; so that in them the

: a6
number of arbitrary constants is 5” (m + 1).

Hence the ratio borne by the number of arbitrary constants in the expres-
sions for stresses to the corresponding number in the expressions for molecular

displacements, is
2n
nm+2°

This ratio is less than, equal to, or greater than unity, according as is less —
than, equal to, or greater than 2; therefore the distribution of stress is inde-
pendent of the co-efficients of elasticity for molecular displacements expressed
by rational functions not exceeding the second degree, and stresses expressed
by constants and by linear functions of the co-ordinates. Q.£.D.

As n increases indefinitely, the above-mentioned ratio approximates to 2,
being its value for irrational functions.

(9.) The Classes of External Pressures which produce stresses answering the
preceding description are the following :—

Homalotatic Pressures, for which the stresses are expressed by constants,
and Antibarytic, Homalocamptic, and Homalostrephic Pressures, for which the
stresses are expressed by linear functions.

It has already been seen, that for the systems of pressures designated as
Homalotatic and Antibarytic, the internal stresses are determined independently
of the co-efficients of elasticity of the body, being in the former case uniform,
and in the latter consisting in a vertical normal stress, which is a linear function
of the vertical ordinate from the horizontal plane of the body’s centre of gravity.
The consideration of these two systems of stresses forms part of the solution of
every problem concerning the equilibrium of an elastic solid.

DeEFIniTIoNs.—Homalocamptic Pressures (or pressures of Uniform Bend-
ing).—A system of external pressures corresponding to a system of normal in-
ternal stresses uniform in direction, whose intensity is a linear. function of an

VOL. XXVI. PART IV. 9B

724 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

ordinate perpendicular to that direction, measured from a surface passing
through the body’s centre of gravity. That is to say, the centre of gravity
being the origin, let O z be any direction in which there is a normal stress N, ,
let b, c, be two constants, and let

N, = by + cz ; : ‘ (11),

which fulfils of itself the differential equations (7) of internal equilibrium. Then
by the equations (4)—
P=n,N, =n, (by + cZ) met 5 (12),
QO] 0; R=0;

will represent a system of Homalocamptic Pressures.

Homalostrephic Pressures, or Pressures of Uniform Twisting.—A system of
pressures corresponding to a system of tangential internal stresses uniform as
to the pair of internal directions in which they act, and whose intensity is a
linear function of an ordinate perpendicular to those directions, and measured
from a plane passing through the body’s centre of gravity ; that is to say, for
example, let
Looe : . . (13),

which fulfils of itself the equations (7). Then by the equations (4)—

P= O50) = ail eae

P= Fl Oe, ce

A system of Homalostrephic Pressures is equivalent to a pair of systems of
Homalocamptic Pressures, making angles of 45° with the directions of the
Homalostrephic Pressures. For let Oy,, Oz,, be a pair of axes in the plane
yz, inclmed at 45° to y and z, so that + y lies between + y, and z, Then is
T, equivalent to a pair of normal stresses,

Neg=— n= + @2,
IN, = l= ee.
Prop. V.—TuHeorEM. Every Homalocamptic System of Pressures is Ar-
rhopte.
For the following are the Rhopimetric co-efficients derived from equa-
tion (12)—
B=. CSD Hv
A=ff(Pae @s=bffa'y.dydzt+effea.dydz;
Ea fP2e@s=bfy 2 .djde eye ajar,
F=fPy@s=bffy’? .dydz+effyz.dydz;

(dy dz being as usual considered as + ” or —™, according to the sign of 7,).

FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. 725

Now each term of A is null, because the origin is the centre of gravity of
the body; and each term of E and F is null, because for each positive
element dy dz of the projection of the body’s surface there is an equal negative
element.

Pee Oe N0 = BOs

and the system of pressures is Arrhopic. Q.£.D.
Corollary 1. Every Homalostrephic System of Pressures is Arrhopic.
Corollary 2. The subtraction from any Arrhopic system of pressures, of
a Homalocamptic or Homalostrephic system, leaves an Arrhopic residual
system.

(10.) ExampLe.—Homalocamptic Pressures, Uniform Bending Stress in a
Prism.—tThe consideration of Homalocamptic and Homalostrephic Pressures
does not, like that of Homalotatic and Antibarytic Pressures, form an essential
part of the solution of every problem of the internal equilibrium of an Elastic
Solid; but is to be employed only when it evidently tends to simplify the
problem.

The most generally useful example of a single system of Homalocamptic
Pressures is the following :—

Let the axis of z be that of a prismatic pillar, traversing its centre of
gravity. Then for the ends of the prism respectively,

OE ae Ais sa Ole e912 = Oi

and for the sides, 7, = 0. Let }z = cy be the equation of any plane passing
through the centre of the prism, and let each element of the ends of the prism
be acted on by normal pressures, proportional to the distance from that plane,
tensile towards + y, and compressive towards — y. Then for

Nea a = Se (by + cz); Gi 0;.R = 0;
and for (15),
= Oe 0 OO. 0;
and the internal stresses are
NS oybeusN w= N= Dye, = Ty 10 j (16).

When a system of normal pressures is distributed im any manner on the
two ends of a prism, the system of Homalocamptic Pressures which approxi-
mates most nearly to the actual system is found by computing the moments of
the pressures on one end relatively to the planes zy and zz, viz.,

ff Pzedydz and —f/Py.dydz,

726 PROFESSOR MACQUORN RANKINE ON THE DECOMPOSITION OF

and make
| _ Pe.dydz ff Py.dydz
a fe .dyde? © ff P .dydze * = (tz).
The denominators of these expressions are the geometrical factors of the
moments of inertia of the cross-section of the prism round y and z respectively.
It is obviously immaterial which end of the prism is chosen for the compu-
tations.

(11.) Example of Homalostrephic Pressures ; Uniform Twisting Stress in an
Elliptic Cylinder.

Let Oa be the longitudinal axis of an Elliptic Cylinder, Oy and O z parallel
to its greatest and least diameters, and let 2 and 2q be those diameters.
Then for the ends of the cylinder,

n, Sat Sapo wa De

and for the elliptic surface

ym zm
i, =O y= 350, =
on 2 y pe? ‘Zz qo?
when
i Saige Eat
peerings

Let there be two Homalostrephic systems of Tangential Stresses, thus re-
presented

Then the external pressures constituting a pair of Homalostrephic systems com-
bined, will be as follows :—
On the ends,

form, = +1; P=0; Q= +e; R= + by,
on the elliptic surface

b
P=2,T, + 0, T, = myz. fart gq }sQ@=0;R=0.

Now let the external pressures be subject to the condition that the pressure on
the elliptic surface shall be everywhere null; then we must have

b
nag es (0

pet

Consequently, let

FORCES EXTERNALLY APPLIED TO AN ELASTIC SOLID. (27

ae ee fe : (18),

and for the external pressures at the ends of the cylinders, which are wholly
tangential ;

fee ey : (19).

The resultant tangential pressure at any point (y, z) of one end of the cylinder,
or of one of its sections,

OE? — — ; being pro. |
and its direction-cosines are
Q eee 18 ae ym
Vein © Be Vqrsn > ue |
showing it to be a tangent to an ellipse similar and concentric to the outline of .
the end of the cylinder, and proportional to the diameter of that ellipse to
which it is parallel.

The total moment of torsion M, that is, the moment about O a of the forces
applied to one end of the cylinder, is as follows :—

y 22
TT (By — Q2) dy dz = aff (by + a) dy de,

(20),

which, because

ee eae
pag hs
becomes
M = a x area of elliptic base = wapq ; iy

which equation serves to determine the constant @ when the moment of torsion
is given, Viz.,
agut
mpg *
The tangential stresses at the extremities of the greatest and least diameters

of the ends are inversely as those diameters, viz.,

a M Q

mpg?

(22).

These results agree with those obtained by Cauchy, but have the peculiarity
of being arrived at independently of the co-efficient of elasticity of the sub-
stance.

VOL. XXVI. PART IV. 9 ¢

( 729 )

XXITI.—On the Geometrical Mean Distance of Two Figures on a Plane.
By Prof. J. CLerk MAxwett, F.R.S.

(Received January 5th ; read January 15th, 1872.)

There are several problems of great practical importance in electro-magnetic
measurements, in which the value of a quantity has to be calculated by taking
the sum of the logarithms of the distances of a system of parallel wires from a
given point. The calculation is in some respects analogous to that in which
we find the potential at a point due to a given system of equal particles, by
adding the reciprocals of the distances of the particles from the given point.
There is this difference, however, that whereas the reciprocal of a line is com-
pletely defined when we know the unit of length, the logarithm of a line has no
meaning till we know not only the unit of length, but the modulus of the system
logarithms.

In both cases, however, an additional clearness may be given to the state-
ment of the result by dividing, by the number of wires in the first case, and by
the number of particles in the second. The result in the first case is the loga-
rithm of a distance, and in the second it is the reciprocal of a distance ; and
in both cases this distance is such that, if the whole system were concentrated
at this distance from the given point, it would produce the same potential as it
actually does. |

In the first case, since the logarithm of the resultant distance is the arith-
metical mean of the logarithms of the distances of the various components of the
system, we may call the resultant distance the geometrical mean distance of
the system from the given point.

In the second case, since the reciprocal of the resultant distance is the
arithmetical mean of the reciprocals of the distances of the particles, we may
call the resultant distance the harmonic mean distance of the system from the
given point.

The practical use of these mean distances may be compared with that of
several artificial lines and distances which are known in Dynamics as the radius
of gyration, the length of the equivalent simple pendulum, and so on. The
result of a process of integration is recorded, and presented to-us in a form
which we cannot misunderstand, and which we may substitute in those ele-
mentary formulz which apply to the case of single particles. If we have any
doubts about the value of the numerical co-efficients, we may test the expression

VOL. XXVI. PART IV. 9D

730 PROFESSOR CLERK MAXWELL ON THE GEOMETRICAL

for the mean distance by taking the point at a great distance from the system,
in which case the mean distance must approximate to the distance of the
centre of gravity.

Thus it is well known that the harmonic mean distance of two spheres, each
of which is external to the other, is the distance between their centres, and that
the harmonic mean distance of any figure from a thin shell which completely
encloses it is equal to the radius of the shell.

I shall not discuss the harmonic mean distance, because the calculations
which lead to it are well known, and because we can do very well without it.
I shall, however, give a few examples of the geometric mean distance, in order
to show its use in electro-magnetic calculations, some of which seem to me to
be rendered both easier to follow and more secure against error by a free use
of this imaginary line.

If the co-ordinates of a point in the first of two plane figures be z and y,
and those of a point in the second € and », and if 7 denote the distance between
these points, then R, the geometrical mean distance of the two figures, is
defined by the equation

‘log BR .fifda dy dé dn = iff log r dx dy dédn .

The following are some examples of the results of this calculation :-—

(1.) Let AB be a uniform line, and O a point
not in the line, and let OP be the perpendicular
from O on the line AB, produced if necessary, then
if R is the geometric mean distance of O from the

E A “line AB,

AB. (log R + 1) = PB. log OB — PA log OA + OP. AOB .

(2.) The geometrical mean distance of P, a point in the line itself, from AB»
is found from the equation

AB (log R + 1) = PB log PB — PAlog PA .

When P lies between A and B, PA must be taken negative, but in taking the
logarithm of PA we regard PA as a positive numerical quantity.

(3.) If R is the geometric mean distance between two finite lines AB and
CD, lying in the same straight line, i
AB.CD (2 log R + 8) = AD’ log AD + BC’ log BC — AC’ log AC

— BD’ log BD .

MEAN DISTANCE OF TWO FIGURES IN A PLANE. 731

(4.) If AB coincides with CD, we find for the geometric mean distance of
all the points of AB from each other

i= A Bees sayy
R P

(5.) If R is the geometric mean distance of the
rectangle ABCD from the point O in its plane, and
POR and QOS are parallel to the sides of the

rectangle through O, C s D

ABCD (2log BR + 3) = 20P.OQ log OA + 20Q. OR log OB
+ 20R.0O8S log OC + 20S . OP log OD
+ OP? .DOA + 0Q? . AOB
+ OR? . BOC + 08? .COD

(6.) If R is the geometric mean of the distances
of all the points of the rectangle ABCD from each 4% “ a

other, :

if AB AC IL Ce AC
log R = log AC — & poz log kB — & ape oS BG

DEABE Oa Eh ant os ; 0
+ 3pq BAC +3 ap ACB — 5 :

When the rectangle is a square, whose side = a,

log R = loga + rlog 2 == ogee
= loga — 0°8050866

R = 044705 a .

(7.) The geometric mean distance of a circular line of radius a, from a point in
its plane at a distance 7 from the centre, is 7 if the point be without the circle,
and a if the point be within the circle. |

(8.) The geometric mean distance of any figure from a circle which completely
encloses it is equal to the radius of the circle. The geometric mean distance of
any figure from the annular space between two concentric circles, both of which
completely enclose it, is R, where

(a," — a,”) (log R + ) = a, log a, — a, log a ,

a, being the radius of the outer circle, and a, that of the inner. The geometric
mean distance of any figure from a circle or an annular space between two con-

732 ‘PROFESSOR CLERK MAXWELL ON THE GEOMETRICAL

centric circles, the ‘figure being completely external to the outer circle, is the
geometric mean distance of the figure from the centre of the circle.
(9.) The geometric mean distance of all the points of the annular space be-
tween two concentric circles from each other is R, where
qu a.
(ay — a,’)’ (log R — log a) = | (Ba,’ — a’) (a,’ — a,”) — a,* log Pi
When a,, the radius of the inner circle, vanishes, we find

R= @e7? .

When a,, the radius of the inner circle, becomes nearly equal to a,, that of the
outer circle,
R= G2
As anexample of the application of this method, let us take the case of a
coil of wire, in which the wires are arranged so that the transverse section of
the coil exhibits the sections of the wires arranged in square order, the distance
between two consecutive wires being D, and the diameter of each wire d.
Let the whole section of the coil be of dimensions which are small com-
pared with the radius of curvature of the wires, and let
OC & © oe Bae ny. mean distance of the section from itself
e R.

1st, If we begin by supposing that the wires fill up
Ololo the whole section of the coil, without any interval of
insulating matter, then if M is the co-efficient of in-

duction of a linear circuit of the same shape as the coil
on a similar parallel circuit at a distance R, the co-efficient of induction of
the coil on itself will be
Te WE.

2d, The current, however, is not uniformly distributed over the section.
It is confined to the wires. Now the co-efficient self-induction of a unit of
length of a conductor is
C—2logR ,

where C is a constant depending on the form of the axis of the conductor, and
R is the mean geometric distance of the section from itself.
Now for a square of side D,

25
log R, = log D +7 log 2 =F 7 12°

MEAN DISTANCE OF TWO FIGURES IN A PLANE. oo
and for a circle of diameter d

log R, = logd — log2 ==

Hence

R D 4 7 11
log =? = log] +sloge2+s5——z ,
Bikes }Oargn an 3 Osta, aim G

and the co-efficient of self-induction of the cylindric wire exceeds that of the
square wire by
2 flog 4 + 0°1380606}

per unit of length.

3d, We must also compare the mutual induction between the cylindric
wire and the other cylindric wires next it with that between the square wire
and the neighbouring square wires. The geometric mean distance of two
squares side by side is to the distance of their centres of gravity as 0°99401 is
to unity.

The geometric mean distance of two squares placed corner to corner is to
the distance between their centres of gravity as 10011 is to unity.

Hence the correction for the eight wires nearest to the wire considered is

—2 x (001971) .

The correction for the wires at a greater distance is less than one-thousandth
per unit of length.
The total self-induction of the coil is therefore

2M + Q flog + 011835} ,

where x is the number of windings, and / the length of wire.
For a circular coil of radius = a,

M = 4za(log 8a — log R — 2) ,

where R is the geometrical mean distance of the section of the coil from itself.

VOL. XXVI. PART IV. 9:

Trans Roy. Soc Edin: Vol XN. PLAAVL

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Gi7s5uy

XXIV.—On the Lunar Diurnal Variation of Magnetic Declination at Trevan-
drum, near the Magnetic Equator, deduced from Observations made in the
Observatory of His Highness the Maharajah of Travancore, G.C.S.T.
By J. A. Broun, F.R.S. (Plates XXVI-XXVIIL)

(Read 6th May 1872.)

1. The lunar diurnal variation of magnetic declination as first discovered by
Kreit, depended on too few observations to be free from the errors introduced
by irregular disturbing causes. The imdependent discovery of the lunar action
on the magnetic needle made afterwards by myself, was lable to the same
criticism ; but the agreement of the results obtained, both for the magnetic
declination and the horizontal force, was sufficiently great to give a consider-
able value to the conclusion, that the magnetic needle obeys a diurnal law,
depending on the moon’s hour angle, both as to its direction and the force with
which it is directed. This conclusion was farther confirmed in the discussion
first made by myself, for the lunar diurnal variation of the vertical magnetic
force, which gave, within an hour, the same epochs of maxima and minima as
those obtained previously by me for the horizontal component.*

The results obtained afterwards from longer series of observations,t while less
affected by the irregularities due to disturbances, still showed variations of so
small a range, that the fact of the existence of a lunar diurnal variation was
not accepted by all men of science without reserve. Whatever doubt may
have existed before has been dissipated, I have every reason to believe, by the
results communicated in a paper to the Royal Society of Edinburgh in 1867,}
where variations were shown due to the lunar action, which equalled those
produced by the sun.

2. This action of the moon is not constant; it not only varies with the
period of the year, but it varies also for the same month in different years ; so
that, in some seasons, the variation is still so small, as to require the combina-
tion of large masses of observations to eliminate the effects of irregular causes,
in the determinations relating to the laws of variation under different circum-
stances.

3. Though observations made in different parts of the world, both in the
northern and southern hemispheres, seemed to prove the existence of a law of

* Trans. Royal Soc. Edin., vol. xvi. p. 143, § 19.

+ The most important of these have been discussed by General Sir E. Sasrve.
¢ Trans., vol. xxiv. p. 669.

VOL. XXVI. PART. IV. QF

736 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

lunar diurnal variation, no conclusions had been drawn either as to the way in
which the law varied in passing from one hemisphere to another, or as to the
way in which it might change near the equator with the position of the sun or
moon in declination. As nothing is known as to the mode in which the sun
and moon produce these variations, nothing could certainly be predicted a priorz
as to the change of the law under the circumstances just mentioned.

4. The first result obtained from a discussion of observations made during
the five years 1854 to 1858 at Trevandrum, was that the law of lunar diurnal
variation for the group of months about January was the inverse of that for the
months about July.* This fact, which held also for the solar diurnal variation,
appeared to relate both variations to the position of the earth in its orbit, rather
than to the passage of the sun from one hemisphere to the other. The similarity
in the change of the law of lunar and solar diurnal variation for the sun north
and south of the equator, led also to the conclusion that the mean lunar diurnal
movement, like the mean solar diurnal movement, should be in opposite direc-
tions in the high latitudes of the two hemispheres: the facts have since then
been found to be in accordance with this conclusion.t

5. The next question of importance was, whether the moon’s passage from
one hemisphere to the other would produce any marked change in the law of
variation, or in its amount. The discussion of five years’ observations seemed
to show a considerable difference in the relative magnitude of the maximum
and minimum, not only with the moon farthest north and south, but also with
the position of the moon on the equator, according as she was moving towards
the north or towards the south.

6. As the action of the moon on the needle may vary from different causes,
it becomes necessary, to be sure in obtaining the results for any given argument,
that we have got rid of the effects dependent on other arguments.

If we assume, in the first instance, that the solar diurnal variation is the
same for each day throughout a month, and that a similar supposition holds for
the moon ; then, as in a month, which is nearly equal to a lunation, the moon
will have been on all the twenty-four meridians at each of the solar hours, the
mean disturbing action of the moon will have been the same at each of the
solar hours, and the mean position of the needle for each solar hour, as derived
from a month’s observations, will be equally affected or unaffected by the lunar
action.{ A similar conclusion may be arrived at relatively to the mean position
of the needle for the.moon on different meridians: thus, if all the observations
made during a lunation for the moon on the principal meridian be summed

* Proceedings of the Royal Society of London, vol. x. p. 475. 1861.
+ Proce. Roy. Soc. Lond., vol. xvi. p. 59.
{ Hence the mean solar diurnal variation sought, in any case, from fewer than a month’s observa-

tions will be more or less in error, according as the lunar action (and the change of its law during the
lunation) is more or less considerable

MAGNETIC DECLINATION AT TREVANDRUM. (al

together, all for the moon on the meridian of one hour, and so on, and the
means be taken, we see in each case that the sun will have been on all the
meridians for each of these hour angles of the moon; and the means thus
obtained will be equally affected by the solar action. The process is the same
as for the solar diurnal variation.

If, however, the suppositions made are inaccurate, and the solar diurnal
action is not constant during a month, the hourly means for the lunar diurnal
variation will not be equally affected by the solar action. A similar conclusion
holds for the solar diurnal variation, if the law of lunar diurnal variation is not
constant throughout the month.

7. Besides this possible source of error, there is at least another which
depends upon unknown causes, the effects of which have been named disturb-
ances. In order to determine the extent of the disturbance at different hours,
each observed position is compared with the monthly mean for the corresponding
hour, and the means of the differences thus obtained give comparative measures
of the disturbance or displacement of the needle at the different hours.

The discussions for this purpose have shown that the disturbance, though
irregular in action, yet on the whole obeys a solar diurnal law, so that in the
mean of a sufficient number of observations the disturbing action, like the
usual solar action, will be eliminated, and this the more easily when the days
of greatest disturbance are omitted.

8. As it is thought probable that the larger disturbances, at least, are inde-
pendent of the lunar action, it has been sought to avoid the irregularities which
they introduce when limited series of observations are discussed. Three
methods have been employed for this end. By one all the observations differ-
ing from the corresponding hourly mean by a certain arbitrary limit have been
suppressed, the hourly means for the month have been recomputed from the
remaining observations, and the differences taken as if the suppressed observa-
tions had not existed.* By the second (employed by me in the first discussions
for the lunar diurnal variation from the Makerstoun observations for 1844—
1845), the observations exceeding a certain arbitrary limit were considered dis-
turbed, and quantities were substituted for them, derived from preceding and
succeeding observations, which were within the limit. In the third method,
which was employed by me in the first discussion of the Trevandrum observa-
tions ;t all the observations in days which were considered days of marked
disturbance were omitted.

* T have touched on the objections to this method in a note which appeared in the Proceedings of
the Royal Society, of London, vol x. p. 479. The Astronomer Royal (Trans. Roy. Soc. Lond., vol.
cliii. p. 617), and Dr Luoyp (Dublin Mag. Observ., vol. i. p. 91, foot note) have also both objected to
this process.

+ Proc. Roy. Soc. Lond., vol. x. p. 481.

¢ The test for determining whether a day is one of marked disturbance has been obtained in
different manners. I believe that the most certain test would be one depending on the value of the

738 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

There are objections to all these methods, in as far as the rules for rejection
or substitution are empirical, and have really no scientific basis. The objections
to the second method are probably the least important, since the interpolated
values depend on those preceding and succeeding; and in this respect the
operation is somewhat similar to that, where the observed quantities are
deduced from photographic registration, by drawing a line among the
points.

9. It has been stated that the third method, that of rejecting all the obser-
vations in days considered disturbed, was that employed in the first discussion
of the Trevandrum observations. It was, however, found, after the discussion
had been performed, that nearly all the days rejected were simply days much
affected by the lunar action; the lunar diurnal. variations probably amounting
on some occasions to upwards of five minutes, while the mean solar diurnal
variation did not exceed the half of that amount. In consequence of this
discovery, the discussion was made finally including all the observations.*

10. The differences, obtained by deducting the hourly means from the obser-
vations for the corresponding hours, which are equally positive and negative
for a given solar hour during a month, are still affected by the whole lunar
action ; so that they will in a given day be positive or negative, more or less
positive, or less or more negative, according as the lunar action tends to increase
or diminish the deviation of the needle. These differences which are required in
the discussion for the solar disturbance serve also for the determination of the
lunar diurnal variation, as well as the observed values from which they are
derived, with the advantage, that having the mean solar diurnal variation
already deducted, they can be combined more readily with reference to any
given argument.

11. The supposition that the law of solar action is constant throughout a
month can be considered only approximately true: near the magnetic equator,
especially in the months near the equinoxes, the law varies rapidly. To avoid
as far as possible any error due to this cause in the discussion of the Trevandrum
observations, the mean solar diurnal variation was calculated corresponding
to each week in each year in the following manner :—The hourly means were
obtained from the observations in the 1st, 2d, 3d, and 4th weeks of the year 1854;
from those in the 2d, 3d, 4th, and 5th weeks; from those in the 3d to 6th weeks;
and so on to the end of the series in 1865. The means obtained from the first
series of four weeks were then combined with those from the second series, and
the means derived from this combination were considered to represent the mean

mean difference obtained by comparing each hourly observation with that immediately following it ;
the characteristic mark of what is termed a disturbance being the irregular movement of the needle.

* With a single exception (in about 80,000 observations) in which the mean of the preceding
and succeeding observations was substituted.

—

MAGNETIC DECLINATION AT TREVANDRUM. 739

solar diurnal variation in the middle (or third) week. These hourly means were
then compared with the observations at the corresponding hours in the third
week, and the differences obtained. The hourly means from the second and
third sets of four weeks served in a similar manner for the observations during
the fourth week. In a like way the differences were found for all the observa-
tions throughout the series.

12. It is still supposed that the hourly means obtained from the observa-
tions in four successive weeks represent the solar diurnal variation unaffected
by any inequality of the lunar action. The following discussions make it pro-
bable that the mean action of the moon at each solar hour during a Iunation
is constant, or so nearly so, that any error due to this cause may be
neglected.

Mean Lunar Diurnal Variation for each Month in the Year.

13. As the diurnal variation of the magnetic needle produced by the moon
has generally been found small, it has been usual to combine the results from
all the lunations throughout the year, in order to destroy the effects of disturb-
ing causes which still remain, even after a considerable portion of the observa-
tions has been rejected for this object. Although this combination may give
approximations to a general law in high latitudes, it cannot be expected to do
so near the equator; the first discussion of the Trevandrum observations
having shown the law to be inverted in the course of six months; the combina-
tion for that locality of all the observations made during the year, gave a result
which is purely arithmetical, without any representative in fact. As nothing was
known as to the mode in which the law changed from one form to another, it
was necessary to determine it for each month in the year. This has now been
done by the discussion of nearly eighty thousand hourly observations made
during the eleven years, 1854 to 1864. The resulting values, which will
be found in the first volume of the Trevandrum Observations, with a more
detailed account of the reductions, are projected in curves in the first half of
Plate X XVI.

14. As these curves present some slight irregularities, it is believed that an
elimination of the effects of disturbance, as far as they are independent of the
lunar action, will be best made in calculating the most probable variation from
the deduced quantities by the formula of sines,

y = 4, cos 0+ b, sin 0 + a, cos 20 + b, sin 20.

The following are the resulting equations for each month, together with the
probable error of each result as derived from the four terms only:
VOL, XXVI. PART IV. 9G

740 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

Jan. y= — 0:0233 cos 6 — 0:0143 sin 0 + 0:2264 cos 20 — 0°0442 sin 26 Prob. error 0-018
Heb, = +0196 oP poe SES ers ketene ‘ ‘O11
Mar. ee ee er ddd ee EE ees é O14
Apel ==. 0026, 4) = GOlO 2 Pe Oseen mee (OO tmee . 009
May 4. «9406, eS “01Se Sores” | 2 o2089 t, i 013
June = (70045) (=) OID, ie NSBR, = Ost L ‘ O13
July = O18 . -h0sis — = 0eo7 Uo a oss0 ‘ 012
Aug. = 0001 , == +0085) “= -pse6 =, “SE =0sye : ‘017
Sept!) HE 0nd Lueeitones 4 sess ee, Va!) oped, i 010
Oct. = A00d6 or =e O06 eV Th ae oOo iii
Nov. SE OUTS By = OOM se W904 4 — 0008 ,, . 014
Dec. 4: HOBBS |g) = OBR a 740! L,,. + ae e0g08 Ay f 012

The following are the equivalent equations for the diurnal and semi-diurnal

periods, or,
y =a, sin (6 + G) + a, sin (20 + ©).
Jan. y = 0:0273 sin (6 + 238 24) + 0-2307 sin (204101 3)
Feb. y= 0151 sin (0 + 236 14) 4+ °1244 sin (20+ 111 50)
Mar. y= +0697 sin (0 + 204 35) +4 +1038 sin (20 + 139 32)
April y= °0126 sin (0+ 167 45)+ -0980 sin (20 + 156 47)
May y= *0428 sin(@+ 108 12)+4 0280 sin (20 + 223 30)
June y= ‘0149 sin (64197 28) + +0538 sin (20 + 261 12)
July y= ‘0343 sin(@+ 338 3)+ ‘0898 sin (20 + 292 55)
Aug. y= ‘0089 sin(9+180 38)+ ‘0681 sin (26 + 303 45)
Sept. y= 0575 sn (90+ 10 25)+ 0624 sin (20 + 324 58)
Oct. y= °0321 sin(@+230 4)+ ‘0459 sin (20+ 64 38)
Nov. y= ‘0631 sin(@+ 165 32) + -0904 sin (264 90 30)
Dec. y= 0481 sin (94+ 132 18)+ +1766 sin (20+ 80 7)

15. The conclusions from the observed and calculated variations are as
follow (see Plate X XVI.) :—

1st, The mean lunar diurnal variation consists of a double maximum and
minimum of easterly declination in each month of the year.

2d, In December and January, the maxima occur near the times of the
moon’s upper and lower passages of the meridian; while in June, they occur
six hours later, the minima then occurring near the time of the two passages of
the meridian.

3d, The change of the law for December and January to that for June and
July, does not occur as in the case of the solar diurnal variation, by leaps in
the course of single months (those of March and October), but more or less
gradually for different maxima and minima.

16. But the change of hours for the maxima and minima will be better seen
in the following table, where the epochs derived from the computed variations,* ©
as well as those deduced from the observed quantities are given.

* The epochs for the diurnal and semi-diurnal periods are obtained directly from the values of c,

and ¢, in the preceding series of formule; those for both terms are obtained from the formule by the
equation

MAGNETIC DECLINATION AT TREVANDRUM. 741

TABLE I.—Epochs of Maxima and Minima of easterly Declination in the Lunar Diurnal
Variation for each Month, as derived from the calculated Variations and by estimation
From the Projection of the observed Means.

Diurnal term. Semi-diurnal term. Both terms.

Monta. Calculated. Estimated from observations.

eens cy || been he ine ems he meio. meh. one neem, Ihe om, | hy m. he mh. om:
Jan. |14 6| 2 6/11 38/17 38/23 38) 5 38)11 42/17 44/23 34) 5 33/11 40/17 44] 23 55
Feb. |14 15} 2 15/11 16/17 16/23 16} 5 16}/11 17/17 21]23 11] 5 12/11 14/17 35/93 5
Mar. |16 22] 4 22/10 21/16 21/22 21; 4 21/11 0/16 21/21 43) 4 21/10 55/16 50/21 45
3 3
1 2

CONnwWRaAM>
or

April |18 49} 6 49) 9 46/15 46]21 46) 3 46/10 0j)15 41/21 41 52/10 5)15 22)21 50

May | 22 48/10 48) 7 25/13 25)19 25 25) 6 O|12 48/20 26 58] 5 152) 11 152) 20 40

June} 9 8/21 8] 6 23/12 23/18 23) 0 23) 6 30/12 0/18 11] 0 30] 6 15/11 20/18 35 3
July | 6 52/18 52; 5 8]11 18/17 18)23 18) 5 25}11 33/17 0)|22 56/ 4 50]12 15/16 20

Aug. |/18 0} 6 O| 4 52/10 52/16 52) 22 52) 4 50/10 45/16 55/23 O| 4 20/12 30/16 40/23
Sept. | 5 18/17 18) 4 10/10 10/16 10)22 10) 4 23/11 7/15 50}21 23) 4 50/11 30/16 10/20 55
Oct. |14 40} 2 40; O 51] 6 51/12 51/18 51} 0 29} 6 8]13 17/19 30/23 40] 6 40]13 30/18 15
Nov. |19 6] 7 6/23 59] 5 59/11 59/17 59|23 22} 6 8/12 40/17 47/23 30] 5 30/13 25/18 30
Dec. |21 11} 9 11] O 20) 6 20/12 20/18 20} 0 9} 6 30/12 32/18 9/23 45) 6 25/12 251/18 35

17. No conclusion of any value can be drawn from the epochs for the
diurnal term; the irregularity shown in the hours for the different months for
this term, probably depends on causes which will be noticed hereafter (35).
The semi-diurnal term which gives the largest oscillation, excepting in May and .
October, shows a rather regular change of the epochs from one month to the
other, the most marked exception being from September to October, when the
change is3h.19m. It is, however, the whole variations that we have at present
especially to consider, and for these the calculated epochs are the most certain.

18. If we commence with the maximum, which occurs in January near the
moon’s upper passage (23 h. 34 m.), and follow the change of hour for this
maximum till June, when it happens six hours before the upper passage, and
thence to December, when it is near the lower passage (thus going through
twelve hours in the twelve months), we find that the greatest leaps occur from
May to June (21 h.), and from September to October (25 h.) In the months
of March and April this maximum occurs about 2 h. 20 m. before the upper
passage, and in September 3 h. 50 m. after the lower passage.

On the other hand, the maximum which happens near the inferior passage
(11 h. 42 m.) in January, changes suddenly four hours from April to May, and
from September to October.

The two minima change hour with considerable regularity, with one marked
exception, which occurs again from September to October.

On the whole, we can conclude that though the maxima occur near the
upper and lower passages in December, and at six hours after these epochs in
June, that they happen nearly midway between the two in April and September.

where a is the correction to an estimated epoch of the maximum or minimum. See footnote to a
paper on the “ Horizontal Form of the Earth’s Magnetism,” Trans. Roy. Soc. Edin., vol. xxii. p. 529.

742 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

19. 4th, While the lunar diurnal variation changes the hours of maxima and
minima more gradually than the solar diurnal variation, it also makes the
greatest changes at different times. Thus the solar diurnal variation changes
completely during the month of March, or from February to April, while the
lunar diurnal variation makes the greatest change from April to May. The
second change which happens for the sun between September and November
occurs earlier, or between September and October for the moon.

Range of the Mean Lunar Diurnal Variation.

20. When we examine the range of the variation in the different months of
the year, we find :—

5th, That the range is greatest in January, and least in May and October ;
the arc, including the mean diurnal variation for January, from eleven years’
observation, being nearly 0°5 (= 30"), while in May the range was 0718
(= 10"8), in October 0°14 (= 8"°4), and in July 026 (= 156).

21. It has been shown in a paper already cited,* that the lunar diurnal varia-
tion is sometimes as large as the solar diurnal variation, amounting in December
and January sometimes to upwards of five minutes of are (5’), which, allowing
for the greater inclination of the needle (or the smaller horizontal force), would
be equivalent to about 12’in England. But this great oscillation, which some-
times occurs within 24 hours, is subjected to different laws of variation which,
when the mean for a week only is taken, diminish the range so much that, for
example, in the lunation 16th December 1858 to 12th January 1859,+ the
greatest range for a week’s observations is reduced to 27, while the range for
the whole lunations is less than half that amount. When the mean diurnal
variation is derived from all the lunations occurring principally in January
during eleven years, the range is still further reduced to 0’5 nearly. This is
partly due to the fact that the lunar action does not appear to be equally
powerful in the same month in different years.{

22. 6th, The ranges of the mean lunar and solar diurnal variations thus obey
different laws relatively to the epoch of the year; the range for the former in
January being nearly double that in any month from May to September, while
the range of the latter (the solar diurnal variation) in January is little more
than half that for August.

23. Although it would be difficult to prove from the ranges of the solar
diurnal variations observed at different stations on the earth’s surface (even

* Trans. Roy. Soc. Edin., vol. xxiv. p. 673.

+ See the projections for this lunation, Plate XLIII., Trans. Roy. Soc. Edin., vol. xxii.

t It is also partially due to the mode of combination—a lunation being considered im January, if
fifteen days were in that month, the other fourteen being in February or December, for which months
the range is considerably less than in January, and the maxima and minima occur at different times.

MAGNETIC DECLINATION AT TREVANDRUM. 743

when reduced to the same directive force) that the solar action is greater in
December than in June (for the whole earth), yet it seems not improbable that
the great lunar action observed at Trevandrum during the months of December
and January is connected with the greater proximity of the earth and moon to
the sun at that time of the year.

Changes of the Lunar Diurnal Variation with the Moon’s Declination.

24. The next question of importance which presented itself was that relating
to the moon’s passage from one hemisphere to the other. The first investiga-
tions in answer to this question seemed to show, as already mentioned, that the
law of lunar diurnal variation not only varied with the moon farthest north and
farthest south, but even with the position on the equator according as she was
moving towards the north or towards the south. There was nothing contrary
to our knowledge in this result, the solar diurnal law for March not resembling
in any way that for September or October. This discussion, which had been
made for the two groups of months, April to September, and October to March,
from the observations in 1854 to 1858, was extended to the eleven years’
observations, and to each month of the year. The results obtained were not
consistent. The change of law which appeared due to the moon’s passage from
the southern to the northern hemisphere in January was not the same as that
shown for the same movement of the moon in February, and resembled still less
that which occurred in March. The discussions (each requiring the combina-
tions of nearly 80,000 quantities) made in a similar manner with reference to the
moon’s phase and distance from the earth threw no light on this discrepancy.

25. In another discussion with reference to the law of disturbance as related
to the moon’s hour angle,* it occurred to me that it would be desirable to com-
pare the result which might be deduced from the night hours, when the sola¥
disturbance is least at Trevandrum, with that derived from the day hours, when
the solar disturbance is greatest. The conclusions arrived at from this discus-
sion, together with a consideration of the curves representing the apparent
variation of the law of lunar diurnal variation with the moon’s declination,
induced me to employ a similar analysis for the lunar diurnal variation. As
the conclusions from this investigation are most important, it is necessary to
describe the process made use of. The apparent effect of the moon’s change of
declination will be considered thereafter.

Change of the Law of Lunar Diurnal Variation with the Day and Night.
26, Having subducted the solar diurnal variation from the hourly observa-

* See Proc. Roy. Soc. Lond., 20th June 1861, The law of lunar disturbance is that relating to
the greater or lesser irregularity of the position of the needle in the diurnal variation for the moon on
the different meridians.

VOL. XXVI. PART IV. 9H

744 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

tions in the different weeks, the differences due to the lunar action (and irre-
gular causes) were arranged in columns under the lunar hours to which they
respectively belong, as in the following scheme, where the short lines indicate
the + or — differences.*

Lunar|#- | 8. |. |8./8./8./8./8./8./a.)8./a.)8.]/a.)/a./a./a./a]/a./a/a}/e.f/af]alasda
day, }9/1/)2/3)4/5)6]7 ) 849 |10/11}12/13)14)15]16}17/18)]19| 20/21) 22/93/24] 0
ira fe | ae pas eae | ea 2| eey (e be ||| ele es ee ee, Me) ee eae ai

ge (ea (eRe MN Oe ere A |
Bt el he ee ee ee
4 —|—}—|—|—]-]|—|-]-]—|-|-|—|—|-,=)-}-—|-/-}-—|-]-}-|-}—
5 se Fa ie be (on! pa efron ee ee elie =)
6 j—|—|—}—j—)—]—|—]—}—} —] -] -= |} —] -]—) — || —}-]-}]-}-J-Je
fa (a ey a ee | 2 PP eS eee ee ee ee
EEEEEREF EERE REESE REESE
.9 J—|—}—}—|—-}-|—|-}—|}-—|=J=]—|-|}-|-|- | — || — |
1 |-/-|-|-|-|-|-/-|-|-I=|-|-|-|-|-|- -|-|-|--=)-|-|-
98 FEE NN JE ee

The first difference under 0 h. with which the lunation begins is that derived
from the observation made half an hour after noon (the lunation always begin-
ning with the new moon), and corresponding to the moon on the meridian.
The difference under 6 h. in the first lunar day corresponds to after sunset, while
that under 17 h. corresponds to before sunrise. Thus the differences before and
after sunset are separated by a light zigzag line, while those before and after
sunrise have a thicker zigzag between them.

It is obvious that, if each horizontal line represents the lunar diurnal varia-
tion during a lunar day, and if this variation obeys the same law, or nearly the
same law, throughout a lunation, it does not matter how we obtain the means
from the vertical columns of differences; and the means obtained from the
night hours (those within the zigzag lines commencing with 6 h. to 17 h.) should
give the same result as those to be obtained from the day hours. If, however,
the law varies with any argument, such as the moon’s declination or phase, this
will be proved by a comparison of the results for different lunations in different
months of the year.

* It will be seen in the scheme that the lunar day has been considered equal to 25 hours (it is equal
to 24 h. 50 m. nearly), and consequently the lunar hour angle is equal to only 14°4. Also 0h. is
repeated in the last column. This was always done in order to find the correction due to changes of
mean declination, which caused the variations to increase or diminish from the Ist to the following 0 h.,
especially in means derived from limited series of observations. It was found ultimately that it would
be preferable to repeat 1 h., and perhaps even 2 h. in order to obtain this correction with more accuracy.
Other details as to the precautions taken in these discussions will be found in the first volume of
Trevandrum Observations.

— Ss =
ee

..
¥

4
|
4
£
Fe
4
i

>
4

MAGNETIC DECLINATION AT TREVANDRUM. 745

Thus, if the scheme just given represents a lunation in December, the moon
will be farthest south in the first days of the lunation, and the night hours will
correspond to the lunar hour angles of 6h. to 17 h. ; while for a lunation in June,
the moon will be farthest north when the night hours correspond to the same
lunar hours ; in the intermediate months, the position of the moon in declina-
tion will be different at the commencement of each lunation, and the night
hours will thus correspond to the same lunar hours for all the different positions
of the moon.

27. It has been, however, supposed that each horizontal line represents the
lunar diurnal variation in the same way ; but we know that the mean position
of the magnetic needle varies from day to day. A separate investigation has
shown that the change of the daily mean is, if not wholly independent of the
varying positions of the moon, so nearly so that its effect on this discussion
may be neglected, and the other changes may be considered as irregular varia-
tions, the effects of which will be eliminated in the discussion of a sufficiently
large number of observations.*

If, now, we obtain the means of the vertical columns for the night hours,
these means will correspond to all the positions of the moon (in declination, &c.),
a similar remark holds for the means obtained from the day hours. The sums .
having been taken for the night hours in the lunations, the whole or greater
part of which occurred in January in each of the eleven years, these were com-
bined, and the means taken; a similar operation was performed for the day
hours ; and in each case this was done for each of the twelve months. The
conclusions from this discussion are as follows. See second half of Plate
XXVI., where the derived means are projected.

28. 7th, The action of the moon on the declination needle is, in every month
of the year, greater during the day than during the night ; the range of the
oscillation in January and June, being between three and four times greater
during the day than during the night, the ratio being less in the intermediate
months.

The following are the ranges, and the areas of the curves (from observation
and computation by formule similar to those given (14) ) for day and night,
with their ratios for each month of the year :—

Jan. Feb. March. April. May. June. July. Aug. Sept.. Oct. Nov. Dec.

Range eae 085 0"47 0749 0°41 0°24 0°36 0740 0°39 0°31 0723 0°41 0769
observed, ( Night, 0°24 0°26 0°23 0°22 0714 O”11 0°22 0%18 0721 021 0°20 0'23

Ratio, — BGr ie lon ite re eo {7180 ot aap) 1 3-0
ight,

* Had the method here considered, or an idea of any probable result to be derived from it, presented
itself at first, differences, independent of the irregularities of the daily means, would have been obtained
by reducing each of the daily means to the monthly mean, so that the sums of the plus and minus
differences in each lunar day would have been equal to zero.

746 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

Jan. Feb. March, April. May. June. “July. Aug. Sept. Oct. Nov. Dec.
Area ae 6:20 3°05 2°96 2°69 1:66 2°25 218 2°28 1°94 1-01 2-34 Bacay

observed, | Night, 1°39 1°45 1°60 1°17 0°85 0°60 0°98 0°85 1°07 1°28 17-26 1744

Day,

Ratio, Nicht AS BLE 2 DO SBS? FO T 26 eS OF OS Giese
ight,

Ranges Day, 0°85 0°38 0°43 0°35 0°20 0°31 0:35 0°27 0°29 0°17 0°37 07-64
calculated, { Night, 0°20 0°22 0°19 0°18 0°13 0°07 0°14 0°09 0°16 0°16 020 0-20

: Day,

Ratio, NGC 42 417 B22 EO Lo 43 24 30 £8 deel
Oo;

Area Day, 6°29, 3:06 2°95 2°60 -1733 2°22 2°16 208 1°86 0°98 2-87 4-39
computed, | Night, 1°25 1°41 1°56 1°20 0°80 0°49 0°97 0°61 0°99 1712 118 1°39
aah Day, 7
Ratio, Nicht» 0 22) 9 BRIT £5 ° B2 34) 9 090-9 eae

fo)

The computed ratios are probably the nearest to the truth, as they are free
from the irregularities due to accidental disturbances. From the ratio for the
areas of the computed curves (for which the sums of the co-ordinates are
employed) the day is to the night as 5 to 1 in January, and as 43 to 1 in June.
The ratio is least in May and October, the two months in which the curve be-
comes inverted, the day and night areas for the latter month being nearly equal.

29. This is a fact which is wholly independent of the question as to its cause,
whether connected with the varying positions of the moon or not. The law of
variation, however, for the day hours in each month of the year is so consistent,
that it becomes exceedingly probable that the change of law previously found
as apparently connected with the moon’s varying declination is chiefly, if not
altogether due to this cause (the different action in sunlight and in the shade
of night). The law deduced from the night hours shows a variation of such
small range, that it is comparatively more affected by the irregularities due to
change of the daily mean declination and other causes.

30. If the law of lunar diurnal variation really depends on the moon’s position
in declination, then the method just described will give results, in which some of
the diurnal curves for any month will be made up of the parts of the curves
belonging to the day hours for the moon’s different positions, and others to the
parts belonging to the night hours. We can, perhaps, best judge to what extent
the moon’s declination (or longitude) is connected with these results by an exami-
nation of the projected curves derived from the discussion for the moon’s
declination.*

Thus, to begin with January, the curve for the week with the moon farthest
north shows the maximum of easterly declination, which happens near the

* Postscript.—When this was written it was intended to give the projected curves for the different
positions of the moon in declination ; this the Postscript (43) and the Plate (XXVILI.), with curves for

eadieies p se ee

MAGNETIC DECLINATION AT TREVANDRUM. 747

moon’s lower passage, to be much greater than the other near the upper pas-
sage; while for the moon farthest south in January it is just the reverse.
Now this difference is connected also with the fact, that in each of these cases
the greatest maximum happened in the day hours—that is to say, when the
moon is farthest north in January the lower passage occurs in the day-time ;
and when farthest south in January, the upper passage occurred in the day-
time.

A similar difference is shown for the weeks in January, when the moon was
near the equator, moving north and moving south ; when moving north, the
most marked minimum occurred about six hours before the upper passage, and
when moving south, the marked minimum happened six hours qa/ter the upper
passage ; both of these epochs corresponded to the day hours.

31. Had, however, the change’in the moon’s declination been a cause of
these differences in the values of the extreme elongations, we should expect
the same cause to produce similar effects in other months. If in January, the
moon being farthest north caused the maximum at the lower passage to be the
greatest, a similar result might be expected in February and March, for which
months the law of the mean diurnal variation is nearly the same. This is not
the case. In February and March the greatest maximum occurs near the
lower passage, not when the moon is farthest north, but when on the equator,
moving south.*

In June, when the moon was farthest north,t the variation may be said to
have been zero near the lower passage, instead of, as in January, being the
most important for this position of the moon. Also, when the moon was
farthest south in June, the movement near the upper passage is very small,{
which is just the reverse of the fact for the same position of the moon in
January.

32. An examination of the results for each month of the year, in the same
way, leads to the conclusion that the variable magnitude of the maxima and
minima depends chiefly, if not wholly, on the hours of the day corresponding
to the hour angles of the moon for which there is a maximum or minimum of
easterly declination. This conclusion will be evident on examining the projected

_ the times of the moon’s phases, render unnecessary ; for the conclusions here drawn the curves for the
phases may be consulted, approximately, as follows :—
In January, for moon farthest north, see curve for full moon.

a op south, ng new moon.
Fs », On equator moving north, see curve for Ist quarter.
a ms 3 south, | 7 3d quarter.
In February, on equator moving south, see 3d quarter.
In March, s Se see full moon.

* See curves for third quarter in February, and for full moon in March.
+ See curve for new moon in June.
{ See curve for full moon in June, Plate XX VII.

VOL. XXVI. PART IV. OI

748 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

curves, where the parts derived chiefly from day-hours are distinguished from
those from the night-hours.*

33. It was still a question, whether there might not be some difference in
the lunar action in the forenoon and in the afternoon, and in the first and
second halves of the night. Though the number of observations to be dis-
cussed did not seem great enough, in comparison with the irregularities to be
neutralised, to allow a quite satisfactory reply to this question, the discussion
was, however, performed for the four parts of the day.

The result for January, the month of greatest lunar diurnal variation, and
consequently, that for which any difference should be most easily perceived,
was found well marked. In January the forenoon hours showed the greatest
minimum six hours before the upper passage, while the afternoon hours gave the
greatest minimum six hours after the upper passage; and the range of the
variation, in each case, was nearly six times greater than that derived from the
night hours. But though differences are found in the results from forenoon and
afternoon hours in the other months of the year, they are never so well marked
nor neither do they resemble those for January. It is probable, then, that
this difference is due to some other cause, or some additional cause which is
felt only in that part of the earth’s orbit.

The differences in the results for the two halves of the night are too small
and irregular to found any conclusion upon them.

34, All the day hours together give variations for each month of the year
resembling those already considered as derived from all the observations day
and night, consisting of a double maximum and minimum in each month of the
year, with varying epochs.

35. The variations derived from all the night hours do not appear to change
epochs from month to month, as in those obtained from the day hours ; the law
of variation appears on the whole to be nearly constant throughout the year ;
that is to say, a maximum of easterly declination occurs in all the months of the
year near the times of the upper and lower passages of the meridian.

36. The mean lunar diurnal variations, as derived from all the observations
made throughout the year at different stations on the earth’s surface, have pre-
sented the common feature of the greatest deviation of the needle towards the
west in the northern hemisphere, and towards the east in the southern hemi-

* It will be obvious that this separation cannot be perfect, since, for example, in the vertical sum-
mation of the first seven horizontal lines in the scheme (26), when the difference of day and night
hours is disregarded (as in the discussions for the moon’s position), the mean for the lunar hour 3
would be obtained from three observations made during the day and four during the night. In the
projected results, six hours on each side of the mean midnight point are marked as night hours ; that
point would correspond to the lunar hour 84 in the means for the first week in the scheme. In this
way, however, the extremes of parts marked as night hours are still affected by the greater action of the
day. An examination of Plate XLIII, Trans. Roy. Soc. Edin., vol. xxiv. (differences under solar
hours), will show the small lunar effect during the night hour throughout a lunation.

MAGNETIC DECLINATION AT TREVANDRUM. 749

sphere, within an hour and a half of the moon’s passages of the meridian. We
see from the present discussion how these means include different causes
of variation, producing absolute inversion of the laws (at least during the day)
near the magnetic equator, and that in such positions the mean for the year
may not represent the fact at any period ; also for different latitudes, it is pro-
bable that the law will vary during the year with the different lengths of the
day and night.

3/7. It has long been known that the solar diurnal variation was greatest
during the day ; we now find this is true, also, for the lunar diurnal variation.
We might suppose that there is a greater amount of the magnetic ether heaped
up over the face of the earth next the sun than on the other side in the earth’s
shadow, and that the moon’s action upon this ether is thus greatest where the
greatest disturbance can be produced. In whatever direction the hypothetical
electrical currents proceed, their intensity (or quantity) diminishes within the
earth’s shadow, and is it not improbable that the result now found for the varia-
tion of the easterly magnetic force will be found true also for the northerly
and vertical forces.* ,

Diurnal Variation with reference to the Moon in the Half Orbits farthest from
and nearest to the Earth.

38. The only other discussion in connection with the lunar diurnal variation
which it seems to me of sufficient importance to give here, is that connected
with the moon’s distance from the earth. For this question the discussion was
divided into two parts: First, for the months October to April; and second,
for the group May to September ; in each case the law of diurnal variation
within the group is sufficiently constant to allow a conclusion from the mean
variations.

The equations of smes computed from the means are as follows :—

October to APRIL.

R y = + 00014 cos — 00249 sin @ + 071049 cos 20 — 00061 sin 20 Prob 0%
poses 1 = + 070249 sin (9 + 176° 52’) + 01050 sin (20 + 93° 19) \ POD PRIUS 10 Hn0e:

é y = — 00024 cos 6 — 0"0316 sind + 0°1114 cos 26 — 0°0494sin 20
Perigee { \

= + ("0317 sin G Soe 21’) + 01218 sin (20 ae 11S€ 54’) 33 = 0013.
May vo Sepremeer.
A y = — 0'0098cos 4 + 0-0118sind — 00446 cos26 + 00046 sin26 ) 0-013
Bes = + 0°0153sin(6 + 320° 4’) + 00448 sin (26 + 275° 57’) eee a OMS.
Peri y = + 0°0118cos6 + 00006 sin 6— 0°0578 cos20 + 0'-0236sin 20 0-008
er1lgee | = 4+ 0°0118 sin (@ + 86° 53’) xe 0’-0624 sin (20 ie 292° 13’) \ _ x

* The calculations are being made with reference to these components of force.

750° * J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

From the means of the observations, and the quantities computed by the
preceding equations, we obtain the following results :—

39. In the group October to April, the areas of the curves representing the
observed and computed variations are in the ratios—

For Apogee : for Perigee : :1: 1:18 by observation ;
a * : :1: 1:15 by computation.

While for the group May to September the ratios are—

For Apogee : for Perigee : :1: 1°31 by observation ;
s és : : 1: 138 by computation.
The ratios of the ranges of the two oscillations (from the observed quan-
tities) are approximately

October to April, : , W Neg) Site lp a L240)
May to September, . : Na tl Sei ini Lai (522)

The total ranges of the oscillations by computation are in the ratios

October to April, : : A:P=1 118
May to September, . : AtesP ae or6
40. The ratios appear greatest for the mean curves representing the group
of months May to September, but an examination of the ratios of the areas of
the curves for the separate months (which vary considerably, and from other
causes than that of distance), shows that this difference is accidental. On
taking the ratios of the areas of the observed curves for each month, and the
means of these ratios for the six months October to March, and April to Sep-
tember, we obtain—

October to March, : ail ee Pe el Oy
April to September, . : AO 3P

—
ie
i)
3c)

This agreement of the ratios is probably accidental, since when the means
of the ratios for the six months January to June, and July to December, are
taken, they are found as 1: 1:14, and as 1: 1:34 respectively. But in whatever
way we obtain the ratios, the mean for the year is always nearly the same, or

A:P=1: 1:24 nearly,

which is probably not far from the truth.

41. The ratio of the moon’s mean distance from the earth in the half orbit
about apogee is to that in the half orbit about perigee nearly as 1:07 is to 1; as
the cube of 1:07=1-23 nearly, we see that the mean ranges of the curves, as
well as the mean areas, for the two distances are in the approximate ratios of
the inverse cubes of the moon’s distance from the earth, as in the theory of the
tides.

MAGNETIC DECLINATION AT TREVANDRUM. 7ol

42. The regular double maximum and minimum shown in all the mean
results obtained for the lunar diurnal variation of the magnetic elements has
given the idea that the variations are due to the action of an attractive force ;
any such idea will now require some modification such as that suggested (37)
in order to satisfy the facts of this paper.*

POSTSCRIPT.+

43. In the preceding paper, no allusion has been made to the results when
related to the moon’s phase ; these phases coinciding nearly in each month with
one of the four positions of the moon in declination, for which the curves had
been projected, it did not seem necessary to give the curves for the phase also.
It has been pointed out (foot note to 32) that the day and night hours overlap
each other when the means for several successive days are taken, and therefore
the separation of the part of the curve derived from day hours, from that
derived from the night hours, is not perfectly definite.

This, however, is the case in these combinations, not only from the mode of
combining several successive days, but also because, for example, in the month of —
January, the moon does not pass the meridian at the same hour for the day she
is farthest north in different years, the difference amounting to upwards of three
hours in the course of the eleven years. It seemed desirable, then, to avoid this
latter complication, in order to show more distinctly the difference betwixt the
movements during the day and during the night hours. This was best done in the
combinations for the moon’s phase, since the moon is on the meridian at noon
nearly on the day of new moon, and at midnight nearly on the day of full
moon ; the only indefiniteness remaining is that common to both combinations,
depending on the overlapping in the means of successive days already noticed.

After communicating the results of the preceding paper to the Royal Society,
I projected the curves for each month corresponding to the new moon, first
quarter, full moon, and third quarter. That is to say, the means derived from

* The mean diurnal variations of the atmospheric pressure due to the sun and moon, show similar
regular double maxima and minima. The consideration of a mass of observations made by me in India
at different heights, induced me several years ago (Phil. Mag., August 1858) to suggest an electro-mag-
netical attraction as a cause of these variations (and I found an analogical phenomenon in the action of
the sun on the gases of comets). The atmosphere having acquired a certain polarity, would by this attrac-
tion assume ellipsoidal forms with the longer axis directed towards the attracting luminaries (or making
nearly constant angles with these directions), thus producing waves or currents as a given meridian
turns round under axes of different lengths. I have in consequence of this idea sought to find some
connection between the laws of terrestrial magnetism and those of atmospheric pressure, but hitherto with-
out any conclusive result. The difference of the lunar action on the magnetic needle during the day and
during the night, suggested the examination of the lunar diurnal variation of atmospheric pressure for a
like difference ; but the result of this investigation was negative, no difference being perceptible ; the lunar
diurnal variation in January and June was shown equally well by the observations during the night
and by those during the day.

+ Added by permission of the Council, June 27th 1872.

VOL. XXVI. PART IV. 9K

752 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

combining the lunar diurnal variations for each of these phases with those for
three days before and three days after them.

44. The curves for the new moon have been projected in thin lines, and
those for full moon in thick lines on the same zero line (see Plate XX VIL) ;
similarly for the curves for the first and third quarters in thin and thick lines
respectively. In each case the night part of the curve is distinguished from
the day part, the former being a dotted, the latter a continuous, line.

45. The principal conclusions to be drawn from these curves are the follow-
ing :—

1st, The great difference of the amount of movement during the day and
during the night is shown distinctly in each case.

2d, If we consider, in the first instance, the curves for the new moon, we
find in the months of January to May, that the north end of the needle changes
its direction of motion, and begins to move rapidly eastward at sunrise (almost
_ exactly at sunrise). This holds true till new moon in May; but in June the
direction is completely reversed, the north end of the needle turning westwards
shortly after sunrise ; this continues in July, August, and September, but at the
new moon in October the reversal again occurs more distinctly than in May :
the needle now moves rapidly eastward after sunrise till June again.

3rd, The same statement holds for full moon, only that the movements in
May and October are combinations of those for the preceding and succeed-
ing months. I have no doubt that the change would be shown equally well and
equally rapidly from one direction to the other, if the full and new moon always
happened at exactly the same points of the earth’s orbit, but these results are
derived from combinations of observation for full moons occurring in the begin-
ning and in the end of May and October, and therefore partaking partly of the
character due to the preceding and following months.

4th, When we examine the curves for the first and third quarters, which are
projected together, we find for both quite similar results to the preceding, only
that the motion after sunrise is towards the west in January and towards the
east in June.

46. Thus, if sunrise happens when the moon is near the meridian of 18 h.
(that is, at new moon, the sun rising always near 18 h. at Trevandrum), or
near the meridian of 6 h. (full moon), the needle then turns from its previous
direction and moves rapidly (relatively speaking) eastwards in the months from
October to May, and westwards from May to October. When the moon is on
the meridian of 12 h., or 0 h. at sunrise, the directions of movement are the
reverse of the preceding. That is to say, whether a maximum or minimum of
easterly declination should happen at sunrise, the movement following takes
place comparatively rapidly.

47. If we had found previously that a minimum or a maximum always hap-

MAGNETIC DECLINATION AT TREVANDRUM. (OS:

pened when the moon was on one of the meridians of 0h., 6h., 12h., and 18h.,
there would have been nothing remarkable in this result, it would have agreed
with previous ideas of the mean law of lunar diurnal variation ; but this is not
the case, the maxima and minima occurring at hour angles differing from the
above by three hours in some months, and changing hour with more or less
regularity from month to month. It could scarcely, therefore, be supposed to
be a coincidence dependent on the true law of variation.

48. At least two questions present themselves in reference to this result :
First, How this constancy of maxima and minima for fixed hour angles of the
moon can agree with the fact of the change of hour shown (18) in the means
for each month? We find an explanation of the apparent contradiction when
we examine the curves for the phases, and consider the maxima and minima
which do not happen at sunrise. Thus, the curve for new moon in the months
of January to May shows that the principal maximum gradually shifts from an
hour before the upper passage in January, to 3 h. before it in April and May.
Similarly the passage of the minimum (in the same curves) for the moon on the
meridian of 5 h. in January, to on the meridian of 0 h., though not equally regu-
larly, is clearlyshown. The change for the corresponding minimum in the third
quarter happening more gradually from 53h. in January to 0 h. in July.

49. The next question has reference to the mode in which the moon acts on
the needle, on days intermediate betwixt two of the quarters; when, for example,
she is on the meridian of 17 h., 16h., 15 h., at sunrise, how does the movement
change from that shown in the thin curve in January (new moon), where the
movement is rapidly eastward after sunrise to that shown in the thin curve
for the first quarter, where the movement is rapidly westwards at the same solar
hour? Does the epoch of maximum or minimum vary at all with sunrise, or
does the coincidence seen in these curves hold only for the four positions of
the moon to which the curves strictly belong ?

50. In order to answer these questions, the consideration of one period of
seven days will suffice ; for this end twelve periods from new moon to the first
quarter in the months of December and January, showing the greatest varia-
tions, were chosen. In each of these periods the sum was taken of the diurnal
variations for each hour in the week, having new moon in the middle; the sums
of the twelve sums thus found having been formed, the means (from seventy-two
days) gave the mean variation corresponding to new moon. A similar calcula-
tion was made for the weeks having the moon, one, two, to seven days old
in the middle. These means are projected, Plate XX VII.

51. It will be seen from these curves, that the sunrise seems to pass from
the minimum near 18 h. to the maximum near 12 h., gradually and without
having any marked influence on the epoch of minimum or maximum. These,
however, are mean results, the curve for each day being derived from three days

754 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

before and three days after the day of the moon’s age which it represents ; and
we have seen in the results of this paper, the danger of trusting in mean results
where different laws may be in question, and one movement overlies another.
It was sought then to determine with more distinctness the change of form
from one day of the moon’s age to another, and for this end, the lunar diurnal
variation for the twelve single days of new moon was obtained, and that for
each of the days for which the moon was, one, two, to seven days old.*

52. Though the number of days is scarcely sufficient to give means free from
irregular disturbances, the results appear little affected by them; they are pro-
jected in the second line of curves, Plate XXVIII. Here we find that the
minimum takes place at sunrise on the day of new moon, and next day; but
when the moon is two days old, the minimum is half an hour after sunrise; on
the third day it is an hour and a half after sunrise, and there appears an inflexion
in the movement westward; on the fourth day the movement has already turned
westward at sunrise, which is the case also on the fifth and sixth days; on the
seventh day the movement westward begins only one and a half hour after sunrise.

53. These results appear then to show, that the turning point is, as it were,
attracted by sunrise, and that the change from an easterly movement to a
westerly movement at sunrise occurs within an interval of about two days.

54. So curious and unexpected a fact required a still more careful arrange-
ment of the observations. On the days of new moon the observation near
noon was always marked as 0 h. for the moon on the upper meridian; the next
observation as the moon on the meridian of 1 h., and so on. As, however, the
moon is not generally on the meridian at the time of the observation nearest
noon at new moon, being sometimes half an hour before or after, an error of an
hour occurred easily in the lunar hour corresponding to the following observa-
tions, so that in the combinations considered previously, the observation nearest
sunrise was not always entered under the true lunar hour angle for a given day’
of the moon’s age. Another method then was now employed, in order to avoid
any error from this cause, and to insure that we have the exact effect produced
at sunrise.

The observations nearest sunrise were made at 5 h. 58 m. a.m. in 1854 and
1855, and at 6 h. 28 m. A.M. in the following years.t In order to avoid any com-
plication due to this difference of hour, the observations in 1854 and 1855 were
not employed in the following combination. All the periods for which the new
moon occurred, between the 10th December and the 20th January (the time

* As no observations were made on Sunday, there were twelve days’ observations for new moon,
and for the moon four and seven days old, eleven days for the moon six days old, ten days for te
moon three days old, and only nine days for the moon one, two, and five days old.

+ The time of sunrise varies at Trevandrum from 6 h. 8 m. .?., on the 15th December, to 6 if
20m. on the 15th January, a change due nearly altogether to the equation of time, the interval betwixt
sunrise and true noon, being (refraction effect included) nearly 5 h. 48 m. during the whole month.

MAGNETIC DECLINATION AT TREVANDRUM. 759

of the greatest lunar action), in the years from January 1856 to January 1865
were taken (with the exception of that having new moon the 16th December
1857, large disturbances having occurred on the 17th, 18th, and 19th).

The moon’s hour angle corresponding to 6 h. 28 m. A.M. (mean time), was first
computed to the nearest minute for each of the days from new moon to the first
quarter, for each of the seventeen periods under discussion. The days on
which the moon’s hour angle was between 18°5 h. and 17°5 h. (at the above
hour) were marked as 1d.; those in which the hour angles were betwixt
17°5 h. and 16°5 h. as 2d.; and so on to 12°5 h. and 11°5 h. The variations
due to the lunar action were now arranged for 1d. under each other, according
to the solar hours (beginning with 12 P.m.), so that all the observations made at
the sunrise hour (183) were under each other; the means of the vertical
columns were then taken, and that at 184 h. corresponded to the mean of the
calculated hour angles of the moon (= 18°06h.) A similar calculation was
made for each of the following days for which the moon’s mean hour angle differed
by an hour at 185 h. mean solar time. In this way any possible error is due to the
variation of the moon’s hour angle (differmg on the average a quarter of an
hour from the mean) corresponding to 184 h., and this error the previous results
have shown to be of less importance than that due to overlapping of the obser- -
vations before and after sunrise. The results of these combinations are given in’
Table II., and they are projected in the third line of curves of Plate XX VIIL.,*
the observation near sunrise (65 A.M.) having an O round the point, and that
near sunset aX.

55. In this more accurate combination, we find that the needle’s movement
westwards on the first and second days ends at sunrise; on the third day, the
greater part of the movement westwards takes place after sunrise; on the fourth
and following days, the movement westward begins at sunrise.

56. There is here, however, an additional fact of importance which presents
itself on the fourth day: the movement is diminished so much that the variation
has the appearance of being nearly obliterated. The moon’s hour angle on this
day at the sunrise observation is 15 h., and it seems as if, in this middle position,
the needle were solicited by opposing forces.

57. The fact then is established, that the direction of the movement changes
at or near sunrise, and that near the time when the needle emerges from the
earth’s shadow into sunlight the moon’s action is reversed. This sudden
reversal bears some resemblance to that shown in the curves from April to May,
or May to June; for example, in the curves for the first quarter (Plate XX VIT.)

* It must be remembered that these curves cannot be compared with those preceding them, day
by day; 1d. (under Od. in the plate) does not correspond to new moon (but to half a day or more after
it), and there are only six days here included between the day for themoon on the meridian near
noon, and the moon on the meridian at 6 h. p.m., whereas seven days were included by the other method,

VOL. XXVI. PART IV. 9L

756 J. A. BROUN ON THE LUNAR DIURNAL VARIATION OF

for April and May. In the former month, the needle moves westwards, in the
latter eastwards from the hour when the moon is on the inferior meridian
(sunrise).

TABLE Il.—WMean Lunar Diurnal Variation on seven successive days, for the Moon on the Meri-
dians of 18h, to 12h. at 64 am, in January and December, derived from seventeen
quarter Lunations, 1856-65.*

Bolen ut Pag: 2d. 3d. 4d. 5d. 6d. 74.
12taw | — 0:05 |. — 013) + 0-07 | = 0-06 | — 0-16.) —0-11 0-05
14 ,, .| — 0-06 | — 0-10 | + 0-13 | + 0:03°| — 0-16 | — 0:09 | — Gas
24, | —.0°07 | — 0:07 | + 0:07 | 4+.0:03 | — 0:07"| — 0:00} + 0-02
34, | — 0:10 | — 0°14 | + 0°16 | + 0:04 | + 0:01 | + 0:09 | — 0:05
Ae | 10:13) = O14 +0708. |* 20-06") 42 0-05) | = 0-10 | a-e aot
bt 4, | — 0:35 | — 0°20 | — 0:06 | + 0°04 | + 0:14 |} + 0-20'] + 0:24
64 ,, | — 0°69 | — 0-55 | — 0-24 | 4+ 0°06 | + 0-40] + 0-48 | + 0°54
TE, | 062) = 0-5L| — O57 | = O29) 09 e037 Vee
84 ,, | — 0°32 | — 0°38 | — 0:82 | — 0-25 | — 0°10 | 4+ 0°16 | + 0°39
92 ,, | + 0:15 | + 0:08 | — 0°72 | — 0°18 | — 0-48 | — 0°35 | — 0-06
104 ,, | + 0°48 | + 0-11 | — 0-40 | + 0°13 | —'0°91 | — 0°58 | — 0°64
112 ,, | + 0-43 | +.0:23 |— 0:17 | + 0°26 | — 0°66 | —10-52 | — 0-79
OL pm. | + 0°31 | + 0-40 | + 0°08 | + 0:25 | — 0°31 | — 0:36 | — 0-61
14 ,, | + 0-11 | +-0-34 | + 0:46 | + 0:09 | — 0-44 | — 0:08 | — 0°55
2k 4, | — 0-20 | + 0°18 | + 0-45 | + 0-11 | — 0°38") + 0°36 | —-0-35
3k | —-0:37 | — 0:23 | + 0-29 | + 0-12 | + 0-22 | 4 0-58 | +4 0-07
4h, | — 0°43 | — 0:44 | — 0°04 | + 0°12 | + 0°47 | 4+ 0°51 | +4 0°47
Bk. | — 0°38] — 0-62'| = 014 | —"0-09"| 4 0-26 | + 0-24 | == 02s
64 ,, | — 019 | — 0:15 | — 0:10 | — 0-12 | + 0:05 | + 0-12 | + 0°10
7 | Or | = 0-20) |, — O11) = Orr G-02) | 10:04 es Os
si ,, | — 0-20 | — 0-11 | — 0°08 | — O11 |! — 0:04 | + 0:05 | + 0-04
94.5, | = 1012 O07 | — (0:07 I= 00s = 0:16) PE O03) | E1002
104 ,, | — 0:09 | — 0:05 | — 0:13 | — 0:09 | — 0°03 | + 0-02 | — 0:05
11%, ‘| —O-T1 | — 0-09, | — 0°09 | — 0-15 | — 0-05 | — 0-070) — O22
124. 5, | —2O*t 18 0°06), = 07044 ="0-189/ = 10-19 |= .0-10 4 | Os
Mean hour
angle of h. h. h. h. h. h. h.
the no 18-06 17-04 16:07 15°05 14:05 13°10 12°05.
at 64 A.M.

58. If we now examine the epochs near sunset, we find that on the first day
the needle has turned (though little) an hour before the sunset observation; on
the second day it turns at the sunset observation; on the third day the move-
ment is arrested, or nearly so, from sunset to sunrise; on the fourth day the
movement westwards ceases at sunset; on the fifth day it has begun to move
eastwards an hour before; on the sixth, two hours before; and on the seventh,
one hour before the observation nearest sunset. I think there is little doubt
that there is a relation betwixt the direction of movement and sunset like that

* The means for 123 a.m., are calculated for the end of each day as well as for the beginning, in

order to connect in the projection the value for the end of each day with that for the beginning of the
next.

MAGNETIC DECLINATION AT TREVANDRUM. 757

for sunrise, but not so well marked. The curves, Plate XX VII., for the moon’s
phase seem to show that the turning point occurs generally within two hours of
sunset, but these are derived from the superposition of successive days, and are,

- in consequence, less fitted to show the facts exactly than the curves from the
single days under consideration.

It is instructive to remark the differences of the results from the various cal-
culations. In the last few single days we see how the needle behaves under
the influence of the moon on different meridians, in different circumstances of
day and night, of sunrise and sunset. In the former, we have results which
always diverge more from the true movements, the greater the number of
successive days or months which are included in the calculations.

Trans. Roy. Soc. Edin*

Vol. XXVI, Plate XXIX.

M° Farlane & Erskine, Lith’ Edin?

P. Thomson, photo.

Vol. XXVI, Plate XXX.

Uy ‘aUrysig 4 aUELeT SW

‘oloyd ‘uosuiouy, J

Trans. Roy. Soc. Edi

[ 759 ]

XXV.—On the Occurrence of Ziphius cavirostris in the Shetland Seas, and a
Comparison of tts Skull with that of Sowerby’s Whale (Mesoplodon Sowerbyi).
By Professor TurNER. (Plates XXIX., XXX.)

(Read 20th May 1872.)

CONTENTS.
PAGE. PAGE,
Historical Sketch of Ziphius cavirostris,. 759 Historical Sketch of Sowerby’s Whale, , 771
Description of Shetland Ziphius, . Ol Description of the Skull in the Museum
Comparison of the Shetland Ziphius with of Science and Art, Hdinburgh, . oo Ur
previously recorded specimens, . . 768 Comparison of this Skull with previously
recorded specimens, . : : 5 HES

The illustrious CuVIER, in his treatise “ Sur les Ossemens fossiles,”* described
and figured an imperfect skull which had been obtained, in 1804, by M. Raymonp.
GorRSSE in the department of Bouches-du-Rhone, near Fos, on the southern
coast of France. It had been found on the sea-shore in the preceding year
by a peasant. Cuvier recognised it to belong to an undescribed genus of
cetaceans, to which he gave the name of Ziphius; and from the deep hollow
which it possessed at the base of the rostrum, he named it Ziphius cavirostris.
From the condition of the bones, and the general characters of the specimen,
he judged it to be a fossil. Cuvrer’s description, though brief, and from a
mutilated specimen, yet clearly states the most salient features of the skull.

This idea of the fossil nature of the cranium prevailed amongst zoologists until
1850, when M. Pavut Gervais carefully re-examined Cuvier’s specimen, and com-
pared it with the somewhat mutilated skull of a Ziphioid whale, which had been
stranded, in the month of May 1850, at Aresquiés, in the department of
Hérault, on the Mediterranean coast of France.t From this re-examination
and comparison he not only pronounced CuvIER’s specimen to be not a fossil,
but that the skull of the animal stranded at Aresquiés was specifically identical
with the cranium from Fos; and since that time the non-fossil nature of the Fos
specimen has been generally admitted. :

* Tome v. premiére partie, 350, fig. 3, pl. xxvii. Paris, 1825.

+ Annales des Sciences Naturelles, 3d series, xiv. 1850; also “ Zoologie et Paléontologie Francaises,”

1te ed. p. 154, et 2™¢ed. p. 287. M. Gervais has, in order to give an opportunity for making a com-

parison, reproduced figures of CuviEr’s specimen, both in his “ Zool. et Pal, Frangaises,” and in pl. xxi.
of the “ Ostéographie des Cétacés.”

VOL. XXVI. PART IV. 9m

760 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

The specific identity of these two crania, however, has been called in ques-
tion by more than one zoologist. M. Duvernoy regarded the skull from
Aresquiés as more closely related to the genus Hyperoodon, and named it
Hyperoodon Gervaisi.* M. Fiscner considered it to be another species of the
genus Ziphius, and named it Z. Gervaisz ;+ and Dr. J. E. Gray has associated it
with the genus Epiodon, and termed it Epiodon Desmarestit.t

In 1864 the cranium ofa Ziphius was found at Lanton, on the shore of the
Bay of Arcachon, on the Atlantic coast of France. It has been preserved in
the Museum of the Scientific Society of Arcachon, and has been carefully
examined by M. Fiscuer,§ who pronounced it to be specifically identical with
Cuvier’s specimen. The skull is in a good state of preservation, but the lower
jaw is wanting.

The attention of naturalists in France having thus been particularly directed
to the occurrence of one or more species of Ziphius in the adjacent seas, a care-
ful examination of zoological literature has been made with the view of ascer-
taining if any cetaceans had been described by other naturalists which could be
referred to the same genus. MM. Gervais, DuvERNoy, and FIScHER agree in
considering that a cetacean, stranded on the shores of Corsica, and described by
M. Dovumet in 1842|| as a Hyperoodon, is really a member of the genus Ziphius.
Fortunately the skeleton has been preserved by M. Doumet at Cette, a
brief description of which has been published by M. Fiscuer, in the form of an
appendix to his memoir, and a drawing of the cranium has been reproduced by
M. GERVAIS, in plate xxi. of the “ Ostéographie des Cétacés,”1 now in course of
publication, and there can be no doubt that it resembles in its configuration the
crania from Fos and Arcachon, Attempts have been made by some naturalists
to show that a cetacean, named by RAFINESQUE Epiodon urganantus, one described
by Risso as Delphinus Desmaresti, and one described by Cocco as Delphinus
Philippi, are specimens of the same animal; but, as M. FiscHer has pointed out,**
the descriptions which have been recorded of these animals are too indefinite to
enable the zoologist to state with certainty that they belong to the genus Ziphius.

In a short paper on Ziphioid Whales, published in “ Nature,”tt Professor W.
H. Fiower, of London, states that a complete skeleton of an adult Ziphius
obtained at Villa Franca in 1867, by Professor HAErcKEL, is mounted in the
Anatomical Museum of the University of Jena. But no description has been
as yet given of this skeleton. All the five European specimens which have

* Ann. des Sciences Nat. xv. 1851, p. 49.

+ Nouvelles Archives du Muséum. Paris, 1867, p. 55.

{ Catalogue of Seals and Whales in the British Museum, London, 1866; and Supplement, 1871.
§ Comptes Rendus, 1866, Aug. 6; and in Nouvelles Archives du Muséum. Paris, 1867.

|| Revue Zool. Soc., Cuvimr, 1842, pp. 207, 208.

| Plate xxi, figs. 8 and 9. Paris.

e-Opucii.p: 08,65)

t+ December 7, 1871.

AND MESOPLODON SOWERBYI. 761

hitherto been recorded, excepting that from Arcachon, have been obtained on
the shores of the Mediterranean.

But from other parts of the world recent crania of Ziphioid whales have
been procured, which are closely allied to, if not specifically identical with the
European species. In 1863, M. Van BENEDEN described by the name of Ziphius
indicus* a perfectly adult skull, brought from the Cape of Good Hope, which
he thought had been obtained from an animal captured in the Indian Ocean;
hence the trivial name indicus by which he designated it. This skull is now
in the Museum of the University of Louvaim. In 1865, Dr J. E. Gray
described by the name of Petrorhynchus capensist the cranium of another Ziphioid
sent from the Cape of Good Hope by Mr E. L. Layarp. An excellent descrip-
tion of the beak of this skull, and the region of the pre-nasal fossa, with illustra-
tive figures, has been given by Professor OwEn,{ who altogether condemns the
attempt made to exalt this specimen into a new genus, and ranks it along with
the skull described by Van BENEDEN as an example of Ziphius indicus.

In August 1865, Dr Burmeister, of Buenos Ayres, obtained a young male
Ziphioid, 13 feet long, which had been stranded near that city. He has
described and beautifully figured the external form of the animal, its visceral
anatomy, and the skeleton.§ Owing to its youth, the teeth were still imbedded
in the gum, and not only were two large teeth observed at the point of the
lower jaw, but from thirty to thirty-two small teeth were counted in the gum
on the mandible, and twenty-five on each side in the gum of the upper jaw.
BuURMEISTER provisionally named the animal Delphinorrhynchus australis, and
shortly afterwards proposed to call it Ziphiorrhynchus cryptodon. Subsequently
he adopted Dr Gray’s generic name Lpiodon, and, after suggesting as specific
names successively cryptodon and patachonicum, finally decided on Epiodon
australe.

Description of the Shetland Ziphius—tIn the autumn of 1870, I purchased
through one of my pupils, Mr M. Coucurrey, from Mr Joun ANDERson of Hills-
wick, Shetland, a number of cetacean bones, chiefly those of a large Rorqual
(Balenoptera Sibbaldit), which had been stranded in October 1869, in Hamna
Voe, on the north-west coast of the main Island. Along with the Rorqual
Mr Anperson sent the skull of a smaller whale, which he informs me was cap-
tured in 1870 off Hamna Voe, out at sea, and towed to the shore. When this
skull came into my possession it was invested by the dried and hardened textures

* Mém. Couronnés de I’ Acad. Royale de Belgique, 6th June 1863. Collection in octavo, vol. xvi.
1864. Plate i.

t Proc. Zool. Soc. 1865, and Catalogue of Seals and Whales, 1866.

{ British Fossil Cetacea of the Red Crag, in the Memoirs of the Paleontographical Society, vol.
xxiii. 1870.

§ Annals of Natural History, 1866, vol. xvii. pp. 94, 303, plates iii., vi.; also in Anales de
Museo Piblico de Buenos Aires, Tomo i. 1868. Pl. xv.—xx.

762 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

of the head, so that although I judged it to be a member of the Ziphioid group of
whales, yet I was unable to submit it to a-sufficiently close examination to deter-
mine the species, until after it had been for some months in the macerating trough.
When the bones were cleaned I had no difficulty in deciding that it resembled
the Ziphius cavirosiris of Cuvier. As the Shetland specimen is the first
example of this rare cetacean which has been met with in the British seas, I
have the satisfaction of adding this animal to the list of British mammals.

From the size of the skull and the condition of many of the sutures, it was
obviously that of an adult, if not an aged animal; and as it had not been injured,
and the lower jaw was preserved, I am enabled to describe and figure a perfect
specimen of this part of the skeleton.

The general outline of the skull was triangular, with curved sides, the base at
the occiput, and the apex at the tip of the beak. The summit of the skull was
formed by the two nasal bones, the frontal and the upper borders of the two
pre-maxille. Its greatest length measured in a straight line was 354 inches; its
greatest breadth between the post-orbital processes of the frontal, 214 inches;
its greatest height, 184 inches.

When regarded from the dorsal surface, the skull was seen to slope rapidly
downwards and backwards, from the summit to the foramen magnum and
occipital condyles. The slope forwards to the tip of the beak was much more
prolonged, so that a far larger proportion of the antero-posterior diameter of
the skull was in front of the summit than behind. The beak was triangular
in form. Its breadth, on a line with the superior maxillary foramina, was 12
inches; its length from that line to the tip was 193 inches, and its breadth at
the tip 14 inch. : ,

The tip of the beak was formed by the two pre-maxillaries and the meso-
rostral bone. The pre-maxille varied much in shape and size in different parts
of their extent. Near the tip they were elongated and almost straight, but
about 6 inches from the tip their upper borders curved inwards, so as partially to
overlap the meso-rostral bone. Here also they diverged from each other and
became more expanded; and as they were traced backwards, this divergence and
expansion became more strongly marked. At their hinder ends they mounted
upwards to assist in the formation of the summit of the skull. Those parts of
the pre-maxillz which lay behind the meso-rostral bone formed the sides and
a portion of the posterior boundary of the great pre-nasal cavity at the base of the
beak, from the presence of which the specific name of cavirostris was applied by
Cuvier. The upper borders of these bones, which in the rostrum were curved
inwards, and separated by an interval of about 2 inches, at the sides of the pre-
nasal fossa were so far everted as to be 84 inches asunder, a measurement
which expresses the transverse diameter of the fossa. The two bones in the pre-
nasal region were far from symmetrical. The right was much more expanded

AND MESOPLODON SOWERBYI. 763

than the left, and its inner surface was directed forwards and inwards, whilst that
of the left was more vertical, and to a great extent directed inwards. The upper
end of the right bone mounted also higher than that of the left, and formed a
thick and strong lobed projection, which overhung the pre-nasal fossa and the
orifice of the right nostril. The summit of the left bone was thinner, less curved,
less projecting, and more ridge-like than lobed. A nervo-vascular canal of some
size opened on the inner concave surface of each pre-maxilla, about 4 inches
behind the posterior end of the meso-rostral bone.

The meso-rostral bone formed one of the most characteristic features of the
skull. It occupied the interval between the anterior ends of the pre-maxille.
Near the tip it was narrow, and so intimately blended with the other bones of
the beak, that a faint superficial groove on each side was the only indication
of their original separation. But where the pre-maxille began to diverge, then
the meso-rostral bone dilated to a thick dense bar, 24 inches wide in its greatest
transverse diameter, which extended backwards to 134 inches behind the tip of:
the beak. At the same time, the grooves marking its separation from the pre-
maxille increased in depth and breadth, more especially on its left side. Hence
this bar was not absolutely mesial, but had a slight inclination to the right
side. At its hinder end it abruptly ended in an almost vertical, truncated
smooth face, which became continuous with the vomer, where that bone formed
the floor of the hollow of the beak. There can be no doubt but this bar
rested on the anterior part of the vomer, and was anchylosed to it. At the tip
of the beak the meso-rostral bone seemed as if subdivided into two lateral
halves by a longitudinal cleft, but I am inclined to think that this cleft was
rather to its left side, and marked the separation of the bar from the left lateral
half of the vomer, the separation of which, again, from the corresponding pre-
maxilla was indicated by a shallow groove.

The two nasal bones were situated between the summits of the two pre-
maxille, and presented flattened surfaces superiorly. They were inclined for-
wards and to the left, and, with the lobe of the right pre-maxilla, overhung, like
the eaves of a house, the apertures of the nose. They were firmly anchylosed to
each other, to the frontal, pre-maxille, and mes-ethmoid, and only faint traces
of the original sutures remained. The right nasal, which was rather the longer
of the two, was 6 inches in its antero-posterior diameter, by 14 inch in its trans-
verse diameter.

The mes-ethmoid portion of the nasal septum was curved to the left, so that
the nasal fossee were unequal in size, the right being 2§ inches in antero-
posterior, and 2 inches in transverse diameter; the left being 23 inches in
antero-posterior, and only 13 inch in transverse diameter. The upper border
was sharp, and prolonged into a spine at its junction with the anterior border.
The anterior border expanded into a smooth surface 1¢ inch broad at its greatest

VOL, XXVI. PART IV. 9N

764 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

transverse diameter. This surface was in the hollow of the beak, and formed
indeed a part of the posterior boundary of the floor of the great pre-nasal fossa.
It was separated by a suture from the vomer, and the distance between this
suture and the posterior truncated end of the meso-rostral bone was 62 inches.
The smoothness, not only of the truncated surface of the meso-rostral bone, but
of the expanded anterior border of the ethmoid, induced me to think that the
bar of cartilage, which had undoubtedly connected them together in the young
state of the skull, had either altogether, or to a large extent, disappeared prior
to maceration. For in the skeletons of the Cetacea when unossified cartilage is
continuous with the end of a bone, the surface of bone in apposition with the
cartilage possesses a faintly tuberculated appearance, such as one is familiar
with on the attached surface of an epiphysis of a human bone.

The vomer formed the anterior part of the floor of the pre-nasal fossa, and
passed backwards to embrace the inferior border and the sides of the mes-
ethmoidal part of the nasal’septum. The backward prolongation of the vomer
on the left side of the mes-ethmoid was partially concealed by short yet strong
bars of bone connecting the left pre-maxilla with the mes-ethmoid. The
posterior or cerebral surface of the mes-ethmoid was expanded laterally, and
instead of being perforated into a cribriform plate, possessed only a single
foramen on each side, in all probability for the transmission of a nasal branch
of the fifth cranial nerve.

The superior maxilla formed the side of the beak, but did not extend to
within 24 inches of the tip. For some distance backward it was a compara-
tively narrow bar of bone, having a deep furrow along its line of articulation
with the pre-maxilla ; but opposite the large maxillary foramen it expanded
both vertically and transversely, overlapped the anterior surface of the frontal,
and formed a deep maxillary fossa immediately to the outer side of the upper
and posterior end of the pre-maxilla. The free surface of this part of the bone
was pitted with irregular shallow depressions, and perforated by a large canal.

An ecto-maxillary ridge, faintly grooved at its free border, extended along
the outer edge of the maxilla, and at the base of the beak was elevated into a —
maxillary tuberosity, sufficiently large to form a noticeable feature in the profile
view of the cranium. The maxillo-premaxillary furrow extended backwards
immediately to the inner side of this tuber to become continuous with the
deep maxillary fossa, and the large maxillary foramen opened into it at the
inner side of the tuber. In Globio-cephalus, and in various others of the
toothed whales, the pre-maxilla does not so completely overlap the superior
maxilla, but that a portion of the latter bone appears on the surface to the
inner side of the pre-maxilla, and intervenes between it and the nasal. In Z. cavi-
rostris, on the other hand, owing to the incurvation of the pre-maxilla, the ~
portion of the superior maxilla above referred to was thrown into the outer wall

AND MESOPLODON SOWERBYI. 7659

of the nose, and, so far as the state of the sutures allowed me to judge, the pre-
maxilla seemed to be directly anchylosed to the nasal bones.

The inferior surface of the beak, triangular in its general outline, was convex
from side to side, and the convexity gradually increased from before backwards.
The tip consisted of the two pre-maxille, behind which the superior maxilli
formed this surface of the beak, except in the mesial line, where a narrow bar
of bone—the lower edge of the vomer, nearly 10 inches long—came to the sur-
face of the palate. No definite anterior palatine foramen was seen, but a distinct
posterior palatine canal was situated on each side at the articulation of the
superior maxilla and palate bones.

The palatine plate of the palate bone was little more than 1 inch in antero-
posterior diameter at its widest part. The anterior margin was convex, and as
the superior maxillz extended backwards for some distance between the two

_ palate bones, the latter only articulated with each other in the mesial line for

about 14 inch. Laterally each palate bone extended backwards and outwards
as far as the lachrymal, a thin scale of the superior maxilla intervening between
it and the malar.

The two palate bones were inserted between the anterior borders of the.
pterygoids, which bones articulated with each other for 7 inches in the mesial
line of the under surface of the skull, and formed on each side of this line a
well-marked curved ridge, which extended as far back as the posterior orifice
of the nose. This orifice was bounded below and at the sides by the two ptery-
goids, and above by the expanded part of the vomer, the latter of which articu-
lated by the margins of its expanded part with the base of the pterygoids. The
vertical and transverse diameters of this orifice were almost equal—between 5 —
and 6 inches. Owing to the great backward development of the pterygoids,
the thick posterior edge of the vertical plate of the vomer, forming the
nasal septum, was 53? inches in front of the posterior margin of the palatal
plates of the pterygoids. The vertical plate of the vomer was in the mesial
plane, so that the want of symmetry displayed at the anterior nares did not exist
at the posterior nostrils. Deeply within the nose a narrow bar of the palate bone
could be seen on each side, intercalated between the pterygoid and the side of the
vomer. The outer surface of the pterygoid was hollowed into a very shallow fossa,
which was not, however, closed in front and externally by a reflected plate of
bone, to form a posterior palatine air sinus, as in Globio-cephalus and most
other cetaceans.

Behind the expanded part of the vomer the basis cranii, deeply concave
from side to side, owing to the lateral elevations of the basi-occipital, extended
backwards to the foramen magnum. All trace of the suture between the basi-
sphenoid and basi-occipital had disappeared, and the anchylosis of the various
elements of the occipital bone with each other was perfect.

766 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

The occipital surface of the skull, broadly triangular, ascended almost ver-
tically to near the summit of the cranium, where it articulated with the frontal.
The foramen magnum, directed backwards, was of almost equal diameter (about
21 inches) vertically and transversely. The occipital condyles were inclined
obliquely downwards and forwards at the sides of the lower half of the foramen,
but did not coalesce. Their faces were smooth, convex, and directed back-
wards and outwards. A vertical, but not quite mesial, ridge ascended from the
foramen to the summit of the supra-occipital. No trace of an interparietal
bone was seen. The ex-occipital, broad and thick, was prolonged at its outer
and lower part into a thick process (par-occipital or jugal), which was separated
by a deep cleft from the lateral elevation of the basi-occipital.

The temporal was an irregular bone, situated in front of the ex-occipital
with which it articulated. The part, jointed with the jugal process of the latter,
had not only the shape of a mastoid process, but was articulated with the |
squamous temporal by a strongly denticulated suture. A strong zygomatic pro-
cess arched forwards, and almost touched the post-orbital process of the frontal.
A thinner, more scale-like portion of the temporal ascended to form a part of the
floor of the temporal fossa, where it articulated with the parietal. The under
surface of the temporal bone presented two concavities : an antero-external, or
glenoid, for articulation with the lower jaw, and a postero-internal, for the lodg-
ment of the petro-tympanic bone.

The recess for the lodgment of the petro-tympanic bone was of small size,
and bounded by the basi and ex-occipitals, and by the squamous and mastoid
parts of the temporal. The petrous bone was not anchylosed to the tympanic ;
it was 21 inches long by 14 inch in its greatest transverse diameter, though its
outline was irregular. Externally it passed for some distance behind the mas-
toid, with which it articulated by a smooth flattened surface, whilst its inner end
was in apposition with a smooth, almost vertical process of the squamosal, 14
inch in length. The canal in the bone for the auditory nerve was relatively
small. The surface of the bone, forming the inner wall of the tympanum, had
the solid, rod-like stapes articulated by a movable joint with the foramen
ovale; the incus and malleus had not been preserved. Opening into the recess
in which the petrous bone was lodged, was a circumscribed canal, communi-
cating with a cranial cavity obviously for the transmission of the seventh nerve.
Between the ex- and basi-occipital were two canals, apparently for the trans-
mission of the eighth and ninth nerves. Somewhat in front of, and internal to
the base of the vertical process of the squamous, was another canal situated in
the ali-sphenoid, which had probably transmitted the inferior maxillary division
of the fifth nerve.

The tympanic bone, 22 inches in length by 1% inch in greatest transverse
diameter, possessed the conchoidal form so characteristic of this bone in the

AND MESOPLODON SOWERBYI. 767

Cetacea. Its convex superficial aspect was comparatively smooth ; from the
antero-external thin border a tongue-like curved process projected forward ;
whilst the opposite rounded border was crenulated. The bone corresponded
closely in form with the tympanic bone of Hyperoodon.

The parietal bone was small in size, and seemed not to extend beyond the
temporal fossa, a portion of the floor of which it formed. The suture along its
upper border was, however, too faint to enable the exact extent of the bone to
be accurately observed.

The great lateral crest of the cranium was formed by the narrow free border
of the frontal appearing between the superior maxilla and the supra-occipital.
Above the orbit, however, the frontal widened out to form an arched roof for
that chamber, and terminated both in front and behind in a process—the pre- and
post-orbital. A large canal, obviously the optic, opened into the deeper part
of the orbit. One inch in front of this canal was an oval opening leading into
a deep fossa, into which two canals opened, one leading backwards into the
cranial cavity, the other forwards to the great maxillary foramen. These canals
doubtless served for the transmission of the ophthalmic and superior maxillary
nerves. |

But a small part of the great wing of the sphenoid appeared on the under
aspect of the skull, as it was extensively overlapped by a thin plate of the
pterygoid. A portion of this wing, however, ascended into the temporal fossa,
and articulated with the squamoso-temporal, parietal and frontal. This fossa
had no great size, but possessed some depth. It was bounded by the frontal
with its post-orbital process, by the zygomatic and by the ex- and supra-occipitals.

The malar consisted of a rough plate of bone intercalated between the superior
maxilla and the lachrymal, and of a flat, smooth, slender process, which passed
backwards for a short distance to form the lower boundary of the orbit. This
process was broken on both sides of the skull, so that its proper length could
not be ascertained.

The lachrymal was a large plate-like bone, which closely articulated with the
orbital surface of the frontal, and entered into the formation of the anterior
part of the roof of the orbit. Its anterior border was wedged in between the
malar, superior maxillary, and frontal bones.

With the exception of the meso-rostral bone, the petro-tympanics, the ane
septum, and the pre-maxille, the bones of the skull were of a loose spongy
texture, even on their surfaces—a circumstance which will, doubtless, account
for the mutilated condition of almost all the crania of this Ziphioid which have
come under the notice of the anatomist.

The lower jaw consisted of two lateral halves anchylosed at the symphysis.
Viewed in profile, the upper and lower borders were concavo-convex. The
concavity on the upper border was partly on a line with and partly behind the

VOL XXVI. PART IV. 90

768 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

symphysis, that on the lower border was altogether behind the symphysis. The
anterior half of the upper border was marked by a shallow groove, as if for the
lodgment of rudimentary teeth, though none were found in the specimen.
Quite at the anterior end of the bone, on each side of the symphysis, was a large
alveolus, about three-fourths of an inch in diameter, in which at one time the
large mandibular tooth had undoubtedly been lodged, but this socket was now
occupied by coarsely spiculated bone. The length of the mandible was 32
inches ; the length of the symphysis, 7 inches ; the width between the articular
condyles, inside measurement, 17 inches. The condyle was situated about the
middle of the posterior border of the bone. When articulated with the skull,
the lower jaw projected so far beyond the rostrum that the mandibular teeth
would, if present, have been altogether in front of the beak.

Comparison of the Shetland Ziphius with previously recorded Specimens.—I
propose now to compare the Shetland cranium with the figures and descrip-
tions which have been published of four of the five European,* and the three
exotic specimens referred to in the historical introduction, with the object
of determining whether they represent different genera, or whether they belong
to one or more species of the genus Ziphius. In their general configuration all
the skulls closely correspond with each other, but as the peculiarly constructed
beak and the pre-nasal fossa with the bones which form its boundaries con-
stitute the most distinctive features of these crania, my attention has more
especially been directed to a comparison of the forms and relations of the bones

which enter into their construction.

In my description I have named the dense, solid bar in the middle of the
beak the meso-rostral bone. This bar corresponds with the ‘‘ vomer” of Cuvier,
GERVAIS, and Gray, with the “anterior tuberosity of the vomer” of FIscHER,
with the “continuation of the pre-frontals forward to near the end of the pre-
maxillaries” of OweEn,t and with the “anterior prolongation of the ethmoid”
of FLowEr.{ Whatever name be applied to it, there can be no doubt that it is
an ossification of the anterior end of the long cartilaginous bar, which in the
cetacea is prolonged forwards to the end of the beak, and in relation to the sides
and lower surface of which the spout-like vomer is formed. In the specimens
recorded by Cuvier, FIscHerR, DoumMEeT, VAN BENEDEN, and Gray and OwEN,
the meso-rostral bar, as in my Shetland specimen, was strongly pronounced.
Slight differences do, however, undoubtedly exist in the shape of this bone in
these crania. In those described by Fiscuzr, Gray and Owen, and myself, the
posterior end is a little more truncated than in those recorded by Cuvizr,

* No figure or description of the Villa Franca specimen in the Jena Museum, so far as I can
ascertain, has yet been published.

+ Report of British Association, 1846, p. 226, and British Fossil Cetacea of the Red Crag, p. 27.

¢ Introduction to the Osteology of the Mammalia, p. 191. London, 1870.

AND MESOPLODON SOWERBYI. 769

Dovumet, and VAN BENEDEN ; and in the two exotic crania it is more swollen, and
projects somewhat higher above the pre-maxille than in the European skulls.
In the Cape cranium, figured by Gray and OWEN, it is more mesial and uni-
formerly tapering from behind forwards than in the Shetland specimen, and in
those described by DoumeT and by Fiscuer. But these are all differences so
trifling in degree, as not to exceed that range of individual variation which one
often meets with in comparing a series of crania of the same species of animal,
and which may easily be accounted for by one skull being a little more advanced
in its ossification than another.

In my account of the pre-nasal fossa in the Shetland skull, and of the
“septum narium,” and its relations to the vomer, I might indeed have adopted
almost verbatim the description which Owen has given of these parts in the
Cape cranium, so well does it express the arrangement. Similarly, the form
and relations of the nasal bones, the configuration of the upper ends of the pre-
maxille and superior maxille, the form of the palatine surface of the beak, and
the relations of the bones which enter into its construction, are identical in all
the specimens in which they have been described.

There can be no doubt, in my opinion, that all these crania, whether exotic or
European, should be referred to the genus Zzphius, and in so far I cordially
concur with the remarks made by Professor Owen, that there should be “ a tacit
burial and oblivion” of the ill-defined generic names with which systematic
zoology has of late been needlessly and unscientifically encumbered. I also
hold that the crania from Fos, Corsica, Arcachon, and Shetland, are specifically
identical, and that the ‘“‘ type” of the species is the Z¢phius cavirostris of CUVIER.
But further, from a comparison of the Shetland cranium with the figures and
descriptions of the two specimens from the Cape, I am of opinion that they
should not be separated from Cuvier’s ‘“ type” species by the distinctive name
of Ziphius indicus. In recommending a new specific name for his Cape skull,
M. VAN BENEDEN appears to have taken GERVAIS’ specimen from Aresquieés, in
which the meso-rostral bone is absent, as the type of cavirostris rather than
Cuvier’s original example, in which it is well developed, just as in the crania
from Corsica, Arcachon, Shetland, and in Van BENEDEN’s own specimen. He
also refers, as another feature of difference, to the absence of teeth in the upper
jaw in his specimen, and their presence in the gum of the upper jaw in the
Aresquiés cranium. But these teeth were quite rudimentary and functionless,
and the presence of such aborted organs ought no more to form a basis for
establishing a specific difference than should the entire absence of teeth, both
in the upper and lower jaw, in the Shetland cranium be a reason for regarding
it as a distinct species. Further, there is no evidence that teeth were present
in the upper jaw in Cuvier’s type-specimen, or in the skulls from Corsica and
Arcachon. Moreover, both in VAN BENEDEN’s Cape skull and in the one from

770 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

Shetland, the lachrymal bone was very distinct, and the mastoid part of the
temporal was a separate ossification. The tympanic bone also, as figured by
Van BENEDEN, closely corresponds in shape to that of the Shetland cranium.

There is, however, greater difficulty in coming to a positive conclusion as to
the specific position of the cranium from Aresquiés described by GeRvats, and
that from Buenos Ayres by Burmetster. These skulls agree in possessing the
characteristic hollow at the base of the beak, but they differ from the specimens
above referred to, in possessing an open meso-rostral canal, extending along the
whole length of the beak, and in the consequent absence of a meso-rostral bone.
GERVAIS’ specimen not only had rudimentary teeth, but it is also stated by
FiscHER that the nasal fosse were not so deflected to the left, and the hollow of
the prenasal fossa was not so great as in the skull from Arcachon. GERVAIS
himself does not regard these differences as sufficient to form a new species,
and refers the Aresquiés cranium to Z. cavirostris. FISCHER, however, con-
siders them to be of specific value, and gives the specimen the name of Z. Ger-
vaist. BURMEISTER’S specimen was admittedly a young male, and not only were
rudimentary teeth, as in GERVAIS’ specimen, present in the gum, but a strong
cylindrical cartilage occupied the canal between the two intermaxillary bones.
Should the non-ossification of the anterior end of this cartilage be a persistent
condition in these animals, even in adult life, then they would undoubtedly have
to be regarded as forming species distinct from cavirostris. But if the want of
ossification of the cartilage is due, like the presence of rudimentary teeth, merely
to the youth of the animals—and as the conversion of the anterior part of this
cartilage into bone is altogether an exceptional occurrence in the cetacea, it is
possible that it may not take place in the genus Zphius until towards the end
of the period of ossification—then these characters cannot be adduced as satis-
factory evidence of a specific difference. I am disposed, until further informa-
tion has been obtained regarding this question, to rank provisionally these
crania also with cavirostris, which will include therefore the following speci-
mens :—

ZIPHIUS CAVIROSTRIS.

Fos, Bouches du Rhone, ; : CUVIER.
Aresquies, Hérault, . : ; ; GERVAIS.
Corsica, : : : f ; DovumMET.

Cape of Good Hope, ‘ VAN BENEDEN.
Arcachon, . ; : FISCHER.

Cape of Good Hope, : ; Gray and Owen.
Buenos Ayres, } ; : BURMEISTER.
Villa Franca, : : : : HAECKEL.

Shetland, : : ; ; ; TURNER.

AND MESOPLODON SOWERBYI. wil

If this mode of regarding the specific unity of these specimens be correct, then
Ziphius cavirostris will have a geographical range equal to that possessed by
the spermaceti whale.

Historical Sketch of Sowerby’s Whale.—Early in the present century, Mr
JAMES Sowerby figured and gave a short description* of a new species of
cetacean cast ashore in 1800, near Brodie House, county of Elgin, which he
termed Physeter bidens, or the two-toothed Cachalot. The animal was a male,
and the beak, with the anterior part of the cranium and the lower jaw, are
preserved in the Oxford University Museum. DE BLAINVILLE associated the
name of SowErRsBy with this animal, and since then the specific name Sowerbyi,
or Sowerbiensis, has been attached to it, although with varying generic appel-
lations. For whilst some zoologists class it as a species of the genus Ziphius, by
others again the generic name of Mesoplodon, given by M. GERVAIS, is not un-
frequently accepted. In 1864 another specimen, also a male, was stranded in
the Bay of Brandon, on the coast of Kerry in Ireland, and the head has been
figured and described by Mr Wit1t14m ANnpDreEws.t The skull is preserved in
the Museum of the Royal Dublin Society. At a meeting of the Royal Irish
Academy, June 23, 1870, Mr AnpRrews mentioned that a second specimen,
17 feet in length, had been captured in the same locality, in the month of May
of that year.{ Other animals belonging to the same species have been stranded
on the coasts of the continent of Europe. A female at Havre, near the mouth
of the Seine, in 1825, described by Cuvier as Delphinorhynchus micropterus, the
skull of which is in the Museum of Natural History, Paris.§ Another female
stranded at Sallenelles, Calvados, in 1825, the skull and part of the skeleton
of which are preserved in the Museum at Caen.|| A young female stranded at
Ostend in 1835, the skeleton of which is in the Brussels Museum.f A
mandible in the Museum at Christiania, found some years ago on the coast of
Norway.**

But from the seas of the southern hemisphere specimens have been pro-
cured which, though differing in some particulars, yet conform in many essential
points with the European examples. In the Museum of Natural History in
Paris is a skull brought from the Seychelles Islands, to which the specific name

* British Miscellany, plate i. p. 1. London, 1806.

t Trans. Roy. Irish Academy, Part X. 1869.

{ Nature, Aug. 11,1870. [Professor Macatister writes me that the bones of this specimen are
still undergoing maceration. Dr J. E. Gray states, “ Annals of Nat, Hist.,” August 1872, that Mr
W. Anprews informs him of the receipt of a perfect male skeleton of this rare whale at the Dublin
Museum, being the third specimen taken on the west coast of Ireland —Note, October 1872.]

§ Hist. Nat. des Cétacés. Paris, 1836; p. 114, plate vii. Figured also by Gurvais in “ Ostéo-
graphie des Cétacés,” plate xxvi.

|| Figured by Gurvais in “ Ostéographie des Cétacés,” plate xxvi.

{| Dumortier. Mém. del Acad. Roy. de Belgique, tom. xii. 1839; and Van Brnepen in Mém.
Couronnés Coll. in Oct., tom. xvi. plate ili. 1864.

** Van BENEDEN in “ Bulletins de Acad. R. de Belgique,” xxii. 1866.

VOL. XXVI. PART IV. 9 P

772 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

denstrostris was applied by DE Buarnvitiz.* <A skeleton from Lord Howe’s
Island, recently obtained by Mr Krerrt, has also been referred to the same
species.t A skull transmitted by Mr Layarp from the Cape of Good Hope to
the British Museum, and named in the first instance Ziphius Layardi after
that naturalist, has recently been described as forming a new genus, Dolichodon,
by Dr Gray. { Mr Krerrr has also obtained a whale, 18 feet long, from Little
Bay near Sydney, the skeleton of which is in the Sydney Museum ;§ he con-
siders it to be a new species, and names it Mesoplodon Giintheri; but Mr
FLower thinks that it may be of the same species as Layardi, but with the
tooth much less developed ;|| Dr Gray, however, regards it as a new genus,
and proposes to call it Callidon.{1 In a paper communicated to the Welling-
ton Philosophical Society of New Zealand, January 1870,** Mr FrepERIcK Knox
and Dr Hecror figure and briefly describe the skull of a young male Ziphioid
whale, 9 feet 3 inches long, killed in 1866 in Titai Bay, Cook’s Strait ; Dr
Gray, though recognising the affinity of this skull to SowrerBy’s whale, yet
because the mandibular teeth are situated at the anterior end of the jaw, refers
it to the genus Berardius, and terms it B. Hectori.tt Lastly, M. Gervais has
described and figured by the name of Dioplodon europeus tt the skull of an
animal frequenting the seas off the department of La Manche on the North
Coast of France, which possesses many affinities with SowERBY’s whale.
Description of the Skull in the Edinburgh Museum.—tThe skull, to which I
next direct the attention of the Society, I recognised in 1869, when examining
the Cetacean crania in the Museum of Science and Art in this city. No label
or mark of any kind was attached to it to show that any attempt had been
made either to identify the species, or even to record the locality from which it
had been obtained. Fortunately the Anatomical Museum of the University
possesses, through the courtesy of Dr AcLAND, a copy of the cast of the original
example of SowERBY’s whale, so that I had no difficulty, on comparing it with

* Duvernoy called it Mesodiodon densirostre (Ann. des Sc. Nat., 1851, xv. p. 58, plate ii.), and
Gervais has figured it in “Ostéographie des Cétacés,” plate xxv., as Dioplodon Sechellense.

+ Proc. Zool. Soc., 1870, p. 426; and Gray’s Synopsis, p. 102.

t Proe. Zool. Soc., 1865; and Synopsis, p. 101. Beautifully figured as Z. Layardi, by Pro-
fessor OWEN in his Memoir in Trans. Paleeontographical Society, vol. xxiii. plate i.

§ Annals of Natural History, 1871, vii. 368.

|| Nature, Dec. 7, 1871, p. 105.

4] Annals of Natural History, 1871, vii. 368.

** Trans. New Zealand Institute, vol. iii. plates xiv. xv. p. 125 es. In Dr Hucror’s notes it
is stated that plate xiv. refers to a specimen captured in Porirua Harbour, 1866, but this is evidently
an error, as Mr Kwox informs us that only a rude sketch and a few measurements of that animal were
preserved. I am indebted to Dr Lauprr Lrpsay for the opportunity of consulting these Transactions.

tt Annals of Natural History, 1871, viii. p. 116.

tt Zoologie et Paléontologie Frangaises, 2d ed. p. 289. Ostéographie des Cétacés, plate xxiv. Also
M. Destonecuamps in Bull. Soc. Linn. Normandie, t. x. 1866. Dr Gray, as if to add one more to
the multitude of generic names he has coined in his classification of the Cetacea, calls this specimen
Neoziphius (Synopsis, p. 101).

AND MESOPLODON SOWERBYI. tia

the cast, in determining it to be a younger skull of that species. As none of the
officials connected with the Museum of Science and Art could give me any
information as to the history of the specimen, I think it very probable that it
had formed a part of the Natural History collection of the University, prior
to its transference to the Department of Science and Art in 1854. As the
skull is almost perfect, and the bones not quite free from oil, it is clear that the
specimen had not been lying about the sea-shore and subjected for a time to
the action of the weather, but had been removed from a newly killed animal.
It is not unlikely that the animal had been captured somewhere on the Scottish
coast, and that the skull had been presented to the late Professor J AMESON.*

The following description of this specimen of the skull of SowERBy’s whale
has been written with the especial object of pointing out the features of resem-
blance and dissimilarity between it and cavirostris. The skull of Sowerbyi was
not only much smaller, but more elegantly formed than that of cavirostris. Its
greatest length in a straight line was 294 inches ; its greatest breadth, between
the post-orbital processes, 112 inches ; its height, 923 inches. It was obviously
not perfectly adult, as the cranial sutures were not obliterated, and the pair of
mandibular teeth projected but slightly from their sockets. The texture of the
bones was not so open and friable as in cavirostris.

The summit of the skull was formed by the frontal and superior maxillaries.
The beak was slender, its sides more nearly parallel, and it was absolutely longer
than that of cavirostris, as the distance from a line drawn across the base between
the maxillary foramina to the tip was 204 inches, whilst its breadth at the same
line was only 10 inches. The tip of the beak was formed by the pre-maxille,
which extended backwards almost horizontally as far as the base of the beak.
Their upper borders were curved inwards so as almost completely to roof in
the elongated meso-rostral canal, and in no part of their extent were they
more than 3ths of an inch asunder. On a line with the base of the beak the
pre-maxille rapidly ascended, formed the sides of the anterior nostrils, and each
terminated superiorly in a roughened, slightly overhanging ridge. The surface
of the ascending part of each bone widened out somewhat, and looked almost
directly forward, so that no hollow existed at the base of the beak. The pre-
maxillee were almost symmetrical, the right bone being a trifle broader than the
left. A foramen opened on the free surface of each bone on a line with the
maxillary foramina.

No meso-rostral bone occupied the canal in the middle of the beak, which
was quite empty, though with the soft parts 7m sitw, it would undoubtedly have
contained the elongated mes-ethmoid cartilage.

* T have from time to time pointed out this cranium to various naturalists, amongst whom I may
mention Dr Gtnrner, Dr Actanp, and Professor Van BrenepEen. From a reference to it in Professor
Fiower’s article in “ Nature,” already quoted, it would appear that M. Van Benepen had supposed
the skull to be at the present time in the University Museum.

774 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

The nasal bones, about 2 inches long, were laterally compressed, almost verti-
calin position, and locked in between the upper overhanging borders of the two
pre-maxillz, which partially concealed them. The nasal septum was mesial, so
that the anterior nares were symmetrical. The vertical diameter of the entire
orifice was 24 inches, the greatest transverse 13 inch. The upper border of the
septum was sharp and concave ; its anterior border closed in the posterior end
of the meso-rostral canal, and was faintly tuberculated as if for attachment to
the mes-ethmoid cartilage.

The vomer formed the floor and in part the sides of the meso-rostral canal.
It gradually tapered off in front, and did not reach to within 64 inches of the
anterior end of the beak; posteriorly it was prolonged along the sides and
inferior border of the mes-ethmoid nasal septum, with which it blended.

The superior maxilla extended only to within 43 inches of the tip of the
beak, and formed a narrow bar of bone, between which and the pre-maxilla only
a shallow furrow was seen. Although it expanded rapidly at the base of the
beak to overlap the frontal, the maxillary fossa was slight, and the surface of the
bone was not pitted. The maxillary foramen was not large and single, but was
subdivided into several smaller openings. The ecto-maxillary ridge had a sharp,
knife-like edge, and the maxillary tuberosity could scarcely be said even to be
indicated.

The inferior surface of the beak flattened at the tip was formed by the pre-
maxille : at and towards the base it was slightly convex, and consisted of the
superior maxille ; but the intermediate part was faintly concave, and its sides
were formed of the superior maxillee, between which the pre-maxille extended
backward for some distance, whilst the lower border of the vomer came to the
surface in the mesial line for a distance of 63 inches.

The palatine plates of the palate bones articulated mesially for 14 inch, and
then diverged in front to permit the superior maxillee to pass backward between
them. Each plate stretched out laterally as far as the lachrymal bone. A
small posterior palatine foramen was situated close to the right palato-maxillary
suture. The two palate bones were in part inserted between the two pterygoids,
which also articulated mesially with each other, and passed back to form the
sides and floor of the posterior orifice of the nose. Each pterygoid curved out-
ward from the mesial and palato-pterygoid sutures to form a ridge, which over-
hung the outer surface of the pterygoid, and formed a deeper fossa than was
seen in cavirostris—an arrangement which presented a closer approximation to
a posterior palatine air-sinus than was seen in that animal, though, as in it a
reflected plate of bone was not developed. The posterior nasal opening was 24
inches in height by 3 inches in transverse diameter. As in cavirostris, the hinder
edge of the vertical plate of the vomer was some distance within the opening,
the roof of which was formed by the expanded part of that bone, whilst a

AND MESOPLODON SOWERBYI. 775

part of the palate bone entered into the construction of the outer wall of the
nares.

The basis cranii was concave, owing to the lateral elevations of the basi-
occipital. The occipital surface of the cranium was not so vertical as in cavi-
rostris, and had a faint mesial ridge. The foramen magnum was 1{ inch high
by 18 inch wide. There was no trace of an interparietal, unless a small pro-
cess anchylosed to the supra-occipital, and projecting into the mesial part of
the frontal, could be thus considered. The ex-occipitals were prolonged exter-
nally into a jugal process, which was separated by a cleft from the lateral
elevation of the basi-occipital. .

The squamoso-zygomatic part of the temporal resembled the same bone in
cavirostris. Unfortunately, the mastoid and petro-tympanic elements of the
temporal had not been preserved in this specimen of Sowerbyz. Three canals
communicating with the cranial cavity opened into the periotic hollow.

The parietal bone formed a large share of the floor and anterior wall of the
temporal fossa, and ascended between the occipital and frontal bones for 14
inch above the temporal crest. The relations of the frontal bone to the great
lateral crest of the cranium were similar to those described in cavirostris, but
at the summit of the skull, owing to the differences in size, shape, and direction
of the two nasals, the frontal passed forwards between the upper ends of the
two superior maxille, and articulated not only with the nasal but with a
narrow process sent backwards from each pre-maxilla. The frontal formed also
the roof of the orbit, and possessed a pre- and post-orbital process. An optic
and a pre-optic foramen opened into the deeper part of the orbit.

A small part only of the great wing of the sphenoid was visible on the
surface of the skull, and it contributed only in a minor degree to the formation
of the floor of the temporal fossa, which was chiefly composed of the parietal
and squamoso-temporal bones. The malar had the same relations as in cavi-

rostris, but the lachrymal was relatively larger than in that animal, for not only

did it assist in the formation of the roof of the orbit, but it extended so far out-
wards and forwards as to articulate with the pre-orbital process of the frontal,
and contributed to form the profile outline of the cranium.

The lateral halves of the lower jaw were not anchylosed at the symphysis.
Both the upper and lower borders were concavo-convex, but the concavity on
the lower border was deeper and proportionally longer than in cavirostris, so
that the bone had a lighter and more elegant appearance. The upper border
possessed in its anterior half a dental groove, and a single large, triangular,
laterally compressed tooth was present on each side, only the curved and
backward directed apex of which projected beyond the socket. The slight pro-
jection of the teeth may, perhaps, be due not merely to the non-adult state of
the animal, but may indicate that it was of the female sex. The socket was

VOL. XXVI. PART IV. 9Q

776 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS

situated 9 inches behind the tip of the jaw, and almost on a line with the hinder
end of the symphysis. The length of the mandible was 274 inches, that of its
symphysial portion 92 inches. The symphysis was therefore not only relatively,
but absolutely longer in Sowerbyi than in cavirostris. In the articulated skull
the lower jaw projected, as in cavirostris, beyond the tip of the beak.

Comparison of this Skull with previously recorded Specimens.—Of the specific
identity of the skull in the Edinburgh Museum with Sowersy’s original speci-
men, and with the crania from Havre, Calvados, Ostend, and Christiania, I
have satisfied myself, by comparing it with the cast of the skull of the first,
and with the published figures of the last-named crania. The Irish specimens
also are in all probability of the same species, although, so far as I can ascertain,
figures of these crania have not been published. Differences, however, of a
very appreciable character exist between the Edinburgh skull, the exotic speci-
mens referred to in the historical sketch, and the skull named Dioplodon
europeus by GERVAIS. In the skull from the Seychelles Islands, to which the
specific name densirostris is usually applied, not only is the want of symmetry
more decided, but the meso-rostral canal instead of being empty is occupied by
an elongated slender bar of dense bone, and the mandible is thicker and deeper
behind the symphysis, where the mandibular pair of teeth project from their
alveoli. The skull from the Cape, named Layardi, also unsymmetrical, pos-
sesses even a more strongly marked meso-rostral bone than densirostris; the
mandible is not, however, thickened and deepened, but contains a pair of
remarkably elongated and curved teeth, which arch upwards and backwards at
the sides of the rostrum to meet each other superiorly. Although various
zoologists have proposed to give a generic value to the differences exhibited by
Sowerbyi, densirostris, and Layardi, and have made each the type of a distinct
genus, yet I agree with Owen that there is nothing in the structural characters
of either of the three to justify more than a specific difference. The New Zealand
specimen figured by Knox and Hector, though referred by Gray to the genus
Berardius, is without question much more closely allied to SowErBy’s whale.
Like Sowerbyi it did not possess a meso-rostral bone, but the beak, judging
from the figure, was not so slender and elongated, the want of symmetry in the
nasal region was greater, and the pair of mandibular teeth were situated close
to the tip of the jaw. Some of these differences may, perhaps, be due to the
youth of the specimen, but the forward position of the mandibular teeth marks,
in all probability, a distinct species.

In the Dioplodon europeus of Gervais, the cranium is longer and wider than
in Sowerbyi; the beak also is wider, owing to the rostral part of the superior max-
ill being more strongly pronounced, and the meso-rostral canal is completely
filled by an elongated meso-rostral bone. Moreover, the mandible is not so
curved at its upper and lower margins, its symphysis is shorter, and the single

AND MESOPLODON SOWERBYI. (awe

tooth on each side, is situated nearer the tip. But these differences also, I would
submit, are of specific and not of generic import. For, although modifications,
such as I have described, occur in the construction of the beak, in the form of
the lower jaw, and in the position of the mandibular teeth; the conformation of
the nasals, pre-maxillaries, and maxillz, in the region of the anterior nares,
presents but slight modifications in Sowerbyi, densirostris, Layardi, and euro-
peus. Hence, I am disposed to consider that it is in the region of the anterior
nares, and in the bones surrounding these orifices, we are to look in the Ziphioid
group of whales for the characters which indicate generic resemblance or dis-
similarity, whilst the beak and lower jaw furnish us with the characters on
which specific relations may be based.

For if we recur to the group of crania, which in the first part of this memoir
I have classed under the head of Ziphius cavirostris, we find that they all
possess, from the peculiar shape of the pre-maxille, a wide and deeply excavated
pre-nasal fossa, at the bottom of which the anterior nares open, and that in all
the unsymmetrical, lobate nasals, with their shelving pent-house-like projection,
overhang the anterior nares. In the construction of the beak modifications
such as I have described occur in various of these crania, but in none can these
modifications, if they are to be regarded as anything more than mere individual
or sexual variations, be considered as marking more than specific differences.

Again, all the crania which I have referred to in the historical sketch of
SowERBy’s whale, agree with that animal in the absence of a pre-nasal fossa ;
for in these skulls the pre-maxillz ascend almost vertically, and with their
anterior surfaces so flattened, and the nasal bones so included between them,
that the anterior nares open directly, if I may so say, on the anterior plane of
the skull, and not at the bottom of a deep pre-nasal fossa. And although in the
construction of the beak, in the conformation of the lower jaw, and in the position
of the mandibular teeth, differences occur which may fairly be regarded as
specific, yet their common naso-premaxillary arrangements unite them, I believe,
into a single genus.

Are we then to consider, as has been done by Professor Owen, that
SowErBy’s whale and its allies belong, like the cavirostris of Cuvier, to the
genus Ziphius, and form distinct species of that genus, or are we to regard
them as forming a distinct genus, having various specific subdivisions? The
value which I am disposed to attach to the conformation of the naso-premaxil-
lary region in the Ziphioid group of whales, as a basis for classification, leads me
to the conclusion that Sowerbyi, with its congeners, should be placed in a genus
distinct from cavirostris. Reserving them for the latter, the name of Ziphius,
which was originally applied to it by Cuvier, I shall adopt Gervais’ name of
Mesoplodon as the generic designation for Sowerbyi and its allies. This genus
may be regarded as including the following species :—

778 PROFESSOR TURNER ON ZIPHIUS CAVIROSTRIS, ETC.

MESOPLODON SOWERBYI.
Examples from coast of Elgin, Scotland; Havre; Calvados; Ostend ;
Norway ; Brandon Bay, Ireland; also in the Edinburgh Museum of
Science and Art.
MESOPLODON DENSIROSTRIS.
Examples from Seychelles Islands ; Lord Howe’s Island.
MESOPLODON LAYARDI.
Example from Cape of Good Hope.
MESOPLODON EUROPAUS.
Example from La Manche, France.

Provisionally also, the specimen captured in Little Bay, near Sydney, and
the animal caught in Titai Bay, New Zealand, may be registered as distinct
species of the genus Mesoplodon.

MESOPLODON GUNTHERI.
Example from Little Bay, Sydney.
MesopLopon HEctort.
Example from Titai Bay, Cook’s Strait, New Zealand.

EXPLANATION OF PLATES XXIX., XXX.

The Plates have been lithographed from photographs of the crania, taken under my superintendence
by Mr Peter Thomson.

Fig. 1. Dorsal surface of cranium of Ziphius cavirostris, reduced nearly one-tenth.

Fig. 2, Anterior nares and pre-nasal fossa of the same animal.

Fig. 3. Dorsal surface of cranium of Mesoplodon Sowerbyi, reduced one-ninth. In placing the skull
before the camera, the photographer had unfortunately turned it over slightly to the left side,
so that the picture is not absolutely a full face view.

Fig. 4. Occipital surface of the skull of Z. cavirostris.

Fig. 5. Occipital surface of the skull of MZ. Sowerbyt.

Fig. 6. Palatal aspect of the cranium of Z. cavirostris.

Fig. 7. Palatal aspect of the cranium of M. Sowerbyt.

Fig. 8. Profile view of cranium of Z. cavirostris,

Fig. 9. Profile view of the lower jaw ; and

Fig. 10. View of the alveolar borders of the lower jaw of the same animal.

Fig. 11. Profile view of the cranium of MZ. Sowerbyi.

Fig. 12. Profile view of lower jaw of same animal.

Fig. 13. Outer surface of tympanic bone ; and

Fig. 14. Outer surface of petrous bone of Z. cavirostris, both the size of nature. ‘The stapes st. in this
specimen is in situ.

APPENDIX.—Novemper 23, 1872.

In drawing up the Historical Sketch of Cuvirr’s Hollow-beaked Ziphius
and of SowrrBy’s Whale, I omitted to refer to a brief description by Pro-
fessor A. W. Mat, in a memoir, entitled ‘“ Hvaldjur i Sveriges Museer
ar 1869,’* of the skeleton of a specimen of each of these rare Cetaceans
preserved in the Natural History Museum in Goteborg, Sweden. Although I
possessed, through Professor MAtm’s great courtesy, a copy of his elaborate
memoir, yet I had not, owing to its being written in the Swedish language,
mastered its contents. My friend, Professor FLower, having within the last
few days referred me to MALm’s description of these specimens, Dr CHARLES
Witson has very kindly translated for me the passages in which they are
described, and as a knowledge of Swedish is by no means general amongst
British zoologists, I think it desirable to append a statement of the more im-
portant particulars which Mam has recorded.

The specimen of Ziphius cavirostris was found stranded at Holma, near
Gullmarsfjard, Sweden, on 22d April, 1867. It was supposed to have been
suffocated by getting under the ice which had formed about Christmas, and
was so putrid that only a hand’s breadth of dark-grey skin remained. It
was a female, and measured 22 feet 2 inches (Swedish) in length. 75 kanns
(7500 cubic inches) of oil were obtained from it. In the stomach a tangled
mass of a transparent worm, 3 feet long, was found, which apparently belonged
to the genus Echinorhynchus. The total length of the skeleton, including the
cranium, was 6140mm., to which an additional 34mm. must be added for
the projection of the lower jaw beyond the beak :—

Length of cranium ; , : : ; , ; ; - 1015mm.
» » Mandible ‘ , : ‘ : : 3 : : 887 ,,
» » Symphysis 5 : : : ; : : E : 205 ,,
» 5, greatest breadth of skull . : : : : : ‘ 570 ,,

Two teeth, similar in size and shape, were in the lower jaw, but only the point
of each was tipped with enamel. The epiphyses were anchylosed to the
vertebral bodies. The vertebral formula was—C, D, Ly Cd, — 46. The
four upper cervical vertebree were anchylosed together. The sternum was
subdivided into five pieces, only the two posterior of which were completely

* Konig. Svenska Vetenskaps—Akad. Handlingar. Band 9, No. 2. Stockholm, 1871.
VOL. XXVI. PART IV. 9A

780 APPENDIX.

anchylosed. There were nine chevron bones. ‘The constitution of the carpus
is represented by MA. in plate v. fig. 51.

The Swedish specimen furnishes, therefore, an additional example to that
which I have obtained from Shetland of the occurrence of this Cetacean in
the North Sea; a habitat which had, indeed, been given for this animal by Dr.
Gray, in the Supplement (p. 98) to his Catalogue of Seals and Whales, although
no example from this sea was particularized by him.

Professor FLOWER writes to me that he has lately seen a fine skeleton of
cavirostris in the Museum at Pisa. As the Mediterranean is evidently one of
the usual habitats of this cetacean, this specimen had in all probability been
obtained from an animal stranded on the coast of Italy. Hence it will be
necessary to add to the list of specimens given on p. 770 the following
examples of
ZIPHIUS CAVIROSTRIS.

Holma, Sweden ; : , : . ; 4 Maim
Pisa (Mediterranean) , , : FLOWER

The specimen of SowERBy’s whale in the Gdteborg Museum was found in 1869
by a fisherman upwards of 100 miles from Kiaringd, Sweden. It was floating
vertically in the sea with its long-poimted snout directed upwards, and was
much decomposed. It was a male. Matm obtained the entire skeleton, with
the pelvic bones zm situ. The total length of the skeleton, including the
cranium, was 4409mm., to which an additional 8mm. must be added for
the projection of the lower jaw beyond the beak :—

Length of cranium. : , : : : : : : . 733mm.
5 mandible. "*, : : ; 5 ‘ : : : + BM 5,
» 3 symphysis . : ’ : ; A : : ‘ < alee

Greatest breadth of skull . : ; : : : 4 : - oS

In addition to the large tooth usually found on each side of the lower jaw in
this animal, Matm’s specimen possesses a second small tooth on each side
behind the large one, and the grooved character of the alveolar edge of the
mandible led him to think that it might at an earlier age have possessed others.
The distance from the point of the longer tooth to the point of the symphysis
was 240mm. ‘The vertebral formula was—C, D, L, Cd, — 46. The atlas
and axis were anchylosed together, and the epiphyses were united to the
bodies of the vertebrae. The sternum consisted of five pieces, only the two
posterior of which were completely anchylosed. Chevron bones ten; pelvic bones
63mm. long, 9mm. broad. Mam represents the constitution of the carpus in
plate v. fig. 52, and names the specimen Micropteron bidens.

guyett)

XXVI.—Remarks on the Ipecacuan Plant (Cephaélis Ipecacuanha, Rich.), as
cultivated in the Royal Botanic Garden, Edinburgh. By Joun Hutton
Batrour, M.D., F.R.S., Sec. R.S.E., F.L.S., Hon. Mem. Pharm. Soc.,
and Professor of Medicine and Botany in the University of Edinburgh.
(Plates XXXI., XXXII.)

(Read 18th March and 3d June 1872.)

The Ipecacuan plant, Cephaélis [pecacuanha of ACHILLE RicHARD, has been
cultivated in the Edinburgh Botanic Garden for upwards of forty years, but
it was not propagated to any extent until 1870, when a proposal was made to
attempt the cultivation of the plant in India. This suggestion was made on
account of the continued destruction of the plant by the collectors in Brazil, and
the risk of scarcity in the supply of this most valuable remedy for dysentery.
The Secretary of State for India (His Grace the Duxe of ARGYLL), under the
recommendation of several medical officers in Bengal, authorised an attempt to
propagate the plant in our Indian possessions, and with that view application
was made to me and others to aid in this important undertaking. Accord-
ingly, I at once set about the propagation of the plant in the Edinburgh Garden,
with the assistance of Mr M‘Nas the curator. He found that the plant could
be multiplied very rapidly by dividing the annulated root, cuttings of which,
though very small, give off young shoots when placed in favourable circum-
stances. By this means, numerous plants were produced very rapidly, and
the method was also followed by the Messrs Lawson, Nurserymen, Edinburgh,
who supplied a large stock of vigorous plants. Mr M‘Nas drew up a report of
his mode of propagation, which was printed, and distributed extensively to
district officers in India and elsewhere. The paper also appeared in the Trans-
actions of the Botanical Society of Edinburgh, vol. x. p. 318.

In Plate XXXII. fig. 6, two portions of a root giving off leaf-buds are
shown.

It appears from a report by Dr Kine, Director of the Calcutta Botanical
Garden, that in 1866 a single plant of Ipecacuan was received at the Calcutta
Garden from Dr Hooxer, but apparently artificial propagation had been
attended with sparing success, as Dr Kine reported at the beginning of 1872
that “the only surviving offspring of the Kew plant amounted to five plants
-in Sikkim, and seven in the Calcutta Garden.” It is understood that cuttings
of the stem were planted, but not of the roots. When, however, the plan pro-
posed by Mr M‘Nas was adopted for the propagation of the plant, much greater

VOL. XXVI. PART IV. 9s

782 PROFESSOR BALFOUR ON THE IPECACUAN PLANT.

success was obtained. In May 1871, a Wardian case was sent from the Edin-
burgh Botanic Garden containing twelve plants, and in October seventy-four
plants were despatched. The greater number of these reached their destination
in a good state.* In a report received since the reading of this paper, it is
stated, with reference to the late consignment, ‘These plants were forwarded
to Sikkim as soon as practicable after their arrival. The Calcutta climate
having proved totally unsuitable to this plant, all attempts to propagate it
there have been abandoned. The plants are at present under the immediate
care of the European gardeners of the Cinchona plantation, and propagation is
being carried on chiefly in one of the hot deep valleys of the Rungbee reserve.

“From what we have been able to learn from observation, Ipecacuanha will
apparently thrive best under deep shade, and in a hot, steamy, equable climate.
These conditions are supplied most fully in the valleys on the outer slopes
_of the Sikkim Himalaya, which open toward the Terai. <A fine small valley
near Sookna, at the point of entrance into the hills of the cart-road from Silli-
goree to Darjeeling, has accordingly been taken up as an Ipecacuanha
reserve. Hitherto the plant has not perfected seed in India, although
flowers have frequently been produced; we must therefore look to increase
by cuttings and other artificial methods.”—(Gardener’s Chronicle, Oct. 5, 1872,
p. 1322.)

As regards the specimens in the Botanic Garden, their propagation was
accomplished in the first instance by taking cuttings from the roots of a plant
sent by Sir W1tL1AM Hooker from the Glasgow Botanic Garden. The original
specimen had been forwarded to him by Mr Maxoy of Liege, and it flowered
at Glasgow in 1843. (See “Botanical Magazine,” tab. 4063.) Propagation
from this single stock, however, was not sufficient to meet the requirements
of India. I therefore applied for an additional supply of plants to my friend
Dr GuNNING, a medical graduate of the University of Edinburgh, who now
resides at Palmeiras, near Rio Janeiro. Sir Ropert CHRISTISON also aided
me in this request, and has continued to take a deep interest in the matter.
By Dr Gunninc’s kind services, we were able to secure a considerable supply
of plants from Brazil—the roots of which, by division, have yielded abundance
of young shoots.

The original plant in the Botanic Garden had produced flowers on several
occasions, but no fruit or seeds. The cuttings taken from its roots grew rapidly,
and at the end of a year’s growth many of them flowered, producing shrubby
stems. Some of the plants assumed a branching habit, and attained a large
size. The dimensions of the largest specimen in the garden are as follows :—

* Since this paper was read a large additional number of plants have been sent from the Botanic

Garden. In July 1872, 112 plants; in November 1872, 68—making in all during 1871-72, 300
plants.

PROFESSOR BALFOUR ON THE IPECACUAN PLANT. 783

Height, 16 inches ; length of leaves, 64 inches; breadth of leaves, 3} inches ;
length of peduncle, 1 inch; circumference of stem, ? of an inch.

The following are the general characters of the plant cultivated in the
garden :—Stem (Plate XX XI. fig. 1, a, a, a), more or less shrubby when fully
grown, simple or branching, with marks of the leaves giving a somewhat
annulated aspect, varying in height from 12 to 16 inches. The young stem is
herbaceous and quadrangular (Plate XXXII. fig. 7), The plants sent from
Brazil had marked rhizomes, and corresponded exactly with the figure given by
Marttvs in his Materia Medica of Brazil, Tab. I., but those in cultivation have
assumed an erect form. This may depend on the latter being restricted by
potting, and being propagated from root-cuttings.

The structure of the young stem, one year old, is shown in Plate XX XII.
figs.2 and 3. In fig. 2 there is a transverse section of the stem magnified
about thirty diameters, showing, externally, cellular hairs on the epidermis (a, @) ;
next, cortical parenchyma (6, b) composed of angular cells; next, bundles of
vessels (c, c), and lastly, cellular pith (d). In fig. 3 there is a longitudinal
section of the same stem, showing a portion of the cellular tissue of the bark
(b, 6); the vascular bundles consisting of spiral, pitted, and woody vessels
(c, c); and the central pith (d). :

The root of the plant is about the size of a writing-quill. It is well charac-
terised by its irregularly contorted and annulated appearance (Plate XX XI.
fig. 3). The roots come off from the lower part of the shrubby stem (Plate
XXXI. fig. 1, b, 6, 6). The roots may be said to combine the usual functions of
the root with those of the stem, inasmuch as they are capable of producing leaf-
buds ; when the root is cut into pieces, each portion producing a leaf-bud, as
shown in Plate XXXII. fig. 6. The outer or cortical part of the root (Plate
XXXI. fig. 3, a, a) is cellular, and has small projecting rings closely applied to
each other; the central part 0, called meditullium, is slender, and has a firm
woody structure. In Plate XX XIT., figs. 4 and 5, the root structure is given ;
fig. 4 shows a transverse section of the root; a, a, cellular epidermal portion ;
b, 6, cellular cortical portion, containing many granules of starch; c, central
fibrous portion, consisting of vascular tissue. Fig. 5, longitudinal section of the
root; a,a, epidermal cells; 0b, b, cortical starch cells; ¢, meditullium, or central
vascular system, consisting of woody vessels marked with dots.

The leaves are entire, often with a wavy margin, opposite, with short petioles,
stipulate, their form varying from oval to elliptico-lanceolate, the apex being
sometimes blunt and sometimes pointed. They vary in length from 2 to 4
inches. In Plate XXXI. fig. 2, a delineation is given of an elliptical, blunt-
pointed leaf of the natural size. The leaves of the plants propagated from the
original specimen received from Sir W. Hooxer and those from the specimens
sent by Dr Gunnin exhibit a difference in character. This seems to be merely

784 PROFESSOR BALFOUR ON THE IPECACUAN PLANT.

a slight variation, although conspicuous in the general aspect of the plants.
The leaves in the Hookerian plant are firmer in texture, somewhat coriaceous,
their form is elliptical or oval, apex rather blunt, and margin wavy. These
are represented in Plate XXXI. fig. 1, and Plate XXXII. fig. 1. In the Rio
Janeiro plants the leaves are thinner and more delicate in texture ; the form is
rather elliptico-lanceolate, the apex pointed, and the margin less wavy ; in the
young state the leaves are fringed with hairs ; the plant grows more freely, and
is less shrubby. This form is seen in Plate XXXII. fig. 7. As the plants get
older, the difference in form and texture is less marked.

The stipules (Plate XX XII. fig. 8) are conspicuous, interpetiolary, opposite,
united at the base, and are cut at the upper part into long narrow segments.
At the base of the stipules there are several ovate-lanceolate glands (Plate
XXXII. fig. 9).

The flowers are, in capitula, surrounded by a four-leaved involucre, and are
supported on a stalk which is about an inch in length, at first erect, and then
bent from its base downwards (Plate XX XI. fig. 4). Each capitulum con-
tains from ten to twelve flowers, which are white, sessile, and sweet-scented.
The calyx is superior, persistent, its limb cut into five divisions (Plate XX XI.
fig. 5). Corolla, funnel-shaped, with a cylindrical tube and a limb divided into
five broadly-ovate, pomted segments (Plate XX XI. fig. 5). Stamens, five,
inserted at the upper part of the corolla, shorter than the limb (Plate XX XI.
figs. 5,6, 7, a). Pollen roundish (Plate XXXII. fig. 14). Pistil, consisting
of an inferior bilocular ovary, with an ovule in each division; stigma bifid
(Plate XX XI. figs. 5, 6,7, c). The stamens and pistil are found to vary in
length in different flowers. The plant is thus dimorphic. In the figure
given by Sir W. Hooker in the “Botanical Magazine,” the stamens are long
and the style short. This character is seen in the specimens propagated from
the plant in the Botanic Garden (Plate XX XI. fig. 5). In the plants produced
from the Rio Janeiro specimens, two forms of flower are seen, viz., one with
a short style and long stamens, as in the Hookerian plant, and another with a
long style and short stamens. These two forms are seen in Plate XX XI. figs.
6 and 7, where a marks the stamens, and 0 the style. It is only within the last
year that fruit has been produced on the plants by artificial fertilisation. In
the Hookerian plant, Mr Linpsay, the house-foreman and propagator, fertilised
the pistil with pollen from the same flower. Fruit was produced, but not so
abundantly as in the dimorphic forms when fertilised by applying pollen from
the long stamens of one flower to the long pistil of another.

In Plate XXXII. fig. 1, a representation is given of a stem (a) bearing
elliptical wavy leaves (4, 6) with short petioles, and a cluster of fruit (c) borne
on a peduncle, which is bent downwards. The bending of the peduncle takes
place after flowering, and gradually increases during fruiting, until it forms a

PROFESSOR BALFOUR ON THE IPECACUAN PLANT. 785

more or less acute angle with the lower portion of the stem. In Plate XXXII.
fig. 10, there is shown the full-grown fruit (natural size) of a Rio Janeiro
plant, artificially fertilised, supported on its peduncle, with the involucre at the
base of the fruit. In fig 11 the fruit is magnified about one-half more than
natural. The fruit is drupaceous, of a deep purple violet colour, and shining
lustre. It consists of a finely coloured epicarp ; a whitish, pasty, and nearly
tasteless mesocarp, enclosing two hard stony nucules, each contaiing a hard
albuminous ovate seed with a minute embryo. The fruit produced by plants
with short styles (as seen in Plate XXXII. fig. 1, c) is short and round ; that
from the plants with long styles (as seen in Plate X XXII. fig. 10) is larger and
slightly narrowed at the apex. In Plate XXXII. fig. 11, the two central
nucules are shown, convex on the outside, and nearly flat on the inner side,
with a ridge. Fig. 12 shows one of the nucules separated, exposing its
inner surface with its single rib. Fig. 13 shows the hard seed, having the
form of the endocarp, the cavity of which it fills completely, and having a groove
on its flat surface. The albumen consists of thick-walled starch cells (Plate
XXXII. fig. 15). There is a minute central embryo.

Memorandum as to the Mode of transmitting Specimens of the Ipecacuan Plant
to India. [Added October 1872.]

In August 1869, Mr M‘Nas made cuttings of the root of the Ipecacuan,
and young plants were produced freely. Early in 1871 there was a large stock
of well-grown plants, which were sent to India in Wardian cases. The con-
struction of these and the mode of packing are detailed in Mr M‘Napz’s paper in
the “Transactions of the Botanical Society of Edinburgh,” vol. x. The plants
were successfully transmitted under the care of forest officers and gardeners who
happened to be going to India. In one instance cases were sent by the Messrs
Lawson without any one to look after them, and the plants arrived in safety.
In one of the cases sent from the Botanic Garden, every plant was in a good
condition when they reached Calcutta.

At first, most of the plants were sent in earth placed in pots, well fastened
down in the case. Afterwards sphagnum moss was employed, and this method
_ is strongly recommended by Mr M‘Naz. With the view of sending a large
supply, the plants were taken out of the pots, wrapped round with fresh moss,
and closely packed, so that a case 24 inches long and 16 broad contained easily
fifty or sixty well-grown plants.

I am disposed to think it is possible to send out the roots of the Ipecacuan
attached to the stem, but without leaves, in dry soil, made up of peat and sand,
and that they may even be transmitted by post in a close box. Boxes with

VOL. XXVI. PART. IV. 9 T

786 PROFESSOR BALFOUR ON THE IPECACUAN PLANT.

plants in a withered state, but with the roots in dry earth, reached the garden
safely from Rio Janeiro. They had been kept in a dry place, and not watered.
Their roots had been dormant, and they were ready to sprout when planted
out and watered. By imitating nature, and allowing the plants to remain dor-
mant for a time, the vitality of the roots had not been destroyed, and much
trouble was avoided in transport. Under-ground stems or sprouting-roots may
be kept for a long time in a dry condition. If this plan were adopted, a far
larger number of Ipecacuan plants, having the upper part of the stem and leaves
cut off, might be transmitted in a state fit for germination and for yielding cut-
tings when placed in favourable circumstances as regards moisture and heat
combined. The vitality of rhizomes even in a dry state is very great. Dr
GEORGE HENDERSON of the Bengal Service, who is about to superintend the
Calcutta Garden during Dr K1ne’s absence, will take with him plants prepared
in this manner. I have also supplied a small box containing roots in a dry state
for transmission to Calcutta through the post. If the method succeeds, there
will be a great saving of trouble and expense.

There is now a good stock of plants in India, and I have no doubt that,
from the roots of those now in cultivation there, a large stock of young plants
may be speedily produced in Sikkim, so as to furnish an abundant supply
of this most important drug for our Indian possessions. Mr AnpREw T. JAF-
FREY, in a letter to Sir Rosert Curistison, dated Darjeeling, 19th September
1872, states that by the end of the year he expects to be able to report that he
has 2000 to 3000 plants of Ipecacuan in cultivation. I may also state that Mr
LinpsAy finds that the leaves, taken from the plant and placed with their
petioles in damp warm sand, and covered by a bell-glass, give off abundance of
roots. It still remains to be determined whether these roots will be developed
sufficiently to furnish cuttigs for propagating the plant.

EXPLANATION OF PLATES XXXI. AND XXXII.
Illustrating the Form and Structure of the Ipecacuan Plant.
Puate XXXII.

Figure 1. Plant of Ipecacuan (Cephaélis Ipecacuanha, Rich.) grown in the Royal Botanic Garden of
Edinburgh. It is represented about half the natural size. The stem (a, a, a, a) is woody and
branching. It is, however, usually simple. The roots (}, b, b) come off from the lower part
of the stem, and are annulated. It can be cut into sections, which produce leaf-buds, and
seem to combine the characters of stem and root. The leaves (¢, c, c) are opposite, and
have oval or elliptical forms with a blunt point. From the upper part of the stem proceed
peduncles bearing capitula of flowers. At first the capitula are erect, as seen in the branch
to the left ; while in that to the right they are bent down by a change in the direction of
the peduncle.

PROFESSOR BALFOUR ON THE IPECACUAN PLANT. 787

Figure 2. Elliptical leaf, about the average natural size, having a blunt apex, with midrib and curved
veins indicated by dotted lines,

Figure 3. An annulated root, slightly enlarged. This is the pharmaceutical part of the plant. The
cortical portion (a, a) composed of cells containing starch grains; this portion is in the form
of rings closely applied to each other. The meditullium (0) is the central woody portion.
Processes are seen projecting from this, which are concerned in the formation of leaf-buds.

Figure 4. Capitulum, or head of gamopetalous funnel-shaped sessile flowers, white and sweet-scented,
having each a five-divided limb, and being all surrounded by a four-leaved involucre. The
capitulum is magnified rather more than twice the natural size.

Figure 5. Flower from the old Hookerian specimen of the plant, magnified. The corolla is split down
to the base, to show the organs of reproduction. The corolla is funnel-shaped, and has a five-
divided limb, with broadly ovate and pointed segments. The ovary (c) is inferior, and is
crowned by the irregularly toothed limb of the calyx. The stamens (a) are five in number,
and in this case are longer than the style (6), which terminates in a two-lobed stigma.

Figures 6 and 7 show the dimorphic flowers of the plant. (Magnified.) The two forms are required
for complete fertilisation.

Figure 6. Flower taken from one of Dr Gunnine’s plants, showing ovary (c) crowned by calyx ;
tubular corolla, with one of the segments cut off to show the long stamens (a) ; and the short
style (6) with stigma.

Figure 7. Another flower taken from one of Dr Gunwine’s plants, with a portion of the corolla
removed to show the short stamens (a) and the long style (0); the ovary (c) is inferior.

Puate XXXII.

Figure 1. Portion of Ipecacuan plant (Cephaélis Ipecacuanha), showing a somewhat shrubby stem (a),
wavy leaves (0, 6), and drupaceous fruit (c), supported on a peduncle, which is bent down-
wards. This is a young plant about the natural size.

Figures 2 and 3 show microscopical sections of the young stem, about one year old. ‘The dissection
made by Mr Joun SapLeEr, my assistant.

Figure 2. Transverse section of young herbaceous stem, the epidermis (a, a) shows delicate cellular
hairs, the cortical portion (0, b) composed of angular cells, vascular bundles in wedges (c, c),
cellular pith (d). Magnified about thirty diameters.

Figure 3. Longitudinal section of the same stem more highly magnified. ‘This section does not extend
to the epidermis. Cortical cells (0, b); vascular bundles (c, ¢c), consisting of spiral, pitted,
and woody tubes ; hexagonal pith (d).

Figures 4 and 5. Sections of the root, also made by Mr Saptzr.

Figure 4. Transverse section of the root (magnified), showing outer epidermal portion (a, a); the
cortical portion (4, 6), composed of cells containing many starch granules; central portion,
meditullium (c), composed of wood tubes.

Figure 5. Longitudinal section of the root (magnified), showing epidermal portion (a, a), containing
starchy cells (0, 6); central woody and pitted vessels (c).

Figure 6. Two portions of the annulated root bearing leaf-buds, showing the mode in which the plant
may be propagated. The root having both the functions of an ordinary root anda stem. <A
very small portion of a root, not larger than the § of an inch, will do for the purpose of
propagation.

Figure 7. Young Rio Janeiro plant, from Dr Gunnine’s specimens, showing somewhat quadrangular
herbaceous stem, opposite, ovate, acute, delicate leaves.

Figure 8. United stipules which embrace the stem ; each is divided at the apex into four long narrow
segments. Magnified about six times.

Figure 9. Glands at the base of the stipules, of a somewhat ovate-lanceolate shape, composed of cells.
Highly magnified.

788 PROFESSOR BALFOUR ON THE IPECACOUAN PLANT.

Figure 10. Fruit of one of the Rio Janeiro plants, of the natural size ; four in a cluster. The plant has
dimorphic forms of flowers. It was artificially fertilised. The fruit is large, slightly narrowed
towards the apex, shining, and of a deep violet purple. The succulent portion is insipid.

Figure 11. One of the fruits magnified about twice the natural size, showing external epicarp, which is
coloured ; mesocarp within, colourless ; and two pale hard nucules, corresponding to divisions
of the endocarp, each of which contains one hard seed.

Figure 12. One of the nucules removed, showing its flattish inner surface, with a ridge in its centre.

Figure 13. Seed shown separately, with its flat surface having a groove in its centre. It consists of
hard horny albumen and a minute embryo. ;

Figure 14. Pollen grains, highly magnified, of a somewhat irregular rounded form.

Figure 15. Section of albumen of the seed, showing thick-walled cells, with starch grains.

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PROCEEDINGS

OF THE

STATUTORY GENERAL MEETINGS ;

AND

LIST OF MEMBERS ELECTED AT THE ORDINARY MEETINGS;
WITH

LIST OF DONATIONS TO THE LIBRARY,

From NovEMBER 1869 To NovEMBER 1872.

VOL. XXVI. PART IV. 9U

PROCEEDINGS, &c.

Monday, 22d November 1869.

At a Statutory General Meeting, Professor KELLAND, Vice-President, in the Chair, the
Minutes of the Statutory Meeting of 23d November 1868 were read and confirmed.

The following Office-Bearers were elected for 1869-70 :—

Professor Curistison, M.D., D.C.L., President.

His Grace the Durer of Araytn, Honorary Vice-President.
Professor Lyon Prayrarr, C.B.,
Davip Mitne Home, LL.D.
Professor KELLAND,

The Hon. Lorp Nzavss,
Professor Sir WiuLiam THomson,
Wiu1am Forses SKENE, Esq.,
Dr Joun Hurron BaxFour, General Secretary.
Professor Tart,
Professor TURNER,
Davin SuitH, Esq., Treasurer.

Dr Mactacan, Curator of Library and Museum.

Vice-Presidents.

\ Secretaries to Ordinary Meetings.

COUNCILLORS.
Grorce Ropertson, Esq., C.E. THomas StEvensoN, Esq., C.E.
Professor Piazzi Smytu. Dr Hanpystne.
Patrick Dupcxon, Esq. of Cargen. ARCHIBALD GEIKIE, Esq.
Dr Hue Ciecuorn. Professor A. Crum Brown.
Dr James M‘Batn, R.N. Principal Sir ALEXANDER GRANT.
Dr Witiiam Ropertson. Rev. W. Linpsay ALexanper, D.D.

The TREASURER gave in his annual printed Report, certified by the Auditor.

GEORGE AULDJO JAMIESON, Esq., was elected Auditor for the year 1869-70.

The SECRETARY reported as follows :—

Number of Ordinary Fellows at November 1868, : : 289

New Fellows, 1868-69, 3 : ; ; 26

Total, 315

Deduct—Deceased, 10 ; resigned, 2, : : é ‘ 12
Number of Ordinary Fellows at November 1869, . : 303 4

(Signed) BR. Curistison, President.

PROCEEDIN GS OF STATUTORY GENERAL MEETINGS. 791

Monday, 28th November 1870.

At a Statutory General Meeting, Dr CuHrisTison, President, in the Chair, the Minutes
of the Statutory Meeting of 22d November 1869 were read and confirmed.

The following Office-Bearers were elected for 1870-71 :—

Professor Curistison, M.D., D.C.L., President.

His Grace the Duxr of Areyit, Honorary Vice-President.
Davip Mitng Home, LL.D.,
Professor KELLAND,

The Hon. Lorp Neaves,
Professor Sir WiLt1aAm THOMSON,
Witt1am Forses Skene, Esq.,
Principal Sir Auex. Grant, Bart.,

Vice-Presidents.

Dr Joun Hurton Batrour, General Secretary.
Professor Tart,

Secretaries to Ordinary Meetings.
Professor TURNER, \ Ta a oe
Davip Situ, Esq., Treasurer.

Dr Mactacan, Curator of Library and Museum.

COUNCILLORS.
Dr James M‘Bary, R.N. - Rev. W. Linpsay ALexanpsr, D.D.
Dr Wittiam Ropertson. Professor FLuEMiIne JENKIN.
Tuomas Stevenson, Esq., C.E. Professor Wrvittr THomson, LL.D.
Dr HanpysiDE. James Donaupson, LL.D.
ARCHIBALD GEIKIE, Esq. Dr THomas R. Frasmr.
Professor A. Crum Brown. Dr ArtHuR GAMGEE.

The TREASURER gave in his annual printed Report, certified by the Auditor.
GEORGE AULDJO JAMIESON, Esq., was elected Auditor for the year 1870-71.

The SrcreTary stated that the Council had memorialised the First Lord of H. M. Treasury
relative to the establishment of a Chair of Geology in the University of Edinburgh, for which
Sir RopErRicK Murcuison had offered the sum of L.6000. The memorial was laid on the
table.

The SECRETARY intimated that, in conformity with the request of the Council, Davin
Mine Home, Esq., had kindly consented to give the Opening Address on Monday, 5th
December.

The SECRETARY reported:as follows :-—

Number of Ordinary Fellows at N: ovember 1869, s ; 303
New Fellows, 1869-70, : : : : 30

Total, 333

Deduct—Deceased, 5 ; resigned, 2, ; p , = 7
Number of Ordinary Fellows at November 1870, : ; 326

(Signed) PHILIP KELLAND, Vice-President.

792 PROCEEDINGS OF STATUTORY GENERAL MEETINGS.

Monday, 27th November 1871.

At a Statutory General Meeting, Professor KELLAND, Vice-President, in the Chair, the
Minutes of the Statutory Meeting of 28th November 1870 were read and confirmed.

The following Office-Bearers were elected for 1871-72 :—

Sir Ropert Curistison, Bart., M.D., D.C.L., President.
His Grace the Dux of Arayi~u, Honorary Vice-President.
Professor K=LLAND,

-The Hon. Lorp Nuaves.

Professor Sir WiLt1amM THomson,
Principal Sir ALEx. Grant, Bart.,
Sir W. Strrtine Maxwet., Bart.,
Professor W. J. Macquorn RankINe,
Dr Joun Hutton Batrour, General Secretary.
Professor Tart,
Professor TURNER,
Davin Situ, Esq., Treasurer.

Dr Mactaean, Curator of Library and Museum.

Vice-Presidents.

Secretaries to Ordinary Meetings.

COUNCILLORS.
Professor GEIKIE. Dr Tuomas R. Fraser.
Professor A. Crum Brown. Dr ArtHur GAMGEE.
Rev. W. Linpsay Atexanper, D.D. Atmxanprr Bucuan, M.A.
Professor FLEEMING JENKIN. Professor A. Dickson.
Professor Wryvitte THomson. Davip Mitne Hons, LL.D.
James Donatpson, LL.D. James Lesuiz, Esq., C.E.

The TREASURER gave in his annual printed Report, certified by the Auditor.
GEORGE AULDso JAMIESON, Esq., was elected Auditor for the year 1871-72.

The SECRETARY reported as follows :—

Number of Ordinary Fellows at November 1870, : : 326
New Fellows, 1870-71, , : . \ 15

Total, 341

Deduct—Deceased, 10; resigned, 3, ; ae : = 13
Total Ordinary Fellows at November 1871, ; ; : 328
Honorary Fellows deceased— British, 2; Foriegn, 1, . Total, 3

(Signed) _R. Curistison, President.

LIST OF MEMBERS ELECTED. 793

LIST OF MEMBERS ELECTED.

December 20, 1869.
Sr Jonn Vincent Day, Esq., C.E. Davip Munn, Esq.
Rozert R. Tatiocr, Esq.

January 3, 1870.

ALEXANDER RusseEL, Esq. Dr James Cricuton Browne.
Dr Jonn Duncan. Witiiam Burns THomson, Esq.
Dr Wittram R. Sanvers. Rev. Dr AnpREw THomson.
Professor JosEePH LisTER. Dr Witiiam ANDERSON.

January 17, 1870.

Dr G. H. B. Macteop. Dr Tuomas A. G. Banrour.
February 7, 1870.

W. E. Heaturimnp, Esq. Dr Epwarp Jamms SHEARMAN,

Parrick Swan, Esq. Dr. H. AtLEynE Nicnorson.

Rev. Dr Hopson (re-admitted).

February 21, 1870.
Dr J. WARBURTON BEGBIE.

March 7, 1870.
Joun Winzer, Esq.

March 21, 1870.

Spencer C. THomson, Esq. Simon 8. Lauris, Esq.
May 2, 1870.
James Sime, Esq. THomas Harvey, Esq.
Joun Youne Bucwanay, Esq. Joun Hunter, Esq.
The Right Hon. The Lord Justice Cumrx. The Hon. Lord Grrrorp.

May 16, 1870.
James Watson, Esq. The Hon. Lord Mackenzie.

December 5, 1870.
Joun AvLp, Esq.

January 16, 1871.
Rev. THomas M. Liypsay. Wittiam Rosertson Situ, Esq.
Stair AcNew, Esq.

January 30, 1871.
Dr CHartes Hayss Hicerns. Dr Aneus Macponatp.
February 6, 1871.

Rev. Wituiam Scorr Moncrierr. Professor A. R. Simpson.
Dr R. J. Buairn CuNYNGHAME. Dr Cosmo Gorpon Login.

VOL. XXVI. PART IV. 9x

794 LIST OF MEMBERS ELECTED.

April 3, 1871.
Jamus Gerkin, Esq. ' Dr Tuomas E. THorps.

April 17, 1871.
Dr Joun Smita, F.R.C.S.E.

May 1, 1871.
Rev. Professor CRAWFORD.

May 15, 1871.
Tuomas J. Boyp, Esq.

December 4, 1871.
ALEXANDER H. Len, Esq., C.E. Rosert Les, Esq., Advocate.
Joun Anprrson, LL.D.

January 15, 1872.

Davip Mactaean, Esq., C.A. Major Rickarp.

Dr Joun Sippap. Dr J. G. Fremine.
Rev. Anprew Tart, LL.D. Davin Grigve, Esq.
The Right Rev. Bishop Correritt. Grorce Barctay. Esq.

February 5, 1872.
GrorcrE Forses, Esq., B.A. Dr J. Linpsay Stewart.
‘Rev. Cuartus R. Tears, M.A.

February 19, 1872.
ARCHIBALD ConsTaBLE, Esq.

March 18, 1872.
GEORGE Seton, Esq. Captain Cuartes Hunter.

April 1, 1872.
James THomson Borromury, Esq. _ ’ Tuomas Knox, Esq.
Dr D. Arcyit Ropertson.

2 April 15, 1872.
Dr Tuomas B. Curistin.

May 20, 1872.
Rev. Hue Macmituan, LL.D.

1846
1871
1868
1866

1867

1848.

1856

1849
1872

1845
1823
1867
1862
1849
1871

1843

~ 1835

1870
1867
1872
1858
1870
1843

1861
1866
1850

1863
1857
1862
1854
1872
1869
1871
1864

1859
1861
1835
1870
1867
1856
1833
1869
1870
1857
1847

1869

1865
1866
1860

1872
1823

1863
1856
1844
1829
1829
1850

ALPHABETICAL LIST OF THE ORDINARY FELLOWS OF THE SOCIETY,
Corrected up to 15th January 1873.

N.B.—Those marked * are Annual Contributors.

Alex. J. Adie, Esq., Rockville, Linlithgow
*Stair Agnew, Esq, 2 Buckingham Terrace
*Rey. Dr David Aitken, 4 Charlotte Square
*Major-General Sir James EH. Alexander of Westerton,
Bridge of Allan
*Rey. Dr W. Lindsay Alexander, Pinkie Burn, Mussel-
burgh
Dr James Allan, Inspector of Hospitals, Portsmouth
Dr G. J. Allman, Emeritus Professor of Natural History,
Wimbledon, London
*David Anderson, Esq., Moredun, Edinburgh
John Anderson, LL.D., 32 Victoria Road, Charlton,
Kent .
Dr Thomas Anderson, Professor of Chemistry, University,
Glasgow 10
Warren Hastings Anderson, Esq., Isle of Wight
*Thomas Annandale, Esq., 34 Charlotte Square
*T, C. Archer, Esq., Director of the Museum of Science
and Art, 5 West Newington Terrace
*His Grace the Duke of Argyll, K.T., (Hon. Vicz-
PRESIDENT), Inverary Castle
*John Auld, Esq., 18 Grosvenor Crescent

Dayid Balfour, Esq., Trenaby
Dr J. H. Balfour (GENERAL SECRETARY), Professor of
Medicine and Botany, 27 Inverleith Row
*Dr Thomas A. G. Balfour, 51 George Square
*George F. Barbour, Esq., 11 George Square
*George Barclay, Esq., 17 Coates Crescent 20
Edmund C. Batten, M.A., Lincoln’s Inn, London
*Dr James Warburton Begbie, 16 Great Stuart Street
Dr Bennett, Professor of Institutes of Medicine, 1 Glen-
finlas Street
*George Berry, Esq., 2 Windsor Terrace, Portobello
*Adam Black, Esq., 38 Drummond Place
*Hugh Blackburn, Esq., Prof. Mathematics, University,
Glasgow
*Professor Blackie, 24 Hill Street
*John Blackwood, Esq., 3 Randolph Crescent
*Rev. Dr W. G. Blaikie, 9 Palmerston Road
Ernest Bonar, Esq. 30
*James Thomson Bottomley, Esq., College, Glaszow
*Robert Henry Bow, Esq., C.E., 7 South Gray Street
*Thomas J. Boyd, Esq., 41 Moray Place
*Dr Alex. Crum Brown, Prof. of Chemistry, 8 Belgrave
Crescent
*Dr John Brown, 23 Rutland Street
*Rey. Thomas Brown, 16 Carlton Street
William Brown, Esq., 25 Dublin Street
Dr James Crichton Browne, Wakefield
*A. H. Bryce, D.C.L., LL.D., 42 Moray Place
*Dayvid Bryce, Esq., Architect, 131 George Street 40
His Grace the Duke of Buccleuch, K.G., Dalkeith Palace
* Alexander Buchan, A.M., 72 Northumberland Street
*John Young Buchanan, Esq., 10 Moray Place
*Dr W. M. Buchanan, 3 Carlton Terrace
J. H. Burton, LL.D., Advocate, Craig House

*Rev. Henry Calderwood, LL.D., Professor of Moral
Philosophy, Craigrowan, Napier Road, Merchiston

*Alfred R. Catton, B.A.

*Dayid Chalmers, Esq., Kate’s Mill, Slateford

*William Chambers, Esq. of Glenormiston, 13 Chester

Street
Dr Thomas B. Christie, Royal India Asylum, Ealing,
London 50

Sir Robert Christison, Bart., D.C.L., Professor of Materia
Medica (PrEsIDENT), 40 Moray Place
Dr H. F.C. Cleghorn, Stravithy, St Andrews
*Thomas Cleghorn, Esq., Advocate, 26 Queen Street
Dr Thomas R. Colledge, Lauriston House, Cheltenham
The Right Honourable Lord Colonsay, London
A. Colyar, Esq.
*Dr James Scarth Combe, 36 York Place

1872
1843

1872
1843
1863
1854
1830
1829
1871

1853
1852
1871
1823

1851
1841
1867
1848
1870

1867
1869
1869

1869
1867
1863
1867
1866
1839

1868
1867
1860
1863
1870
1851
1859
1866

1869
1856
1855
1866
1863

1866
1859
1868
1858
1852
1872
1872

1859
1828
1864
1858
1867
1867

1867
1867
1868
1861

1871
1870
1868
1846

*Archibald Constable, Esq., 11 Thistle Street
Sir A ohn Rose Cormack, M.D., 7 Rue d’Aguesseau,
aris
*The Right Rev. Bishop Cotterill, 24 Rutland Square 60
Andrew Coventry, Esq., Advocate, 29 Moray Place
*Charles Cowan, Esq., Westerlea, Murrayfield
*Sir James Coxe, M.D., Kinellan
J. T. Gibson-Craig, Esq., W.S., 24 York Place
Sir William Gibson-Craig, Bart,, Riccarton
oRey, Dr Crawford, Professor of Divinity, 18 Great King
treet ‘
Rey. John Cumming, D.D., London
*James Cunningham, Esq., W.S., 50 Queen Street
*Dr R. J. Blair Cunyninghame, 6 Walker Street
Liscombe J. Curtis, Esq., Ingsdown House, Devonshire 70

*K. W. Dallas, Esq., 34 Hanover Street
James Dalmahoy, Esq., 9 Forres Street
*David Davidson, Esq., Bank of Scotland
Henry Davidson, Esq., Muirhouse
*St John Vincent Day, Esq., C.E., 4 Hamilton Park
Terrace, Glasgow
*Francis Deas, LL.B., Advocate, 9 St Colme Street
*James Dewar, Esq., 15 Gilmore Place
*Alexander Dickson, M.D., Professor of Botany, University
of Glasgow
*William Dickson, Esq., 38 York Place
Henry Dircks, LL.D., C.E., London 80
*W. Dittmar, Esq.
*James Donaldson, LL.D., 20 Great King Street
*David Douglas, Esq., 41 Castle Street :
Menge Brgwn Douglas, Esq., Advocate, 21. Moray
ace.
*Rey. D. T. K. Drummond, B.A.y 6 Montpelier
*G. Stirling Home Drummond, Esq., Blair-Drummond
*Patrick Dudgeon, Esq. of Cargen
*Dr J. Matthews Duncan, 30 Charlotte Square
*Dr John Duncan, 8 Ainslie Place
*Sir David Dundas, Bart. of Dunira 90
*Rey. Dr John Duns, 4 Mansion-House Road, Grange
*Dr James Dunsmure, 53 Queen Street

*George Elder, Esq., Knock Castle, Wemyss Bay
*W. Mitchell Ellis, Esq., Wellington Lodge, Portobello
Robert Etheridge, Esq., Clifton, Bristol
*William Euing, Esq., Glasgow
J. D. Everett, LL.D., Prof. Nat. Phil., Queen’s College,
Belfast

*James Falshaw, Esq., C.E., 26 Castle Street
*Dr Fayrer, Professor of Surgery, Calcutta :
*Robert M. Ferguson, Ph.D., 12 Moray Place 100
Frederick Field, Esq., Chili
Dr Andrew Fleming, H.M.I.S., Bengal
*Dr J. G. Fleming, B.A., 155 Bath Street, Glasgow
*George Forbes, Hsq., Lecturer on Natural Philosophy,
Anderson Institution, Glasgow, 4 Coates Crescent
Major James George Forlong, Bombay
John Forster, Esq., Liverpool
*Dr John Foulerton, Manila
*Professor Fraser, M.A., 20 Chester Street
*Dr Thomas R. Fraser, 3 Grosvenor Street
*Frederick Fuller, Esq., Professor of Mathematics, Uni-
versity, Aberdeen 110

Dr Charles Gayner, Oxford
*Dr Arthur Gamgee, 27 Alva Street
J. Samson Gamgee, Esq., Birmingham
*A. Geikie, Esq., Professor of Geology, Geological Survey
Office, India Buildings, George IV. Bridge
*James Geikie, Esq., 16 Duncan Street, Newington
*Hon. Lord Gifford, Granton House
*Rey. Joseph Taylor Goodsir, 11 Danube Street
L. D. B. Gordon, Esq., C.E., London

1850
1867
1869

1851
1824

1872
1860

1868
1868

1867
1867
1867
1833

1837
1854
1869

1867
1870
1859
1855
1870

1862
1869

1871
1859
1828

1870
1869
1872

1870
1864
1855
1858

1840
1863
1860
1825

1869
1865
1863

1869
1867

1867
1866
1839

1868
1872

1868
1870
1863
1865
1856

1872
1872
1863
1858
1871

1861

( 796 )

*Lieut.-Col. W. D. Gosset, R.E., Portsmouth

*Dr Andrew Graham, R.N., 35 Melville Street. 120

*Principal Sir Alex. Grant, Bart., (VICE-PRESIDENT), 21
Lansdowne Crescent

*Rey. Dr James Grant, D.C.L., 15 Palmerston Place

Dr Robert E. Grant, Prof. Comp. Anat., Univ. Coll.,

London

*David Grieve, Esq., 13 Lochend Road, Leith

*Dr Frederick Guthrie, M.A., Prof. of Physics, School
of Mines, London

*Col. Seton Guthrie, Thurso

*Rey. Dr Thomas Guthrie, 1 Salisbury Road

*Dr D. R. Haldane, 22 Charlotte Square
*Frederick Hallard, Esq., Advocate, 61 York Placé
*James H. B. Hallen, Esq., Canada 130
Alexander Hamilton, LL.B., W.S., The Elms, Whitehouse
Loan
Dr P. D. Handyside, 11 Hope Street
Professor Robert Harkness, (Queen’s College, Cork
Sir Charles A. Hartley, C.E., Sulina, Mouth of the
Danube
*Sir George Harvey, 21 Regent Terrace
*Thomas Harvey, Esq., LL.D., 32 George Square
*G. W. Hay, Esq. of Whiterigg
*James Hay, Esq., 3 Links Place, Leith
W. E. Heathfield, Esq., 20 King Street, St James,
London
*Dr James Hector, Wellington, New Zealand 140
*Isaac Anderson-Henry, Esq. of Woodend, Hay Lodge,
Trinity
Dr Charles Hayes Higgins, Alfred House, Birkenhead
Lieut. John Hills, Bombay Engineers
David Milne Home, Esq. of Wedderburn, LL.D. (Vicr-
PRESIDENT), 10 York Place
*Rey. Dr Hodson, St Andrew’s College, Broadfield, Reading
*Alexander Howe, Esq., W.S., 17 Mray Place
*Captain Charles Hunter, Glencarse, Junior Naval and
Military Club, London
John Hunter, Esq., Professor of Mathematics, King’s
College, Windsor, Halifax
*Robert Hutchison, Hsq., Carlowrie Castle

*The Right Hon. John Inglis, D.C.L., LL.D., Lord Justice-
General, 30 Abercromby Place 150
*Professor Innes, M.A., Inverleith House

Edward J. Jackson, Esq., 6 Coates Crescent
William Jameson, Hsq., Surgeon-Major, Saharunpore
*George A. Jamieson, Esq., 58 Melville Street
Sir William Jardine, Bart., LL.D., of Applegarth, Jardine
Hall, Lockerby
*Protessor H. C. Fleeming Jenkin, 5 Fettes Row
*Charles Jenner, Esq., Haster Duddingston Lodge
*“Hon. Charles Baillie, LL.D., Lord Jerviswoode, 10
Strathearn Road
Dr John Wilson Johnston, India
*T, B. Johnston, Esq., 9 Claremont Crescent 160
*William Keddie, Esq., 5 India Street, Glasgow
*Dr Alexander Keiller, 21 Queen Street
Rey. Prof. Kelland, M.A. (Vicz-PrusIDENT), 20 Claren-
don Crescent
*Thomas Key, Esq., 42 George Square
*Thomas Knox, Esq., 2 Dick Place

*J. W. Laidlay, Esq., Seacliff

*Simon S. Laurie, Esq., Brunstane House, Portobello

*Charles Lawson, Hsq., 35 George Square

*Charles Lawson, jun., Esq., 35 George Square

*Dr Laycock, Professor of the Practice of Medicine, 13
Walker Street 170

*Alexander H. Lee, Esq., C.E., 45 Moray Place

*Robert Lee, Esq., Advocate, 26 Charlotte Square

*Hon. G. Waldegrave Leslie, Leslie House, Leslie

*James Leslie, Esq., C.E., 2 Charlotte Square

*Rev. Thomas M. Lindsay, Professor of Divinity and
Church History, Free Church College, Glasgow

*Dr W. Lauder Lindsay, Gilgal, Perth i

1864
1870
1871

1857
1861
1869
1849

1855

1861
1868
1867
1866
1871
1820

1847
1869
1840
1870
1843

1853
1869
1870

1869
1872
1864
1869
1868
1872

1866
1840
1858
1869
1864
1866
1856

1849
1863
1853
1841
1869
1852
1833
1866
1843
1865
1870

1871

1868
1866
1861
1870
1857

1856

1866
1870
1847
1863
1837
1863
1868
1869
1849

1859
1834

*William Lindsay, Esq., Hermitage-Hill House, Leith
*Professor Lister, 9 Charlotte Square '
*Dr Cosmo Garden Logie, Surgeon Major, Royal Horse
Guards

Thomas Login, Esq., C.E., India
*Professor Lorimer, Advocate, 21 Hill Street
*Maurice Lothian, Esq. of St Catherine’s, 54 Queen Street
*Dr W. H. Lowe, Balgreen, Murrayfield

180

*Dr Stevenson Macadam, 11 East Brighton Crescent, Porto-
bello
*Dr James M‘Bain, R.N., Logie Villa, York Road, Trinity
*Dr Thomas Smith Maccall, Polmont
*John M. M‘Candlish, Esq., 4 Doune Terrace
*John M‘Culloch, Esq., Banker, 11 Duke Street
*Dr Angus Macdonald, 41 Northumberland Street
Dr Wm. Macdonald; Prof. Civ. and Nat. Hist., St
Andrews 190
W. Macdonald Macdonald, Esq., St Martins
*David MacGibbon, Esq., Architect, 89 George Street
John Mackenzie, Esq., 11 Abercromby Place
*Hon. Lord Mackenzie, 12 Great Stuart Street
Dr Maclagan (Curator), Prof. of Medical Jurisprudence,
28 Heriot Row
Lieut.-Col. R. Maclagan, Royal Engineers, Bengal
*Dr R. Craig Maclagan, 5 Coates Crescent
*Dr G. H. B. Macleod, Professor of Surgery, University,
Glasgow
*Dr William C. M‘Intosh, Murthly
*David Maclagan, Esq., C.A., 9 Royal Circus 200
*Peter M‘Lagan. Esq. of Pumpherston, M.P.
*John M‘Laren, Esq., Advocate, 5 Rutland Square
*John F. M‘Lennan, Esq., Advocate, 81 Princes Street
*Rev. Hugh Macmillan, LL.D., 30 Hamilton Park Ter-
race, Glasgow
*John Macnair, Esq., 33 Moray Place
Sir John M‘Neill, G.C.B.
*Dr R. B. Malcolm, 126 George Street
Dr Henry Marshall, Clifton, Bristol
*J. D. Marwick, Esq., 10 Bellevue Crescent
*Professor David Masson, 10 Regent Terrace 210
*James Clerk Maxwell, Esq., Prof. Exp. Phys., Cambridge,
Glenlair, Dalbeattie
*Sir William Stirling-Maxwell, Bart., Keir
*Edward Meldrum, Esq., Dechmont, Broxburn
*Greme Reid Mercer, Ksq., Ceylon Civil Service
John Miller, Esq., C.E., M.P., 2 Melville Crescent
*Oliver G. Miller, Esq., Panmure House, Forfarshire
*Thomas Miller, Esq., A.M., LL.D., Rector, Perth Academy
Admiral Sir Alexander Milne, G.C.B., Inveresk
*Dr Arthur Mitchell, 5 East Claremont Street
Joseph Mitchell, Esq., C.E., Viewill, Inverness
*Dr John Moir, 52 Castle Street
*The Right Hon. James Moncreiff, Lord Justice-Clerk, 15
Great Stuart Street
*Rev. William Scott Moncrieff, of Fossaway, 14 George
Square
*Rey. James F. Montgomery, 17 Atholl Crescent
*Dr Charles Morehead, 11 North Manor Place
*John Muir, D.C.L., LL. D., 10 Merchiston Avenue
*Dayvid Munn, Esq., 11 Gayfield Square
Dr John Ivor Murray, The Knowle, near Tunbridge Wells

220

*Hon. Lord Neaves, LL.D. (Vicz-PresiDENT), 7 Char-
lotte Square
*Thomas Nelson, Esq., Arthursley 230
*Dr Henry A. Nicholson, Prof. of Nat. Hist., Toronto
James Nicol, Esq., Prof. Nat. Hist., Aberdeen
*Hon. Lord Ormidale, 14 Moray Place
Dr Richard Parnell, Melrose
*Dr Alexander Peddie, 15 Rutland Street
*John Dick Peddie, Esq., Architect, 33 Buckingham Ter. —
John Pender, Esq., Manchester :
*W. Pirrie, Esq., Professor of Surgery, Marischal College, —
Aberdeen
*Lyon Playfair, C.B., LL.D., M.P., 4 Queensberry Place,
South Kensington, London

Mungo Ponton, Esq., W.S., Clifton, Bristol 240

1852

1865
1849

1868
1869
1865
1836
1872

1840
1872
1859
1832
1860
1862
1870
1852
1837

1869

1870

1863
1864
1849
1846
1853
1864
1872
1870

1834
1872
1870
1871
1829
1859
1868

839

863
1866
1871
1855
1871
1846
1866
1850
1843
1844
1868
1848
1858
1872

1868
1869

1866
1848

( 797 -)

Eyre B. Powell, Esq., Director of Public Instruction,
Madras

*James Powrie, Esq., Reswallie, Forfar
*Hon. B. F. Primrose, 22 Moray Place

*Samuel Raleigh, Esq., Park House, Dick Place
Rev. Thos. Melville Raven, M.A., Crakehall, Bedale
*Rey. Francis Redford, M.A., Rectory, Silloth
David Rhind, Esq., Architect, 54 Great King Street
Major F. Ignacio Rickard, Government Inspector-General
of Mines, Argentine Republic, Buenos Ayres, South
America
Martyn J. Roberts, Esq., Crickhowell, South Wales
*Dr D. Argyll Robertson, 40 Queen Street 250
*George Robertson, Esq., C.E., 47 Albany Street
Dr Montgomery Robertson, Mortlake, Surrey
*Dr William Robertson, 28 Albany Street
*Dr E. Ronalds, Bonnington Road
*Alexander Russel, Esq., 9 Chester Street
*Alex. James Russell, Hsq., C.S., 9 Shandwick Place
J. Scott Russell, Esq, 5 + Westminster Chambers,
London
*Dr William Rutherford, Professor of Physiology, King’s
College, London

*Dr William R. Sanders, Prof. General Pathology, 11
Walker Street
*James Sanderson, Esq., Surg.-Major, 41 Manor Place 260
*Reyv. D. F. Sandford, 19 Rutland Street
*Hdward Sang, Hsq., 2 George Street
Dr Schmitz, International Institution, London
*Hugh Scott, Esq. of Gala, Galashiels
*Professor Sellar, LL.D., 15 Buckingham Terrace
*George Seton, Esq., Advocate, 42 Greenhill Gardens
Dr Edward James Shearman, Moorgate, Rotherham,
Yorkshire
Dr Sharpey, Prof. Anatomy, Univ. Coll., London
*Dr John Sibbald. 16 Dalrymple Crescent
*James Sime, Esq., Craigmount House, Dick Place 270
*Dr A. R. Simpson, Prof. of Midwifery, 52 Queen St.
Ven. Archdeacon Sinclair, Kensington
*William F. Skene, LL.D., W.S., 20 Inverleith Row
*Adam Gillies Smith, Esq., C.A., 5 Lennox Street
David Smith, Esq., W.S. (TREASURER), 10 Eton Ter.
*Dr John Alexander Smith, 7 West Maitland Street
*Dr John Smith, F.R.C.P.E., 20 Charlotte Square
*Dr John Smith, F.R.C.S.E., 11 Wemyss Place
*R. M. Smith, Esq., 4 Bellevue Crescent
*W. R. Smith, Esq., Free Church Coll., Aberdeen 280
Professor Piazzi Smyth, 15 Royal Terrace
*Professor Spence, 21 Ainslie Place
*Dr James Stark, 21 Rutland Street
Henry Stephens, Esq., Red Braes Cottage, Bonnington
David Stevenson, Esq., C.E., 45 Melville Street
John J. Stevenson, Esq., Hyde Park, London
Thomas Stevenson, Esq., C.E., 17 Heriot Row
*Rev. Dr Stevenson, 37 Royal Terrace
Dr J. Lindsay Stewart, Conservator of Forests, Punjab,
India
Major J. H. M. Shaw Stewart, R. Engineers, Madras 290
*John L. Douglas Stewart, Esq. of Glenogil, 7 Grosvenor
Crescent
*Dr T. Grainger Stewart, 19 Charlotte Square
*Patrick J. Stirling, Esq., LL.D., KippendavieHouse

1823
1870
1848

1844

1872
1861

1870
1846
1872
1843
1870
1842

1863
1864
1870
1847

1870
1849
1855

1871

1822
1867
1861
1849

1867
1869
1829
1864
1853
1870
1866
1866
1862

1840

1869
1868
1858
1834
1847
1863
1870

1864
1864
1855
1864

1861
1863

Captain T. D. Stuart, H.M.1.S.
*Patrick D. Swan, Esq., Kirkcaldy
William Swan, Esq., Professor of Natural Philosophy
St Andrews
Archibald Campbell Swinton, Esq., Kimmerghame,
Dunse

Rey. Andrew Tait, Rector of Kilkerrin, Ireland
*Professor P. Guthrie Tait, M.A. (SecrETARY), 17 Drum-
mond Place
*Robert R. Tatlock, Esq., 151 George Street, Glasgow 300
Dr Taylor, Pau, France
*Rev. Charles R. Teape, 15 Findhorn Place
Dr Allen Thomson, Prof. Anatomy, Univ., Glasgow
*Rey. Dr Andrew Thomson, 63 Northumberland Street
James Thomson, Esq., C.E., Norfolk Square, Hyde Park,
London .
*Dr Murray Thomson, Roorkee, Kast Indies
*R. W. Thomson, Esq., C.E., 3 Moray Place
*Spencer C. Thomson, Esq., 10 Chester Street.
Sir William Thomson, Prof. Nat. Phil. (Vicr-Pre-
SIDENT), Glasgow
*William Burns Thomson, Esq., 11 St John Street 310
*William Thomas Thomson, Esq., Bonaly
le Thomson, LL.D., Prof. Nat. Hist., 20 Palmerston
ace
*Thomas E. Thorpe, Ph.D., Lecturer on Chemistry, Ander-
son Institution, Glasgow
Sir W. C. Trevelyan, Bart., Wallington, Morpeth
*William Turnbull, Esq., 14 Lansdowne Crescent
*Professor Turner, M.B. (SrcrErary), 6 Eton Terrace
*Most Noble the Marquis of Tweeddale, K.T., Yester
House, Haddington

*Peter Waddell, Esq., 5 Claremont Park, Leith
*Viscount Walden, Yester House, Haddington
James Walker, Esq., W.S., Tunbridge Wells 320
*William Wallace, Ph.D., Glasgow
Dr James Watson, Bath
*James Watson, Esq., 45 Charlotte Square
*John K. Watson, Hsq., 14 Blackford Road
*Dr Patrick Heron Watson, 16 Charlotte Square
*Rey. Robt. Boog Watson, Madeira, 4 Bruntsfield Place,
Edinburgh
Allan A. Maconochie Welwood, Esq. of Meadowbank
and Pitliver
*Captain T. P. White, Royal Engineers, 1 Drummond Place
*W. Williams, Esq., Veterinary College, Clyde Street
*Dr Thomas Williamson, 28 Charlotte Street, Leith 330
Dr Isaac Wilson
Professor John Wilson, College
*Dr J. G. Wilson, 9 Woodside Crescent, Glasgow
John Winzer, Esq., Assistant Surveyor, Civil Service,
Ceylon
*Dr Alexander Wood, 36 Moray Place
*Dr Andrew Wood, 9 Darnaway Street
Dr Wright, Cheltenham
*Robert S. Wyld, Esq., W.S., 19 Inverleith Row

*James Young, Esq., of Kelly, Wemyss Bay
*Dr John Young, Professor of Natural History, Glas-
gow 340

Fellows elected between the commencement of the Session and the 1st January of the following year are entered under the latter
date, by which their Subscriptions wre regulated :—Thus, Fellows elected in December 1871 have the date of 1872 vrefixed

to their names.

VOL. XXVI. PART IV.

( 798 )

LIST OF THE PRESENT ORDINARY MEMBERS,

Corrected up to January 15, 1873.

IN THE ORDER OF THEIR ELECTION.

PRESIDENT.

Str ROBERT CHRISTISON, Barr.

HONORARY VICE-PRESIDENT, HAVING FILLED THE OFFICE OF PRESIDENT.

His Grack THE DUKE OF ARGYLL, K.T.
Date of
Election.

1820 William Macdonald, M.D., F.R.C.P.E., Professor of Natural History, St Andrews.
1822 Sir W. C. Trevelyan, Bart., Wallington, Northumberland.
1823 Captain Thomas David Stuart, of the Hon. East India Company's Service.
Warren Hastings Anderson, Esq.
Liscombe John Curtis, Esq., Ingsdon-House, Devonshire.
Sir Robert Christison, Bart., M.D., Professor of Materia Medica.
1824 Robert E. Grant, M.D., Professor of Comparative Anatomy, University College, London.
1828 John Forster, Esq., Architect, Liverpool,
David Milne Home, LL.D., Advocate, of Milne-Graden and Wedderburn.
1829 A. Colyar, Esq. ;
Right Hon. Sir William Gibson-Craig, Bart. of Riccarton.
Right Hon. Lord Colonsay.
Venerable Archdeacon Sinclair, Kensington.
James Walker, Esq., W.S.
1830 J. T. Gibson-Craig, Esq., W.S.
1832 Montgomery Robertson, M.D.
1833 Admiral Sir Alexander Milne, R.N., G.C.B.
His Grace the Duke of Buccleuch, K.G., Dalkeith Palace.
Alexander Hamilton, LL.B., W.S.
1834 Mungo Ponton, Esq., W.S., Clifton, Bristol.
Isaac Wilson, M.D., F.R.S., Lond.
William Sharpey, M.D., LL.D., F.R.S., Professor of Anatomy, University College, London.
1835 John Hutton Balfour, A.M., M.D., F.R.S., Professor of Medicine and Botany.
William Brown, Esq., F.R.C.S.E.
1836 David Rhind, Esq., Architect.

Date of

LIST OF ORDINARY MEMBERS.

Election.

1837

1839

1840

1841

1842
1843

1844

1845
1846

1847

1848

1849

John Scott Russell, A.M., London.

Richard Parnell, M.D. ;

Peter D. Handyside, M.D., F.R.C.S.E.

David Smith, Esq., W.S.

Rev. Philip Kelland, A.M., F.R.S., Professor of Mathematics.
Francis Brown Douglas, Esq., Advocate.

Alan A. Maconochie Wellwood, Esq., of Meadowbank and Pitliver.
Martyn J. Roberts, Esq., Crickhowell, South Wales.

Sir John M‘Neil, G.C.B., LL.D.

Edward J. Jackson, Esq.

James Mackenzie, Esq.

John Miller, Esq., of Leithen.

James Dalmahoy, Esq.

James Thomson, Esq., Civil Engineer, London.

A. D. Maclagan, M.D., Professor of Medical Jurisprudence.
Sir John Rose Cormack, M.D., F.R.C.P.E., 7 Rue d’ Aguesseau, Paris.
Allen Thomson, M.D., F.R.S., Professor of Anatomy, Glasgow.
Joseph Mitchell, Esq., Civil Engineer, Viewhill, Inverness.
Andrew Coventry, Esq., Advocate.

John Hughes Bennett, M.D., Professor of Physiology.

D. Balfour, Esq., of Trenaby.

Henry Stephens, Esq.

Archibald Campbell Swinton, Esq., of Kimmerghame.

David Stevenson, Esq., Civil Engineer.

Thomas R. Colledge, M.D., F.R.C.P.E.

Thomas Anderson, M.D., Professor of Chemistry, Glasgow.

A. Taylor, M.D., Pau.

Alexander J. Adie, Esq., Civil Engineer.

L. D. B. Gordon, Esq., C.E., London.

L. Schmitz, LL.D., Ph.D., International Institution, London. ,
Charles Piazzi Smyth, Esq., F.R.S., Professor of Practical Astronomy.

799

Sir William Thomson, M.A. Camb., LL.D., F.R.S., Professor of Natural Philosophy, Glasgow.

John Hill Burton, LL.D., Advocate.

James Nicol, Esq., Professor of Natural History, Aberdeen.
William Macdonald Macdonald, Esq., of St Martins.

John Wilson, Esq., Professor of Agriculture.

Thomas Stevenson, Esq., C.E.

James Allan, M.D., Inspector of Hospitals, Portsmouth.

Henry Davidson, Esq.

William Swan, Esq., Professor of Natural Philosophy, St Andrews.
Patrick James Stirling, LL.D.

Sir William Stirling-Maxwell, Bart., of Keir and Pollok.

1849 William Thomas Thomson, Esq.

W. H. Lowe, M.D., F.R.C.P.E., Balgreen.

800

Date of

LIST OF ORDINARY MEMBERS.

Election.

1849

1850

1851

1852

1853

1854

1855

1856

1857

Honourable Bouverie F. Primrose.

David Anderson, Esq., of Moredun.

W. Rk. Pirrie, M.D., Professor of Surgery, Aberdeen.

His Grace the Duke of Argyll, K.T., Inverary Castle.

The Most Noble the Marquis of Tweeddale, K.T., Yester House.
Edward Sang, Esq.

James Stark, M.D., F.R.C.P.E. (Re-admitted.)

Lieutenant-Colonel W. Driscoll Gosset, R.E.

Hugh Blackburn, Esq., Professor of Mathematics, Glasgow.

James Scarth Combe, M.D., F.R.C.S.E.

Sir David Dundas, Bart., of Dunira.

E. W. Dallas, Esq.

Rev. James Grant, D.D., D.C.L., one of the Ministers of Edinburgh.
Eyre B. Powell, Esq., Madras.

Thomas Miller, A.M., LL.D., Rector, Perth Academy.

James Cunningham, Esq., W.S.

Alexander James Russell, Esq., C.S.

Andrew Fleming, M.D., Bengal.

James Watson, M.D., Bath.

Lieutenant-Colonel Robert Maclagan, Bengal Engineers.

Rev. John Cumming, D.D., London.

Hugh Scott, Esq., of Gala.

Greme Reid Mercer., Esq.
Robert Harkness, Esq., Professor of Mineralogy and Geology, Queen's College, Cork.
Sir James Coxe, M.D., F.R.C.P.E.

Ernest Bonar, Esq.

Stevenson Macadam, Ph.D.

Robert Etheridge, Esq.. Clifton, Bristol.

Right Honourable John Inglis, D.C.L., LL.D., Lord Justice-General.
Wyville T. C: Thomson, LL.D., Professor of Natural History.
Thomas Wright, M.D., Cheltenham.

James Hay, Esq.

R. M. Smith, Esq.

David Bryce, Esq.

William Mitchell Ellis. Esq.

George J. Allman, M.D., F.R.S., Emeritus Professor of Natural History.
Honourable Lord Neaves, LL.D.

Thomas Laycock, M.D., Professor of the Practice of Medicine.
Thomas Cleghorn, Esq., Advocate, Sheriff of Aryyleshire.

James Clerk Maxwell, Esq., F.R.S., Professor of Experimental Physics, Cambridge.
John Ivor Murray, M.D., F.R.C.S.E.

John Blackwood, Esq.

W. M. Buchanan, M.D.

Thomas Login, Esq., C.E.

LIST OF ORDINARY MEMBERS. 80L
Date of
Election.
1857 Edmund C. Batten, M.A., Lincoln’s Inn, London.
1858 Thomas Williamson, M.D., F.R.C.S.E., Leith.
Robert B. Malcolm, M.D., F.R.C.P.E.
Frederick Field, Esq., Chili.
James Leslie, Esq., C.E.
Cosmo Innes, Esq., Professor of History.
Alexander Campbell Fraser, M.A., Professor of Logic.
Rev. William Stevenson, D.D.
1859 William F. Skene, LL.D.
G. W. Hay, Esq., of Whiterigg.
Joseph Fayrer, M.D., F.R.C.S.E., Professor of Surgery, Calcutta.
George Robertson, Esq., C.E.
Lyon Playfair, C.B., Ph.D., F.R.S., M.P., 4 Queensberry Place, South Kensington, London, W-
John Brown, M.D., F.R.C.P.E.
Rey. John Duns, D.D.
Lieutenant John Hills, Bombay Engineers.
Major James George Forlong.
1860 William Robertson, M.D., F.R.C.P.E.
Frederick Guthrie, M.D., Professor of Physics, School of Mines, London.
George A, Jamieson, Esq.
Patrick Dudgeon, Esq., of Cargen.
William Chambers, Esq., of Glenormiston.
1861 Rey. Thomas Brown.
James M‘Bain, M.D., R.N.
Peter Guthrie Tait, A.M., Professor of Natural Philosophy.
John Muir, D.C.L., LL.D.
William Turner, M.B., Professor of Anatomy.
William Lauder Lindsay, M.D.
James Lorimer, A.M., Professor of Public Law.
Archibald Geikie, Esq., F.R.S., Director of the Geological Survey, Scotland.
George Berry, Esq.
James Young, Esq.
1862 Rev. William G. Blaikie, D.D.
Edmund Ronalds, Ph.D.
Thomas C. Archer, Esq., Director of Museum of Science and Art.
James Hector, M.D., Wellington, New Zealand.
Rev. Robert Boog Watson, Madeira.
1863 H. F. C. Cleghorn, M.D., Stravithy, St Andrews.
John Stuart Blackie, Esq., Professor of Greek.
Edward Meldrum, Esq.
1863 Charles Lawson, Esq., of Borthwick Hall.
Alexander Peddie, M.D., F.R.C.P.E.
William Jameson, Esq., Surgeon-Major, Saharunpore.
Murray Thomson, M.D., Roorkie, India.
VOL. XXVI. PART IV. 9Z

802 LIST OF ORDINARY MEMBERS.
Date of
Election.
1863 John Young, M.D., Professor of Natural History, University of Glasgow.
J. G. Wilson, M.D., F.R.C.S,E.
J. Matthews Duncan, M.D., F.R.C.P.E.
W. Dittmar, Esq.
Honourable Lord Ormidale.
Joseph D. Everett, D.C.L., Professor of Natural Philosophy, Queen’s College, Belfast.
Honourable G. Waldegrave Leslie, Leslie House.
Honourable Charles Baillie, Lord Jerviswoode.
James Sanderson, Esq., Surgeon-Major.
Charles Cowan, Esq.
John Alexander Smith, M.D., F.R.C.P.E.
1864 Alex. Crum Brown, M.D., D.Sc., Professor of Chemistry.
Alex. Wood, M.D., F.R.C.P.E.
Andrew Wood, M.D., F.R.C.S.E.
Robert William Thomson, Esq., C.E.
James David Marwick, Esq.
Rey. Daniel F. Sandford.
Robert S. Wyld, Esq., W.S.
Peter M‘Lagan, Esq., of Pumpherston, M.P.
William Lindsay, Esq.
W. Y. Sellar, M.A., Professor of Humanity.
Robert Hutchison, Esq., Carlowrie Castle.
William Wallace, Ph.D. .
John Foulerton, M.D., F.R.C.S.E., Manilla.
1865 Alfred R. Catton, M.A. Camb.
Rev. Francis Redford, M.A., Rector of Silloth.
John Moir, M.D., F.R.C.P.E.
James Powrie, Esq., of Reswallie, Forfar.
Charles Jenner, Esq.
Charles Lawson, jun., Esq.
1866 Alexander Keiller, M.D., F.R.C.P.E.
William Euing, Esq.
John M‘Culloch, Esq.
T. Grainger Stewart, M.D., F.R.C.P.E.
Major-General Sir James E. Alexander, of Westerton.
Charles Morehead, M.D. —
David Masson, M.A., Professor of Rhetoric and English Literature.
David Douglas, Esq.
John Macnair, Esq.
James Spence, Esq., F.R.C.S.E., Professor of Surgery.
Thomas Nelson, Esq.
Adam Black, Esq.
James Dunsmure, M.D., F.R.C.S.E. —
Arthur Mitchell, M.D.

Date of
Election

1866

1867

1868

LIST OF ORDINARY MEMBERS.

Patrick Heron Watson, M.D., F.R.C.S.E.
John Smith, M.D., F.R.C.P.E.

John Falshaw, Esq., C.E.

John K. Watson, Esq.

David Chalmers, Esq.

T. B. Johnston, Esq.

George F. Barbour, Esq., of Bonskeid.
David Davidson, Esq.

Peter Waddell, Esq.

Sir George Harvey.

George Stirling Home Drummond, Esq., of Blair-Drummond.
Frederick Fuller, Professor of Mathematics, Aberdeen.
Andrew Graham, M.D., R.N.

William Turnbull, Esq.

Archibald Hamilton Bryce, D.C.L., LL.D.
Francis Deas, LL.B., Advocate.

Arthur Gamgee, M.D..

Sheriff Hallard.

Thomas R. Fraser, M.D.

Thomas Annandale, Esq., F.R.C.S.E.

D. R. Haldane, M.D., F.R.C.P.E.

John M. M‘Candlish, Esq.

James Donaldson, LL.D., Rector of the High School.
James H. B. Hallen, Esq., India.

Henry Dircks, Esq., C.E., London.

Charles Gayner, M.D., Oxford.

William Keddie, Esq., Glasgow.

Rev. W. Lindsay Alexander, D.D.’

John F. M‘Lennan, Esq., Advocate.

Rev. David Aitken, D.D.

Robert M. Ferguson, Ph.D.

J. W. Laidlay, Esq., of Seacliff.

W. Williams, Esq., Veterinary College.

J. Samson Gamgee, Esq., Birmingham.
Rev. D. T. K. Drummond, B.A. Oxon.
Rev. Joseph Taylor Goodsir.

Major J. H. M. Shaw Stewart, Royal Engineers, Madras.
John J. Stevenson, Esq.

Very Rev. Dean Montgomery.

John Dick Peddie, Esq., Architect.
Colonel Seaton Guthrie.

Samuel Raleigh, Esq.

Thomas Smith Maccall, M.D., Polmont.
Rev. Thomas Guthrie, D.D.

803.

804 LIST OF ORDINARY MEMBERS.

Date of
Election.

1868 Thomas Key, Esq.
Adam Gillies Smith, Esq., C.A.
1869 Oliver G. Miller, Esq.
John Leveson Douglas Stewart, Esq., of Nateby Hall.
Alexander Buchan, Esq. :
H. C. Fleeming Jenkin, Esq., Professor of Engineering.
William Dickson, Esq.
John Pender, Esq., Manchester.
Isaac Anderson-Henry, Esq., of Woodend.
George Elder, Esq., Knock Castle, Wemyss Bay.
Sir Charles A. Hartley, C.E., Sulina, Mouth of the Danube.
David MacGibbon, Esq., Architect.
Rev. Thomas Melville Raven, M.A., Crakehall, Bedale.
Alexander Howe, Esq., W.S.
Viscount Walden, Yester House.
Alexander Dickson, M.D., Professor of Botany, University of Glasgow.
William C. M‘Intosh, M.D., Murthly.
Henry Marshall, M.D., Clifton, Bristol.
William Rutherford, M.D., Professor of Physiology, King’s College, London.
R. Craig Maclagan, M.D.
James Dewar, Esq.
Rev. Henry Calderwood, LL.D., Professor of Moral Philosophy.
Sir Alexander Grant, Bart., LL.D., Principal of the University of Edinburgh.
Captain T. P. White, Royal Engineer's.
John Wilson Johnston, M.D., India.
Robert Henry Bow, Esq., C.E.
Maurice Lothian, Esq., of St Catherine’s.
John M‘Laren, Esq., Advocate.
1870 St John Vincent Day, Esq., C.E., Glasgow.
David Munn, Esq.
Robert R. Tatlock, Esq., F.C.S., Glasgow.
Alexander Russel, Esq.
James Crichton Browne, M.D., West Riding Asylum, Wakefield.
John Duncan, M.D.
William Burns Thomson, Esq., F.R.C.S.
William R. Sanders, M.D., Professor of General Pathology.
Rev. Andrew Thomson, D.D.
Joseph Lister, Esq., F.R.S., Professor of Clinical Surgery.
G. H. B. Macleod, M.D., Regius Professor of Surgery, University, Glasgow.
1870 Thomas A. G. Balfour, M.D., F.R.C.P.E.:
W. E. Heathfield, Esq., F.R.G.S., F.C.S., London.
Edward James Shearman, M.D., F.R.C.P., F.R.C.S. Ene.
Patrick D. Swan, Esq., Kirkcaldy.
H. Alleyne Nicholson, M.D., D.Sc., M.A., Toronto.

LIST OF ORDINARY MEMBERS. 805

Date of
Election.

1870 Rev. James S. Hodson, D.D., St Andrew’s College, Broadfield, Reading. (Re-admitted
James Warburton Begbie, M.D.
John Winzer, Esq., Assistant Surveyor, Ceylon Civil Service, Galle, Ceylon
Spencer C. Thomson, Esq., B.A., Actuary.
Simon S. Laurie, Esq.
James Sime, Esq.
Thomas Harvey, LL.D., M.A., Rector, Edinburgh Academy.
John Young Buchanan, Esq., M.A.
John Hunter, Esq., M.A., F.C.S.
Sir James Moncreiff, Lord Justice-Clerk.
Hon. Lord Gifford.
James Watson, Esq.
Hon. Lord Mackenzie.

1871 John Auld, Esq., W.S.
Rev. Thomas M. Lindsay, Glasgow.
Rev. William Robertson Smith, M.A., Aberdeen.
Stair Agnew, Esq.
Charles Hayes Higgins, M.D., M.R.C.P., F.R.C.S., Birkenhead, Cheshire.
Angus Macdonald, M.D.
Rev. William Scott Moncrieff, of Fossaway.
Alexander R. Simpson, Professor of Midwifery.
R. J. Blair Cunynghame, M.D.
Cosmo Gordon Logie, M.D., Surgeon-Major, Royal Horse Guards.
James Geikie, Esq., Surveyor of Geological Survey of Scotland.
Thomas Ed. Thorpe, Ph.D., Lecturer on Chemistry, Anderson’s Institution, Glasgow.
John Smith, M.D., F.R.C.S.E.
Rev. Thomas J. Crawford, Professor of Divinity.
Thomas J. Boyd, Esq.

1872 Alexander H. Lee, Esq., Civil Engineer.
Robert Lee, Esq., Advocate.
John Anderson, LL.D., Victoria ‘Road, Charlton, Kent.
David Maclagan, Esq., C.A., F.S.S.
Major F. Ignacio Rickard, F.G.S., Inspector-General of Mines, Buenos Ayres.
John Sibbald, M.D.
J. G. Fleming, M.D., Glasgow.
Rev. Andrew Tait, LL.D., Rector of Kilkerrin, Ireland.
David Grieve, Esq.
Right Rev. Bishop Cotterill.
George Barclay, Esq.
George Forbes, Esq., B.A., Lecturer on Natural Philosophy, Anderson’s Institution, Glasacw,
J. Lindsay Stewart, M.D., F.R.C.S., F.LS., F.B.G.S., Punjaub, India.
Rev. Charles R. Teape, M.A., Ph.D.
Archibald Constable, Esq.
George Seton, Esq., M.A. Oxon., Advocate.

VOL. XXVI. PART IV. 10 4

806 LIST OF ORDINARY MEMBERS.

Date of
- Election.

1872 Captain Charles Hunter, London.
James Thomson Bottomley, Esq., M.A., Lecturer on Natural Philosophy, University, Glasgow.
Thomas Knox, Esq.
D. Argyll Robertson, M.D.
Thomas B..Christie, M.D., M.R.C.P. Lond., F.R.C.P. Edin. , Ealing, London.
Rev. Hugh Macmillan, LL.D., Glasgow.

(. 807...)

NON-RESIDENT MEMBER,

ELECTED UNDER THE OLD LAWS.

Sir Richard Griffiths, Bart., Dublin.

LESTOF HONORARY KELL Ow Ss.

His Royal Highness the Prince of Wales.

‘ FOREIGNERS (LIMITED TO THIRTY-SIX BY LAW X.)

Louis Agassiz, Cambridge, Massachusetts.
J. B. A. L. Léonce Elie de Beaumont, Paris.

Claude Bernard, Do.
Robert Wilhelm Bunsen, Heidelberg.
5. Michael Eugene Chevreul, Paris.

James D. Dana, LL.D.,

Newhaven, Connecticut.

Jean Baptiste Dumas, Paris.
Charles Dupin, Do.
Christian Gottfried Ehrenberg, Berlin.

10 Elias Fries, Upsala.
Francois Pierre Guillaume Guizot, Paris,
Christopher Hansteen, Christiania.
Herman Helmholtz, Heidelberg.
Gustav Robert Kirchhoff, Do.

15 Albert Kolliker, Wurzburg.
Johann von Lamont, Munich.
Richard Lepsius, Berlin.
Rudolph Leuckart, Leipzig.
Urbain Jean Joseph Leverrier, Paris.

20 Baron Justus von Liebig, Munich.
Henry Milne-Edwards, Poris.
Theodore Mommsen, Berlin.

Prof. Benjamin Peirce,

Adolphe Pictet,
Lambert Adolphe Jacques Quetelet

United States Survey.
Geneva.
Brussels.

808 LIST OF HONORARY FELLOWS.

_ M. Le Comte De Remusat, ) a Baris:
Henry Victor Regnault, Do.
Auguste De la Rive, Geneva.
Gustav Rose, Berlin.

30 Angelo Secchi, Rome.
Karl Theodor von Siebold, Munich. °
Bernard Studer, Berne.
Rudolph Virchow, Berlin.

34 Friedrich Wohler Gottingen.

BRITISH SUBJECTS (LIMITED TO TWENTY BY LAW x.)

John Couch Adams, Esq.,

Sir George Biddell Airy,

Thomas Andrews, M.D.,

Thomas Carlyle, Esq.,

Arthur Cayley, Esq.,

Charles Darwin, Esq.,

James Prescott Joule, LL.D.,

William Lassell, Esq.,

Rev. Dr Humphrey Lloyd,

10 Sir William E. Logan,
Sir Charles Lyell, Bart.,
John Stuart Mill, Esq.,
Richard Owen, Esq.,
Lieut.-General Edward Sabine, R.A.,

15 George Gabriel Stokes, Esq.,
William Henry Fox Talbot, Esq.,
Alfred Tennyson, Esq.,

18 Sir Charles Wheatstone, D.C.L.,

Cambridge.
Greenwich.
Belfast ( Queen's College).
London.
Cambridge.
Down, Bromley, Kent.
Clifpoint, Higher Broughton, Manchester.
Liverpool.
Dublin.
London.

Do.

Do.
Cambridge.
Lacock Abbey , Wiltshire.
Freshwater, Isle of Wight.
London.

( 809 )

LIST OF FELLOWS DECEASED AND RESIGNED,

From NoveMBER 1869 To NovEMBER 1872.

HONORARY FELLOWS DECEASED (FOREIGN).

Wilhelm Carl Haidinger, Vienna.
Hugo von Mohl, Tubingen.

HONORARY FELLOWS DECEASED (BRITISH).
Sir John Frederick William Herschel, Bart.
Sir Roderick Impey Murchison, K.C.B.

ORDINARY FELLOWS DECEASED.

William Anderson, LL.D.
Charles Babbage, Esq.
Honourable Lord Barcaple.
Thomas Barnes, M.D.
Robert Chambers, LL.D.
Robert Daun, M.D.

Adam Hunter, M.D.
Alexander Keith Johnston, LL.D.
Patrick Miller, M.D.
Sheridan Muspratt, M.D.
Robert Nasmyth, Ksq.

Sir William Scott, Bart.

Sir James Simpson, Bart., Professor of Midwifery.

Moses Steven, Esq., of Bellahouston.

James Syme, Esq., Professor of Clinical Surgery.

John Addington Symonds, M.D.
Robert Russell, Esq.

Right Rev. Bishop Terrot.
Fraser Thomson, M.D.

ORDINARY FELLOWS RESIGNED.

W. A. F. Browne, Esq.

Nicholas Alexander Dalzell, Esq.
Rev. John Hannah, D.D.

John Macmillan, Esq.

David Page, LL.D.

Arthur Abney Walker, Esq.

VOL. XXVJ. PART IV.

10 B

( 810 )

The following Public Institutions and Indwiduals are entitled to receive Copies of
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Royal Asiatic Society.

Zoological Society.

Royal Society of Literature.

Royal Horticultural Society.

Royal Institution.

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Statistical Society.

Institution of Civil Engineers.

Institute of British Architects,
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Medico-Chirurgical Society.

Athenzum Club.

Cambridge Philosophical Society.
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(eco Ws aah)

Munich, Royal Academy of Sciences of Bavaria
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Paris, Royal Academy of Sciences.
Geographical Society.
Royal Society of Agriculture.
Society for encouragement of Industry.
Geological Society of France.
Ecole des Mines.
Marine Depét.
.... Museum of Jardin des Plantes.
Rotterdam, Batavian Society of Experimental
Philosophy.
St Petersburg, Imperial Academy of Sciences.
Archeological Society.
Pulkowa Observatory.
Stockholm, Royal Academy of Sciences.
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M. Michelotti.
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Vienna, Imperial Academy of Sciences.
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Academy of Arts and Sciences.

Society of Natural History.
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Academy of Natural Sciences.

United States Naval Observatory.
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Yale College, United States.

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Proceedings.)

The following Institutions and Individuals receive the Proceedings only :-—

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Whitby Philosophical Society.
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Ashmolean Society of Oxford.
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Meteorological Office, 116 Victoria Street, London.
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IRELAND.
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Paris, Editor of L’Institut.

Abbé Moigno, Paris.

Em. Alglave, Directeur de la Revue des Cours
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Cherbourg, Society of Natural Sciences.

Belgium, the University of Ghent.

Sicily, Catania, Academia Gionia de Scienze
Natural.

UNITED STATES.

Peabody Academy of Science, Salem, Massachu-
setts.

( 813 )

LIST OF DONATIONS.

(Continued from Vol. XXV., p. 780.)

DONATIONS.
TRANSACTIONS AND PROCEEDINGS OF SociETIns, ACADEMIES, &c.—
Amsterdam.—Jaarboek van der Koninklijke Akademie van Wettenschappen
gevestigd te Amsterdam, 1868-1870. 8vo.
Processen-verbaal van de Gewone vergadering der Koninklijke Akademie
van Wettenschappen, 1869-1871. 8vo.
Verhandelingen der Koninklijke Akademie van Wettenschappen. __Let-
terkunde, Deel iv., v., vi.; Natuurkunde, Deel xii.
Verslagen en Mededeelingen der Koninklijke Akademie van Wetten-
schappen. Natuurkunde, Deel i., iv., v.; Letterkunde, Deel
ail. SO: .
Flora Batava, Nos. 211-217. 4to.

Augusta (U. S.).—Third Report of the Commissioner of Fisheries of the State
of Maine, 1869. 8vo.

Baltimore.—Address of the President to the Board of Trustees of the Peabody
Institute, 1870. 8vo.

Proceedings of the Board of Trustees of the Peabody Institute, Nov.
1870. 8vo.

Third, Fourth, and Fifth Annual Reports of the Provost of the Peabody
Institute to the Board of Trustees. 8vo.

Basle.—V erhandlungen der Naturforschenden Gesellschaft in Basel. Fiinfter
Theil, Zweites Heft. 8vo.

Batavia.—Observations made at the Magnetical and Meteorological Obser-
vatory at Batavia. Vol. i. Fol.

Berlin.—Abhandlungen der Koniglichen Akademie der Wissenschaften,
1868-1870. 4to.

Die Fortschritte der Physik im Jahre 1866, 1867, dargestellt von der
Physikalischen Gesellschaft zu Berlin. Jahrgang xxii., xxiii.
8vo.

Monatsbericht der Koniglich Preussischen Akademie der Wissenschaften,

1869-1871, Jan.—April 1872. 8vo.

Verzeichniss der Abhandlungen der Koniglich Preussischen Akademie

der Wissenschaften von 1710-1870. 8vo.

Berne.—Beitrege sur Geologischen Karte der Schweiz herausgegeben von der
Geologischen Commission der Schweiz. Naturforsch. Gesellschaft
auf Kosten der Eidgenossenschaft, 1872. 4to.

Mittheilungen der Naturforschenden Gesellschaft in Bern, aus dem
Jahre 1868-1870. 8vo.

Materiaux pour la Carte Geologique de la Suisse. Liv. 7,8. 4to.

Birmingham.—Reports of the Free Libraries’ Committee, Birmingham, for
1869, 1870. 8vo.

VOL. XXVI. PART IV.

DONORS.
The Academy.
Ditto.
Ditto.
Ditto.
The King of Hol-
land.
The Commissioner.
The Institute.
Ditto.
Ditto.
The Society.
The Government.
The Academy.
The Society.

The Academy.
The Society.

The Commission.

The Society.
The Natural His-

tory Society.
The Committee.

10 ¢c

814 LIST OF DONATIONS.

DONATIONS.
TRANSACTIONS AND PROCEEDINGS oF Sociptres, &c.—cuntinued.
Bologna.—Archivio per la Zoologia, Anatomia, e la Fisiologia.
Vol. i., Vol. 11.; Fase. 1. 8vo.

Memorie dell Accademia delle Scienze dell Instituto di Bologna.
Serie i. Tomo v. Fasc. 3, 4.; Tomo vi., vil., vili., ix., x. Serie ii.
Tomo i., 1. Fase. 1. 4to.

Rendiconto delle Sessioni dell Accademia delle Scienze dell Instituto di
Bologna. Ann. Accademic. 1865-66, 1866-67, 1867-68, 1868-69,
1870-71, 1871-72. 8yo.

Bordeauz.—Mémoires de la Société des Sciences Physiques et Naturelles de
Bordeaux. Tome v. No.4; Tome vi. No. 3.; Tome vii.; Tome
Vil, IN@; Ne 2a thio.

Boston.—Annual Report of the Trustees of the Museum of Comparative
Zoology, 1868. 8vo.

Bulletin of the Public Library. Nos. 10-20. 8vo.

Memoirs of the Society of Natural History. Vol. i. Part 4. 4to.

Occasional Papers of the Society of Natural History. No. 1, 1869. 8vo.

Proceedings of the Society of Natural History. Vol. xii., xiii.

Brussels.—Annales de Observatoire Royale de Bruxelles publiés aux frais
de l’Etat, par le directeur A. Quetelet. Tome xix., xx. 4to.

Annuaire de l’Académie Royale des Sciences, des Lettres et des Beaux-
Arts de Belgique, 1870, 1871. 12mo.

Annuaire de l’Observatoire Royal de Bruxelles, par A. Quetelet, 1870,
1871. 12mo.

Biographie Nationale publice par l’Académie Royale des Sciences, des
Lettres et des Beaux-Arts de Belgique. Tome iii. Part 1. 8vo.

Bulletin de Académie Royale des Sciences des Lettres et des Beaux-
Arts de Belgique. Tome xxvii., xxvili., Xxix., XXX., XXxi., xxxii,
XXXlli., xxxiv. 8vo.

Mémoires de l’Académie Royale des Sciences, des Lettres et des Beaux-
Arts de Belgique. Tome xxxvili. 4to.

Mémoires couronnés et Mémoires des Savants Etrangers publices par
VYAcadémie Royale des Sciences, des Lettres et des Beaux-Arts de
Belgique. Tome xxxv., xxxvi. 4to.

Mémoires couronnés et autres Mémoires, publics par Académie Royale
des Sciences des Lettres et des Beaux-Arts de Belgique. xxi. 8vo.

Observations des Phénoménes Périodiques pendant les Années 1867 et

Serie 11.

1868. to.
Calcutta.—Account of the Operations of the great Trigonometrical Survey of
India. Vol. i. 4to.

Annual Report of the Geological Survey of India, and of the Museum
of Geology for 1867. 8vo.
Report of the Commissioners appointed to inquire into the Origin,
Nature, &c., of Indian Cattle Plagues, with Appendices. 1871. Fol.
Journal of the Asiatic Society of Bengal. Parts i. and ii., 1869. Parts
i, and i. Nos.‘1, 2, 3; 1870! Part 1. Nos. 1-3. Part 1, Nos:
1-4, 1871. Parti. No. 1; Part ii. No. 1, 1872. 8vo.
Memoirs of the Geological Survey of India, Palzontologia.
Nos. 1-13; Vol. v. Parts 5-10; Vols. vi., vii. 4to.
Memoirs of the Geological Survey of India. Vol. vi. Part 3. Vol. vii.
Parts 1-3. 8vo.
Records of the Geological Survey of India. Vol. i. Parts 1-3; Vol.
ii, Parts 1-4; Vol. iii., Vol iv. Parts 1-4. 8vo.
Proceedings of the Asiatic Society of Bengal. Nos. 2-11, 1869;
No. 11, 1870; Nos. 1-13, 1871 ; No. 1-5, 1872. 8vo.
California.— Memoirs of the Academy of Sciences. Vol. i. Parts 1,2. Ato.
‘““ Proceedings of the Academy of Sciences. Vol. iv. Parts 1-4. 8vo.
Cambridge.—Proceedings of the Philosophical Society. Parts 3-6. 8vo.
Transactions of the Philosophical Society. Vol. xi. Part 2. 4to.
Cambridge (U.S.).—Addresses at the Inauguration of Charles William Eliot
as President of Harvard College, 1869. 8vo.

Vol. i.

DONORS.
The Editors

The Academy.
Ditto.
The Society.

The Trustees.
The Library.
The Society.
Ditto.
Ditto.
The Observatory.
The Academy.
The Observatory.
The Academy.

Ditto.

Ditto.
Ditto.
The Survey.
Ditto.
The Indian
Government.
The Society.
The Survey.
Ditto.
Ditto.
The Society.
The Academy.
Ditto.
The Society.

Ditto.
The College.

LIST OF DONATIONS.

DONATIONS.

TRANSACTIONS AND PROCEEDINGS OF Socintips, &¢.—continied.

Cambridge (U.S.).—Annual Report of the Librarian of Harvard University,

1863, 1864, and 1869. 8vo.

Annual Reports of the President and Treasurer of Harvard College,
1868, 1869. 8vo.

Annual Report of the Trustees of the Museum of Comparative Zoology
at Harvard College for 1870, 1871. 8vo.

Catalogue of the Collection of Engravings bequeathed to Harvard Col-
lege by Francis Calley Grey. By Louis Thies. 4to.

New Catalogue of Harvard College Library. 8vo.

Catalogue of Officers and Students of Harvard University for 1869,
1870. 8vo.

Catalogus Senatus Academici Collegii Harvardiani, 1869. 8vo.

Illustrated Catalogue of the Museum of Comparative Zoology at Harvard
College. Nos. 3-6. 8vo.

Bulletin of the Museum of Comparative Zoology at Harvard College,
Cambridge, Mass. Vol. ii. Nos. 1-3; Vol. ui. No. 1. 8vo.

The Complete Works of Count Rumferd, published by the American
Academy of Arts and Sciences. Vol. i. 1870. 8vo.

Memoirs of the American Academy of Arts and Sciences. Vol. ii.
Part 2; Vol. iv. Part 1; Vol. x. Part 1. Ato.

Proceedings of the American Academy of Arts and Sciences. Vols. u.—
vill. 8vo.

Proceedings of the American Association for the Advancement of
Science. 1867-1870. 8vo.

Canada.—Report of Progress of Geological Survey of, for 1866-1869. 8vo.
Cape of Good Hope.—Results of Astronomical Observations made at the
Royal Observatory, Cape of Good Hope, in the year 1856. 8vo.
Catania.—Atti dell Accademia Gioenia dé Scienze Naturali de Catania.
Serie Terza. Tomo ii., 1868; Tomo ii., 1869. 4to.
Cherbourg.—Catalogue de la Bibliothéque de la Société Impériale des Sci-
ences Naturelles. Parti. 8vo.

Mémoires de la Société Impeériale des Sciences Naturelles.
Kiva, KV., XVI. OVO.

Christiania.— Annexe a la Statistique Officielle du Royaume de Norvége
pour l’année 1869. Ato.

Beretning Rigets Oeconomiske Tilstand, Aarene 1861-1865.
Hefte. to.

Beretning om Skolevesenets Tilstand i Kongeriget Norges Landdistrikt
for Aarene 1864-66, og Rigets Kjbstader og Ladesteder for Aaret
1867. Ato.

Beretninger om Norges Fiskerier, i Aaret 1868, 1869. Ato.

Beretning den Hoiere Landbrugsskole i Aas, i Aarene fra April 1867
til April 1870. Ato.

Criminalstatistiske Tabeller for Kongeriget Norge for Aaret 1866,
samt den Kongelige Norske Regjerings Underdanigste Indstilling
af 3 Juni 1870. 4to.

Det Kongelige Norste Frederiks-Universitets Aarsberetning for 1868—
1870. 8vo.

Den Norske Statstelegrafs Statistik for 1869.

Tome xii.,

Andet

Ato.

Det Norske Meteorologiske Instituts Storm Atlas udgivet med Bestand
af Videnskabs-Selskabet i Christiania. ol.
Driftsberetning for Kongsvinger-Lillestrom Jernbane, i Aaret 1869. 4to.

Driftsberetning for Hamar-Elverum-Jernbane, i Aaret 1869. Ato.
Driftsberetning for Norsk Hovid-Jernbane, i Aaret 1869. 4to.
Fattig-Statistik for 1867. 4to.

Flateyjarbok en Samling af Norske Kongl. Sagaer, &c. 1868. 8vo.

Forhandlinger ved de Skandinaviske Naturforskeres, Tiende mode, fra
den 4%, til den 10° Juli 1868. 8vo. .

Forhandlinger i Videnskabs-Selskabet. » Aar 1868-1870. 8vo.

815

DONORS.
The University.
The College.
Ditto.
Ditto.

Ditto.
The University.

The College.
Ditto.

Ditto.

The Academy.
Ditto.
Ditto.

The Association.

The Survey.
The Observatory.

The Academy.

The Society.
Ditto.

The Government
of Norway.
Ditto.

Ditto.
Ditto.
Ditto.

Ditto.

The University.

The Government of
Norway.
The Institute.

The Government of
Norway.
Ditto.
Ditto.
The Society.
Ditto.

Ditto.

LiST OF DONATIONS.

DONATIONS.

TRANSACTIONS AND PROCEEDINGS OF SOCIETIES, &c.—continued.

Ato.
4to.

Christiania.—Le Névé de Justede et ses Glaciers par le de Sene.
Norsk Meteorologisk Aarbog for 1868-1870. Aargang. II.
‘Norske Universitets-og-Skole, Annaler udgivne af Universitets Secre-

tair, Mai 1869. 8vo.

Nyt Magazin for Naturvidenskaberne. Bind. xvi., xvii, xviii. . 1869.
8vo.

Tabeller vedkommende Norges Handel og Skibsfait, i Aaret 1869.
4to.

Tabeller vedkommende Skiftevoesenet i Norge, Aaret 1868. Tilligemed
opgave ov e de efter Overformynder-Regnskaberne for Aaret 1868—
1869, under rigets Overformynderiers Bestyrelse Henstaaende Midler
saint den Kongelige Norske Regjerings Underdanigste Indstilling
af 15 Juli 1870, 12 Sept. 1871. 4to.

Cincinnati.—Annual ‘Address, delivered in 1845, before the Astronomical
Society, by E. D. Mansfield, Esq. 8vo.

Annual Report of the Director of the Observatory. 1869, 1870. 8vo.

An Oration delivered before the Astronomical Society, by J. Quincy
Adams. 8vo.

Connecticut.—Transactions of the Connecticut Academy of Arts and Sci-
ences. Vol. i. Part 2; Vol. ii. Part 1. 8vo.
Copenhagen.—Det Kongelige danske Videnskabernes Selskabs,

femte Rekke. 1869-70. 4to.

Oversigt over det Kongelige danske Videnskabernes Selskabs Forhand-
linger og dets Medlemmers Arbeider i Aaret, 1867-1870, 1871,
Nos. 1, 2. Kjobenhavn. 8vo.

Dorpat.—Meteorologische Beobachtungen 1866-1871. 8vo.

Dresden.—Nova Acta Academie Czsarez Leopoldino-Caroline Germanic
Nature Curiosorum. Vol. xxxv. 4to.

Dublin.—Astronomical Observations and Researches made at Dunsink.
Part i., 1870. 4to.

Journal of the Royal Dublin Society. No. 39. 8vo.

Journal of the Royal Geological Society of Ireland. Vol. i. Parts 1,
2. 8vo.

Observations made at the Magnetical and Meteorological Observatory
at Trinity College. Vol. ii, 1844-1850. Dublin, 1869. 4to.

Proceedings of the Royal Irish Academy. Vol. x. Parts 1-3. 8vo.

Transactions of the Royal Irish Academy. Vol. xxiv. ; Science, Parts
9-15 ; Polite Literature, Part 4; Antiquities, Part 8. 4to.

Tables of Tris, computed with regard to the Perturbations of Jupiter,
Mars, and Saturn, including the perturbations depending on the
square of the mass of Jupiter. By Francis Briinnow, Ph.D.,
F.R.AS. 4to.

Edinburgh.—Astronomical Observations made at the Royal Observatory,
Edinburgh, by Charles Piazzi Smyth, F.R.SS.L. and E., F.R.A.S,
F.R.S.8.A., Professor of Practical Astronomy, and Astronomer
Royal for Scotland. Vol. xiii., for 1860-1869, with additions to
1871. Ato.

Report presented to, and read before, the Board of Visitors, Aime
by Government for the Royal Observatory, at their Visitation held
on Thursday, 27th July 1871. to.

Scottish Meteorology, 1856-1871, computed at the Royal Observatory.
Ato.

Forty-Second and Forty-Third Annual Reports of the Council of the
Royal Scottish Academy of Painting. 8vo.

Thirteenth and Fourteenth Detailed Annual Reports of the Registrar-
General of Births, Deaths, and Marriages in Scotland. 8vo.

Quarterly Return of the Births, Deaths, and Marriages Registered in
the Divisions, Counties, and Districts of Scotland. Nos. 58 to 65.
Monthly Returns of the same, for 1869-1872. 8vo.

Skrifter,

DONORS. .
The University.
The Meteorological
Institute.
The University.
The Government
of Norway.
Ditto.

Ditto.

The Society.

The Observatory.
The Society.

The Academy.
The Royal Academy

of Sciences.
Ditto.

Univ. of Dorpat.

The Academy.

The Board of Trinity
College.

The Society.

The College.

The Academy.
Ditto.

Royal Astronomical
Society.

Royal Observatory.

Ditto.

Ditto.
The Academy.
Registrar-General.

Ditto.

LIST OF DONATIONS.

DONATIONS.

TRANSACTIONS AND PROCEEDINGS or Socterins, &c.—continued.

Edinburgh.—Journal of the Scottish Meteorological Society. Nos. 21-35.

8vo.
Eighth Decennial Census of the Population of Scotland, taken 3d April
LS Als WViolaie Hol:

Supplement to Catalogue of the Library of the Royal College of Physi-
cians, 1863-1870. 4to.

Transactions and Proceedings of the Botanical Society. Vol. x., Vol. xi.
Part 1. 8vo.

Transactions of the Geological Society. Vol. i. Part 3. 8vo.

Transactions of the Highland and Agricultural Society of Scotland.

Vol. iv. 8vo.
Transactions of the Royal Scottish Society of Arts. Vol. vii. Parts 1, 2.
8vo.
Erlangen.—Sitzungsberichte der Physicalisch-Medicinischen Societét zu
Erlangen. Heft 3. 8vo.

Frankfort.—Abhandlungen herausgegeben von der Senckenbergischen Na-
turforschenden Gesellschaft. Band vu. Heft 1-4; Band vii.
Heft 1, 2. Ato.
Bericht iiber die Senckenbergische Naturforschende Gesellschaft, 1869-
1871. 8vo.
Geneva.—Mémoires de la Société de Physique et d’Histoire Naturelle de
Genéve. Tome xx. Partie 1,2; Tome xxi. Part 1. 4to. ‘Table
des Mémoires, Tomes i.—xx.
Glasgow.—Proceedings of the Philosophical Society.
Vol. viii. No. 1. 8vo.
Transactions of the Geological Society. Vol. ii., and Supplement. 8vo.
Gottingen.—Abhandlungen der Koniglichen Gesellschaft der Wissenschaften
Band xiv., xv., xvi. Ato.
Astronomische Mittheilungen von der Konigl. Sternwarte zu Gottingen.
Erster Theil. 4to.
Nachrichten von der K. Gesellschaft der Wissenschaften und der
Georg-Augusts-Universitit, aus dem Jahre 1869-1871. 12mo.
Greenwich.—Astronomical and Magnetical and Meteorological Observations
made at the Royal Observatory in the year 1867, 1868, 1870.
London, 1869. 4to.
Haarlem.—Archives du Musée Teyler. Vol. ii., Vol. iii. Fasc. 1, 2. 8vo.
Archives Néerlandaises des Sciences Exactes et Naturelles publiées par
la Scciété Hollandaise 4 Haarlem. Tome iii, Tome iv., Tome v.,
Tome vi., Tome vii. Liv. 1-3. 8vo.
Halifax, Nova Scotia.—Proceedings and Transactions of the Nova Scotian
Institute of Natural Science. Vol. 11. Part 2. 8vo.
Helsingfors.—Bidrag till Finlands Officiela Statistik v. Temperaturforhallan-
den i Finland 1846-1865. Heft. 1. to.
Bidrag till Kannedom af Finlands Natur och Folk utgifna af Finska
Vetenskaps-Societeten Sjuttonde Hiaftet. 8vo.
Acta Societatis Scientiarum Fennice. Tomusix. 4to.
Ofversight af Finska Vetenskaps-Societetens Forhandlingar.
8vo.
Innsbruck.—Berichte des Naturwissenschaftlich-Medizinischen Vereines in
Innsbruck, Jahrgang i. Heft 1, 2; ii. 1-3. 8vo.
Jena.—Jenaische Zeitschrift fiir Medicin und Naturwissenschaft herausgege-
ben von der Medicinisch Naturwissenschaftlichen Gesellschaft zu
Jena. Bands i., ii., i., iv. Heft 3; v. Heft 1-4; vi. Heft 1-4.
8vo.
Jerusclem.—Ordnance Survey of 1865.

Vol. vu. No. 3;

1870-71.

Maps. Fol.

Kasan.—Reports of the University of Kasan, 1864-1869. 8vo.

Kiel.—Schriften der Universitat zu Kiel, aus dem Jahre 1868. Band xv.,
1869 ; Band xvi, 1870; Band xvii., 1871 ; Band xviii. Ato.

Lausanne.—Bulletin de la Société Vaudoise des Sciences Naturelles. Vol. x.
No. 62. 8vo.

VOL. XXVI. PART IV.

817

DONORS.
The Society.
Registrar-General.
The College.
The Society.

The Society.
The Society.

The Society.
The Society.
The Society.

Ditto.

The Society.

The Society.
The Society.

Ditto.
Ditto.
The Observatory.
The Museum.
The Society.
The Society.
The Society of
Science.
The Society.

Ditto.
Ditto.

The Society.
The Society.

The Secretary of
State for War.

The University.

The Society.
10D

818 LIST OF DONATIONS.

DONATIONS. DONORS.
TRANSACTIONS AND PROCEEDINGS OF SocintTins, &c.—continued.
Lausanne.—Feuille Centrale de la Société de Zofingue. Huititme Année The Society.
No. 8. 8vo.
Leeds.—Reports of the Proceedings of the Geological and Polytechnic Society The Society.
of the West Riding of Yorkshire, 1869, 1870. 8vo.
Reports of the Philosophical and Literary Society, 1868-1871. 8vo. The Society.
Leewwarden.—Nederlandsch Kruidkundig Archief, Vijfde deel. Viorde The Editors.
Stuk. 1870. 8vo.
Leipsig.—Berichte tiber die Verhandlungen der Koniglich Siachsischen The Royal
Gesellschaft der Wissenschaften zu Leipzig. Math. Phys. Classe. Academy.
1867, Nos. 3,4; 1868, Nos. 1-3; 1869, Nos. 1-4; 1870, Nos.
1-4; 1871, Nos. 1-3.—Phil. Hist. Classe. 1868, Nos. 2, 3;
1869, Nos. 1-3. | 8vo.
Bestimmung der Sonnenparallaxe durch Venusvoriiberginge vor der Ditto.
Sonnenscheibe mit Besonderer Beriicksichtigung des im Jahre 1874
eintreffenden voruberganges von P. A. Hansen. Band ix., No. 5.
8vo.
Elektrische Untersuchungen ueber die Thermo-elektrichen Eigenschaften Ditto.
des Topases. Band viii., ix. No. 4. W.G. Hankel. 8vo. ‘
Elektrodynamische Maassbestimmungen Insbesendere iiber das Princip Ditto.
der Erhaltung der Energie, von Wilhelm Weber. Band x. No. 1.
8vo.
Erophile Vulgaergriechische Tragoedie von Georgios Chortatzes aus Kreta. Ditto.
Ein Beitrag zur Geschichte der Neugriechischen und der Italianischen
Litteratur von Conrad Bursian. Band v. No. 7. 8vo.
Entwickelung eimes neuen veranderten Verfahrens zur Ausgleichung Ditto.
eines Dreiecksnetzes mit besonderer Betrachtung des Falles in
welchem Gewisse Winkel voraus bestimmte Werthe bekommen
sollen, von P. A. Hansen. No. 2. 8vo.
Fortgesetzte geoditsche Untersuchungen bestehend in zehn Supplementen Ditto.
zur Abhandlung von der Methode der kleinsten Quadrate im All-
gemeinen und in ihrer Anwendung aaf die Geodisie. Von P. A.
Hansen. 8vo. ,
Supplement zu der Geodiitische Untersuchungen benannten Abhandlung Ditto.
die Reduction der Winkel eines Sephiroidischen Dreiecks betreffend
von P. A. Hansen. 8vo.
Preisschriften gekront und herausgegeben von der fiirstlich Jablonowskis- Ditto.
chen Gesellschaft zu Leipzig. xiv., xv., xvi. 8vo.

Saxon

Tafeln der Amphitrite mit Beriicksichtigung der Storungen durch The Astronomical

Jupiter, Saturn, and Mars, entworfen von Dr E. Becker. 4to. - Society.
Tafeln der Pomona mit Beriicksichtigung der Stérungen durch Jupiter, Ditto.
Saturn, und Mars berechnet von D. Otto Lesser. No.9. Ato.

XV. Tafeln zu H. Engelhardt Flora der Braunkohlenformation in The Royal Saxon
Konigreich Sachsen. Preisschriften der Furstl Jablonowskischen Academy.
Gesellschaft xvi. 8vo.

Untersuchung des Weges eines Lichtstrahls durch eine beliebige Anzahl Ditto.
von brechenden sphirischen Oberflachen. P. A. Hansen. 8vo.

Vierteljahrsschrift der Astronomischen Gesellschaft; Jahrgang iv. The Society.

Heft 2-4; Jahrgang v. Heft 1-4; Jahrgang vi. Heft 1-4. ; vii.
Heft 1. 8vo.

Zur Experimentalen Aesthetik, Von Gustay Theodor Fechner. Band The Royal Saxon

ix. No. 6. 8vo. Academy.
Leyden.—Annalen der Sternwarte. Zweiter Band. 1870. 4to. The Observatory.

Lisbon.—Catalogo das Publicacoes da Academia Real das Sciencias de The Academy.

Lisboa. 8vo.
Memorias da Academia Real das Sciencias de Lisboa, Classe de Ditto.
Sciencias Mathematicas, Physicas e Naturaes, Nova Serie. Tomo
TV.) Parte 1,2) 4a;
Liverpool.—Proceedings of the Literary and Philosophical Society of Liver- The Society.
pool. Nos. 23,24. 8vo.
Transactions of the Historic Society of Lancashire and Cheshire. ‘The Society.

Vols. vili.-x. 8vo. e

LIST OF DONATIONS. 819

DONATIONS. DONORS.
TRANSACTIONS AND Procenpines or Socrerins, &c.—continued.
London.—Proceedings of the Society of Antiquaries. Vol. iv. Nos. 3-9; The Society.
Vol. v. Nos. 1-3. 8vo.
Transactions of the Society of Antiquaries. Vol. xl. Part 2; xli. Parts Ditto.
1,2; xlin. Part 1. Ato.
Journal of the Society of Arts for 1869-1872. 8vo. The Society.
Journal of the Royal Asiatic Society of Great Britain and Ireland. Vol. The Society.
IV., Ve, vi. Part 1.) 8vo:
A General Index to the first Twenty-Nine Volumes of the Monthly ‘The Society.
Notices of the Royal Astronomical Society. 8vo.
Astronomical, and Magnetical, and Meteorological Observations, made at The Society.
the Royal Observatory in the year 1869. London, 1871. 4to.
A General Index to the First Thirty-Kight Volumes of the Memoirs of Ditto.
the Royal Astronomical Society. 8vo.
Memoirs of the Royal Astronomical Society. Vol. xxxvii. Parts 1, 2 ; Ditto.
Vols. xxxvil., xxxix. Part 1. to.
Monthly Notices of the Royal Astronomical Society for 1869-1872. 8vo. Ditto.
Barometer Manual (1871). 8vo. The Board of Trade.
A Descriptive Catalogue of the Calculi and other Animal Concretions, The College.
contained in the Museum of the Royal College of Surgeons of
England. Supplementi. 4to.
Journal of the Chemical Society. 1869-1871; 1872, Jan—Nov. 8vo. The Society.
Reports on Experiments made with the Bashforth Chronograph to H.M. Stationery

determine the Resistance of the Air-to the Motion of Projectiles. Office.
1865-1870. 8vo.
Transactions of the Clinical Society, Vol. iii, iv.,v. 8vo. The Society.
Journal of the East India Association. No. ii. 8vo. The Association. —

Address at the Anniversary Meeting of the Royal Geographical Society, The Society.
1871, by Sir Roderick Impey Murchison, Bart. 8vo.

Journal of the Royal Geographical Society. Vols. xxxviil., xxxix., xl. Ditto.
8vo.

Proceedings of the Royal Geographical Society. Vol. xiii. No. 5; Vol. Ditto.
“xiv. Parts 1-5 ; Vol. xv. 1-5; Vol. xvi. Parts 1-3. 8vo.

Catalogue of the Published Maps, Sections, Memoirs, and other Publica- The Survey.
tions of the Geological Survey of the United Kingdom to June,
1870. 8vo.

Explanation of Quarter Sheet, 93° S.W., of the One-Inch Geological Ditto.
Survey Map of England. 8vo.

Report of the Geologists’ Association and Excursions for 1869-1871. 8vo. The Association.

Proceedings of the Geologists’ Association. Vol. ii. Nos. 1-6. Annual Ditto.
Report for 1871. 8vo.

Geology of the Country between Liverpool and Southport, and Ex- The Geological
planation of Geological Map, 90° S.E. 8vo. Survey.

Quarterly Journal of the Geological Society. Vol. xxv. Parts 3,4; The Society.
Vol. xxvi. Parts 1-4; Vol. xxvii. Parts 3,4; Vol. xxviii. Parts
1-3. 8vo.

Memoirs of the Geological Survey of Great Britain. London, 1869- The Survey.
1870. 8vo.

Memoirs of the Geological Survey of England and Wales. Vol. iv. 8vo. Ditto.

Mineral Statistics of the United Kingdom of Great Britain and Ireland The Geological

for 1869. 8vo. Survey.
The Journal of the Royal Horticultural Society. Vol. iii, Parts 9,10. The Society.
8vo.

Catalogue of the Library of the Institution of Civil Engineers. Supple- The Library.
ment to Second Edition, 1870. 8vo.

Education and Status of Civil Engineers. 8vo. The Society.

Proceedings of the Institution of Civil Engineers. Vols. xxvii.-xxxii.; The Institution.
Vol. xxxiii. Part 1; Vol. xxxiv. Part 2. 8vo.

Index to Proceedings of the Institution of Civil Engineers. Vols. Ditto.
XXi-xxx. 8vo.

Journal of the London Institution. Vol. i. Nos. 1-15. 8vo. The Institution.

List of the Linnean Society. 1870-71. 8vo. The Society.

820 LIST OF DONATIONS.

DONATIONS. DONORS.
TRANSACTIONS AND PROCEEDINGS OF SociEtins, &c.—continued.
London.—Journal of the Linnean Society. Vols. xi., xii. (Botany); Vol. xiii. The Society.
Nos. 65-67; Vols. x., xi. (Zoology), Nos. 52-54. 8vo.
Proceedings of the Linnean Society, Sessions 1869-70, 1870-71, Ditto.
1871-72. 8vo.
Transactions of the Linnean Society. Vol. xxvi. Parts 3,4; Vol. xxvii. Ditto.
Parts 1-4; Vol. xxvii. Parts 1,2; Vol. xxix. Part 1. Ato.
Proceedings of the Mathematical Society. Nos. 16-47. 8vo. The Society.
General Index to the first Fifty-three Volumes of the Medico-Chirurgical The Society.

Transactions. 8vo.

Proceedings of the Royal Medical and Chirurgical Society.
Nos. 4-8; Vol. vi. Nos. 1, 2. 8vo.

Transactions of the Royal Medical and Chirurgical Society. Vols. lii.-

Vol. vi.

liv. 8vo.
Quarterly Journal of the Meteorological Society. Vol. i. New Series.
Proceedings of the Meteorological Society. Nos. 42-56. 8vo.

The President’s Address, delivered before the Royal Microscopical
Society, February 10, 1869. 8vo.

Charts showing the Surface Temperature of the South Atlantic Ocean
in each Month of the Year. London, 1869. Fol.

Quarterly Weather Report of the Meteorological Office, 1869 ; Parts 1—
4,1870; Part 1, 1871. to.

Transactions of the Pathological Society. Vols. xx.-xxii. 8vo.

Lists of Members of the Royal Institution of Great Britain. 8vo.

Proceedings of the Royal Institution of Great Britain, Vol. v. Parts
5-7 ; Vol vi. Parts 1-4. 8vo.

Royal Society Catalogue of Transactions, Journals, &c. 8vo.

Royal Society Catalogue of Scientific Papers. Vols. iii.-v. 4to. 8vo.

A Discussion of the Meteorology of the Part of the Atlantic lying

The Society.
Ditto.

The Society.
Ditto.
The Society.

The Meteorological
Office.

The Meteorological
Committee of the
Royal Society.

The Society.

Ditto.

The Royal Society.
Ditto.
Ditto.

north of 30° N. for the Eleven Days ending 8th February 1870; ~

with Chart and Diagrams. 4to.

Correspondence concerning the Great Melbourne Telescope.
Parts. 1852-1870. 8vo.

Contributions to our knowledge of the Meteorology of Cape Horn and
the West Coast of South America. 1871. 4to.

In three

Currents and Surface Temperature of the North Atlantic Ocean, from
the Equator to Latitude 40° N. for each Month of the Year; with
a General Current Chart. 4to.

List of the Royal Society. 1869. Ato.

Proceedings of the Royal Society. Vol. xvii. Nos. 121-136. 8vo.

Reports of the Meteorological Committee of the Royal Society for
the Years 1868-1871. 8vo.

Transactions of the Royal Society of Literature.
Vol. x. Part 1. 8vo.

Transactions of the Royal Society of London.
Vol. elx. Parts 1, 2; Vol. clxi. Part 1. 4to.

Statistical Report of the Health of the Navy, for the year 1869. 8vo.

Journal of the Statistical Society. Vol. xxxii. Parts 2-4; Vol. xxxiii.
Parts 1-4; Vol. xxxiv. Parts 1-4; Vol. xxxv. Parts 1-3. 8vo.

Catalogue of the Library of the Zoological Society. 8vo.

Revised List of the Vertebrated Animals now or lately living in the
Gardens of the Zoological Society. 1872. 8vo.

Proceedings of the Zoological Society. 1868, Part 3; 1869, Parts 1-3;
1870, Parts 1-3; 1871, Parts 1-3; 1872, Part 1. 8vo.

Transactions of the Zoological Society. Vol. vi. Part 8; Vol. vil.
Parts 1-8; Vol. vii. Parts 1, 2. 4to.

Vol. ix. Part 3;
Vol. clix. Parts 1, 2 ;

Lyons.—Annales de la Société Impériale d’Agriculture, Histoire Naturelle et

Arts Utiles de Lyon. Quatriéme Série. Tomei. ii. 8vo.
Annales des Sciences Physiques et Naturelles d’Agriculture et d’Indus-
trie. Tome xi. 8vo.

Ditto.
The Meteorological
Committee of the

Royal Society.
The Royal Society.

Ditto.
Ditto.
Ditto.
The Society.
The Royal Society.

The Admiralty.
The Society.

The Society.
Ditto.

_ Ditto.
Ditto.
The Society.

Ditto.

LIST OF DONATIONS.

DONATIONS.
TRANSACTIONS AND PROCEEDINGS OF SociEtiEs, &c.—continued.
Lyons.—Mémoires de l’Académie Impériale des Sciences Belles-Lettres et
> Arts de Lyon. Classe des Lettres. Tome xiv.—Classe des
Sciences. Tome xvii, xvili. 8vo.
Madrid.—Censo de la Ganaderia de Espana segun el recuento verificado en
24 de Setiembre de 1865 por la Junta General de Estadistica. 8vo.
; : _ Maine.—Reports of the Commissioners of Fisheries of the State of Maine
for the years 1867 and 1868, 1870. 8vo.

Manchester.—Proceedings of the Literary and Philosophical Society.
lii., v., vi., vu.; Vol. xi..No..1. 8vo.

Milan.—Atte della Societé Italiana di Scienze Naturali. Vol. xii. Fase. 4;
Vol. xiii. Fase. 1-3; Vol. xiv. Fasc. 1-4; Vol. xv. Fase. 1. 8vo.

Annuario del Instituto Lombardo di Scienze e Lettere 1868. 12mo.
Memorie del Reale Istituto Lombardo di Scienze e Lettere. Classe di
Lettere e Scienze Morale e Politiche. Vol. xi. Della ii. Serie iii.
Fase. 3; Vol. xii. Fasc. 1-4.—Classe di Scienze Matematiche e
Naturali. Vol. xi. Fasc. 1-3; Vol. xii. Fase. 1, 2. 4to.
' Rendiconti Reali Istituto Lombardo di Scienze e Lettere. Serie ii.
Vol. 1. Fasc. 11-20; Vol. ii. Fase. 1-20; Vol. ii. Fase. 1-20;
Vol. iv. ; Vol. v. Fase. 1-7. 8vo.
Solenni- Adunanze del R. Istituto Lombardo di
Vol. i. Fase. 5. 8vo.
Moscow.—Bulletin de la Société des Naturalistes. 1860, Nos. 2-4; 1868,
' Nos. 3,4; 1869, Nos. 1-4; 1870, Nos. 3-4; 1871, Nos. 1-4. 8vo.
Nouveaux Mémoires de la Société Impériale des Naturalistes de Mos-
cow.. Tome xiii. Liv. 2, 3. to.

Munich.—Abhandlungen der kiniglich, bayerischen Akademie der Wissen-
schaften. Historischen Classe. Band xi. Abth. 1-3.—Mathema-
tisch-Physikalischen Classe. Band x. Abth. 2, 3.—Philosophisch-
Philologischen Classe. Band xi. Abth.3; Band xu. Abth. 1, 2. Ato.

Almanach der kéniglich. bayerischen Akademie der Wissenschaften fur
das Jahr 1871. 16mo.

Annalen der Koniglichen Sternwarte bei Miinchen.
xvii. 8vo.

Catalogus Codicum Manu Scriptorum Bibliothece Regie Monacensis.
Tome ii. Pars 2. 8vo.

Sitzungsberichte der konigl. bayer. Akademie der Wissenschaften.
1869, Band i. Heft 1-4; Band ii, Heft 1-4; 1870, Band i. Heft
1-4; Band it. Heft 1-4.—Philosophisch-Philologischen und
Historischen Classe. 1871, Heft 1-6; 1872, Heft 1.—Mathe-
matisch-Physikalischen Classe. 1871, Heft 1-3; 1872, Heft 1. 8vo.

Verzeichniss von telescopischen Sternen, Sup. Band vill, ix., xi. 8vo.

Vols.
Scienze e Lettere.

Band xvii., Band

Naples.—Rendiconto delle Tornate e dei Lavori dell’ Accademia di Scienze
Morali e Politiche. 1869, January to May, September to Decem-
ber; 1870, January to March. 8vo. ‘
: Neuchatel.—Bulletin de la Société des Sciences Naturelles de Neuchatel.
Tome viii. Nos. 2, 3; Tome ix. Part 1. 8vo.
New Haven (U.S.).—Journal (American) of Science and Art, conducted by
Benjamin Silliman. Vol. i., Vol. iz, Vol. ui, New Haven. 8vo.
: New York.—Annual Reports of the Regents of the University of the State
of New York, on the Condition of the State Cabinet of Natural
History. 8vo.
: 81st, 82d, and 83d Annual Reports of the Regents of the University of
the State of New York. 8vo.
: 50th, 51st, 52d, and 53d Annual Reports of the Trustees of the New
York State Library. 8vo.
Monthly Report of the Deputy Special Commissioner of the Revenue
in charge of the Bureau of Statistics, Treasury Department. 1869-
70. Ato.
Natural History of New le (Paleontology).
4to.

VOL. XXVI. PART IV.

By James Hall. Vol.

Iv.

DONORS.

The Academy.

The Junta.

The Commissioners.

The Society.
The Society.

The Institute.
Ditto.

Ditto.

. Ditto.
The Society.
Ditto.

The Academy.

Ditto.
The Royal
Observatory.
The Compilers.

The Academy.

The Royal
Observatory.
The Academy.
The Society.
The Editor.

The University.

Ditto.

The Library.

The Commissioner

The State of New

York.
10 5B

822

LIST OF DONATIONS.

DONATIONS.

TRANSACTIONS AND PROCEEDINGS OF Societies, &¢.—cuntinued.

New York.—Protiles, Sections, and other Illustrations designed to accompany
the final Report of the Chief Geologist of the Survey, F. V. Hayden.
Ato.
New Zealand.—Statistics of New Zealand for 1868. Wellington, 1869.
Fol.
Ohio.—Report (22d, 23d, 24th) of the State Board of Agriculture for 1867—
1870, Columbus. 8vo.
Orleans.—Archives of Science, and Transactions of the Orleans County
Society of Natural Sciences. Vol. i. No. 3. 8vo.
Oxford.—Astronomical and Meteorological Observations made at the Rad-
cliffe Observatory, Oxford, in the year 1866. Vols. xxvi., xxvil.,
LOGpully, So-db<, fh}
Second Radcliffe Catalogue, containing 2386 Stars deduced from Obser-
vations extending from 1854 to 1861 at the Radcliffe Observatory,
Oxford. 8vo.
Palermo.—Giornale de Scienze Naturali ed Economiche. Vol. i. Fase.
3,4; Vol. ii. Fase. 1; Vol. \iii. Fase. 1-3; Vol. iv. Fase. 4;
Vol. v. Fase. 1-4. 8vo.
Paris.—Annales Hydrographiques. No. 4, 1868; Nos. 1-4, 1869; No. 2,
1870. 8vo.
Annuaire des Marés des Cotes de France. 1871, 1872. 12mo.
Publications of the Depédt de la Marine, with Charts. Nos. 448, 449,
452, 454, 455, 456, 458, 459, 461, 462, 463, 464, 465, 467, 468,
470, 472, 473, 474, 476, 490. 8vo.

Annales des Mines. Tome xv. Liv. 2°, 3°; xvi. Liv. 4°, 5°, 6°; xvii.’

Dive SP 3es) evan. lay. 45) Deedes xi inva Lee 2? oe aes
Liv. 4°, 5°, 6°; Septiéme Serie, Tome xxi. Liv. 1°, 2%. 8vo.
Bulletin de la Société de Géographie; Mai, Juin, Juillet, Aout, Sep-

tembre, Octobre, Novembre, Decembre, 1869; 1870; 1871;
Janvier, Fevrier, Mars, Avril, Mai, Juin, 1872. 8vo.
Comptes-Rendus Hebdomadaires des Séances de J Académie des
Sciences, 1869-70, 1870-71, 1871-72. 4to.
Nouvelles Archives du Muséum d’Histoire Naturelle de Paris. Tome
v. Fasc. 3,4; vi. Fasc. 1-4; vii. Fase. 1-4. 4to.
Pesth.—A Magyar Tudomanyos Akadémie Ertesitoje; Szam 9-20, 1868 ;
Szam 1-20, 1869 ; Sz4m 1-12, 1870. 8vo.
Ertekezések a Mathematikai Osztaly Korébol Kiadja A. M. Tudomanyés
Akadémia. Sz4m 3, 4, 1868-69. 8vo.
Ertekezések a Termeszettudomanyok Korébol Kiadja A. M. Tudomanyés
Akadémia. Szam 13-19, 1868-69 ; Sz4m 1, 2, 1870. 8vo.
Philadelphia.—Announcement of the Wagner Free Institute of Science for
the Collegiate year 1870-71. 8vo.
Journal of the Academy of Natural Sciences. New Series. Vol. vi.
Parts 3,4; Vol. vii. 4to.
Proceedings of the Academy of Natural Sciences. Nos. 1-6, 1868
Nos. 1-4, 1869; Nos. 1, 2, 3, 1870; Nos. 1, 2, 3, 1871. 8vo.
Proceedings of the American Philosophical Society. Vol. x. Nos. 78,
79; Vol. xi. Nos. 81-85 ; Vol. xii. Nos. 86, 87. 8vo.
Transactions of the American Philosophical Society. Vol. xii. Part
3; Vol. xiv. Parts 1-3 Ato. .
Portland. —Proceedings of the Portland Society of Natural History. Vol. 1.
Part 2. 8vo.

Quebec.—Manusctipts relating to the Early History of Canada. 8vo.
Report of the Council of the Literary and Historical Society, 1869. 8vo.
Transactions of the Literary and Historical Society. New Series.

Parts 5-8. 8vo.

Rotterdam.—Nieuwe Verhandelingen van het Bataafsch Genootschap der
Proefondervindelijke Wijsbegeerte. Deel ii. Stuk 1. 4to.

St Andrews.— University Calendar for 1870-71. 12mo.

St Petersburg.—Annales de l’Observatoire Physique Central de Russia.
Année 1865-1868. 4to.

DONORS.

The Geological
Survey. —

‘The New Zealand ;

Government. -
The Board.

The Society.

The Observatory.
Ditto.
The Institute.

The Depdt de la
Marine.
Ditto.

Ditto.

The Ecole des
Mines.

The Society.

The Academy.

The Museum.

The Academy.
Ditto.
Ditto.

The Institute.

The Academy.
Ditto.

The Society.
Ditto.

The Society.

The Literary and

Historical Society:

The Society.

The University.

The Russian Go-
vernment.

LIST OF DONATIONS.

DONATIONS.
TRANSACTIONS AND PRocEEDINGS OF SociETIES, &c.—continued.
St Petersburg.—Bulletin de l Académie Impériale des Sciences de St Peters-
bourg. Tome xii. Nos. 4,5; Tome xiv. Nos. 1-6; Tome xv.
Nos. 1-5 ; Tome xvi. Nos. 1-16; Tome xvii. Nos. 1-3. Ato.
Compte-Rendu de la Commission Impériale Archéologique pour |’ Année
1867-1869. 4to. (Atlas Fol.)
Jahresbericht des Physikalischen Central-Observatoriums for 1869, 1870.
Ato.
Mélanges Physiques et Chémiques tirés du Bulletin de Académie Im-
périale des Sciences. Tome vill. 8vo.
Mémoires de |’Académie Impériale des Sciences de St Petersbourg.
vii.° Série. Tome xii. Nos. 4, 5; Tome xiii. Nos. 1-8; Tome
xiv. Nos. 1-9; Tome xy. Nos. 1-8; Tome xvi. Nos, 1-14;
Tome xvii. Nos. 1-12 ; Tome xviii. Nos. 1-7. 4to.
Observations faites 4 la Lunette Méridienne. Vols. i. i, 1869. to.
Observations de Poulkova. Vol. iii. 4to.
Repertorium fiir Meteorologie. Bandi. Heft 1, 2; Bd. ii. Heft 1,2. 4to.
Salem (Mass.).—The American Naturalist. Vol. i, Vol. iii., Vol. iv., Vol.
ve No. 1. 8y¥o.
Memoirs of the Peabody Academy of Science. Vol. i. No. 1. 4to.
First, Second, and Third Annual Reports of the Trustees of the Peabody
Academy of Science 1869-70. 8vo.
Bulletin of the Essex Institute. Vols. i., 1, 11., Nos. 1-12. 8vo.
Proceedings of the Essex Institute. Vols. i-iv., v. Nos. 7, 8. 8vo.
Southampton.—Ordnance Survey of the Peninsula of Sinai, made under the
Direction of Colonel Sir Henry James ; with Maps and Illustra-
tions. 5 vols. 1869. Fol.
Stockholm.—Icones Selectze Hymenomycetum nondum delineatorum ; sub
auspiciis Regiz Acad. Scientiarum Holmiensis, Editze ab Elia
Fries. Parts 1-6. Fol.
Kongliga Svenska Fregatten Eugenies Resa Omkring Jorden under
befal af C. A. Virgin Aren, 1851-1853. Haft 12. to.
Kongliga Svenska Vetenskaps-Akademiens Handlingar. NyFoljd. Bd.v.
Heft 2,1864; Bd.vi. Heft 1,2,1865-66 ; Bd. vii. Heft 1,1867. 4to.
Lefnadsteckningar ofver Kongl. Svenska Vetenskaps-Akademiens efter
ar 1854 aflidna Ledamoter. Bandi. Heft 1. 1869. 8vo.
Meteorologiska Iakttagelser i Sverige utgifna af Kongl. Svenska Veten-
skaps-Akademien anstiiallda och bearbetade under Inseende af Er.
- Edlund. Band vi. 1864; Band vi. 1865; Band viii. 1866. 4to.
Ofversight af Kongl. Vetenskaps-Akademiens Forhandlingar, 1865-
1868. 8vo.
Sveriges Geologiska Undersokning ; with Charts. Livs. 31-34. 8vo.

Switzerland.—Verhandlungen der Schweizerischen Naturforschenden Ge-
sellschaft in Einsiedeln. 1868. 8vo.
Throndhjem.—Det Kongelige Norske Videnskabers-Selskabs, Skrifter i det
19% Aarhundrede. Bind v. Heft 2. 8vo.
Toronto.—Canadian Journal of Science, Literature, and History. Vol. xii.
Nos. 3-6 ; Vol. xiii. Nos. 1-4. 8vo.
Turin.—Atlante di Carte Celesti Contenenti le 634 stelle principali visibili
alla latitudine Boreale di 45°. Fol.
Atti della Reale Accademia delle Scienze. Vol. iv., Vol. v. Disp. 1-7;
Vol. vi. Disp. 1-6. 8vo.
Memorie della Reale Accademia delle Scienze di Torino. Serie
Seconda. Tomo xxv., xxvi. Ato.
Reale Accademia delle Scienze de Torino Regio Osservatorio Atlant
di Carte Celesti. Fol.
Bollettino Meteorologico dell’ Osservatorio Astronomico dell’ Universita,
1868-1872. 4to.
R. Osservatorio dell’ Universita di Torino. Supplemento al V. Bollet-
tino Annuale 1870, dell’ Osservatorio. 8vo.

823

DONORS.

The Academy.

The Commission.

The Royal Aca-
demy.
Ditto.

Ditto.

The Poulkowa
Observatory.
The Royal Academy.
The Peabody Aca-
demy of Science.
Ditto.
Ditto.

The Institute.
Ditto.
The Rt. Hon. the
First Commissioner

of H.M. Works.
The Academy.

Ditto.
Ditto.

Ditto.

The Bureau de la Re-
cherche Geolo-
gique de la Suede.

The Society.

The Canadian Insti-
tute.

The Royal Aca-
demy.
Ditto.
Ditto.
Ditto.

The University.

Ditto.

824 LIST OF DONATIONS.

DONATIONS. DONORS.
TRANSACTIONS AND PROCEEDINGS OF SoctEtins, &c.—continued.
Ulm.—Verhandlungen der Verein fiir Kunst und Alterthum in Ulm und The Editor.
Oberschwaben. Heft 1, 1869. Ato.
Upsala.—Bulletin Météorologique Mensuel de l’Observatoire de Université. The University.
Vol. u. Nos. 1-6; Vol. ii. Nos. 1-12. Ato.
Nova Acta Regiz Societatis Scientiarum Upsaliensis. Vol. vii. Fase. The Society.
1, 2; Vol. vii. Fase..1. 4to.
Utrecht.—Nederlandsch Meteorologisch Jaarboek 1867-68, 1868-69, 1869- Meteorological In-
5 ebiroy stitute of Utrecht.
Nederlandsch Kruidkundig Archief. Deeli. Stak. 1. 8vo. The Editors.
Aanteekeniiigen van het Verhandelde in de Sectivergaderingen van het The Society.
Provinciaal Utrechtsch Genootschap van Kunsten en Wetenschap-
pen, 1868-1870. 8vo.

Catalogus der Archeologische Verzameling van het Provinciaal Utrechtsch Ditto.
Genootschap van Kunsten en Wetenschappen. 1868. 8vo.
Verslag van het Verhandelde in de algemeene Vergadering van het Ditto.

Provinciaal Utrechtsch Genootschap van Kunsten en Wetenschap-
pen, 1868-1871. 8vo.
Mémoire sur le genre Potérion par P. Harting. 4to. ; Society of Arts and
Sciences, Utrecht.
Venice.—Atti del Reale Istituto Veneto di Scienze, Lettere ed Arti. Tomo The Institute.
xii. Dispenso 10; Tomo xiil., xiv. ae 1-10; Tomo xv. Disp.
1-10 ; Tomo xvi. Disp. 1-10; Serie iv. ; Tomo i. Disp. 1-4. 8vo.
Victoria (Austr alia).—Abstracts of Specifications of Patents applied for The Registrar-

from 1854 to 1866. Metals, Parti. Melbourne, 1872. 4to. General.
Patents and Patentees. Vol. iv. Melbourne, 1871. 4to. Ditto.
Agricultural Statistics of the Colony for 1869-70. Fol. Ditto.
Statistics of the Colony for 1868-1870. Population. Fol. Ditto.
Mineral Statistics of the Colony for 1871. Fol. Ditto.
Census of Victoria for 1871. Parti. Inhabitants and Houses. Fol. The Australian Gov.
Reports of the Mining Surveyors and Registrars for Quarter ending Ditio.
31st March 1872. Fol.
Report of the Board on Coal-Fields, Western Port. No. 19. Fol. Ditto.
Transactions and Proceedings of the Royal Society, Victoria. Vol. x. The Society.
Part 2. 8vo.
Vienna.—Almanack der kaiserlichen Akademie der Wissenschaften, 1869— The Academy.
ISv1. Syo:

Denkschriften der kaiserlichen Akademie der Wissenschaften. Math. Ditto.
Nat. Classe. Band xxix., xxx., xxxii—Phil. Hist. Classe. Bands
RVa,, XVI MaKe eee AOL
Jahrbuch der kaiserlich-koniglichen Geologischen Reichsanstalt. B. xix. The Society.
Nos. 1-4 ; B.xx. Nos. 1-4 ; B. xxi. Nos. 1-4 ; B. xxii. Nos. 1,2. 8vo.
Phanologische Beobachtungen aus dem Pflanzen und Thierreiche von The Academy.
Karl Fritsch. Heft 8. Jahrgang 1857. 4to.
Register zu den Banden 51 bis 60 der Sitzungsberichte der Philos.-His- Ditto.
_ tor. Classe.
Sitzungsberichte der kaiserlichen Akademie der Wissenschaften. Phil. Ditto.
Hist. Classe. Band viii. Heft 1,2; Bandix. Heft 3-5 ; Band xxvii.
Heft 2,3; Band xxx. Heft]; Band xxxvi. Heft 2; Band lix. Heft
1-4 ; Band lx. Heft 1-3; Band lxi. Heft 1-3 ; Band lxu. Heft 1—
4,—Mat. Nat. Classe. Band xxvii. Heft 2; Band xxx. Heft 16, 17 ;
Band xxxv. Heft 7-9 ; Band xxxix. Heft 2; Band lvii. Heft 4, 5 ;
Band lviii., Heft 1-5 ; Band lix. Heft 1-5 ; Band lx. Heft 1, 2.—
Mineralogie-Botanik, &c. Band lvii. Heft 4,5; Band lviii. Heft
1-5 ; Band lix. Heft 1-5; Band Ix. Heft 1, 2. 8vo.
Sitzungsberichte der kaiserlichen Akademie der Wissenschaften. The Academy.
Phil. Hist. Band lxiii. ; Band lxiv. ; Band lxv. ; Band Ixvi. Heft
2,3; Band lxvii. Heft 1-3; Band ‘Ixviil, H eft, 1-4; Band Ixix.
Heft 1- 3.—Mat. Nat. Classe, Band lx. Heft 3.5; Band 1x1.
Heft 1-5 ; Band Ixii. Hert 1-5 ; Band Ixiii.; Band Ixiv.—Botanili,
Zoologie, &e. Band lx. Heft 35; Band Ixi. Heft 1-5 ; Band lxu.
Heft 1-5 ; Band Ixiii.; Band Ixiv. 8vo0.

LIST OF DONATIONS.

DONATIONS.
TRANSACTIONS AND ProcrrpDiInGs or Societies, &c.—continued.
Vienna.—Die Echinoiden der Oesterreichisch-Ungarischen oberen Tertiaera-
blagerungen, von Dr Gustav C. Laube. Band v. Heft 3. Ato.
Die Fossilen Mollusken des Terticer-beckens von Wien, von Dr Hornes.
Band ii. Nos. 9,10. 4to.
Die Reptilfauna der Gosau—Formation in der Nuen Welt bei Weinner-
Neustadt, von Dr Emanuel Bunzel. Band v. Nos. 1, 2. Ato.
Verhandlungen der kaiserlich-koniglichen zoologisch-botanischen Gesell-
schaft in Wien. Band xix., xx., xxi. 8vo.
Verhandlungen der kaiserlich- kéniglichen geologischen Reichsanstalt.
1869, Nos. 1- 5, 10-18 ; 1870, Nos. 1- a 1871, Nos. 1-5, 7-10;
1872, Nos. 1-6. 8vo.
Warwick,—Thirty-fourth, Thirty-fifth, and Thirty-sixth Annual Reports of
Natural History and Archeological Society, 1870-1872. 8vo.
Washington.—Astronomical and Meteorological Observations made at the
United States Naval Observatory during 1866, 1867, 1869. 4to.
Congressional Directory for the Third Session of the Forty-first Congress
of the United States of America. 8vo.
Reports of the Commissioner of Agriculture for 1868, 1869, and 1870.
8vo.
Monthly Reports of the Department of Agriculture for Tee? 1870,
and 1871. Edited by J. R. Dodge. 8vo.
Twelfth Annual Report of the Columbia Institution for the Deaf and
Dumb, 1869. 8vo.
Annual Reports of the Commissioner of Patents for 1867 and 1868.
8vo.
Report of the Superintendent of the United States Coast Survey for
1866, 1867, and 1868. 4to.
Special Report on Immigration. 1872. 8vo.
Report of the United States Geological Survey of Montana. 1872. 8vo.
Reports of the National Academy of Sciences for 1867 and 1868. 8vo.
Reports of Surgical Cases in the Army. No. 3, 1871. Ato.

Annual Report of the Board of Regents of tie Smithsonian Institution
for 1867, 1868, 1869, and 1870. 8vo.

Smithsonian Contributions to Knowledge. Vols. xvi., xvii. 4to.

The Transatlantic Longitude as determined by the Coast Survey Expe-
dition for 1866. By Benjamin Apthorp Gould, 1869. 4to.

Smithsonian Miscellaneous Collections. Vols. viii., ix. 8vo.

Catalogue of Orthoptera of North America described previous to 1867.
8vo.

Wellington (New Zealand).—Statistics of New Zealand for 1867, 1869,
1870, and 1872. Fol.

Whitby.—Forty-eighth Report of the Literary and Philosophical Society,
1870. 8vo.

York.—Communications to the Monthly Meetings of the Yorkshire Philo-
sophical Society. 1870, 1871. 8vo.

Zurich.—Neue Denkschriften der allgemeinen schweizerischen Gessellschaft
fiir die gesammten-Naturwissenschaften—(Nouveaux Mémoires de
la Société Helvétique des Sciences Naturelles). Band xxiii. mit
26 Tafeln ; Band xxiv. mit 11 Tafeln. 4to.

VOL. XXVI. PART Iv.

825

DONORS.

The Society.
Ditto.
Ditto.
Ditto.

The Society.

The Society.
The United States

Government.
The Congress.

The United States
Government.
The Editor.

The Institution.

The United States
Patent Office.
The Survey. °

The Bureau.

The Survey.

The Academy.

The Surgeon-Gene-
ral’s Office.

The Institution.

Ditto.
Ditto.

The New Zealand
Government.

The Society.

10 F

826 - LIST OF DONATIONS.

AUTHORS’ WORKS OR DONATIONS.

Agassiz (Alexander). Application of Photography to Illustrations of Natural
History; with Two Figures printed by the Albert and Woodbury
processes. 8vo.

Agassiz (Louis). Address delivered on the Centennial Anniversary of the Birth
of Alexander von Humboldt, under the auspices of the Boston Society of
Natural History. Boston, 1869. 8vo.

Contributions to the Fauna of the Gulf Stream at Great Depths. Cam-
bridge, Mass. 8vo.

Report upon Deep Sea Dredgings. Cambridge, Mass. 8vo.

Allen (J. A.). Mammalia of Massachusetts. Cambridge, Mass. 8vo.

Anderson (Benjamin). Narrative of a Journey to Musadu, the Capital of the
Western Mandingoes. New York, 1870. 8vo.

Anderson (John), M.D. Note on Occurrence of Sacculina in the Bay of Bengal.

8vo.

— On some Indian Reptiles. 8vo.

—— Description of a New Genus of Newts from Western Yunan. 8vo.

—— Note on Testuda Phayrit. 8vo.

—— Description of a New Cetacean from the Irrawaddy River, Burmah. 8vo.
— On three New Species of Squirrels from Upper Burmah and the Kakhyen

Hills, between Burmah and Yunan. §8vo.

‘—— On Eight New Species of Birds from Western Yunan, China. 8vo.

Notes on some Rodents from Yarkand. 8vo.

Description of a New Species of Scincus. 8vo.

A Report on the Expedition to Western Yunan. 4to.

Asman (Dr. P. H.). Proeve eener Geneeskundige Plaatsbeschrijving ven de
Gemeente Leeuwarden en. Utrecht, 1870. 8vo.

Balfour (Professor). Description of Hieraciwm collinum of Fries, anew British
Plant. 8vo.

Barclay (Joseph Gurney). Astronomical Observations taken during the years
1865-1869, at his Private Observatory. Vol. iii London, 1870. 4to.

Baudet (P. J. H.). Leven en Werken, van Willem Janz, Blaeu. Utrecht,
1871. 8vo.

Benson (Prof. Lawrence 8.). Dissertation on the Principles and Science of
Geometry. New York, 1871. 8vo.

Bergman (Jo. Theod.). Memoria Ludovici Caspari Valckenarii. Rheno-Trajecti,
1871. 8vo.

Bert (M. P.). Influence des diverses couleurs sur la Vegetation. 4to.

Blade (M. Jean Francois). Etudes sur Origine des Basques. 8vo.

Defense des Etudes sur l’Origine des Basques. 8vo.

Blandford (W. T.). Observations on the Geology and Zoology of Abyssinia, made
during the progress of the British Expedition to that Country in 1867-68.
8vo.

Blyden (Rev. Edward W.). Appendix to Benj. Anderson’s Journey to Musadu.
New York, 1870. 12mo.

Blytt (A.). Christiania, Omegns Phanerogamer og Bregner. 8vo.

Bonnel (J. F.). Essai surles Definitions Géometriques. Paris, 1870. 8vo.

Boott (Francis), M.D. TIlustrations of the Genus Carex. Part iv. London,
1867. Fol.

Botten-Hansen (Paul). La Norvége Littéraire. Christiania, 1868. 8vo.

Boyle (W. R. A.). The Tribute of Assyria to Biblical History. London, 1868. 8vo.

Literature under the Shade of Great Britain. In a Letter to the Right

Hon. W. E. Gladstone. London, 1870. 8vo.
Breen (Hugh). Corrections of Bouvard’s Elements of Jupiter and Saturn. Paris,

1821.
Brigham (W.T.). Historical Notes on the Earthquakes of New England, 1638-
1869. Ato.

Notes on the Eruption of the Hawaiian Volcanoes, 1868. Boston, 1869. 4to.

—— The Colony of New Plymouth and its relation to Massachusetts. Boston,
1869. 8vo.

Contributions of a Venerable Savage to the Ancient History of the
Hawaiian Islands. Boston, 1868. 8vo.

DONORS.
The Author.

Ditto.

Ditto.
Ditto.
Ditto.

Ditto.

Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto. .
Ditto.
Indian Govern-
ment.
The Author.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.

Ditto.

LIST OF DONATIONS.

- AUTHORS’ WORKS OR DONATIONS.
Brink (B. Ten). Levensbeschrijving van Rijklof Michaél van Goens. Uirecht,

1869. 8vo.
Bristow (H. W.), and Whitaker (Wm.). On the Formation of the Chesil Bank,
Dorset. 8vo.

Brown (Robert), Ph.D., A.M. Descriptions of some new or little known Species of
Oaks from North-West America. (From Ann. Mag. Nat. Hist., April
1871.) 8vo.

— On the Physics of Arctic Ice, as Explanatory of the Glacial remains in
Scotland. (From Quart. Jour. Geol. Soc., Feb. 1871.) 8vo.

Caspari (Dr le P.). Ungedruckte unbeachtete und wenig beachtete Quellen zur
Geschichte des Taufsymbols und der Glaubensregel. - Christiania. 8vo.

Chatelier (M. L. Le). Railway Economy. Translated by Lewis D. B. Gordon.
Edinburgh, 1869. 8vo.

Colding (A.). Om Stroemningsforholdene i almindelige Ledningerog i Havet.
Kjcebenhavn. 4to.

Cox (E. T.). First Annual Report of the Geological Survey of Indiana during
the year 1869. 8vo.

Day (St John Vincent). On Patents for Inventions. Glasgow, 1870. 8vo.
On some Evidences as to the very early use of Iron. Edinburgh, 1871. 8vo.
—C.E. On Asbestos, with special reference to its Use as Steam-Engine
Packing. Glasgow, 1872. 8vo.
Delesse (M.). Revue de Géologie pour les Années 1867 et 1868. Tome vii.
Paris, 1871. 8vo.
Dircks (Henry), C.E., LL.D. Patent Monopoly, as represented by Patent Law
Abolitionists, impartially examined. London, 1869. 8vo.
Scientific Studies, two Popular Lectures. 1. Marquis of Worcester.
2. Chimeras of Science. London, 1869. 8vo.
—— Nature Study. London, 1869. 8vo.
The Policy of a Patent Law. London, 1869. 8vo.
Dole (Sandford ze). A Synopsis of the Birds of the Hawaiian Islands. Boston,
1869. 8vo.

Erlenmeyer (Dr Emil). Die Aufgabe des Chemishen Unterrichts gegeniiber den
Auforderungen der Wissenschaft und Technick. Munchen, 1871. 4to.

Everett (Prof. J. D.). On the General Circulation and Distribution of the
Atmosphere. 8vo.

Fayrer (J.), M.D., C.S.I. The Thanatophidia of India ; being a Description of
the Venomous Snakes of the Indian Peninsula, with an Account of the
of their Poison on Life. London, 1872. Fol.

H.R.H. The Duke of Edinburgh in India.. Calcutta, 1870. to.

Frauenfeld (George Ritter Von). Die Grundlagen des Vogelschutzgesetzes.

Wien, 1871. 8vo.
Friis (Professor J. A.) .Salbmagirje (Lappisk Salmebog). Christiania, 1871. 12mo.
Fuchs (Dr C. W.C.). Die Kiinstlich dargestellten Mineralien nach G. Rose’s
Krystallo-chemischen Mineralsysteme geordnet. Haarlem, 1872. 4to.

Gabba (Luigi). Rapporti sui Progressi delle Scienze. Milano. 1870. 8vo.

Gamgee (Dr Arthur). Researches on the Blood.—On the Action of Nitrites
on Blood. 4to.

—— On Force and) Matter in Relation to Organisation. Edinburgh, 1869. 8vo.

Gamgee (Sampson). On the Treatment of Fractures of the Limbs. 8vo.

Geikie (James). On Changes of Climate during the Glacial Epoch. 8vo.

Ghirardini (Alessandro). Studi sulla Lingua Umana sopra alcune Antiche
Inscrizioni, e sulla Ortografia Italiana. Milano, 1869. 8vo.

Giltay (Dr K. M.). Gedachtenisviering von het honderdjarig bestaan von het
Bataafsch Genootschap der Proefondervindelijke Wijsbegeerte te Rotterdam
1769-1869. Rotterdam, 1869. 4to.

Gore (G.), F.R.S. On Hydrofluoric Acid. From the Transactions of the Royal
Society for 1868. 4to.

DONORS.
The Author.

The Authors.

The Author.
Ditto.

Ditto.

The Translator.

The Author. —
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.

827

828 LIST OF DONATIONS.

AUTHORS’ WORKS OR DONATIONS.

Gorresio (Gaspare). Sunti dei Lavori Scientifici letti e discussi nella Classe di
Scienze Morali, Storiche e Filologiche. Torino, 1868. 8vo.

Gould (Augustus A.), M.D. Report on the Invertebrata of Massachusetts.
Boston, 1870. 8vo.

Gould (Benjamin Apthorp). Investigations in the Military and Anthropological
Statistics of American Soldiers. New York, 1869. 8vo.

Grant (Robert E.), M.D. Umrisse der Vergleichenden Anatomie. Leipzig,
1842. 8vo.

Grundfjeldet (I.). On Skuringsmeeker Glacialformationen og Terrasser. Kris-
tiania, 1871. Ato.

Haeckel (Dr Ernst). Entwickelungsgeschichte der Siphonophoren. Utrecht,
1869. Ato.

Hall (Townshend M.), F.G.S. Topographical Index to the Fellows of the
Geological Society of London. 8vo.

Harris (Thaddeus William), M.D. Entomological Correspondence of. Edited by
S. H. Scudder. Boston, 1869. 8vo.

Hasskarl (Carolo). Commelinacee Indicae, imprimis Archipelagi Indici.
Vindobonae, 1870. 8vo.

Haswell (James). On Columnar Structure developed in Mica Schist, from a
Vitrified Fort in the Kyles of Bute. 8vo.

-—— Notice of Sandstone, now in the course of formation at Elie, Fifeshire. 8vo.

Hauer (Franz Ritter v.). Zur. Ermnerung an Wilhelm Haidinger. Vienna,
1871. 8vo.

Haug (Dr Martin). Brahma und die Brahmanen. Munich, 1871. 4to.

Heller (Prof. Cam). Die Zoophyten und Echinodermen des Adriatischen
Meeres. Vienna, 1868. 8vo.

Henwood (William Tory), F.R.S. Address to the Royal Institution of Cornwall.
Penzance, 1869. 8vo.

Hertzberg (Ebbe). En fremstilling af de norske Aristokratis histoire. Christiania,
1869. 8yo. .

Hoeufft (Jacobi Henrici). Urani, Carmen Didascalicum Petri Esseiva. Am-
stelodami, 1870. 8vo.

Hoffman (Dr C. K.), und H. Weyenbergh (J.). Die osteologie und myologie
von Sciurus vulgaris L. Harrlem, 1870. 4to.

Jervis (Cav. Guglielmo). R. Museo Industriale Italiano Ilustraziari delle,
Collizioné Didattica. Parte Prima. 8vo.

“ Julian.” Biology versus Theology; or, Life on the Basisof Hylozoism. Lewes,
1870. 8vo.

Koérosi (Josef). Vorlanfiger Bericht uber die Resultate der Pester. Volkszahlung
vom Jahre, 1870. 8vo.

Kuntsler (Gustav). Die unseren Kulturpflanzen Schidlichen Insokten. Wien,
1871. 8vo.

Lea (Isaac), LL.D. Observations on the Genus Unio, together with Descriptions
of new Species in the Family Unionidx, and Descriptions of new Species
of the Melanide and Paludine, with 26 Plates. Vol. xii. Philadelphia.

Ato.
—— Index to Vol. xii. of Observations on the Genus Unio. Philadelphia,
1869. to.

A Synopsis of the Family Unionide. Philadelphia, 1870. 4to.

Lévéque (G.). Recherches sur /’Origine des Gaulois. Paris, 1869. 8vo.

Linnarsson (J. G. O.). On some Fossils found in the Eophyton Sandstone at
Lugnas in Sweden. Stockholm, 1869. 8vo.

Littrow (Carl von). Ueber das Zuriickbleiben der Alten in den Naturwissen-
schaften. Wien, 1869. 8vo.

Logan (Sir W. E.). Geological Map of Canada. 1866.

Loven (Af. S.). Om en marklig i Nordsjén lefvande art af Spongia. Stockholm. 8vo.

Lowe (E. J.). Natural Phenomena and Chronology of the Seasons. London,

1870. 8vo.

DONORS.
The Author.

The Boston Society
of Natural History.
The United States.
Sanitary Commission
The Author.

Ditto.

Ditto.
The Boston Society
of Natural History.
The Author.

Ditto.

Ditto.
Ditto.

Ditto.
Ditto.
Dita.

The Authors.

The Author.

Ditto.

Ditto.
Ditto.
Ditto.

Ditto.
Ditto.

LIST OF DONATIONS.

AUTHORS’ WORKS OR DONATIONS.
Lubbock (Sir John), Bart. Note on some Stone Implements from Africa and
Syria. 8vo.
— On the Development of Relationships. 8vo.

Mackinder (D.), M.D. Clinical Notes. 8vo.

M‘Farlane (Patrick). Antidote against the Unscriptural and Unscientific Tend-
ency of Modern Geology ; with remarks on several Cognate Subjects. 8vo.

Martins (Ch.), et Chancel (G.). Des Phénoménes Physiques qui accompagnent
la rupture par la Congelation de Eau des Projectiles Creux de divers
calibres. Montpellier, 1870. Ato.

Maxwell (J. Clerk), LL.D. Theory of Heat. 12mo.

Meissner (C. F.). Denkschrift auf Carl Friedr. von Martius. Munich, 1869. 4to.

Miller (Rev. Jas. N.). The true Direction and Velocity of Wind observed from
Ships while sailing. London, 1870. 8vo.

Mohn (H.). ‘Température de la mer entre l'Islande, l’Ecosse et la Norvége. Chris-
tiania, 1870. 8vo.

Morris (John). Lead-bearing Districts of the North of England. London, 1869.
8yvo.

Mueller (Ferdinand von), M.D. New Vegetable Fossils of Victoria. Fol.

Fragmenta Phytographiz Australie. Vol. vi. Melbourne. 8vo.

— The Principal Timber Trees readily eligible for Victorian Industrial Cul-

ture. 8vo.

Forest Culture in its relation to Industrial Pursuits. 8vo.

Muir (J.), D.C.L., LL.D. Original Sanskrit Texts on the Origin and History of
the People of India. Vols. ii—v. 8vo.

Mullins (J. D.). Catalogue of the Reference Department of the Birmingham
Free Libraries. Birmingham, 1869. 8vo.

Neilreich (Dr August). Die Vegetationsverhaltnisse von Croatien. Vienna, 1868.

8vo.

Nicholson (H. Alleyne), M.D. Monograph of the British Graptolitide. Part 1.
8vo.

Nordenskiold (A. E.). Sketch of the Geology of Spitzbergen. Stockholm, 1867.
8vo.

Nowicki (Prof. Dr Max.). Ueber die Weizenverwiisteri Chlorops Tzniopus
Meig und die Mittel zu ihrer Bekimpfung. Wien, 1871. 8vo.

Orlandini (C. C.). Rivelazioni Astronomiche aggiunte alla Declamazione Filosofica.
Bologna, 1869. 8vo.

Pacini (Prof. Filippo). Sull’ Ultimo Stadio del Colera Asiatico. Firenze, 1871.
3 :

vo.

Packard (A. 8.), M.D. Record of American Entomology for 1868 and 1869.
Salem, 1869. S8vo.

Parrish (R. A.), Jun. Details on an Unpaid Claim on France for 24,000,000
Francs, guaranteed by the Parole of Napoleon III. Philadelphia, 1869.
8vo.

Pascucci (Prof. Luigi). Brevi Cenni sulle Specialité Mattei con sunto delle Malatte
Senate nella Citté di Roma 1869. Rome, 1870. 8vo.

Peacock (R. A.). Changes on the Earth’s Physical Geography, and consequent
Changes of Climate. London, 1871. 8vo.

Peters (Dr). Report on the Longitude of the Western Boundary Line of the State
of New York. Albany, 1868. 8vo.

Plantamour (E.). Résumé Météorologique de année 1868, pour Geneve et le
Grand Saint Bernard. Geneve, 1869. 8vo.

—— Résumé Météorologique de V’année 1869-70. Geneve et le Grand Saint
Bernard. 8vo.

—— Nivellement de Précision de la Suisse. Geneve, 1870. 8vo.

— Détermination Télégraphique de la Différence de Longitude, par E. Planta-

' mour, R. Wolf, et A. Hirsch. 1871. Ato.

—— Nouvelles Expériences faites avec le Pendule Réversion et Determination

de la Pesanteur 4 Genéve et an Righi. Kulm, 1872. 4to.
VOL. XXVI. PART. IV. y

829

DONORS.
The Author.

Ditto.

Ditto.
Ditto.

The Authors.
The Author.
Ditto.
Ditto.
Ditto.
The Geologists’ As-
sociation.
The Author.
Ditto.
Ditto.

Ditto.
Ditto.

Ditto.

Ditto.
Ditto.
Ditto.

Ditto.

Ditto.
Ditto.

Ditto.

Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.

Ditto.

106

830 LIST OF DONATIONS.

AUTHORS’ WORKS OR DONATIONS.

Plaseller (Dr J.). Compendium Stenographie Latine. iniponte, 1868. 8vo.

Pourtales (L. F. de). Contributions to the Fauna of the Gulf Stream at Great
Depths. (Second Series.) Cambridge, Mass., 1868. 8vo.

Preger (Wilhelm). Die Entfaltung der Idee des Menschen durch die Welt-
geschichte. 4to.

Prestel (Dr M. A. F.). Das Gesetz der Winde abgeleitet aus dem Auftretender-
selben iiber Nordwest-Europa. Emden, 1869. 4to.

Quatrefages (A. de). La Race Prussienne. Paris, 1871. 12mo.

Quetelet (Ad.). Note sur Aurore Boréale du 6° Octobre et les Orages de 1869.
Brussels. 8vo.

—— Physique Sociale ou Essai sur le Développement des Facultés de ! Homme.
Brussels, 1869. 8vo.

—— Sur les Orages observés en Belgique pendant Vannée 1868, et le premier

Trimestre de 1869. Brussels. 8vo.

Sur les Etoiles Filantes du mois d’Aout 1869, observées & Bruxelles. 8vo.

—— Anthropométrie ou Mesure des Différentes Facultés de Homme. Brus-

sels, 1870. 8vo. :

—— Observations des Phénoménes Périodiques pendant l’année 1869. to.

— Loi de Périodicité de 1Espéce Humaine. 8vo.

Notice of Sir John F. W. Herschel. 8vo.

Quetelet (Ern.). Notices sur les Aurores Boréales des 15 Avril et 13 Mai 1869.
Brussels, 1869. 8vo.

Realis (M. 8.). Note sur le Nombre. Paris, 1869. 8vo.

Regnault (M. V.). Relation des Expériences pour detérminer les lois et les
données Physiques necessaires au calcul des Machines a Feu. Paris, 1870.
Ato.

Reid (Hugo). Memoir of the late David Boswell Reid, M.D., F.R.S.E. Edin-
burgh, 1863. 8vo.

Rein (Dr. J. J.). Bericht ttber die Senckenbergische Naturforschende Gesellschaft
in Frankfurt om Main. 1869. 8vo.

“Research.” Earth, True Theory of the. Edinburgh, 1869. 8vo.

Report on Measures adopted for Sanitary Improvements in India during the year
1868 and up to the month of June 1869. London, 1869. Fol.

Risfen (Hartvig). Stoleveefenets Ordnung i Massachusetts. Christiania, 1868. 8vo.

Rive (Prof. A. de la). Recherches sur la Polarisation rotatoire magnétique des
Pe _8yvo.

Dees. eee Geneva, 1871. 8vo.
Notice sur Emile Verdet.. Paris, 1870. 8vo. .
Roy (Alphonse le). L’Université de Liége depuis sa fondation. Liége, 1869. 8vo.

Settimanni (Capt. César). D’une seconde Nouvelle Méthode pour déterminer la
Parallaxe du Soleil. Florence, 1870. 8vo.

— Nouvelle Théorie des principaux Eléments de la Lune et du Soleil. Flo-
rence, 1871. Ato,

Sexe (S. A.). Le Glacier de Boium en Juillet 1868. Christiania, 1869. 4to.

Simpson (Martin). A Guide to the Geology of the Yorkshire Coast. 4th Edition.
London, 1868. 8vo.

Smith (Dr John Alexander). Notice of Remains of the Reindeer (Cervus tarandus),
found in Ross:shire, &., with Notes of its occurrence throughout Scot-
land. Edinburgh, 1869. 8vo.

Snellaert (F. A.). Nederlandsche Gedichten uit de veertiende eeuw van Jan
Boendale, Hein van Aken, en andaren. Brussels, 1869. 8vo.

Sobrero (Ascanio), Notizia Storica dei Lavori fartti della Classe di Scienze
Fisiche Matematiche della Reale Accademia delle Scienze di Torino negli
anni 1864 e 1865. 8vo.

Stal (Carolus). Hemiptera Africana. Tom.i-iv. Holmiz, 1854. 8vo.

Steen (Adolph). Om Integrationen af Differentialligninger, der fore til Addi-
tionstheoremer for transcendente Funktioner. Copenhagen, 1868. 4to.

DONORS.
The Author.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.

Ditto.
Ditto.
Ditto.
Ditto.

Ditto.
Dr Morehead

The Author.
Ditto.

The Authors.

The Author.
Ditto.

Ditto.
Ditto.

Ditto.

Ditto.
The Royal Academy
of Sciences,
Copenhagen.

LIST OF DONATIONS.

AUTHORS’ WORKS OR DONATIONS.
Stevenson (David), F.R.S.E. Altered Regulations of British and Foreign Indus-
tries and Manufactures ; the Cause and the Cure. An Address to the
Royal Scottish Society of Arts on 8th November, 1869. Edinburgh,
1869. 8vo.
— The Principles and Practice of Canal and River Engineering. Second
Edition. 1872. 8vo.
Stewart (B.). Account of Certain Experiments on Aneroid Barometers made at
Kew Observatory. 8vo.
Stirling-Maxwell (Sir Wm.), Bart. Address to the Students of the School of
Arts, Edinburgh, under charge of the Hon. the Commissioners of the
Board of Manufactures, at the delivery of Prizes, January 13, 1870. 8vo.
Stratton (Thomas), M.D., R.N. The Affinity between the Hebrew Language and
the Celtic. Edinburgh, 1872. 8vo.
Strecker (Adolph). Jahresbericht iiber die Fortschritte der Chemie, &c., fiir
1868. Heft 2. Giessen. 8vo.
Jahresbericht tiber die Fortschritte der Chemie, &c., fir 1868. Heft 3.
Giessen. 8vo.
Jahresbericht iiber die Fortschritte der Chemie, &c., fir 1869. Heft 1-3.
Giessen. 8vo.
Struve (Otto). Jahresbericht am 5 Juni 1869 dem Comité der Nicolai-Haupt-
sternwarte. St Petersburg, 1869. 8vo.
Tabule Quantitatum Besselianarum pro annis 1850 ad 1840 computate.
Petropoli, 1869. 8vo.
Jahresbericht am 29 Mai 1870 dem Comité der Nicolai-Hauptsternwarte.
St Petersburg, 1870. 8vo.
Studer (B.). Erliuterungen zur zweiten Ausgabe der Geologischen Karte der
Schweiz von B. Studer und A. Escher. Winterthur, 1869. 8vo.
— Index der Petrographie und Stratigraphie der Schweiz und ihrer Ungebungen,
Bern. 1872. 8vo.
Sundevall (Carl J.). Die Thierarten des Aristoteles von den Klassen den Siuge-
thiere, Vogel, Reptilien und Insekten. Stockholm. 8vo.
Conspectus Avium Picinarum. Stockholm, 1866. 8vo.
Suringar (W. F. R.). Algze Japonicee Musei Botanici Lugduno. Batavi. 8vo.

Thayer (C. F.), and Buswell (H. T.). Address and Ode delivered at the Dedica-
tion of Memorial Hall, Lancaster, 17th June 1868. Boston, 1868. 8vo.
Thomsen (Julius). Thermochemiske underscegelsen. Kjcebenhavn. 4to.
Undersgelser over Basernes Neutralisationsvarme. Kjcebenhayn, 1871. 4to.
Topinard (Dr Paul). Etude sur les Races Indigénes de l’Australie. Paris, 1872.
8vo.
Toynbee (Capt. Henry). On the Meteorology of the North Atlantic between the
Parallels of 40° and 50° North. London, 1869. 8vo.
On the Use of Isobaric Curves. London, 1869. 8vo.
Tschermak (Gustav). Mineralogische Mittheilungen, Jahrgang. 1871. Heft 1.
8vo.
‘Turbiglio (Sébestien). L’Empire de la Logique, Essai @’un Nouveau Systeme de
Philosophie. Turin, 1870. 8vo.

Unger (C. R.). Thomas Saga Erkibyskups-Fortelling om Thomas Becket Erke-
biskop af Canterbury to Bearbeidelser Saint fragmenter af en Fredie.
Christiania, 1869. 8vo.

Vignoles (C. B.). Address on his Election as President of tke Institution of
Civil Engineers, Session 1869-70. London, 1870. 8vo.

Vigorniensis. An Historical Review of the Nature and Results of Vaccination
as unfolded in Dr Baron’s Life of Jenner. Cheltenham, 1869. 8vo.

Vogel (August). Uber die Entwicklung der Agriculturchemie. Munich, 1869. 4to.

Vollenhoven (S. C. Snellen van), Ph.D. Laatste Lijst van Nederlandsche Schil-
daleugelige Insecten (Insecta Coleoptera). Haarlem, 1870. 4to.

Wallis (S. T.). Discourse on the Life and Character of George Peabody. Balti-
more, 1870. 8vo.

831

DONORS.
The Author.

Ditto.
Ditto.

Ditto.

Ditto.
The Editor.
Ditto.
Ditto.
The Author.
Ditto.
Ditto.
The Authors.
The Author.
Ditto.

Ditto.
Ditto.

The Authors.

The Author.
Ditto.
Ditto.
Ditto.

Ditto.
Ditto.

Ditto.

Ditto.
Ditto.
Ditto.
Ditto.

The Peabody Insti-
tute.

S32 LIST OF DONATIONS. —

AUTHORS’ WORKS OR DONATIONS.
Waterhouse (Lieut. J.). Report on the Cartographic Applications of Photo-
graphy. Calcutta, 1870. vo.
Watson-Wemyss (Alexander), M.D. On the Construction of Hospitals for the
Sick and Hurt. Edinburgh, 1870. 8vo.
Wells (Walter). The Water Power of Maine. Augusta, 1869. 8vo.

Will (H.). Jahresbericht iiber die Fortschritte der Chemie, &e., fir 1867. Heft
2,3; 1868, Heft 1,2. Giessen. 8vo. ~

Wilson (Robert). The Screw Propeller, who Invented it? Glasgow, 1860. 8vo.

Wiltshire (Rev. Thos.). On the Chief Groups of the Cephalopoda. 1869.
8vo. ; :

Zittel (Carl Alfred). | Denschrift auf Christ. Erich Hermann von ~ Meyer.
Munich. to. ‘

DONORS.
The Author.

Ditto.
The Hydrographic
Survey. :
The Editor.
The Author. .
The Geologists’ As-
sociation, London.

The Author.

. (883: )

INDEX TO VOL. XXVI.

Acids and Bases, Heat Developed in the Combination of, 85.
Attman (Professor). The Genetic Succession of Zooids in the Hydroida, 97.
On the Homological Relations of the Celenterata, 459.
AnpreEws (Dr Tuomas). Heat Developed in the Combination of Acids and Bases, 85.
Atropia and Physostigma, An Experimental Research on the Antagonism between the Actions of, 529.

B
Balenoptera Sibbaldii, 197.

Batrour (Joun Hurron, M.D.). Remarks on the Ipecacuan Plant (Cephaélis Ipecacuanha, Rich.), with
Plates, 779.

Barnes (Dr THomas). On the Average Quantity of Rain in Carlisle and the Neighbourhood, 313.

Buackie (Professor). Scientific Method in the Interpretation of Popular Myths, with Special
References to Greek Mythology, 41.

On the Place and Power of Accent in Language, 269.

Broun (J. A.). On the Lunar Diurnal Variation of Magnetic Declination at Trevandrum, near the
Magnetic Equator, 735,

Brouncker’s Method, Extension of, 59.

Brown (Rev. Toomas). On the Old River Terraces of the Earn and Teith, viewed in Connection with
certain Proofs of the Antiquity of Man, 149.

C
Cophailis Ipecacuanha, Rich., 779.

Cetacea, Gravid Uterus and Arrangement of Feetal Membranes in the, 467.
Coelenterata, The Homological Relations of the, 459.
Crystals, Refracting, Spectra formed by the Passage of Polarised Light through, 177.

Deas (Francis). On Spectra formed by the Passage of Polarised Light through Refracting Crystals, 177.
Dewar (James). On the Oxidation of Products of Picoline, 189.
Dickson (Professor ALEXANDER). On some Abnormal Cones of Pinus Pinaster, 505.

E
Earn and Teith, Old River Terraces of, 149.
Elastic Solid, Forces Externally applied to an, 715.

VOL. XXVI. PART IV. 10 H

834. m, INDEX.

F
Forces, Reciprocal Figures, Frames, and Diagrams of le
Forces Externally applied to an Elastic Solid, 715.

Fraser (Dr Tuomas R.). An Experimental Research on the Antagonism between the Actions of
Physostigma and Atropia, 529.

Geometrical Mean Distance of Two Figures on a Plane, 729.

H
Hydroida, The Genetic Succession of Zooids in the, 97.

I
Ipecacuan Plant, 779.

Language, On the Place and Power of Accent in, 269.

L
Logarithms, Account of the New Table of, 521.

M‘Intosu (Dr W. C.). On some Points in the Structure of Tubifex, 253.

Magnetic Declination, Lunar Diurnal Variation of, 735.

Maxwe tt (J. Cuerk). Reciprocal Figures, Frames, and Diagrams of Forces, 1.
On the Geometrical Mean Distance of Two Figures on a Plane, 729.

Mesoplodon Sowerbyi, 759.

Motion of a Heavy Body, Additional Note on the, along the Circumference of a Circle, 449.
Mythology, Greek, 41.

Myths, Popular Scientific Method in the Interpretation of, 41.

ied
Prrricrew (Dr JAMEs Bext). On the Physiology of Wings, being an Analysis of the Movements by
which Flight is produced in the Insect, Bat, and Bird, 321.

Physostigma and Atropia, An Experimental Research on the Antagonism between the Actions of, 529.
Picoline, On the Oxidation of the Products of, 189.
Pinus Pinaster, On some Abnormal Cones of, 505.

Ruin, On the Average Quantity of, in Carlisle and the Neighbourhood, 313.

Ranxinz (W. J. Macquorn, C.E., LL.D.). Decomposition of Forces externally applied to an Elastic
Solid, 715.

RuruerrorD (Dr Wittiam). Influence of the Vagus upon the Vascular System, 107.

INDEX. 835

Sane (Epwarp). On the Extension of Brouncker’s Method to the Comparison of several Magnitudes,
59.
Additional Note on the Motion of a Heavy Body along the Circumference of a Circle, 449.
Account of the New Table of Logarithms to 200000, 521.
Spectra formed by the Passage of Polarised Light through Refracting Crystals, 177.

Tarr (Professor). Green's and other Allied Theorems, 69.

Teith and Earn, Old River Terraces of, 149.

Terraces, Old River, of the Earn and Teith, 149.

Theorems, Green’s and other Allied, 69.

Trevandrum, Lunar Diurnal Variation of Magnetic Dechnation at, 735.

Tubifex, On some Points in the Structure of, 253.

Turner (Professor). An Account of the Great Finner Whale (Balenoptera Sibbaldii) stranded at

Longniddry, 197. ;

On the Gravid Uterus and on the Arrangement of the Foetal Membranes in the Cetacea, 467.
Occurrence of Ziphius cavirostris in the Shetland Seas, 759.

V
Vayus, Influence of the, upon the Vascular System, 107.

W
Whale, Account of Great Finner, 197.
Sowerby’s, 759.
Wings, The Physiology of, 321.
Z

Ziphius cavirostris, 759.
Zooids, The Genetic Sucession of the, 97.

END OF VOLUME TWENTY-SIXTH.

PRINTED BY NEILL AND COMPANY, EDINBURGH.

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