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THE 


27OU RN A Ii 


OF 


ANATOMY AND PHYSIOLOGY 


CONDUCTED BY 


G. M. HUMPHRY, M.D. FE.RS. 
PROFESSOR OF ANATOMY IN THE UNIVERSITY OF CAMBRIDGE, 
HONORARY FELLOW OF DOWNING COLLEGE; 


AND 


WM. TURNER, MB. 


PROFESSOR OF ANATOMY IN THE UNIVERSITY OF EDINBURGH, 


Ly ise 


VOLUME VII. 


(SECOND SERIES, VOL. V1) 


MACMILLAN AND CO. 
Cambridge and London. 
’ F873. 


Cambridge: 


PRINTED BY ©. J. CLAY, M.A. 


AT THE UNIVERSITY PRESS. 


a 


CONTENTS. 


FIRST PART. NOVEMBER, 1872. 


Proressor SrruTHers, on the Cervical Vertebre and their Articulations 
in Fin-Whales . 

Dr P. D. Hanpysrpez, Notiee’ of Quadruple ‘Mamie: lhe Lower two 
Beery in two Adult Brothers 

Dr M. Warson, Contributions to the Anatomy of aie ean iéphant 
(Elephas Indicus), Part IJ. Urinary and Generative Organs 

Proressor Turner, Some Observations on the Dentition of the Nar- 
whal (Monodon Monoceros) 

Dr W. Arnsuie Hoxuis, Tissue MethBoliam, or ifs Artificial Thdhetion 
of Structural changes in Living Organisms . . 

Mr D. J. CunnineHam, Observations on the Distribution of some of 
the Nerves of the Head and Neck . : 5 

Mr A. H, Garrop, on Sphygmography . 

Dr T. Currrorp Auueutrr, The Effect of Hrorcive on the Bodily Tem- 
perature 

Proresson TURNER, Observations on the atuatiixe of the Hae 
Placenta 

Dr T. Lauper Denton! Retin of Digitalis « on the Blood- veaer 

Dr Tuomas R. Fraser, On the Kombé Arrow-Poison (Strophanthus 
Hispidus, D. C.) of Africa : . : 

Mr W. W. Waesrarre, Peculiar Honnabon of tite Leg and Foot 

Mr P. Burier Stoney, Effect of Stimuli on the Secretion of the Pa- 
rotid Gland : ‘ = 4 

Mr Waurer Rivineton, eee in tite heat Weis 

Dr J. Wickuam Lrece, Experiments as to the Causes of tie Presence 
of Bile Pigment in the Urine . 

Reviews and Notices of Books : : : 

Report on the Progress of Anatomy, by meas TURNER . 

Report on Physiology, by Dr T. Lauper Brunton, and Dr Davin Fer. 
RIER . , : 

Report on the Physioloiéal Aetion of Medicinal “ana ‘Poivenieen Sub. 
stances, by Dr Fraser 


PAGE 


56 


106 


lv CONTENTS. 


SECOND PART. JUNE, 1873. 


Dr J. Brake, on the Action of Inorganic Substances when introduced 
directly into the Blood 

Mr Atrrep Hurcuison Smee, on the Phy eal nears of the Coagnlation 
of the Blood 

Mr A. H. Garrop, on the Tray a beatae the Beouaanen of ae 
Pulse : 

Proressorn Turner, A Contribution is the tyaeral ies of the 
Greenland Shark 

Mr A. H. Garrop, on the Source ‘of Narva free. iy aneory 

Proressor W. H, Friowrr, Note on the Carpus of the Sloths 

Dr J. Barry Tuxe, on a Case of Hypertrophy of the Right Cerebral 
Hemisphere with Coexistent Atrophy of the Left Side of the Body 

Mr A. H. Garrop, on the Order Dinocerata 

Prorrssor Turner, on the so-called Prickle or Claw at the ord of fhe 


tail of the Lion and other Felines . . ‘ 
Proressor Turner, on an Edentulous Condition of ae Skull of ihe 
Grey Seal 


Mr James Dewar AND DR ae G. M°Kenpricr, on the Phasing 

Action of Light. No. I. 
” ” ” No. IL. 

Proressor RurHerrorp, Cause of the Bepetatad of ie Bales whisk 
follows Artificial or Voluntary Closure of the Nostrils in the Rab- 
bit. A Reply to certain Criticisms - 3 ‘% 

Dr J. J. Cuarues, Notes of Some Cases of Abnormal ame yaces of 
the Arteries of the Upper Extremity : 

Proressor Turner, on the Placentation of the Sloths . 

Dr Joun Curnow, Notes of Some Irregularities in Muscles and Nemes 

Proressor Dracumann, Case of Congenital Absence of the Quadriceps 
Extensor Cruris Muscle, translated by Dr J. W. Moorr 

Reviews and Notices of Books 

Report on the Progress of Anatomy, by Baan ee R 

Report on Physiology, by Prorrssor RurHEeRFoRD 

INDEX 


PAGE 
201 
210 
219 
233 
251 
255 


257 
267 


271 


273 


275 
278 


283 


300 
302 
304 


310 
312 
326 
339 
369 


Hournal of Anatomp and Phpstology. 


ON THE CERVICAL VERTEBRAE AND THEIR 
ARTICULATIONS IN FIN-WHALES. By JOHN 
SrruTHERS, M.D., Professor of Anatomy in the Unaver- 
sity of Aberdeen. (Plates I. and IL) 


THE great diversity presented by the cervical vertebrae in 
Whales gives a special interest to this part of Cetacean ana- 
tomy. The differences relate chiefly to the amount of ankylosis 
and the extent to which the transverse processes are developed. 
These differences have been a good deal relied on in endea- 
vouring to distinguish genera and species, and have been re- 
garded mostly from that point of view. Yet these various 
conditions of the cervical vertebrae do not follow the natural 
affinities within the order, nor can we say that the circum- 
stances which determine them are understood. Sufficient 
allowance has not always been made for difference of age and 
for individual variation, which the study of a series of specimens 
from the same species alone can teach us; and these vertebrae 
have been but little examined in the light of their relation to 
the soft parts, although without an examination from this 
point of view it is impossible to interpret the modifications 
which bones present. The following remarks are founded on 
the observation of a series of osteological specimens and on the 
results of the dissection of the soft parts. I shall first consider 
the neck in Fin-Whales, arranging my remarks in the following 
order. 


VO. Vil. 1 


2 PROFESSOR STRUTHERS. 


Fin-Whales examined. 


(A) In Great Fin-Whales : 


Transverse Processes viewed in relation to function. 

Ligaments of the Transverse Processes. 

Ligaments of the Spines, Laminae, and Articular pro- 
cesses. 

5. Articulations of the Bodies of the vertebrae. 

6. Articulations between the Axis, Atlas, and Occipital 
bone. 


go be 


The Cervical Vertebrae serially considered. 


7. Table of measurements. 

8. Bodies. 

9, Spinal Canal, Laminae, Spines. 
10. Articular Processes. 

~11. Inferior Transverse Processes. 

12. Superior Transverse Processes. 
13. The lateral Rings. 
14, Recognition of the five posterior vertebrae. 
15. The Axis. 
16. The Atlas. 


(B) In the Lesser Fin- Whale: 


17. Transverse Processes and their Ligaments. 
18. Bodies and their Fibro-Cartilages. 

19. Articulations of the Axis and Atlas. 

20. Occipito-Atlantal surfaces. 

21. Explanation of the drawings. 


1. Frin-WHALES EXAMINED.—The specimens to be con- 
sidered belong to the following Whales. 

(a2) Great Fin-Whale (Balaenoptera musculus, Ptero-ba- 
laena communis, Razorback) stranded alive near Wick, Caith- 
nessshire, June 1869. Male 65 or 66 feet in length. Mature 
or aged. Soft parts dissected’. 


1 T am indebted for information regarding this whale, and for the parts of it, 
to the kind exertions at Wick of Captain C. Cox, and Dr R. MacCalman, and 
afterwards, at Golspie, to Dr John Gunn, and Dr Soutar of Golspie. The 
carcase after being flensed near Wick, drifted south to Golspie. The informa- 
tion kindly furnished me by Dr Soutar and Captain Cox leaves no doubt that 
it was the same carcase, The total length according to one account was 72 to 
73 feet, but Captain Cox’s careful measurement, with a tape line, from the tip 
of the upper jaw straight along to the middle of the hinder edge of the tail, 
reduced this to 65 or 66 feet. I am indebted to the kindness of that gentleman 
for the following information. It was alive when stranded, and he saw it an 
hour after, quite fresh and uninjured, Skin on the under half plaited and 
white, or just a shade darker than white. The bristly part of the whalebone 
was white, and the solid part adjoining it had nearly the same colour, becoming 
slaty and then dark at the outer part. Length along the outside of the solid 


CERVICAL VERTEBRAE IN FIN-WHALES. 3 


(b) Great Fin-Whale (B. musculus), November 1871, at 
Stornoway, Lewis, Western Islands of Scotland. Male 60} feet 
in length. Mature or aged’. 

(c) Great Fin-Whale (B. musculus), June 1871, at Peter- 
head, Aberdeenshire. Male 64 feet in length. Not quite 
mature. Soft parts dissected *. 

(d) Atlas and Axis of another Great Fin-Whale (B. mus- 
culus), Norway, 1872. Mature’®. 


part of a plate sketched by him for me, 21 inches, of the longest bristles 10 
inches, of the shortest 5 inches. Tail-fin 15 feet or more. The lower jaw, now 
in my possession, is 14 feet 8 inches in length straight, along the outer side 15 
feet 10 inches; greatest depth of curve 2 feet; coronoid process high and curved, 
height along middle 7 to 8 inches, height of bone to tip of coronoid 22 inches. 
(The upper half of the right coronoid process is fractured, with ligamentous 
union, the fracture crossing obliquely and breaking off more of the outside than 
of the inside, with irregular bony surfaces at the fracture. How had this 
fracture been produced?) The pectoral fin had the usual lance-shape, and was 
8 feet 8 inches in length from the head of the humerus. These characters 
determine this Whale to have been B. musculus. As to age, even the distal 
epiphyses of the radius and ulna are united, an irregular and incomplete furrow 
3 inch from the end marking the place of union, The os magnum and unci- 
form have coalesced on the two surfaces but not deeply, and there is a small 
trapezoid bone concealed in the cartilage. This Whale was therefore mature or 
probably aged. The cervical vertebrae are large. 

1 This Whale was found dead about 14 miles off Stornoway, into which it 
was towed by the fishermen. I am indebted for obtaining the parts to my 
brother, Dr James Struthers of Leith, and to Mr Methuen of Leith, and for 
information regarding it to Mr A. Mackenzie and Dr Millar of Stornoway. 
Their kind attention and replies to my inquiries leave no doubt that it was a 
characteristic specimen of B. musculus. Length according to a public statement 
63 feet, according to Mr Mackenzie’s measurement, taken along the side, 60 feet 
5 inches. Length of pectoral fin 6 feet (taken I infer along the upper border); 
length of bones of left paddle from head of humerus to tip 7 feet 14 inches, but 
tip is malformed apparently from an old injury, somewhat shortening the 
paddle, Tail-fin from tip to tip 11 feet. Tail part of trunk thin, like a double- 
edged knife. Dorsal fin faleate and well marked. The usual furrows on the 
belly and sides. Belly and sides white, with dark patches of cuticle still 
adhering on the side. Whalebone, longest plates 30 inches; colour dark 
externally, cream-coloured on the internal bristly surface. Length of lower 
jaw 15 feet. As to age, all trace of the line of union of the epiphyses has 
disappeared on the cervical and three anterior dorsal vertebrae, in my possession, 
.The same of both epiphyses of the humerus. The pisiform is partially ossified. 
This whale was therefore mature, if not aged. 

2 J gave an account of this Whale, and of certain rudimentary structures 
which I found in it, in this Journal, for November 1871. I was indebted to the 
kind assistance of Dr Jamieson of Peterhead in obtaining the parts of this 
Whale and for his help when I was engaged on it there. It was a well-marked 
B. musculus. As to age, the epiphyses of the humerus were united, no traces of 
the line of union remaining. The epiphyses of the vertebrae are united, the 
traces of the line of union being variously visible on the following vertebrae 
which I have—on hinder ends of 6th and 7th cervical (slightly also on fore end of 
7th) and 1st dorsal; both ends of three succeeding dorsal, of a middle dorsal, and 
middle lumbar; while all trace has disappeared on an anterior caudal vertebra. 
Therefore, although longer than the Stornoway one, this Razorback was less 
mature than it, and the state of the cervical vertebrae agrees with this. 

3 These were among some of the bones of two Great Fin-Whales, brought 


1—2 


+ PROFESSOR STRUTHERS. 


(e) Lesser Fin-Whale (Balaenoptera rostrata, Pike-Whale) 
stranded alive at Aberdeen, July 1870. Young female, 143 
feet in length. Soft parts dissected”. 


I shall first consider the great Finners, distinguishing them 
by their localities (Wick, Stornoway, Peterhead, Norway), 
noticing afterwards the peculiarities of the young Pike-Whale. 


(A) In Great Fin-WHALES. 


2. TRANSVERSE PROCESSES IN THE GREAT FINNERS VIEWED 
IN RELATION TO FUNCTION. The most striking feature in these 
vertebrae is the enormous mass of the transverse processes, 
completing a great lateral foramen (see Fig. 4) so large that 
it is more than half the size of the body of the vertebra, amd 
is twice as capacious as the spinal canal; at once suggesting to 
the observer that a complete vertebrate segment is entitled to 
be regarded as presenting not merely two, but four rings. 
What is the function of these great rings? Contained within 
them is the rete mirabile representing the vertebral artery. This 
is a vast plexus. To realize the bulk of it, the block of verte- 
brae now empty should be turned up, the atlas resting on the 
ground. The series of rings, together with their connecting hga- 
ments, are then seen to form on each side of the vertebral bodies 
a great lateral canal, like a deep well. This canal is completely 
filled by the vascular rete, supported by connective tissue and 
some fat, except the small space occupied by the nerves which 
traverse it. “The canal is continued backwards, diminishing, 
along the anterior dorsal vertebrae by the rings, or spaces, be- 
tween the superior transverse processes and the necks of the 
ribs, or the ligaments representing the ribs. This really wonder- 
ful plexus occupying the lateral canal forms communications in 
various directions, downwards to the carotid region by the 


this year from Aalesund, Norway, for which I was indebted to the kindness 
of Messrs J. and G, Miller of this city. Although there was no history they are 
characteristic of B. musculus. Both lower jaws so like each other that they 
cannot be distinguished. Scapula and radius exactly the size of those of the 
Peterhead Razorback, Distal epiphysis of radius united, a furrow remaining. 
The state of the wings of the axis shows it to be more mature than the Peterhead 
specimen. 

1 This Whale was alive when stranded on our beach. The external charac- 
ters, measurements, and results of the dissection were noted. The skeleton 
and various portions of the soft parts are preserved. 


CERVICAL VERTEBRAE IN FIN-WHALES. 5) 


passages for the inferior division of the nerves, and also- by the 
sides of the vertebral bodies; upwards, by the passages for the 
superior division of the nerves; and inwards by the interverte- 
bral foramina, with the primary nerves, giving continuity with 
the rete within the spinal canal. The spinal canal and its 
lateral openings, the intervertebral foramina, are much more 
occupied by vascular rete, with its supporting connective tissue 
and some fat, than by spinal cord and nerves. While the 
spimal canal averages 6 to 7 inches in width by about 3 in 
height, I found in the Peterhead Razorback the tube of dura 
mater to have a diameter at the fore part of the atlas of about 
two inches, at the hinder edge of the atlas of about 14, and at 
the middle of the neck of about one inch. The rete fills the 
whole of the rest of the spinal canal, and is therefore many times 
bulkier than the spinal cord and its membranes. But great as 
this spinal canal rete is, it is small compared with the rete of 
each lateral canal. The intervertebral foramina, large enough to 
admit three or four fingers, are in like manner chiefly occupied 
by the rich communications between the two lateral and the 
spinal vascular networks. The nerves are comparatively small, 
the inferior division about the size of the little finger, the 
superior several times smaller. Having escaped by the inter- 
vertebral foramina, the nerves divide, the superior or dorsal 
divisions pass immediately up to the dorsal spaces; the inferior 
or ventral divisions sweep outwards across the upper part. of 
the lateral canal, surrounded by rete, as far as I could decide 
after it had been removed, and curving downwards a little, 
escape by the ventral spaces. 

Although the protection of this wonderful network may 
seem a function sufficient to account for the presence of these 
great rings, this view may be as far from satisfactory as it would 
be now to regard the double transverse process in man as a 
provision for the protection of the vertebral artery. In either 
ease the interpretation must be sought not in the idea of a 
protecting ring, but primarily in the locomotive system, in that 
of outstanding processes furnishing points of attachment for the 
muscles and ligaments, the spaces within which are, secondarily, 
more or less occupied by parts of the vascular system. 

The study of the relation of these processes to the soft parts 


6 PROFESSOR STRUTHERS. 


in great Fin-Whales has satisfied me that both upper and 
lower transverse processes, in their various degrees of develop- 
ment, may be interpreted by dividing them into three stages. 
(See Fig. 4, also Figs. 1, 2 and 3.) Lower transverse processes. 
(a) First or root stage, short, directed outwards and down- 
wards, alone present in some. Thick, smooth before and behind. 
Portions of rete mirabile lie here. (b) Second or tubercular 
stage. Directed outwards, extensive, being opposite the inner 
half or more of the ring; begins by an internal angular pro- 
tuberance or process, and terminates by an external protuber- 
ance; thinner at the upper edge where it bounds the ring, 
thick and rough below. Besides muscles, this stage attaches 
a series of strong intertransverse ligaments. (¢) Third, or 
nerve-groove stage, corresponding to the space left for the pas- 
sage of the inferior division of the spinal nerve, accompanied 
by communication of rete mirabile. Process at this stage turns 
upwards and outwards; groove is on anterior surface, directed 
obliquely downwards and outwards, broad enough to receive 
the hand laid flat. The very different thickness of these two 
stages at their lower part gives, especially where they meet, the 
twisted appearance which the processes present on their anterior 
surface. 

Upper Transverse Processes. (a) First, or nerve-groove stage, 
corresponding to the space for the passage of the upper division 
of the spinal nerve, accompanied by dorsal communication 
of rete mirabile. Is opposite the inner third of the ring, 
reaching from the articular process for three or four inches 
outwards, broad enough to receive the hand flat. Groove most 
marked on the posterior surface of the anterior vertebrae and 
on the anterior surface of the posterior vertebrae. (b) Second 
or tubercular stage, opposite the outer two-thirds of the ring ; 
beginning by a marked rough projection on the superior edge, 
and continuing rough outwards. It attaches a series of superior 
intertransverse ligaments. (¢c) Third, or terminal stage; situ- 
ated to the outer side of the ring, and curving downwards and 
inwards a little to unite with the inferior process. It forms the 
extreme part of the transverse process, is scarcely broader than 
the processes in adolescence, but in maturity forms a tabular 
expansion beyond the foramen. 


CERVICAL VERTEBRAE IN FIN-WHALES. t 


These characters will be recognised if one set of these ver- 
tebrae be piled in their natural relation on the table and 
another set arranged on the floor. Then if the observer, bear- 
ing in mind the relation of the human transverse processes to 
their soft parts, will take an articulated set of human cervical 
vertebrae and also a separate human cervical vertebra, and 
compare them in detail with those of the Rorqual, he will 
recognise an interesting correspondence to the stages above 
defined, small as the foramen is in man compared with the 
magnificent rings in the Rorqual. The anterior process presents 
first the root, springing from the body, next the “anterior 
tubercle,” and then comes the groove for the anterior division 
of the nerve. The successional and functional correspondence 
is evident though the proportions are different. So also with 
the posterior process. Springing from the pedicle there 1s, first, 
a stage across which the posterior division of the spinal nerve 
passes, corresponding to the nerve-groove stage in the Rorqual ; 
and beyond it the “posterior tubercle,” greatly extended in the 
Rorqual and expanded at the end in the terminal plate. 

3. LIGAMENTS OF THE TRANSVERSE PROCESSES.—These 
processes are very strongly knit together by three series of 
ligaments. (a) Inferior series, uniting the inferior processes ; 
and (6) those uniting the superior processes, divisible into 
superior portion above the rings, and external portion between 
the parts external to the rings. These are interrupted, or in- 
terosseous, ligaments, not longitudinally continuous, although 
those above and below, especially the latter, are seen on the 
surface. 

(a) Inferior Inter-transverse Ligaments’ (Inter-parapophy- 
sial) pass between the tubercular stages of the processes. 
A series of strong ligaments, broader than and as thick as the 
hand, increasing in breadth forwards and outwards, as the 
tubercular stages on the bones are seen to do, from the sixth 
vertebra to the third; from about 5 inches broad on the fifth 
vertebra to 8 inches broad on the third. They pass between 
the lower parts of the processes, which are correspondingly 

—rough, while the upper part of the processes is smooth. Be- 
tween the third and the axis the ligament is larger and 


1 The parts to which these ligaments are attached are seen in Figs. 1 and 2, 


8 PROFESSOR STRUTHERS. 


changes its direction, passing very obliquely inwards and 
forwards to the process and body of the axis; forming a great 
mass of ligament, 8 inches broad and 1 inch thick. The at- 
tachments of this great sloping ligament account for the breadth 
and prominence of especially the outer part of the tubercular 
stage of the 3rd vertebra, and for the roughness on the inferior 
process of the axis opposite the foramen (tubercular stage, 
but not defined as on the vertebrae behind) and inwards to 
where it joins the body. 

On either side of this series of ligaments spaces are left. 
The internal spaces, between the ligaments and the bodies of 
the vertebrae, opposite the root stages of the processes, admitting 
one or two fingers, but in the putrid state easily enlarged as the 
ligament is thinner here. The rete 1s seen bulging against 
this thinner part, and, as far as one could judge after the ex- 
ternal parts had been cleared away, appears to have sent com- 
munications through the passages. The external spaces are 
the nerve-passages, corresponding to the third stage of the 
processes. They are about 3 inches in breadth and about an 
inch longitudinally, diminishing outwards as the processes con- 
verge, easily admitting three or four fingers flat. The one be- 
tween the axis and third vertebra is smaller than the others (24 
inches). Their inner boundary is some way internal to the outer 
part of the tubercular stage, owing to the obliquity of the 
eroove and the position of the ligaments. Their outer boundary, 
formed by the lowest part of the hgaments which connect the 
external part of the processes, is about an inch internal to the 
outer end of the foramina. Besides the nerve, which is not 
larger than the little finger, they are occupied by communica- 
tions of the rete. 

(b) Superior Inter-transverse Ligaments and Nerve-spaces. 
The dorsal nerve-spaces are between the ligaments externally 
and the zygomal processes and their ligaments internally. Be- 
tween the 6th and 7th vertebrae their breadth is 2} inches, 
increasing forwards to a breadth of 4 inches between the third 
and the axis; but the spaces of the posterior are wider. The 
spaces have a compressed triangular form, tapering outwards, 
and admit four fingers easily. Besides the nerves, which are 
small in proportion to the passages (though the disproportion 


CERVICAL VERTEBRAE IN FIN-WHALES. 9 


is not so great as, for instance, in the posterior foramina of the 
human sacrum), they are occupied by rete, sending communi- 
cations through here, as far as one could judge after the ex- 
ternal parts were cleaned away. The superior inter-transverse 
ligaments (inter-diapophysial) may be conveniently divided 
into superior and external portions. The superior inter-trans- 
verse ligaments, commencing a hand’s breadth from the zygomal 
processes, occupy the processes opposite the outer 3 of the 
ring (tubercular stage). They are attached to about the upper 
half of the processes, which is rough accordingly, and bevelled 
so as to turn this part of the surface upwards. Where they 
commence, at the outer end of the nerve-spaces, is well marked 
on the bones’. The bundles as seen on the dorsal surface are 
both longitudinal and oblique, while the deeper fibres, more 
interosseous in position, are oblique. The external inter-trans- 
verse ligaments are continuous with the last, corresponding to 
the fact that the outer part of the transverse processes is a 
continuation of the upper process. They are the strongest 
ligaments of the processes, and so placed in between the pro- 
cesses, here very close together, that it is only after their division 
that their extent and attachments can be distinctly made out. 
Stated generally, these enormous ligaments occupy about the 
upper and outer half of the breadth of the more or less ex- 
panded part of the processes external to the rings, but coming 
down far enough to form the outer boundary of the ventral 
nerve-passages, the inner half being occupied by loose connective 
tissue lying on the reticular periosteum. Corresponding rough 
and smoother parts are seen on the surfaces of the bones. Be- 
tween the 7th and 6th processes I found a synovial cavity, in 
both of the great Finners dissected, nearly two inches in dia- 
meter, situated on the lower part of the plate, presenting a peri- 
osteal surface on the convex 7th process and a reticular fibrous 
or cushioned surface on the concave 6th process, surrounded by 
a capsular ligament. The 6th and 5th were very close, but there 
was no cavity proper between them (or between any of the other 
transverse processes). The ligament attached to the upper, 


? A smaller and variable rough mark is seen about the middle of the upper 
edge of the nerve-groove stage on the three posterior vertebrae. It happens to be 
strongly developed on the left side in the vertebra represented in Fig. 4. 


10 PROFESSOR STRUTHERS. 


outer, and under parts of the plates was 1 to 14 inch thick, 
and about the same between the 5th and 4th, and between the 
4th and 3rd. 
The ligaments attached to the hinder surface of the great 
wing-like transverse process of the axis are of enormous strength. 
It is not very easy to separate any of these vertebrae, but to 
separate the axis and third is a matter both of art and strength. 
With a forcible sawing motion the long slicing knife, with 
barely room to work, at length makes its way through the dense 
mass. Besides the ligament from the third, the axis has an 
external compound ligamentous mass from the converged tips 
of the 4th, 5th, and 6th processes. (1) That from the third 
passes downwards and outwards to be attached to the wing of 
the axis. Taking it and the dorsal interosseous ligament as 
one, the attachment to the axis is over an extent of 12 inches 
in breadth by two in thickness, following the curve of the 
process of the 3rd, its position on the axis being external to the 
middle of the broad plate and along the superior process above 
the outer half of the rmg. The deeper part of this ligament is 
disposed differently, its much longer fibres converging forwards 
and inwards to near the ring of the axis; thus lining this part 
of the lateral canal, which receives a funnel shape here from 
the comparatively small size of the ring of the axis, while the 
two parts of the ligament receive different obliquities. (2) The 
external ligamentous mass is a compound ligament proceeding 
from the converged tips of the four vertebrae behind the axis 
outwards and forwards to be attached, for about five inches, 
to the most external and inferior part of the wing. This is the 
part of the great wing which forms the extreme triangular 
projection, beyond the rest of the outer edge of the wing. 
Whatever may be the meaning of this convergence of the 
transverse processes in the Rorquals, we see not only that, 
the bones converge here, forming the apex of the pyramidal 
framework, but that the plates in which they terminate, and 
by which they come almost in contact, are firmly tied together 
by these strong ligaments. Viewing the ligaments of the trans- 
verse processes as a whole, while, locally, they enable the pro- 
cesses to strengthen each other as parts for muscular resistance, 
the firm binding together of the whole must co-operate with 


CERVICAL VERTEBRAE IN FIN-WHALES. 11 


the firm binding of the bodies in giving strength to the unan- 
kylosed neck. The convergence must impede especially lateral 
motion, and the ligaments between the converged plates must 
act not merely as binders but as interposed cushions. Viewed 
in relation to the contents of the canal, if this can be considered 
a function, the ligaments complete the walls of the canal all 
round, except at the dorsal and ventral nerve-passages. 

I regret that the circumstances were such as to prevent me 
examining the muscles attached to these processes. ‘The neck 
having been roughly cleaned, I could only recognise the remains 
of various strong tendons. One can hardly doubt that the pri- 
mary function of these processes is to furnish points for mus- 
cular attachment, the more essential parts being the superior 
processes and the tubercular stage of the inferior, while their 
ligamentous, and especially their nerve and blood-vessel rela- 
tions, are subsequent. We must look to the muscles for an 
explanation of the thick single transverse process of the atlas ; 
of the vast and strong-rooted wing of the axis, and its backward 
slope ; to the muscles, and perhaps to the mode of-attachment of 
the first rib, for an explanation of the absence of a bony inferior 
transverse process to the 7th vertebra; and to the adaptation 
of the processes which support the anterior ribs, for an explana- 
tion of the size and forward slope of the superior transverse 
process of the 7th vertebra. Between these two great converg- 
ing and dominating processes, the 2nd and 7th, the outer parts 
of the intervening transverse processes are packed as best they 
may in the available space; the 5th is level and the most pro- 
jecting, the 6th must slope forwards, the 3rd and 4th must 
slope backwards, while their imner parts are adapted to muscu- 
lar attachments. If the part where the nerve crosses is ossified, 
then a ring results. The processes are joined by strong liga- 
ments, giving the various functional results above indicated ; 
and in the space thus left by the adaptations to the locomo- 
tive functions, part of the vascular system has been enclosed or 
has been developed. 

4, LIGAMENTS OF THE SPINES, LAMINAE, AND ARTICULAR 
PROcESSES.—In the specimen (Peterhead) in which cervical spi- 
nous processes are present, there was a supra-spinous ligament ; 
and in between the spines pretty strong inter-spinous ligaments, 


13 PROFESSOR STRUTHERS. 


about half an inch in thickness. The laminae are connected on 
both aspects by inter-laminar ligaments. As seen on the dor- 
sal aspect, the ligament passes from the hinder edge (the thick 
and overlapping edge) of the lamina to the dorsal surface of the 
lamina behind.’ From within the canal a thinner ligament is 
seen passing from the anterior edge of the lamina forwards to 
the inner surface of the lamina in front. The fibres of the two 
seem continuous. The latter corresponds in attachment to the 
ligamentum subflavum of man, but it is white. Also within the 
canal there was, at least between the axis and 3rd (the only 
two vertebrae which had not then been separated), a strap-like 
interlaminar ligament, # inch broad and { inch thick, separated 
from its fellow by a distance of an inch. IJnter-zygomal liga- 
ments continuous internally with the interlaminar ligaments 
pass from process to process, covering the processes, and ex- 
ternal to them form a considerable longitudinal ligament. 

5. ARTICULATIONS BETWEEN THE BODIES OF THE VERTE- 
BRAE.—The inferior and superior common ligaments of the 
bodies are each about an inch in breadth, and therefore not 
ereat ligaments for such bodies. The corresponding marks on 
the bones are well seen above within the canal, while below 
there is rather a narrow ridge not running the whole length of 
the body. There is not much difference in the fibro-cartilages 
between the different cervical vertebrae, though they may di- 
minish a little forwards. Their apparent thickness (length) on 
the surface is deceptive owing to the bevelling of the edges of 
the vertebrae, this being exaggerated at the middle line where 
the bevelling goes so far as to form a notch. This median notch 
is sometimes nearly filled up by ossification. It is strongly 
marked on the hinder edge of the axis, while the rest of this 
edge is less bevelled than the edges of the other vertebrae. 
The thickness of the fibro-cartilages on the surface is 1} inch, 
the real thickness deeply between the bones is less than half 
this, being about 4 inch; perhaps slightly less between the 4th 
and 3rd, and 4 inch less between the 3rd and axis. 

On section, the fibro-cartilages are seen to correspond very 
closely in all the spaces, except between the 8rd and axis, where 
there is a little more of the ligamentous and less of the pulpy 
part. With a body-surface averaging 12 inches in breadth and 


CERVICAL VERTEBRAE IN FIN-WHALES. 13 


8 in height, the ligamentous or capsular part has an average 
depth of 14 inch, dipping in to a depth of 2 inches at the mid- 
dle line above and below, giving the pulp cavity a slightly 
figure of 8 form. The print of this attachment is well seen on 
the bones, with parallel limes marking the attachment of the 
concentric capsular ligaments. The pulp cavity averages 9 
inches in breadth, 4 to 5 in height. An interesting transition 
is seen from the fibrous to the pulpy, in the form of a floating 
wedge, projecting + to # inch into the pulp from the fibrous 
part, and ending in fringes which grade into a tenacious pulp. 
The bodies where they form the walls of the pulp-cavity are 
lined by a thin layer of pearly cartilage; the anterior surface of 
the vertebra slightly convex, the posterior slightly concave. 
The pulp itself is white and glairy, in some parts tenacious 
enough to lift with the fingers. It had in part an oily appear- 
ance, like soft blubber. Under the microscope it showed chiefly 
groups of cartilage cells of various sizes, bundles of wavy con- 
nective tissue, some free oil-globules, fatty crystals, and crystals 
resembling those of phosphates, all immersed in a fluid gluti- 
nous medium. 

The proportion of the fibrous part to the pulpy part, though 
quite mammalian in contrast with the proportions in the fish, is 
much less than in man, and still less than in long-necked quad- 
rupeds; but a circular ligament in reality over 30 inches in 
length and 14 in breadth is a structure capable of great resist- 
ance. The amount of motion permitted between the cervical 
vertebrae, either at the bodies or transverse processes, before the 
fibro-cartilages have become softened by putrefaction is very 
limited. The transverse processes may be made to move a 
little on each other, giving a slight rotatory motion of the ver- 
tebrae, and the bodies may be moved on each other a little in 
any direction. Lateral motion is the least, as it is at once ar- 
rested by the transverse processes coming in contact; rotatory 
motion comes next, and vertical motion is the greatest, but 
is very little. With the block of the five middle vertebrae still 
attached (including therefore the united motion of four fibro- 
cartilages), I could give one end of the block a range of only 
2 inch of vertical motion. It is then firmly and softly checked. 
On then slicing through the connections between the transverse 


14 PROFESSOR STRUTHERS. 


processes, there was no increase in the extent of the motions at 
the fibro-cartilages, except a very little in the lateral direction. 
Although, therefore, the intertransverse ligaments must greatly 
assist to strengthen the neck, they do not limit the extent of 
movement at the bodies. Viewing the vertebrae of the Fin- 
whale’s neck as a whole, one could scarcely conceive of any 
parts more thoroughly bound together than they are, both at 
the bodies and the transverse processes, into a great fibro- 
osseous unyielding lump. What then is the functional adapta- 
tion? It cannot be strength, as the vertebrae in the ankylosed 
neck are still more firmly united. When vertebrae are separate 
a strong binding medium is, of course, rendered necessary, but 
it would appear that the primary functional adaptation is im the 
soft cushioning, and that there must be some difference in the 
actions of the head in the Finners, as compared with the Right- 
whales, to account for ankylosis not taking place’. 

6. ARTICULATIONS BETWEEN THE AXIS, ATLAS, AND Oc- 
CIPITAL BoNE.—Besides the ligaments between the spines 
above, and the continuation of the inferior common ligament 
of the bodies below, there are strong capsular ligaments en- 
closing the articular surfaces, and certain ternal or central 
ligaments. The capsular ligament between the atlas and occiput 
(condylo-capsular) are strong ligaments, about } inch in thick- 
ness, entirely surrounding each articular surface. Coming in 
contact in the middle, they form a septum between the two 
joints, where a median groove marking their attachment is 
seen on the atlas from its canal to its lower edge. This septum 
has a corresponding attachment on the occipital bone, between 
the lower ends of the occipital condyles, with corresponding 
mark on the bones, which but for this would here run into 
each other. The septum was imperfect and stringy, but this 


1 Ankylosis had taken place between the bodies of two of the vertebrae, the 
3rd and 4th, in the Stornoway specimen, all the other vertebrae in these Fin- 
whales being free. The ankylosis had been broken up by foree before they 
reached me. A plate of the 3rd had torn off, remaining with the 4th, and 
exposing the spongy tissue of the 3rd. The ankylosis occupies less than the 
middle half of the body-surfaces, about 5 inches across and 34 high, being the 
central parts of the pulp surfaces. The impression at the attachment of the 
fibrous part of the fibro-cartilage, and other markings on the unankylosed part of 
the surfaces, are as usual. Notwithstanding the ankylosis, the articular sur- 
faces between the articular processes of these two vertebrae are not only as 
well but better marked than those between the 4th and 5th. 


CERVICAL VERTEBRAE IN FIN-WHALES. 15 


may have been from the giving way of parts. In all of the 
four atlases this median groove is well marked, of varying 
breadth widening below into a triangular space into which 
the inferior common ligament dips. The articular surfaces 
between the atlas and axis, on the contrary, run into each other 
inferiorly, forming one great horse-shoe articular surface on 
both vertebrae, more or less notched at the middle line in- 
feriorly’. The cartilage on the articular surface of the front 
of the axis, and on both aspects of the atlas, was about 4 inch 
in thickness. In the Peterhead specimen, and there was an 
appearance of the same in the Wick specimen, the cartilage 
along the rim of the cups of the atlas assumed a fibrous ap- 
pearance, and gave off fringes, half an inch to an inch in length, 
like a fringed marginal fibro-cartilage, the cartilaginous surface 
being again smooth for half an inch to an inch beyond the 
margin, until the attachment of the capsular ligament was 
reached. 

The central ligaments, made out with considerable difficulty, 
are—(a) The transverse ligament of the atlas; (6) pair of 
check ligaments, interosseous between axis and atlas, and two 
longitudinal ligaments ; (c) an inferior, the ligamentum suspen- 
sorium dentis; and (d) a superior, a prolongation of the superior 
common ligament of the bodies. 

(a) Transverse Ligament (see Fig. 5). This great ligament 
is attached as in man, but has no contact with the odontoid 
process, and is-flattened in the opposite direction. It divides 
the canal into two parts, the upper and larger part for the 
spinal canal, while the lower, narrower and pointed inferiorly, 
is occupied by ligaments. This explains the peculiar form of 
the canal of the atlas, the most constricted part corresponding 


1 In the Wick specimen, however, there is an appearance (see Fig. 5) of a 
median separation, by a narrow furrow in the lower part, continued along the 
upper half as a slight irregular depression, still less marked on the axis. In 
dissecting this joint, as the cartilage had begun to peel off, I could not be certain 
that it was continuous across the middle line. In the atlas of a fifth great 
Finner in my possession (afterwards referred to), a larger one than any of the 
others, and presenting a very wide median groove between the anterior articu- 
lating surfaces, there is no trace of median separation of the two joints on the 
posterior surface. In the axis of a fifth (young) great Finner (afterwards re- 
ferred to) the horse-shoe surface appears as if divided by a faint median eleva- 
tion. These appearances I believe to be deceptive, not implying that the 
cartilage and synovial membrane were not continued across in these as in the 
other specimens. 


16 PROFESSOR STRUTHERS. 


to where the attachment of the ligament begins superiorly. It 
is a thick flat ligament, measuring when fresh 2 to 23 inches 
from border to border, half an inch to an inch in thickness, as 
thick and nearly as broad as three fingers laid flat; its breadth 
is about 24 inches, corresponding to the width of this part of 
the foramen. Its upper border is concave, bounding the spinal 
canal, its lower border bounding the canal through which the 
suspensory ligament passes. Its attachment to the atlas is as 
far back as possible on the wall of the canal, but there is still 
left a space between it and the odontoid large enough to admit 
the fingers flat, though its upper edge comes close to the 
summit of the odontoid process, when, as in the Peterhead 
specimen, that process is better marked. It is composed of 
dense transversely arranged fibres, passes straight across, and 
is a tight strong resisting structure. Functionally viewed, this 
ligament is here adapted to serve as a great fibrous beam, pre- 
senting its posterior surface as a continuation of the area for 
the attachment of the check ligaments, while its edges afford 
attachment to part of the longitudinal ligaments. 

(b) Check Ligaments (Alto-odontoid). This pair of liga- 
ments forms the chief retaining structure between the axis and 
atlas; attached behind to the side of the odontoid area, in front 
to the crescentic depression on the atlas internal to its articular 
surface. An examination of the bony surfaces will enable their 
attachments to be readily understood. On the axis, bounded 
by the articular surfaces at the sides and below, and by the 
spinal canal above, is a large non-articular area, from 5 to 6 
inches across and about 4 inches vertically. The whole of 
this area forms a gentle elevation, rising below the middle 
into a low conical eminence. While the whole may be 
termed odontoid area, the latter may be distinguished as the 
odontoid process. The summit of the process is situated at the 
junction of the lower and middle thirds of the area. To this 
process, and to the rough slope below it, is attached the liga- 
mentum suspensorium ; while, laterally, along the base of the 
process and the side of the area, is attached this great check 
ligament, the convexity of its semilunar attachment not going 
to the outer part of the area, and its horns approaching those 
of its fellow above and below. On the posterior aspect of the 


CERVICAL VERTEBRAE IN FIN-WHALES. 17 


atlas the attachment is well marked, as a depressed crescentic 
surface between the articular surface and the edge of the canal 
(see Fig. 5), varying from 1 to 1} inch in breadth, tapering 
upwards and downwards. The check lgament, itself cres- 
centic in section, is attached to this rough crescentic surface ; 
its lower part converges to join its fellow at the middle line 
below the canal, and is attached also to the neighbouring 
part of the wall of the canal; its upper part reaches inwards 
upon the transverse ligament, to which it has a true and ex- 
tensive attachment. The check ligament is over 3 inches in 
height, its greatest thickness about 1} inch. The fibres are 
longitudinal, direction forwards and a little outwards, length 
half an inch to an inch. The shortness, size, and interosseous 
position of this great ligament, explain why the motions be- 
tween the atlas and axis are so very limited. The dissection of 
these check ligaments in the great Finners is very difficult. 
When the atlas is separated forcibly from the axis, they tear 
off from the axis, leaving the odontoid area bare and a few 
tufts attached to the process; and the lower part of the canal 
of the atlas is seen to be blocked by a dense pre-odontoid fibrous 
mass, below the middle of which is a conical recess inta which 
the tip of the odontoid had sunk, the recess being now the 
hollow base of the suspensory ligament. 

(c) Ligamentum suspensorium dentis (Occipito-odontoid), 
This ligament, about the size of a thumb, arises from the tip 
and lower slope of the odontoid, passes through the aperture 
below the transverse ligament, shows itself free for about 
3 inches between the atlas and occiput, is here compressed 
laterally (vertically 1 inch, transversely 4 to 8 inch), and passes 
forwards to be inserted into the triangular fossa between the 
occipital condyles. The appearance of the lower part of this 
ligament being attached to the atlas, on its way forward, is due 
to its fibres being continuous with those of the check ligaments 
where they meet in the middle line below. There is the 
appearance as if it afterwards received an accession of fibres 
at the sides from the atlas and above from the lower edge of 
the transverse ligament, but this is not easily determined as 
its circumference is not isolated as it passes through the aper- 


ture below the transverse ligament. The occipital condyles 


VOL. VII. 2 


18 PROFESSOR STRUTHERS. 


are for about 5 inches separated only by a deep narrow fissure, 
attaching the septum formed by the condylo-capsular ligaments, 
and above this they diverge upwards, leaving a triangular 
fossa 3 inches in length, ? inch deep, and 1} to 2 inches wide 
at the base where it is bounded by the foramen magnum. In 
this triangular fossa the suspensory ligament and the superior 
longitudinal ligament have a continuous insertion. 

(d) The Superior longitudinal ligament (Occipito-axoid), 
a prolongation of the superior common ligament of the bodies 
of the vertebrae, a pretty strong flat ligament, passes from the 
upper surface of the body of the axis forward, partly joming 
the upper part of the transverse ligament but mainly con- 
tinued on above that ligament, and now expanding passes to 
be inserted into the intercondyloid fossa with the suspensory 
ligament, to which it has previously adhered. Tracing these 
two ligaments backwards, they are separated by the vertically 
extended transverse ligament, to the edges of which they 
partly adhere, the lower passing to the odontoid, while the 
upper passes to the axis proper. The soft parts which occupy 
the middle line between the occipito-atlantal joints are the fol- 
lowing, from the lower edge of the body of the atlas up to the 
spinal canal. 1, Median septum for 5 inches, formed by the 
close-together condylo-capsular ligaments. 2, Ligamentum sus- 
pensorium. 3, Mass of rete mirabile, in the triangular space 
between this and the next ligament and between the now 
diverging capsular ligaments on each side, giving this part a 
bulky appearance. 4, Superior longitudinal ligament. 

Viewing these ligaments homologically and functionally, 
the transverse ligament is recognised, fully developed, and 
adapted to assist in binding the axis to the atlas by the at- 
tachment which it affords to other ligaments ; the superior longi- 
tudinal ligament is as in man, and partly adheres, as it also does 
in man, to the transverse ligament; the inferior longitudinal 
ligament is a true ligamentum suspensorium dentis, connecting 
the odontoid and occipital centra; the check ligaments corre- 
spond to the lower check ligaments of man greatly developed ; 
while the upper, or occipito-odontoid, check ligaments are sup- 
pressed, or converged and united with the suspensory liga- 
ment. 


CERVICAL VERTEBRAE IN FIN-WHALES. 19 


THE CERVICAL VERTEBRAE SERIALLY CONSIDERED. 


I have examined these vertebrae closely with the view on 
the one hand of determining their more essential characters, 
and on the other hand of observing the differences which they 
present according to age or individual variation. The series of 
specimens were compared when arranged in position, as seen in 
Figs. 1, 2 and 3, and when arranged separately, and it will be 
observed when the remarks refer to them specially when in 
position and when separate. It is to be borne in mind that of 
the three whales to which these observations chiefly refer, 
the Peterhead one was scarcely mature, the other two being 
mature or aged. From the much greater size of its vertebrae 
the Wick whale might have been supposed to have been a 
much larger animal, but its length was only one or two feet 
greater than that of the Peterhead whale (65 to 66 against 64), 
while the mature Stornoway whale was only 614 feet. Hven 
when the length is ascertained by careful measurement with a 
tape line, there may be different results. Had I gone by the 
measurement along the back, the length of the Peterhead 
whale would have been stated as 68 instead of 64 feet; but 
the measurement ought always to be straight along the side, 
giving the length of the ground over which the animal extends. 
But making due allowance for this source of variation in the 
statements as to length, no reason appears why full-grown 
whales of the same species should not vary as well as men, 
some inches in the one being the equivalent of some feet in the 
other. Although variation in length in man may depend chiefly 
on the lower limbs, the enormous elongation of the caudal part 
of the trunk, the locomotive equivalent, in whales gives ample 
scope for trunk variation. 

7. MEASUREMENTS.—The measurements given in the table 
were made with care, and may be useful for consultation. They 
show points of correspondence and variation besides those re- 
ferred to in the remarks. The transverse measurement of the 
transverse processes (No. 2) is taken from the middle of the side 
of the body; taken from the anterior edge of the body would 
give nearly half an inch less. The length of the spinous pro- 


2—2 


20 


TABLE OF MEASUREMENTS OF VERTEBRAE OF FOU 


1. Greatest width 


2. Width of transverse 
processes. 


3. Breadth of plate be- 
yond ring. 


4. Transverse diameter 
of rings. 


5. Greatest height of 
rings, vertically. 


7. Length of body. 


. Width of body : 
At anterior surface. 
At middle of side. 


. Height of body. 


. Height of spinal canal. 


. Width of spinal canal. 


canal, 


cess. 


tebra. 


. Weight. 


Pounds. 
Ounces. 


Be Pe) ae) ee 


6. Circumference of rings. = 


. Circumference of spinal 
. Length of spinous pro- 


. Greatest height of ver- 


vs) 
top 


aa 
S| 5 
a 
3|¢ 
3) ° 
am ea) 
26 | 28 
63 | 73 
43 | 4 
lic 
ce 
4} | 43 
ice 
I Nig 
143 |14} 
19 |19 


i113 | 8 


PROFESSOR STRUTHERS. 


4 
43 


18 


i) 


31 | 33 
33 

153 |17 
16 |, 
33 | 32 
31 | 28 


23 


54 


2 15 
8. 16s 
8 | 8 


CERVICAL VERTEBRAE IN FIN-WHALES. 21 


GREAT FIN-WHALES OF SAME SPECIES. (B. Muscutus.) 


4th. 5th, 6th. 7th. 1st. Dorsal. 


im bo 
Wl col 


ist) 
is*) 
loo 
(JU) 
oar 


12 | 113/193 |/113 [112 
11 | 108/113 111 [108 


16 [173 {178 ||16 163 |18 


1 
Bil |Z Wl a 


22 PROFESSOR STRUTHERS. 


cesses (No. 13) is taken from within the canal, afterwards deduct- 
ing the thickness of the lamina; the difficulty of fixing on the 
point of commencement of the spine rendering all other methods 
liable to fallacy. The distinction of right and left in the lateral 
measurements (the right always given first) shows the frequent 
a-symmetry, and that, when there is a difference on the two 
sides, there is no general preference of one side more than the 
other, as will appear farther from the remarks on parts not 
noticed in the table. 

8. Bopres.—The bodies diminish in length (thickness) for- 
wards, from the 7th to the 3rd. They gain in width, and lose 
in height, forwards, from the 6th to the 3rd. Longitudinally, 
they are grooved all round, except where the groove is inter- 
rupted by the median ridges and filled up by the roots of the 
inferior transverse processes. If the measurement of the width 
is taken at the middle of the groove it will generally give from 
¢ to 1 inch less than if taken at the edges. The bodies are 
marked below, above, and on the sides by vascular foramina, 
largest below and smallest on the sides, more or less arranged 
in two rows, especially on the sides, one row going before the 
other behind the roots of the transverse processes. The breadth 
of the articular surfaces greatly exceeds the height, the pro- 
portion averaging that of 3 to 2 (see Fig. 4). The form of the 
surfaces varies in the three series, suggesting the semilunar in 
the Peterhead, the square in the Stornoway, and the oval form 
in the Wick series. The fundamentally square form arises from 
the projections where the inferior transverse processes and pedi- 
cles spring, forming lower and upper lateral angles. By the 
former being placed farther out, a greater breadth is given to 
the lower than to the upper part of the body, but the broadest 
part is generally a little below the middle. If the body is con- 
cave to the spinal canal, and the pedicles more internal, the 
semilunar form is given; if it rises up so as to be convex to the 
spinal canal, and the lower median ridge be also broadly deve- 
loped, as in the Wick vertebrae, the form is changed to the 
oval. The posterior surfaces are, transversely, a little concave 
from the axis to the 5th, decreasing backwards ; while the 6th 
is flat, the 7th and Ist dorsal a little convex. The anterior 
surfaces are convex transversely (and also a little convex verti- 


CERVICAL VERTEBRAE IN FIN-WHALES. 23 


cally), diminishing backwards, but distinct on the 6th and 7th. 
The striated ring, about 14 inch broad, for the attachment of 
the capsular part of the fibro-cartilages, is well marked for an 
inch in breadth, and is convex, owing chiefly to the bevelling at 
the edges of the bodies; the inner half inch is concave and the 
concentric limes are less distinct. The contained somewhat 
figure-of-8 space, corresponding to the pulp, presents generally 
a central elevation, with a large shallow depression on each side 
bounded by a raised enclosure (see Fig. 4). The central eleva- 
tion is seen on both surfaces, best marked perhaps on the front 
surfaces especially of the anterior vertebrae, while on the hinder 
surfaces it is better marked on the posterior vertebrae. Hence 
the bodies are thickest in the centre, where the measurements 
given in the table were taken. They measure 1 inch less at the 
ring just within the bevelling, and about 3 inch less at the ex- 
treme bevelled margins. 

9. SPINAL CANAL, LAMINAE, ANAPOPHYSES, SPINOUS Pro- 
CEssES.—In my account of the Peterhead Razorback’, I alluded 
to the high arches and well marked spines in it, compared with 
the low arches and scarcely present spines in the Wick speci- 
men. Also in the latter, to the extreme thinness of the ante- 
rior border of the laminae, indicating atrophy. In both these 
respects the Stornoway specimen is intermediate, although in 
regard to shortness of spinous processes it more resembles the 
Wick specimen. The difference in the shape of the spinal 
canal is great, being triangular in the Peterhead and semi- 
lunar in the Wick series. The semilunar form is owing partly 
to the raising of the floor of the canal, the bodies (at their 
edges and median ridge) having become convex instead of 
concave to the canal; partly to the neural arches being very 
low. But the capacity of the canal, as the measurements of 
circumference show, is not less, being extended laterally by 
the pedicles being placed farther out on the bodies. On the 
four posterior vertebrae, while in the Peterhead specimen the 
height is more than half the breadth, in the Wick specimen 
the height averages less than a third of the breadth. The one 
is fully 14 inch greater in height, while the other is 1} to 1} 
greater in breadth and 14 to 2 inches greater in circumference. 


1 In this Journal, Noy. 1871, p. 120. 


24 ' PROFESSOR STRUTHERS. 


The lowness of the arch is not owing to shortening of the 
pedicles, but to the laminae turning across with very little rise, 
and forming no angle at the middle. The semilunar form is 
less marked on the 8rd, the body being a little concave and the 
arch a little raised at the middle; behind it all the bodies are 
convex, increasingly so backwards; the arches rise a little at 
the 6th and 7th, being lowest at the 4th and 5th; at the 5th 
the curves of the body and of the arch are almost parallel. 
In the Peterhead series all the bodies are concave and all the 
arches triangular. 

The laminae of the posterior vertebrae do not overlap, 
leaving more or less open, in the Wick specimen the two, in 
the Peterhead three, and in the Stornoway specimen the four 
posterior spaces; most open in the Peterhead, elliptical and 
unsymmetrical in the other two specimens; and in the Peter- 
head and Wick specimens there is a median space between the 
laminae of the axis and third. In each there is a series of 
well marked processes projecting backwards (anapophyses) from 
the outer part of the laminae, near the articular processes. In 
the Wick specimen (see Fig. 3) they are 1 to 1} inch in length, 
about two inches in breadth (about one-third of the lamina 
transversely), directed backwards and a little upwards, taper 
to a blunt rough point, and evidently receive their soft attach- 
ments from behind. On the 3rd, 4th, and 5th vertebrae they 
are much larger and longer than the atrophied spines, which 
are mere narrow median roughnesses, with a slight peak 
posteriorly. On the three hinder vertebrae they are consider- 
ably longer on the left than on the right side. In the Peter- 
head specimen (in which the spines are well marked) these 
anapophysial processes are more internal and less marked, 
though quite distinct, except on the 6th and 7th, on which 
they are obscure. In the Stornoway specimen they are 
intermediate. In front, they commence strongly marked on 
the axis, while posteriorly, on the 7th, they merge with the 
posterior articular process; but along the neck they are in- 
ternal to and quite distinct from the articular processes. 

10. ARTICULAR PROCESSES.—These processes are in a very 
reduced condition for vertebrae of this size. The anterior face 
nearly straight up, but with a little inclination outwards and 


CERVICAL VERTEBRAE IN FIN-WHALES. 25 


also forwards, the posterior the reverse. The size of the carti- 
laginous surfaces varies a good deal from the 3rd to the 6th, 
averaging 1} to 2 inches by I, the ellipse or oval being placed 
transversely, or rather obliquely upwards inwards and backwards, 
nearly in the direction of the laminae, on which most of the 
facet of the posterior is placed, the anterior being more on the 
“process,” which projects from about 3 inch on the 8rd and 4th 
to from 1 to 1} on the 6th and 7th vertebrae. They become 
longer and more oblique between the 6th and 7th, and still 
more so between the 7th and Ist dorsal. Between the axis and 
3rd they are nearly twice the size of those of the succeeding 
vertebrae, and present in all the four specimens irregularities 
and pits, giving a worm-eaten appearance to the surfaces. 
When well formed, the surfaces of the anterior processes are 
from within outwards first convex, then concave; those of the 
posterior the reverse. They are best marked in the Peter- 
head, and least in the Stornoway series. 

11. INFERIOR TRANSVERSE ProcessEs.—Those of the 8rd, 
4th and 5th, complete in all the specimens, may be first com- 
pared. The root springs from the lower part of the side of the 
body, placed nearer the anterior than the posterior surface, 
varying in the amount of forward and backward expansion 
which it undergoes in joining the body, the 4th being thinner 
than the 8rd and 5th. The root has a broad attachment 
vertically, like the pedicle which supports the superior process, 
and as broad as it, but thicker in accordance with the greater 
thickness of the inferior processes. The roots are concave and 
smooth on both surfaces. The tubercular stage, nearly on the 
same line internally, increases in length forwards to the 3rd. 
The projection of the inner angles, besides being downwards is 
also forwards on the 3rd and 4th, especially on the former, but 
backwards on the 5th (and on the 6th also, if the process is 
present), except in the Wick specimen, in which the projection 
on the 5th is forwards. The tubercular part rather diminishes 
outwards on the 4th and 5th, but increases on the 8rd, termi- 
nating in a great though variable projection, giving ‘the 3rd 
the longest and largest tubercular stage. The nerve-groove 
stage is well marked in all, best on the 5th. The twist seen on 
the anterior surface of the processes is owing, internally, to the 


26 PROFESSOR STRUTHERS. 


tubercular stage being much thicker below than above, while 
the grooved stage is thin below as well as above; and, externally, 
to increased thickness and less inclination of the terminal plate. 
The posterior surface of the processes is inclined obliquely 
upwards to the lateral canal, nearly uniformly so on both the 
tubercular and the grooved stages. 

When in position, the process of the third is seen, after 
rising forwards a little at the root, to slant very obliquely back- 
wards, parallel to the lower edge of the process and wing of the 
axis, and to bulge more downwards than the axis in the Peter- 
head specimen and at the outer part on the left side in both of 
the other specimens. The fourth has less obliquity backwards, 
and is more slender throughout, than the 3rd. The fifth is 
nearly horizontal, is the process to which the others converge, 
and is the stoutest, especially externally, having to support the 
widest and thickest terminal plate. The conditions of the siath 
in these three necks illustrate well the lability of the inferior 
transverse process of this vertebra to be more or less deficient. 
In the Wick specimen, it is complete on the left side, the stages 
well marked, but on the right side it is wanting at the nerve- 
groove stage; in the Peterhead specimen it is wanting at the 
nerve-groove stage on both s'des; and in the Stornoway specimen 
a large part of the tubercular stage also is wanting, on both 
sides, on the left side little more than the root-stage being 
present. When this process is incomplete in length, it also 
wants the upper part (that which gives breadth to the other 
processes) partially at the root, but more especially at the tuber- 
cular stage. Hence the process is flattened in a direction the 
reverse of those in front of it, and appears to spring from the 
body farther in than the others, and the capacity of the ring is 
thereby incidentally increased. The incomplete process gener- 
ally tapers outwards to a narrow round point. The other end 
of the gap is formed by a flattened pointed process which the 
upper transverse process sends inwards more or less. The 
appearance as if the two ends would not meet if prolonged, is 
owing partly to the natural twist of the grooved stage, partly to 
the two pointed ends not belonging to corresponding parts of 
the plate which would have united them had it been developed. 
In the Peterhead and Wick specimens, in which I had the 


CERVICAL VERTEBRAE IN FIN-WHALES. 27 


opportunity of dissecting the soft parts, the processes were 
represented at these gaps by ligament between their cartilagi- 
nous tips’. 

The seventh shows, as usual, at most a mere rudiment of the 
root stage of the inferior process. It is seen in the Wick and 
Stornoway specimens; is better marked on the Wick 1st dorsal, 
less marked on the Stornoway Ist dorsal; not at all on the 
Peterhead 7th, but well enough marked on the Peterhead 1st 
dorsal. It is placed where the lateral and lower surfaces of the 
body turn round to each other, and is little more than a slight 
rough projection where the ligament was attached. It may 
occupy most of the length of the body, but is mainly on its 
posterior half. 

12. SUPERIOR TRANSVERSE PROCESSES.—The pedicles from 
which these processes arise, about an inch farther in than the 
lower process, are opposite the fore part of the bodies, coming 
quite to the level of the anterior surface, and expanding back- 
wards so as to occupy $ (Peterhead specimen) to 3 (in the other 
two specimens) of the length of the body. They average 
14 inch in length along the middle to the zygomal process; 
increase in width backwards from the 38rd to the 7th; are 
grooved before and behind to form the intervertebral foramina; 
are directed upwards and outwards; and the inner half may be 
said to belong to the support of the neural arch and articular 
process, while the outer half sweeps outwards into the superior 
transverse process, serving as its root. Viewed in position, the 
‘superior processes converge outwards to the 5th, which is nearly 
level. Unlike the inferior processes, they gradually increase in 
strength backwards, the increase becoming more marked on the 
6th, and greatly more on the 7th. At first, at the nerve-groove 


1 In the Wick specimen the right inferior process is 4} inches in length; 
after forming a forward and then a backward-projecting inner angle, it tapers 
rapidly outwards; gap now 31 inches. Terminal expansion of upper process 
not so broad as on left side; pointed process sent in 4 inch internal to outer 
end of ring. In the Peterhead specimen both processes are 4} inches long; 
inner angles developed backwards, left less than right; left more flattened, 
from greater deficiency of upper part, than right. Gap on right side 1, on left 
linch. Plate of upper process most expanded on right side; extent to which 
narrow process turns in, right side, 13, left 2} inches. In the Stornoway speci- 
men the inferior process is 2} inches long on right side, 14 on left; left process 
flattened and tapering to blunt point; right much thicker than left, rounded 
and does not taper much; gap on right side 7}, on left side 8}. Plates of 
superior processes turn in for about } inch, process on right side the most 
distinct. 


28 PROFESSOR STRUTHERS. 


stage, the surfaces are more directly forwards and backwards, 
giving the processes a slender appearance here, and leaving wide 
spaces between. The length of this stage increases forwards 
from the 7th to the 2nd. The tubercular stage, from the rough 
prominences which mark its commencement to opposite the 
outer end of the ring, has its surfaces inclined, the posterior 
looking obliquely downwards to the lateral canal. The anterior, 
looking obliquely upwards, is, in its upper half, rough and 
bevelled so as to look very much upwards, and is broadened so 
as to overlap the process behind it before the outer end of the 
foramen is reached. The inclination of these processes is most 
strongly marked on the 3rd and 4th, extending also to their 
nerve-groove stage; a little less on the 5th; on the 6th to a 
variable extent in the different specimens, in the Peterhead 
specimen throughout, in the Wick specimen scarcely at any 
part. On the 7th, on the contrary, the surface which looks 
obliquely to the canal is the anterior. This process is so thick 
as to present a third surface, looking upwards and rough, cor- 
responding to the rough bevelled part on the anterior surfaces 
of the processes in front of it. 

The general inclination of the posterior surfaces of both 
upper and lower processes, from the 2nd to the 6th inclusive, 
continued externally at their junction, gives the double trans- 
verse process the appearance of the section of a cone, the inner 
circumference, at the ring, being farther forwards than the 
greater circumference. It is seen on a large scale on the pro- 
cesses and wing of the axis. Although the most striking result 
of this is the presentation of a series of oblique surfaces, instead 
of narrow edges, as a bony wall to the lateral canal, it is to be 
regarded rather as the result of adaptation to the attachment 
of the ligaments and muscles on the under and upper aspects 
of the neck. 

Viewed in series, the superior processes in the Peterhead 
and Wick specimens, from the 4th to the 6th, are nearly on the 
same level in point of height; the 3rd rises a little higher, the 
7th rises higher throughout, and prominently so at its outer 
end. In the Stornoway specimen, the five posterior are on the 
same level, the 7th scarcely rising, except at the outer end. 

The terminal plates are a little inclined towards the canal 


CERVICAL VERTEBRAE IN FIN-WHALES. 29 


in their inner portion, continuing the inclination of the posterior 
surfaces of the processes above and below, but the inclination 
is less. The outer portion, when expanded, is less inclined, but 
the whole posterior surface has more or less of a concavity, in- 
creasing backwards, especially marked on the 6th and 7th; the 
anterior surfaces being correspondingly convex. The size of the 
terminal plates varies much according to age and the vertebra. 
They are much less expanded in the less mature Peterhead 
‘ specimen than in the more aged Wick specimen. They increase 
in width backwards to the 5th (except in the Stornoway specimen 
im which the 3rd are as broad), which is in all the most pro- 
jecting ; and they diminish backward from it. In the Peterhead 
specimen the 3rd and 4th are blunt-pointed triangles, not much 
broader ‘than the processes, the 5th forms a larger triangle. 
The 3rd, 4th, and 5th, im the Stornoway and Wick specimens, 
have expanded vertically as well as transversely to form more or 
less square-shaped plates (Fig. 4), the outer edge oblique, the 
lower angle the more prominent,—as seen on a large scale in 
the wing of the axis. The upper angle may not be developed, 
the plate retaining the triangular form, or so much developed 
that there is a hollow between the two angles. The 6th in all 
tends to broaden upwards, as a relation to the 7th, though the 
lower part is still the most prominent point. The extent to 
which the terminal plates may vary is well illustrated on the 
third vertebra; in the Peterhead specimen their breadth is 24 
inches (only 4 inch broader than the processes near them), 
while in the other two specimens they have expanded so as to 
present a breadth of 5 to 5} inches, and a height, vertically at 
the inner part, of 64 in the Stornoway, and 8 inches in the 
Wick specimen. The transverse process of the 7th forms a 
more or less square-shaped terminal expansion. In the Storno- 
way specimen it is very square-shaped, nearly at right angles 
to the rest of the process, the upper angle being, at least on 
the right side, the most extreme point. In the other two 
specimens it is rhomboidal, directed obliquely downwards and 
outwards, the lower angle the most extreme point, and less ex- 
panded in the Wick than in the Peterhead specimen. The 
outer edges present the rough unfinished appearance of ossifying 
bone; in the Peterhead specimen at the broadening points; in 


30 PROFESSOR STRUTHERS. 


the more expanded Stornoway and Wick specimens, all along 
the outer edge and round the upper and lower angles, most 
marked in the Wick specimen’. 

Viewed in position, the 5th vertebra is, next to the axis, the 
most projecting, and its terminal plate is not only usually the 
broadest but is thicker, especially its lower part, than those of 
the 3rd, 4th, and 6th. It is the horizontal process, to which the 
others converge. The 7th is the least projecting in all. Next 
to the 5th in projection are the two next it; in the Stornoway 
specimen these two are equal; in the Peterhead it is the 4th 
on the left side, the 6th on the right; while in the Wick speci- 
men it is the 4th, but in it the 3rd projects as far as the 4th. 
When the bodies are separated to the same extent as they are 
naturally by the fibro-cartilages, the terminal plates, though 
near each other, are not in contact; the thin wedge-shaped 
spaces diminish outwards until there may be only from § to 4 
inch between the tips; but they may be made to touch by a 
little lateral flexion, or the more slender ones by their flexibility 
when the tips are pressed with the fingers. The following 
measurements of the Wick and Stornoway specimens indicate 
the amount of the convergence. At the roots of the inferior 
processes the extreme distance between the third and sixth is 
10 to 104 inches; between the tips, including the thickness of 
the four plates, 24 inches. Including the axis and the 7th, the 
extreme distance between their superior roots averages 18 
inches, while externally they reach the same level. 

13. THe Rrincs.—The characters of the rings (foramina) are 
seen when the vertebrae are laid in series on the floor. In form 
they are between a triangle and a semi-oval, the inner boundary 
obliquely convex, the upper and lower concave; the upper 
angle acute, the lower obtuse, the outer rounded off (Fig. 4). 


1 The thickness of the terminal plates, when expanded, as in the Wick and 
Stornoway specimens, from the 3rd to the 6th, averages about half an inch, 
at the middle; the 4th is the thinnest, the 5th the thickest. They are gene- 
rally rather thinner internally towards the ring, and thicker towards either the 
upper or lower margin, towards the upper in the 3rd and 4th, towards the lower 
in the 5th. The fifth in the Wick specimen is 14 to 14 inch below, } to} inch 
above; in the Stornoway specimen there is marked a-symmetry (Fig. 4), 
on the right 7 below, 4 above; on left side 1 inch below, }{ above. The 6th in 
the Wick specimen is # inch thick on the side on which the inferior process 
is complete, and much thicker below (1}) than above (8), while on the right the 
plate is under } inch and nearly uniform. The 7th is thicker (1 to 14) and less 
expanded than the others. 


CERVICAL VERTEBRAE IN FIN-WHALES, 31 


The oval form is more marked in the anterior, the triangular 
form in the posterior vertebrae. The upper margin is the 
longest, partly from the pedicle being set on the body about 
an inch farther in than the inferior transverse process is, partly 
from the outer end of the ring being below the level of the 
transverse axis of the body. This margin does not rise higher 
than where it begins, but curves gradually outwards and down- 
wards. The lower margin is generally the most bent, varying 
a good deal (from ? to 14 inch) in the degree to which it is 
bent down at the tubercular stage; and the outer half varies 
in the degree of its curvature, in some turning up more abruptly 
so as to give the appearance of an angle on the lower edge of 
the rig, in others having a more continued concavity. The 
greatest vertical height of the rings (as given in the table) is 
at about the junction of the inner and middle thirds, and is 
about 4 to $ inch greater than at the middle of the ring. If 
the line of the transverse axis of the bodies be prolonged, it 
intersects the rings variously; in the 2nd, 2 are above the line; 
in the 3rd, there is rather more above than below (except in 
Stornoway right, most below); the 4th is about equally divided 
(except in Stornoway left, most above); in the 5th most below 
(except in Stornoway left, rather most above); in the 6th most 
(from % to 2) below. A line intersecting the outer ends of the 
rings (foramina) leaves on an average 2 of the rings above it, 
and intersects the bodies so as to leave 2 above (on the sixth 2) 
the line. The extreme tips of the transverse processes are be- 
low the line of the transverse axis of the bodies, and are mostly 
below even the transverse line intersecting the outer ends of the 
rings, but the tips generally are on a line with the general axis 
of the foramina and double transverse process, the direction of 
which is outwards and downwards. The size of the rings gene- 
rally decreases a little backwards from the 3rd to the 5th 
(except in Stornoway 8rd, in which it is not so large as in the 
4th), as the measurements given in the table show. The slight 
increase in capacity at the 6th is chiefly owing to the deficiency 
in the height of the root of its inferior transverse process. It 
will be observed, from the measurements given, that the ex- 
pansion of the terminal plates is not, as in the case of the axis, 
accomplished at the expense of the foramina; for, although the 


32 PROFESSOR STRUTHERS. 


foramina are a little less in the Wick than in the Peterhead 
specimen, they are (except in the 3rd) larger all through in the 
Stornoway than in the Peterhead specimen, while in the latter 
the plates are much less expanded than in the other two. The 
gain is by outward growth beyond the rings. The table of 
measurements shows frequent want of symmetry in the diame- 
ters and capacity of the rings on the two sides of the same 
vertebra. 

14. RECOGNITION OF THE FIVE POSTERIOR VERTEBRAE.— 
These vertebrae may be distinguished from each other by - 
following characters. 

The third and fourth, from the others, by their transverse 
processes slanting obliquely backwards. The articular pro- 
cesses indicate front and back, the anterior facing upwards. 
The third is known from the fourth by the greater slant of the 
transverse processes, making it like a bow when resting on the 
floor, and by the far out position and great development of the 
outer end of the tubercular stage of the lower transverse process. 
The fifth is known by its transverse processes being directed 
nearly horizontally outwards. The sixth by the transverse pro- 
cesses being directed a little forwards, but more readily by the 
inferior transverse processes being usually more or less incom- 
plete. The seventh is known, from the sixth, by its robust 
superior transverse process, and the almost entire absence of a 
bony inferior transverse process; and from the first dorsal, by 
its transverse process being less robust, and being flattened, 
especially at the outer end, while the ends of the first dorsal 
are thick and rounded, 

15. THe Axis.—(a) Transverse processes. This enormous 
process, when fully developed, may be divided into three parts 
of nearly equal length,—the processes and foramen, the broad 
square part of the wing, and the tapering part of the wing. 
The ring is so small as to be scarcely equal in circumference to 
the spinal canal of the vertebra, except in the Stornoway speci- 
men, in which it slightly exceeds it. It is ovoid, the lower 
boundary the most curved, the outer end rather the most 
pointed. The smaller size of the ring of the axis is owing to the 
increase of the processes and inner part of the wing at the expense 
of the ring. The extreme height of the processes opposite about 


CERVICAL VERTEBRAE IN FIN-WHALES. : 33 


the middle of the ring is very little greater than that of those 
of the 3rd vertebra, but they are so developed both in height 
and thickness that they are twice the breadth (height), and 
two or three times greater in circumference. The increase is 
greater relatively on the superior process, but the inferior 
process is actually the greatest, corresponding to the greater 
extent of the lower part of the wing. The wing corresponds to 
the terminal plate of the vertebrae behind, enormously ex- 
panded. Although it presents very various forms with age and 
individual variation, definite characters may be recognised. 
From having been square-shaped, it has become prolonged at 
the lower part, giving the outer border a very oblique direction, 
and rather a triangular appearance to the wmg. In a young 
specimen in my possession’ the processes are as yet flat, the 
lower twice the height of the upper, the plate beyond the ring 
is only half the length of the ring, and a line is seen running 
across it, from the outer end of the ring, where the two have 
united, leaving much the broadest part of the wing opposite the 
inferior process. In the four grown specimens the prolongation 
of the axis of the ring leaves 2 or 2 of the breadth of the wing 
opposite the lower process, except in the Peterhead specimen, 
in which the division is about equal. 

When the wing is developed, the outer border, oblique and 
undulating, presents more or less of a concavity which may (as 
in the Peterhead specimen) be partially subdivided by a promi- 
nence. The inferior border is thick and rough on the process 
opposite the tubercular stage of the vertebrae behind, and then 
sweeps outwards to the tip with a general convexity downward. 
The superior border presents first a well-marked stage corre- 
sponding to the nerve-groove stage of the other vertebrae, termi- 
nated externally by a tubercle where the border is rolled 
upwards and forwards, corresponding to the series of tubercles 
on the processes behind, and before to the hinder projection of the 
transverse process of the atlas. The atlo-axoid intertransverse 


1 Without history, but it is evidently that of a young great Finner. ‘ It has 
the following dimensions. Greatest height 12 inches; greatest width 233; 
width of lateral ring, right 32, left 35; height of lateral ring, right 2, left 2}; 
circumference of lateral ring, right 9, left 94; width of plate beyond ring, 
right 21, left 2; width of spinal canal, anteriorly 5, posteriorly 6; height of 
spinal canal, anteriorly 45, posteriorly 41; circumference of spinal canal, 152. 


VOL. VII. 3 


34 PROFESSOR STRUTHERS. 


ligament was found to be attached here in the dissection of the 
lesser Fin-whale. This tubercle is at the broadest (highest) 
part of the wing, and is nearly opposite the outer end of the 
ring. From the tubercle to the upper angle is the tubercular 
stage of the process and wing; it is rough and bent backwards, 
forming a considerable concavity transversely, giving a very 
undulating appearance along the upper edge-of the process and 
wing (see Fig. 3). 

The extent to which the wing is expanded varies in both 
directions. The length outwards to the superior angle is nearly 
the same (12 to 13 inches, from the edge of the inferior surface 
of the body) in all the four grown specimens, except on the 
right side of the Wick specimen, in which it is 2 inches more. 
The length to the inferior angle varies in these four, from 15 
inches in the Peterhead (right side, left 174) to 20 in the Stor- 
noway specimen. ‘The broadest part of the wing is nearly 
opposite the outer part of the ring, where the processes are 
tubercular, especially the upper; but the breadth (height) is not 
very much greater than that of the processes farther in, being 
about 2 inches greater in the Wick (height of wing 123), 1 to14 
in the Peterhead (right wing 103, left 11), 1 in the Norway (10), 
4 to 4 im the young (83, 83), and in the Stornoway specimen 
1 inch on the left side, none at all on the right (101, 93). From 
this point, to opposite the upper angle, being along the mner 
half of the wing, the breadth diminishes somewhat at both 
borders, and on the outer third rapidly by the obliquity and 
concavity of the outer edge. The outer 5 inches in the fully 
grown Stornoway specimen is abruptly marked off as a nearly 
equilateral triangle, rounded at the tip, and is bent backwards 
so as to give this part a much greater slant than the inner part. 
It is at the same time twisted, so that its posterior surface looks 
partly upwards. The processes and their wing are curved 
transversely, concavity backwards, the depth of the curve, from 
the ring outwards, being about an inch when the tips are much 
bent. The wing is also curved vertically, depth of curve 1 to 2 
inches, influenced by the amount of bending of the margins, 
but well marked over both the surfaces, convex in front, concave 
behind. The thickness of the wing at the middle is about an 
inch, increasing inwards, diminishing outwards to from # to 4 
inch at the tip. 


CERVICAL VERTEBRAE IN FIN-WHALES. 35 


When tn position, the wings of the axis, in the Stornoway 
specimen, are seen to pass outwards beyond the tip of the trans- 
* verse process of the 5th more than two inches; and backwards 
as far as fully to the level of the tip of the transverse process of 
the 7th, and to the level of the junction of the anterior and 
middle thirds of the body of the 6th. In the Peterhead speci- 
men the wings extend two to three inches (over 2 on the 
right, over 3 on the left) outwards beyond the 5th; but back- 
wards only to the level of the tips of the transverse processes 
of the 4th, and to the level of the hinder edge of the body of the 
4th. In the Wick specimen (as accurately as can be deter- 
mined in the partially injured condition of the extreme tips) 
they reach outwards four inches beyond the 5th; and back- 
wards to the level of between the tips of the transverse processes 
of the 6th and 7th, and to the level of the hinder part of the 
body of the 5th, but they may have been longer. Taking all 
the five specimens, the distance to which the slant of the wings 
carries their tips back from the level of the hinder surface of the 
body, is, in the Stornoway specimen 10 inches, Wick apparently 
8, Norway 8, Peterhead 54, young specimen 1?inch. Vertically 
the inferior transverse process is seen to project less downwards 
than the tubercular stage of the process behind it, and the 
superior process scarcely if at all projects above the level of 
the inner stage of the superior processes behind it, but all be- 
yond these points the wing and upper process of the axis pro- 
ject beyond the processes behind them, upwards, downwards 
and outwards, forming a great sloping shield in front of them, 

(b) The region of the spinous process of the axis presents 
very great variety. The Wick and Stornoway specimens re- 
semble each other in presenting a large square-shaped mass, 
partially bifurcated in the former, while, in the Stornoway and 
Norway specimens, this part is flat with two low widely-se- 
parated longitudinal ridges, or crests, in the valley between 
which there is a low median ridge. The mass is apt to be 
regarded as a greatly developed and more or less bifurcated 
spine, but the central ridge must be regarded as the true spine, 
the lateral ridges being processes on the laminae, serial behind 
with the anapophysial processes, and anteriorly forming pre- 
jections on which there may be true articular facets for articula- 

3—2 


36 PROFESSOR STRUTHERS. 


tion with the atlas. Hence the spine appears in the table of 
measurements, in one, } an inch, in another, 34 inches in 
length. We have here a striking illustration of how easily one 
might be misled in endeavouring to found distinctions of species 
on the conditions of the bony processes’. 

(c) Anterior aspect of the body of the axis. There is con- 
siderable variation in the depth of the articular surfaces, in the 
form of the odontoid area between them, and in the form of 
the odontoid process. The greater depth of the articular cavi- 
ties is owing to the greater rising up of the outer sides. In 
the Wick and Peterhead specimens the greatest depth on 
each side, as given by a line laid on between the outer edges 
at their fore part, is 1¢ inch; in the Stornoway specimen ?, in 


1 We can see how the one form may grow into the other. Im the young 
specimen, the lateral ridges, or crests, rise about an inch, are 4 inches apart, 
diverge backwards, and in the valley between them there is a low median spine. 
In the Nerway specimen the lateral crests are two low irregular convex ridges, 
5 inches apart, rising scareely half an inch, except forwards, where they support 
articular facets, while the median ridge, in the very shallow valley, rises at 
most half an inch. The anterior part of the crests and valley begin to meet the 
posterior part at an angle, and the posterior part is the roughest. In the 
Stornoway specimen, the crests, 44 mcehes apart, have mcreased beth m height 
and thickness, especially forwards, the valley is an inch in depth at the fore 
part, much less behind, and there is a low median ridge; and the angle between 
the fore and back parts is increased, but is still very obtuse. In the Wick 
specimen (see Fig. 3) the crests are largely developed forwards, but still more 
backwards, the valley between them bemg at the same time weil filled up. 
The angle between the fore and back parts of the crests and valley has risen up 
to aright angle. The valley in the fore part is nearly filled up, while in the back 
part it presents a very rough excavation. The whole has the appearance of a 
great square-shaped mass partially bifurcated hackwards and with a tendency to 
bifurcate upwards, without median ridge anywhere. In the Peterhead specimen 
the change is carried farther, the angle is carried upwards and backwards to aa 
acute angle, the fore part is quite filled up, the back part concaye and rough. 
The square Iump of bone thus formed presents slopmg lateral surfaces, giving & 
width of 6 inghes to the process at the middle; a square-shaped superior 
surface looking forwards as well as upwards, about 4 inches square, projecting 
more on the left side than on the right; a thick anterior border; and poste- 
riorly, marked off from the fore part by @ sharp transverse overhanging edge, 
an excavated surface, looking backwards and a little upwards, 5 inches across 
by #4 vertically, excavated to a depth of 1} inch, very rough, and with a median 
ridge. 

This great variation isnot a matter of age, for the form presented by the 
young specimen is retained by two of the mature specimens, while the more 
developed form is presented by the mature Wick and the scarcely mature Peter- 
head specimen; unless we suppose that this part having begun as in the young 
specimen, progresses under muscular action to the condition of the square-shaped 
mass presented in the Peterhead specimen (in which the other cervical vertebrae 
have the spines most fully developed), and afterwards becomes reduced with age 
to its early condition, the three other specimens showing stages of that reduc- 
tion. But in the Wick specimen, in which the spines of the vertebrae behind 
have nearly disappeared, this part of the axis presents the next best instances 
of massive development. These differences must therefore be regarded mainly 
as exhibitions of individual variation. 


CERVICAL VERTEBRAE IN FIN-WHALES. 37 


the Norway specimen intermediate. This difference is strik- 
ingly seen from below when the bodies are in position, the atlas 
appearing in the two former to sink into a deep cup in the 
axis, while the depression is comparatively shallow in the Stor- 
noway specimen. This gives the appearance of considerable 
difference in the length of the body at the sides, while at the 
middle below they are mostly the same, as seen in the table 
of measurements. Together with greater depth of the lateral 
cavities, there is a sharpness of finish all round the edges of 
the horse-shoe surface, in marked contrast with the Stornoway 
specimen. The articular edges in the latter are over half an 
inch lower than the odontoid, in the Norway specimen 2 inch 
below it, in the Peterhead and Wick specimens nearly (Peter- 
head scarcely, Wick rather above) level with the odontoid, but 
the odontoid is actually the longest in the Peterhead specimen. 
The upper of the two series of measurements of the length of 
the body given in the table are taken at the odontoid, and 
show it to be 3? inch each in the Wick and Stornoway, and 
4 in the Peterhead specimen. 

The odontoid appears to the eye to vary a good deal in 
length, but this is mainly owing to its form. In the Peterhead 
specimen, in which it appears long, it rises to a height of an 
inch from the edge of the odontoid area; in the other specimens 
about 4+ inch less. In the Peterhead specimen the area and 
process together form a cone, rising well but not abruptly at the 
process ; in the Wick specimen the area is less raised, the pro- 
- cess rather more defined at its base; in the other two grown 
specimens there is only a low general cone, rising to a blunt 
summit, lowest in the Stornoway specimen. In the young spe- 
cimen the cone rises better, and is terminated by a blunt exca- 
vated apex. In all of them the summit is below the middle of 
the area (at about the junction of the lower and middle thirds) 
but above the middle (nearly as high as the junction of the 
upper and middle thirds) of the general body of the axis. In 
height, the odontoid area is nearly the same (nearly 3? inch) in 
all, except in the Norway specimen in which it is fully 43, but 
it varies a good deal in breadth. The following are the breadths 
of this area, and, given within brackets, the breadth of the en- 
tire anterior surface of the body—Peterhead 43 (15), Stornoway 


38 PROFESSOR STRUTHERS. 


5 (134), Wick 52 (15), Norway 6} (15), young specimen 6} 
(123). The area is much rougher on the inner half or two- 
thirds than on the outer part, and is especially rough on the 
process and below it. In the Stornoway specimen, in which the 
articular surfaces are shallow, the edge between them and the 
odontoid area is not so sharply defined as in the others. 

16. ArLas.—(a) The posterior surface presents differences 
corresponding to those noticed on the anterior surface of the 
axis. The transverse convexity of the lateral articular surfaces 
is strongly marked in the Wick and Peterhead specimens, the 
surfaces of the latter being specially prolonged outwards and 
terminated by a raised edge, so that they become concave ex- 
ternally. In the Stornoway specimen, the surfaces are much 
more flat, the internal convexity is low, and the external con- 
cavity is distinct, although the outer edge, instead of being 
prolonged, is so deficient that its upper half is bounded 
by a concave instead of a convex line, and is so low as to be 
nearly on a level with the transverse process. The crescentic 
ligamentous surface, internal to each lateral articular surface, 
varies in breadth with that of the odontoid area, to which the 
two crescentic surfaces and the odontoid division of the canal 
correspond, and also in sharpness of definition. In the Storno- 
way specimen it reaches 1 inch in breadth, and is not much 
depressed ; in the Wick specimen (see Fig. 5) 13 in breadth, 
and is abruptly depressed more than } inch; in the Norway 
specimen fully 1? in breadth, and is obliquely depressed. 

(b) The anterior articulating surfaces vary in the state of 
their edges, in their depth, and in the breadth of the median 
groove. In the Peterhead and Wick specimens the edges are 
sharp, giving the cavity a depth of 3 inches; in the other two 
specimens the cavity is nearly half an inch less in depth, the 
edges being lower and more rounded, sinking especially at the 
sides to the level of the transverse processes. All show the 
furrow round the outer side for the attachment of the capsular 
ligament. ‘The median groove is narrowest in the Wick and 
Stornoway specimens, broader (2 inch) and more defined in the 
Peterhead, broadest (3 inch) in the Norway specimen. In all it 
widens out triangularly below, and also a little above, being 
narrowest at or above the middle, but in the Norway specimen 


CERVICAL VERTEBRAE IN FIN-WHALES. 39 


it remains widely open, and presents a double furrow in its 
floor, indicating the separate attachment of the two capsular 
ligaments. The measurements of the cups of the atlas are; 
greatest height and breadth (diameters) of each, Peterhead spe- 
cimen 11 by 6 inches, Stornoway 10} by 54 (wanting an inch 
in height at the upper part from not extending up over the 
roof of the transverse foramen, as the others do), Wick 11} by 
6, Norway 11 by 53; distance between the outer edges of both 
cups, given in the same order, 134, 123, 135, 123. 

(c) Parts on the neural arch. Occurrence of articular pro- 
cesses between the axis and atlas. The spinous process is distinct 
in all as a median ridge, most marked posteriorly, very little 
marked on the anterior half in the Stornoway and Wick speci- 
mens. It is most developed in the Peterhead specimen in 
which the axis is so greatly developed here, but it is on the 
whole scarcely more pronounced in the Wick than in the Nor- 
way specimen, in which the spine of the axis is so differently 
formed. On each side of the spine, about the middle of the 
lamina, rough lateral ridges occur, serial with the crests on the 
laminae of the axis, increasing and diverging forwards (see Fig. 3), 
The ridge runs forwards into the process which arches over the 
nerve-notch of the atlas and usually converts it into a foramen, 
and backwards more or less into a posterior articular process 
for the axis. True articular processes between the axis and 
atlas, situated above the nerve-escape, existing normally in 
reptiles and birds but not in mammals, are present, more or 
less, in at least four of these five great Fin-whales*. They are 
present on both sides in the Norway and Stornoway specimens, 
on the right side in the Wick (see Fig. 5) and the left side in 
the Peterhead specimen, and seem to have existed on both sides 
on the young axis. On the axis they are situated on the lower 
part of the anterior end of the crest-like ridge, above the middle 
of the lamina, and from their position the facet is very liable 
to be rubbed off and overlooked. The facets on the atlas in 
the Norway specimen are symmetrical, 14 to 1? inch trans- 
versely by about 1 inch longitudinally ; in the Stornoway spe- 
cimen they are elliptical pits, that of the left side divided into 


1 I find them still better developed in some male Narwhals, on one or on 
both sides. 


40 PROFESSOR STRUTHERS. 


two, into which corresponding projections of the axis sink. 
From the greater development of the crest on the axis making 
it overlap the atlas, the facet is seen on the upper aspect of the 
lamina of the atlas, but in the Peterhead specimen it is on the 
under aspect, as the atlas is here the overlapping bone. In the 
Peterhead specimen this functional articular facet (14 by 4 inch) 
is situated high up where the spine and lamina join and partly 
on the spine, on the left side, owing to the great upward pro- 
jection of especially the left side of the square-shaped mass of 
the axis. On both sides in the Peterhead specimen, and on 
the left side in the Wick specimen, there are low projections 
on the laminae of both bones, not meeting, corresponding to 
the ligamentous boundary superiorly of the intervertebral fora- 
men. This foramen thus marked off above, of an oval form, 
admitting from three to four fingers, is rather larger than the 
two succeeding foramina, but not larger than the foramina be- 
tween the three posterior cervical vertebrae. 

(d) The transverse foramen of the atlas is incomplete on 
both sides in the Stornoway specimen; on the left side half an 
inch is wanting, on the right side } inch. The posterior process 
is short especially on the left side, four-fifths of the nearly com- 
pleted foramen being opposite the anterior process, which curves 
back from the end of the articular cavity. It is triangular in 
form, the deficiency in the roof being on the inner side, leaving 
the internal aperture unformed; and the articular cavity does 
not reach upon this part as it does in the other specimens’. 
The breadth of the arch of bone roofing over the foramen 
(giving also the length of the foramen or canal) is, in the Wick 
specimen, 2 inches on the right side, 2} on the left; Norway 
specimen, right 1}, left 2; Peterhead specimen, right 14, left 13. 
The thickness in the Wick specimen is 1 to 14 inch, in the 
other two specimens + inch less’. In the Stornoway specimen 


1 This variety resembles that often seen in the human atlas by the more or 
less complete ossification of the ligament which normally arches over the 
nerve and artery. The serial correspondence of the posterior process to an 
articular process is evident, but it joins in front with a process of the same 
vertebra. The same arrangement is seen in the axis of some mammals, the 
anterior notch being converted into a foramen. In some the posterior notch 
in the dorsal region is converted into a foramen, the articular processes being 
also present. 

2 The articular eup being prolonged on this arch as far up as to opposite the 
middle of the foramen, the edge of the bone receives a forward curve in its upper 


CERVICAL VERTEBRAE IN FIN-WHALES, 41 


the recurved process is an inch in breadth at the middle, and 
3 inch thick. In dissecting the Peterhead specimen I found 
the roof to be naturally continued outwards for over half an 
inch by a ligamentous arch, thick where attached to the bony 
edge, becoming thinner to the crescentic margin in which it 
terminated externally. The broad groove issuing from this 
foramen is continued outwards to the anterior surface of the 
root of the transverse process, and a narrower groove is seen to 
pass nearly straight backwards from the outlet of the foramen 
to a notch on the lower edge of the lamina at the intervertebral 
foramen between the atlas and axis (see Fig. 3). The foramen 
contained, besides the atlantal nerve (about the size of the 
human great sciatic), a plexus of small vessels, one as large as a 
crow-quill, but no large vessel. The foramen is less than a 
third the size of the intervertebral foramina; roughly, it will 
admit a thumb. It is smallest at the imner end, which is oval 
in the Wick specimen (long axis longitudinal and nearly 1 inch), 
smaller and round in the Peterhead, intermediate in the Norway 
specimen. In the Stornoway specimen, in which the roof is 
incomplete, the oval is vertical and the capacity greater. 

(e) The transverse process of the atlas is in series with the 
superior process of the axis. Internally, the process is flattened, 
the surfaces forwards and backwards, but thick enough to pre- 
sent upper and lower. surfaces, the expanded root situated 
opposite the upper half or two-thirds of the articular surface, or 
so-called body, of the atlas, the downward extent of this attach- 
- ment varying, and being at first so gradual that it is not easy to 
define its commencement. Externally, it is flattened in the 
opposite direction, the surfaces inferior and superior, with upper, 
lower, and external borders. Variations are seen in the different 
specimens, as observed both in vertical and antero-posterior 
views. Observed in front, the processes in the Wick specimen 
(as seen in Fig. 5, representing a posterior view) stand out 
transversely, as stout triangular or conical processes, the right 
broadest (highest) externally, the left broadest at the root, which 
passes down 1} inch below the middle of the body. In the 


third, making the length between the posterior and anterior surfaces about an 
inch more above than below in the specimens in which the arch is complete. 
When the vertebrae are built up vertically, the upper third of the cup is con- 
sequently seen to rise to a higher level than the rest. 


42 PROFESSOR STRUTHERS. 


Peterhead specimen they pass straight out, are much thinner 
in the outer half, the upper margin forming a general concavity 
along its outer three-fourths, and the root is opposite very 
little more than the upper half of the cup. In the Stornoway 
specimen the root is opposite the upper 3 of the cup, passing 
14 to 2 inches below the middle, but, as the upper inch 
of the cup is not developed, the middle is a lower point 
in this specimen than in the others, and the processes there- 
fore extend fully to the beginning of the lower third of the 
eup. It widens out sooner on the left than on the right 
side. A little more filling up about the middle of the lower 
margin would give these processes the form of triangular 
masses set opposite the upper 3 of the cup; and I should have 
great hesitation in accepting the extent of this attachment as 
a character in distinguishing species. The upper margin of 
the process is sigmoid, first broadly convex, where the other 
two specimens show only the internal tubercle, and then con- 
cave on the outer % to the turned up tips. Thus in the antero- 
posterior view of the Stornoway specimen the processes are 
stouter at the root than in the other two, are more compressed 
in the outer half than in the Wick specimen, are tumed up at 
the tips, and are about an inch longer than in the other two 
specimens. A line between the middle of the outer parts of 
the transverse processes intersects the cup so as to eut off, in 
the Wick and Peterhead specimens, the upper 8 of the 11 
inches, in the Stornoway specimen the upper 2 of the 10 inches, 
and intersects the base of the process, so that in the Peterhead 
specimen about 3 inches are above and a little more below, 
while in the Stornoway specimen about 3 inches are above and 
5 below. The following are the height and girth at the middle 
of the transverse process in the three specimens—Wick, 44 and 
11} inches; Stornoway, right 34 and 10}, left 4 and 114; 
Peterhead, 2? and 10. The process in the Norway atlas is too 
much injured to admit of accurate conclusions being drawn. 
Observed from above, differences are seen (see Figs 1, 
Zand 3). At the back part of the root there is a rough 
tubercle (internal tubercle) from which a ridge passes obliquely 
across the process towards the fore part of the tip. This 
ridge is much more developed in the Wick specimen, render- 


CERVICAL VERTEBRAE IN FIN=-WHALES. 43 


ing the process much less flat than in the Peterhead speci- 
men. Another tubercle (external inter-transverse tubercle) 
is developed behind the tip in each, rendering the outer 
part, antero-posteriorly, an inch more than the inner part, 
and making the posterior margin internally much more con- 
eave than the anterior. The outer margin is convex in 
the Wick, nearly straight in the Peterhead specimen, and 
the extreme point in both is the anterior angle. In the Stor- 
noway specimen the extreme point is the posterior angle, 
which is prolonged backwards and outwards, the outer margin 
rounding off to continuity with the anterior margin, giving the 
processes a backward droop externally. Farther, the surfaces 
of the process are much twisted beyond the root, so that the 
inferior surface looks obliquely forwards, the superior obliquely 
backwards, and the oblique ridge, noticed on the upper surface 
in the other two specimens, remains as the upper border of 
the twisted process’. In the Peterhead specimen the surfaces 
are directly up and down, superior flat, inferior convex ; but in 
the Wick specimen, especially on the right side, an approach 
to the twisted form is seen, the increase downwards at the root 
giving the anterior surface some obliquity, and prolonging the 
inferior surface of the root outwards, so as to give the process 
a somewhat triangular figure. The differences in form between 
the transverse processes in the Peterhead and Stornoway spe- 
cimens are certainly remarkable. 

Seen in position, the processes stand nearly straight out in 
the Wick and Peterhead specimens, but in the Stornoway spe- 
cimen they have a backward droop externally. The distance 
between the nearest part of the tips of the transverse processes 
of the atlas and the surface of the wing of the axis behind them 
is, in the Wick specimen 6 inches, Peterhead 5, Stornoway 24 
to 3; the distance is nearly an inch less to the tubercle (inter- 
transverse) of the axis, in the Wick and Peterhead specimens, 
from its being turned forwards towards the atlas, while the 
little development of this tubercle in the Stornoway specimen 
is probably related to the unusual prolongation, above noticed, 


1 This twist of the transverse process is an approach to the condition in 
the lesser Fin-whale, in which this character is strongly marked. The same 
obliquity may be seen in man and in various other mammals. 


44 PROFESSOR STRUTHERS, 


of the tip of the process of the atlas. The tips pass out be- 
yond the foramen of the axis for an inch or more in the Peter- 
head and Wick, for about two inches in the Stornoway speci- 
men. Their lower margins, except quite at the base, are nearly 
on a level with the upper edge of the foramen, except in the 
Stornoway specimen, in which they project down in front of the 
upper fourth of the foramen, and more internally towards the 
base. 

(f) Canal of the Atlas. The constriction indicating the 
division into two parts varies. The width at the constriction in 
the several specimens is, Peterhead 1}, Wick 24, Stornoway 24, 
Norway 23, but the appearance of constriction is most marked 
in the Stornoway specimen. The general form of the odontoid 
division of the canal is, in the Wick specimen, that of a blunt- 
pointed triangle with gently concave sides (greatest breadth 2%) ; 
in the Stornoway specimen it is bulged at the sides and more 
pointed below (breadth 3); in the Norway specimen it is like 
the lower two-thirds of an ovoid (breadth 3); in the Peterhead 
specimen the ligaments still remain filling it up, but it most 
resembles the Wick specimen. The projection sustains, as in 
man, the most internally projecting part of the articular surface 
behind it, but more immediately attaches the upper part of the 
transverse ligament, the attachment of the ligament continuing 
down to where the opening begins to contract, leaving a tri- 
angular odontoid opening, bounded above by the concave edge 
of the ligament. The neural division of the canal varies a 
little in width in the different specimens (see table), and its 
form above is influenced by the curvature of the neural arch, 
which is less in the Norway specimen than the others, giving it 
more distinct upper lateral angles. The general form may be 
defined as square-shaped, with rounded angles, and contracting 
downwards, or as triangular with a very blunt apex below. 
Vertically, it measures about 44 inches to the concave edge of 
the ligament, 44 to the bony projections, which are situated 
below the middle of the general opening; its greatest breadth 
is about the same when taken anteriorly (as given in the table), 
but posteriorly the canal is much wider ; near the intervertebral 
foramen in the Wick and Norway specimens it is 6 inches, in 
the Peterhead 5}, in the Stornoway specimen 54. 


CERVICAL VERTEBRAE IN FIN-WHALES. 45 


(g) Sub-aaxial process. The peak on the hinder part of the 
atlas, the development of which has been regarded as a cha- 
racter of specific import, is present in various degrees in these 
specimens. The horse-shoe articular surface recedes here for 
? inch in the Norway and Peterhead specimens, 14 in the Wick 
and Stornoway specimens, leaving a median triangular notch. 
The upper part of this notch is a rough depression, while from 
the lower part a pointed process projects more or less backwards. 
In the Peterhead specimen, though forming a projection from 
where it begins, it does not reach so far as the edge of the atlas 
itself; in the Stornoway specimen it projects half an inch be- 
hind the rest of the atlas and in below the axis, as a median 
conical process, occupying the whole height of the notch; in 
the Wick specimen it projects transversely, as a tongue-like 
process, from the lower part of the notch, $ inch in length, 
24 inches in breadth, but, although larger than in the Storno- 
way specimen, it barely reaches to below the axis’. 


(B). In THE LeEssER FIN-WHALE? (B. Rostrata). 


17. TRANSVERSE PROCESSES.—(a@) Completion of the rings. 
It may be considered as determined that the various degrees of 


1 This process is much more developed in a portion of the atlas of a great 
Finner in my possession, which came I believe from Orkney. It is a larger 
atlas than either of the above. The lateral portions have unfortunately been 
sawn off through the outer parts of the articular surfaces. It has the follow- 
ing dimensions. Greatest height, spine being away, 15¢ inches; height of 
canal 94; greatest width of canal, anteriorly 43, posteriorly 7}; width at con- 
striction 22; below this the canal slightly diminishes downwards to a very 
blunt rounded lower end. Length of articular cup, now 114, probably 12; 
breadth 63; median groove between cups very broad, narrowest (at an inch 
from lower end) 14, at middle 1%, near canal 1%. Transverse canal small, 
length 34; inner end oval antero-posteriorly, long axis scarcely $; outer end 
vertically oval, long axis 1}; thickness of its bony roof 13. On posterior as- 
pect, crescentic ligamentous surfaces scarcely 14 in breadth and moderately 
depressed; lateral articular surfaces, length 103, moderately convex, united in 
one great horse-shoe surface. Swb-axial process greatly developed, amount of 
projection as seen from above, 1 inch, from below 1} inch, breadth at base 51, 
rising gradually, greatest to right side; thickness 1}, coming close up to horse- 
shoe surface, which is 3? inches in height at the middle line. 

2 These vertebrae in B. rostrata have been described in a nearly full-grown 
specimen by Prof. Flower (On a Lesser Fin-Whale, recently stranded on the 
Norfolk coast: Pro. Zoo. Soc. May, 1864), who has also given notices of spe- 
cimens which he examined in the Museums of Leyden, Brussels and Louvain 
(Notes on the Skeletons of Whales in the principal museums of Holland and 
Belgium, P. Z.S. Noy. 1864); and by Drs Carte and Macalister of Dublin 
(On the Anatomy of Balaeroptera rostrata, Trans. Roy. Soc. 1868) in a young 
specimen nearly the same length as mine, Their various conditions of ossifi¢ 


46 PROFESSOR STRUTHERS. 


ossific development of the rings is a matter chiefly of age, 
partly of individual variation. In this 144 feet long specimen 
none of the rings are ossifically complete. That of the axis 
wants 3 inch at the outer edge of the rmg. The terminal plate 
of cartilage by which it is completed is only an inch in breadth, 
the ovoid ring almost two inches. In the fetus, Eschricht 
found the 5th and 6th with ¢omplete rings; and Van Beneden 
and Gervais, speaking of B. musculus, remark: “On peut admet- 
tre que dans cette espece, comme dans la Lalenoptera rostrata, 
ces anneaux sont toujours complets, & létat de cartilage, dans 
le jeune animal, et que les différences ne sont que le résultat 
d'une ossification plus ou moins compléte.” Drs Carte and Mac- 
alister mention “ fibro-cartilage” as completing the rings in their 
young specimen, and in my notes of the dissection the completing 
structure has been recorded as ligamentous. The bony processes 
were succeeded by a cartilaginous stage, and this again by a 
fibrous stage, completing the canal. The cartilagimous stages 
varied in length; on the superior processes of the 4th and 5th they 
were 4 inch; on the inferior processes of the 5th, the most ossi- 
fied of all the inferior processes, they were on the left side $ inch, 
and 3? inch on the less ossified right side. The ligamentous pro- 
longations, beyond the cartilages, were quite distinct from each 
other for some distance, but, owing to the convergence of the 
processes, they became confluent at their outermost part, and 
could not be traced round to meet the corresponding process. 
This was more especially the case with the third, the fibrous 
part of which became intimately connected with the thick peri- 
osteum of the overlapping wing of the axis; and with the 
sixth, owing to the superior process of the 7th slanting forwards 
so much against it. Even the 4th and 5th could not be traced 
round, the fibrous part of the lower process of the 4th appear- 
ing to end on the (farther out) cartilage of the lower process of 
the 5th. The fibrous parts of the processes may be looked on 
as portions of that thick periosteum which encloses the bones 


development are remarked on in the Ost@ographie des Cétacés, of Van Bene- 
den and Gervais, p. 160, and they are figured in P]. 12 and 13 of that valua- 
ble work. Dr J. E. Gray has also figured some of them in P. Z.5S., May, 
1864, and in Cat. Seals and Whales, Brit. Mus. 1866. My remarks relate 
to some farther points in the osteology illustrated by this young specimen, 
and to the articulations. 


CERVICAL VERTEBRAE IN FIN-WHALES, 47 


and cartilages in the cetacea, from within which foetal cartilage 
has been absorbed, or within which ossification is advancing, 

The lower transverse process of the 7th vertebra was nearly 
altogether represented by a ligament, attached internally to a 
short pointed bony process (over 4 inch in length), externally 
to the cartilaginous tip of the superior process; the ligament 
13 inches in length, and flattened in the same direction as the 
bony processes’. This fibrous representative of the inferior 
transverse processes of the 7th vertebra is serial with the liga- 
ments sent in from the heads of the anterior ribs, and from the 
tips of the corresponding dorsal transverse processes, to the 
sides of the bodies of their vertebrae’. 


1 Prof. Turner found in the fcetus of one of the great Fin-whales (B. 
Sibbaldii) the inferior transverse processes represented by cartilage, completing 
a cartilaginous ring. See this Journal, 1871. 

2 These were examined on the three anterior dorsal segments, and are 
still seen on the dried preparation, and I may here note the arrangement. 
The 1st rib rested generally on a fibrous cushion interposed between it and 
the converged tips of the transverse processes, from the axis to the first 
dorsal, more precisely against the 7th cervical and 1st dorsal, but also on 
the 6th cervical (end of its lower transverse process), its chief ligamentous 
connection being with the 6th. From its short capitular process a ligament 
passed in to join the outer part of the ligamentous representative of the inferior 
transverse process of the 7th cervical vertebra, while from the tip of the 1st 
dorsal transverse process a separate ligament passed in to the body of the 
first dorsal vertebra, where it is attached to a conical bony parapophysis, like 
that on the 7th cervical. The 2nd rib, after being connected by a ligamentous 
cushion to the second dorsal transverse process, sends in a pretty long capi- 
tular process, going about half way in to the body, from which a strong liga- 
ment is prolonged to the 2nd and 3rd bodies, and to the fibro-cartilage between. 
The deeper part of this ligament is a prolongation from the second trans- 
verse process along the upper edge of the capitular process of the rib, so that 
the ligament passing in to the bodies is in its upper part parapophysial and 
in its lower part pleurapophysial. The 3rd rib repeats this, but having scarcely 
‘any capitular process, the two parts of the ligament are identified earlier. 
These parapophysial ligaments prolong the lower wall of the lateral canal of 
the neck backwards into the thorax. 

In both of the great Fin-Whales this part was so mutilated, as it is very 
apt to be in dividing the carcase, that I could only infer the presence of a 
ligamentous representative of the inferior transverse process of the 7th cer- 
vical vertebra from portions which remained, showing a ligament as thick as 
two fingers laid together. From the dissection of the Peterhead Razorback I 
inferred that such a ligament, connected to the body of the seventh cervical 
vertebra, 7 to 9 inches of it remaining, had been connected externally to the 
movable capitular process of the first rib, and along it to the transverse process 
of the 7th cervical vertebra. These ligaments will vary with the extent to 
which the ribs develop capitular processes. I may note here that in the 
Stornoway Razorback the three anterior pairs of ribs all develop long 
capitular processes, and that if the movable capitular process which I figured 
on the first rib of the Peterhead Razorback (see this Journal, 1871) were 
ankylosed and a little enlarged, it would represent very well the form which 
both first ribs present in the Stornoway specimen. I was able to examine 
the connection of one of the first pair of ribs to the transverse processes in the 
Wick specimen. It rested in a socket on the transverse processes, formed in 


48 PROFESSOR STRUTHERS. 


(b). The bony inferior transverse processes differ from those 
of the great Finner in being relatively stronger and in having a 
more downward direction. They are present as half-inch-long 
conical processes on the 7th cervical and Ist dorsal vertebrae. 
The tubercular stages of the next four (6th, 5th, 4th, 3rd) 
increase in length forwards to the 3rd, on which they are twice 
the length of those of the 6th, and are directed outwards and 
downwards, unbent, as far as their outer prominence, which 
is strongly marked. ‘These processes, therefore, descend below 
the level of the bodies more than in the great Finner forming 
the sides of a sub-vertebral space, the depth of which at the 
3rd is twice that at the 6th, while in the great Finner it may 
scarcely increase in depth forwards. The nerve-groove stage 
then turns upwards and outwards, tapering, and is variously 
ossified—on the 6th full-length on the right side, half-length 
on the left; on the fifth, nearly full-length on the left, under 
two-thirds length on the right; on the 4th, just beginning on the 
left side, not begun on the right; on the 3rd the process con- 
tinues robust to the end, no part of it turned up. The pro- 
cesses of the 4th are the most horizontal, those behind them 
inclining forwards. Those of the 4th and 3rd are not directed 
backwards, as they are in the great Finner by the greater slant 
of the wing of the axis, but slant a little forwards, that of the 
3rd consequently coming very close to the wing of the axis, and 
it is the most robust of all the inferior transverse processes. 
The different states of the inferior transverse processes of the 
6th vertebra in the two groups forms a marked contrast, the 
grooved stage being more or less unossified in the great 
Finners, while in this lesser Fin-whale this stage is better 
developed than in any of the other cervical vertebra, shooting 
out to near the end of the superior process of the 7th, the 


front by the 7th and partly by the 6th cervical, behind by the Ist dorsal. Its 
chief ligamentous connection was to the 7th cervical. Between the rib and the 
cervical part of its socket was a great fibrous cushion on which it rested, 
while between it and the dorsal part of the socket was a quasi-synovial cavity about 
2 inches in diameter, with periosteal surfaces on the rib and on the first dorsal 
transverse process, the ligaments forming a kind of capsule around it. On the 
2nd dorsal transverse process the attachment of the 2nd rib was indicated by a 
thick capsular cushion with central cavity bounded by soft irregular walls. No 
synovial cavity existed in connection with any of the ribs in the young Pike 
Whale, although the separation of the perichondrium is at first apt to deceive 
the dissector in regard to this. 


CERVICAL VERTEBRAE IN FIN-WHALES. 49 


two forming a strong arch for the ‘support of the first rib. 
The ligaments between the inferior transverse processes (inter- 
parapophysial) are oblique in different directions ; an external 
series running between the tubercles forwards and outwards, 
strongest behind ; and an internal series, the largest, passing 
from the tubercles forwards and inwards, increasing forwards, 
and of great size between the 3rd and axis and between the 
axis and atlas. 

(c) The superior transverse processes are well ossified ; the 
7th fully, like the first dorsal though not so robust ; the 3rd, 
4th, 5th and 6th, in their first two stages, of nearly equal 
length, and much more slender than the corresponding inferior 
processes. The distinction between the nerve-groove and the 
tubercular stages is marked on these four more distinctly 
than in the great Finner, by the presence here of a series of 
triangular processes directed forwards, between which the inner 
part of the superior inter-transverse ligaments passes. On the 
7th cervical and 1st dorsal vertebrae these processes (meta- 
pophysial) are placed more internally, close to, or almost on, 
the anterior articular processes. 

(dq) The lateral canal formed by the rings of the transverse 
processes differs in this young lesser Fin-whale from that of the 
great Finner both in relative capacity and in form. Taken at 
the fourth vertebra, it is transversely less than half the breadth 
of the body, and its capacity is not much greater than that of 
the spinal canal, their circumferences being respectively 7 inches 
and 8. In form, instead of being transversely triangular or 
ovoid, it is vertically ovoid or rather semilunar, the blunt end 
downwards and outwards, owing to the more downward direction 
of the transverse processes, a line prolonged from the transverse 
axis of the bodies leaving a much larger proportion of the ring 
below than in the great Finner. 

The neural arches are high and triangular, with rounded 
lateral angles. At the 4th, the breadth of the spinal canal is 
28, the height 111. Below the 4th it increases in breadth and 
diminishes in height; at the third with about the same breadth 
as at the fourth the height is 2 inches; while at the axis the 
height and breadth are each 24. The laminae are thin on their 
anterior, thick on their posterior edges, overlapping a little but 


VOL. VII. 4 


50 PROFESSOR STRUTHERS. 


uowhere in contact. The spines are mere rudiments on the 
3rd, 4th, and 5th, on the 6th half an inch in length, on the 7th 
Jonger. 

18. Bopies AND THEIR FIBRO-CARTILAGES.—There was 
very little motion between the cervical vertebrae in any direc- 
tion, the motion, small as it is, becoming more limited forwards. 
The rotatory motion taken at the zygomal processes was not 
over ;+ inch in extent. The thickness (length) of the fibro-car- 
tilages, not including the rim of cartilage belonging to the epi- 
physes, was—behind 2nd dorsal, } inch; behind 1st dorsal and 
7th cervical, each } inch; behind 6th, 5th, and 4th cervical, 4th; 
behind 8rd, over 4; behind 2nd, } inch. The bodies of the 3rd, 
4th, and 5th vertebrae are from 4 to 2 inch in thickness, those 
of the 6th and 7th are about } inch more. When divided, the 
fibrous part was seen to reach half an inch inwards above, a 
little more below, dipping in to ? inch at the middle line above 
and below, the measurements of the body surfaces being,— 
breadth, at the third 44 inch, at the sixth $ ich less; height, 
at the third 23, at the sixth + inch more. Before and behind 
the 3rd, although the pulp surfaces are as extensive as at the 
other spaces, there was very little pulp, owing to the nearness 
of this vertebra to those next it. The body surfaces are, espe- 
cially in the transverse direction, a little convex in front, a little 
concave behind, better marked the farther forwards. The mid- 
dle of the anterior surface of the bodies, corresponding to the 
centre of the pulp, is rough, rising into a faint prominence on 
the posterior vertebrae. The epiphyses want from % to $4 inch 
of reaching the edge of the bodies. The thinnest part of the 
body is at each side of the pulp space, so that, in the macerated 
bones, the sides of the bodies come in contact, leaving thin 
spaces between the surfaces, deepest at the middle line, both 
above and below; this being also due in part to the curvature 
of the bodies just mentioned. 

19. ARTICULATIONS OF THE ATLAS AND Axis.—(a) The 
articular surfaces are continuous across the middle line below, 
forming one great horse-shoe cartilaginous and synovial surface. 
There was a faint median depression in the cartilage, though 
no interruption to its continuity, and through the dried carti- 
lage is now seen a median furrow on both bones, which in fully 


CERVICAL VERTEBRAE IN FIN-WHALES. ot 


macerated bones has led to the surfaces being reckoned dis- 
tinct. The odontoid rises higher than in the Razorback and is 
at the same time broader and less abrupt, the whole area rising 
gently to a rounded prominence, the summit of which is near 
the upper part, towards the transverse ligament. (b) The 
transverse ligament (see Fig. 6) is, in this whale, flattened in 
the same direction as in man, but is somewhat prismatic ; upper 
surface flat and a little concave transversely, forming part of 
the floor of the spinal canal, in line with' the body of the axis; 
anterior border thick; lower surface applied obliquely against 
the odontoid process, which in this whale rises a little above 
_ the level of the ligament. But the transverse ligament is not 
free; its hinder edge joins the periosteum on the upper surface 
of the axis, and its lower surface is attached to the odontoid 
as it crosses. 

(c) The check ligaments (Atlo-odontoid) have essentially the 
same connections and function as I have described in the Razor- 
back, but from the greater prominence of the odontoid, and the 
greater width of this division of the ring of the atlas, they are 
less interosseous in position, and their fibres have less of a for- 
ward direction. Its attachment on each side to the atlas is to 
the narrow crescentic surface between the posterior articular 
surface and the edge of the ring, meeting its fellow in the 
middle line below; while its upper fibres, attached inwardly 
to the upper aspect of the odontoid, outwardly to the atlas, 
are continuous with the deeper fibres of the transverse liga- 
ment, giving that ligament its connection to the odontoid. 
In the Razorback the lowness of the odontoid required these 
fibres to pass forwards some distance to reach the posterior sur- 
face of the transverse ligament; while here the transverse liga- 
ment is so close that, besides attaching ligamentous fibres from 
the odontoid, it gives direct support to that process, its surface 
being adapted accordingly. Concealing, from before, the lower 
part of the check ligaments, there is a fibrous structure like an 
inferior transverse ligament, embracing semicircularly the odon- 
toid below as the transverse ligament does above. The deeper 
part of this inferior fibrous girdle is continuous with the lower 
part of the check ligaments, as the transverse ligament is with 
their upper part, and superficially it forms a fibrous cushion, 


4—2 


52 PROFESSOR STRUTHERS. 


levelling up the lower part of the odontoid division of the ring 
of the atlas and giving a soft flooring for the inner part of the 
occipital condyles, which here project inwards beyond the inner 
edge of the atlantal cups. Between these two fibrous girdles 
the odontoid is seen for a breadth of ? inch, with the remains 
of the ligamentum suspensorium attached to it. 

(d) I was able to examine the superficial ligaments be- 
tween the atlas and axis more satisfactorily here than in the 
great Finner in which they had been injured. Below, besides 
the inferior longitudinal ligament of the bodies, a strong inferior 
oblique ligament, serial with the inferior inter-transverse liga- 
ments but much stronger, passed inwards and forwards to the 
atlas from the transverse process of the axis. There was a well 
marked capsular ligament round the outer side of the articular 
surfaces, also well marked on the inner side of these surfaces, 
above the position of the transverse ligament, but below this 
identified with the check ligaments. Above, besides the inter- 
spinous ligament, the interlaminar ligament was a strong mem- 
brane, leaving between it and the upper end of the articular 
surfaces a space for the passage of the second nerve; and a 
strong ligament (superior inter-transverse) passed from the up- 
per transverse process of the axis to the transverse process of 
the atlas, serial with the superior inter-transverse ligaments 
behind. On either side of this ligament was a space from 
which the contents had been removed, apparently the spaces 
through which the superior and inferior divisions of the second 
nerve had passed. 

Notwithstanding the extent of the synovial articular sur- 
faces between them, the motions of the atlas on the axis were 
very limited. Vertical and lateral gliding motions were not 
over ;1, inch in extent. Rotatory motion was checked before 
the tip of the transverse process (which is 4 inches from the 
centre of rotation) had moved 4 inch. All the external lga- 
ments help to check, but after their division the rotatory mo- 
tion remained as limited in extent as before, and as long as the 
check ligaments were undivided the atlas could not be lifted 
3 inch off the axis. These extensive synovial surfaces, usually 
in other animals an adaptation to extensive motion, must, 
therefore, be regarded as rudimentary here, so far as their 


CERVICAL VERTEBRAE IN FIN-WHALES. 53 


synovial condition is concerned. The amount of yielding and 
elasticity is not greater than that which fibro-cartilages allow, 
and their retentive power is inferior. 

20. OccIPITO-ATLANTAL SURFACES.—On applying the atlas 
on the occipital condyles, the lower end of the condyle is seen 
to project about an inch (1 part of the whole length) below the 
cup, and this part of the condyle is more abruptly curved. The 
atlantal cups are seen to project a little laterally beyond the 
condyles, while the condyles approach each other internal to the 
cups, so that over half an inch of the breadth of each condyle is 
seen through the odontoid division of the ring of the atlas. 
The distance to be traversed by the ligamentum suspensorium, 
from the odontoid to the fissure in the floor of the inter-condy- 
loid fossa, does not exceed half an inch. The neural division of 
the canal of the atlas corresponds in form to the foramen mag- 
num but is somewhat larger, being transversely nearly 23, verti- 
cally 12, while the foramen magnum measures transversely 23, 
vertically 18. The odontoid division of the canal is relatively 
much wider than in the great Finner, measuring transversely 
at its widest part near the transverse ligament 18, vertically 12 
inch; total height of canal of atlas 32. The position of the 
transverse ligament is a little below where the distinction of the 
two parts of the canal appears on the bone. The transverse 
foramen for the atlantal nerve, between the lamina and the 
upper end of the cup, is completed by bone on both sides, and 
is about the size of a goose-quill. 

21. EXPLANATION OF THE DRAWINGS. Plates I and II.—The 
drawings are from photographs kindly taken by my pupil, Mr J. 
Shearer. The outlines taken from these were carefully filled in 
and shaded from nature, in my presence, by Mr Gibb. In 
taking the first three views the vertebrae, built up on the table, 
were separated to the extent to which they are naturally sepa- 
rated by their fibro-cartilages. The two front views are placed 
above and below for more ready comparison. 


Fig. 1. Under-aspect of the cervical vertebrae of the Peterhead 
Razorback. The distinction between the tubercular and nerve-groove 
stages of the inferior transverse processes is well marked. It shows 
a deep atlo-axoid articulation ; greater length of the left than of the 


D4 PROFESSOR STRUTHERS. D 


right wing of the axis; a great development of the tubercular stage 
of the inferior transverse process of the 3rd; a nearly symmetrical 
deficiency at the rerve-groove stage of the inferior transverse pro- 
cesses of the 6th; traces of the posterior body epiphyses of the 6th 
and 7th vertebrae, indicating that the animal, though of full length, 
was not quite mature; We. 


Fig. 2. Under aspect of the cervical vertebrae of the Stornoway 
Razorback. Comparing this fig. with Fig. 1, it shows a different 
form of transverse processes of atlas ; a shallow atlo-axoid articula- 
tion ; a more projecting sub-central process of atlas ; the wings of the 
axis greatly developed, backwards to the level of the tip of the trans- 
verse processes of the 7th, and also downwards so as to show part of 
the surface of the wing, while in Fig. 1 only the border is seen. The 
undulations of the wing are seen, the internal eminence, opposite the 
tip of the transverse process of the atlas, is just external to the ring. 
The rudimentary state of the inferior transverse processes of the 6th, 
although it was a mature animal, will be observed ; also the distinc- 
tion between the tubercular and nerve-groove stages on the inferior 
processes of the 3rd, 4th, and 5th vertebrae, though not so strongly 
marked as in Fig. 1; and the transverse process of the 5th is seen to 
be horizontal, and, after that of the axis, the longest and the most 
projecting. 


Fig. 3. Upper aspect of the cervical vertebrae of the Wick Ra- 
zorback. On the atlas is seen the oblique ridge roofing the foramen 
in front, and supporting an articular process behind; the two grooves 
proceeding from the foramen ; and some a-symmetry of the transverse 
processes. The axis shows great development of the crests in the 
region of the spine (still more developed in the Peterhead specimen), 
the right crest articulating with the atlas. On the wings, opposite 
the tip of the transverse process of the atlas, is seen the strongly 
marked and turned forward inter-transverse tubercle, and, bounding 
the concavity external to it, the upper angle of the wing, farther out 
on the right than on the left side in this specimen. The superior 
transverse processes appear thin at their inner third (nerve-groove 
stage) from the direction of their surfaces, and external to this (tuber- 
cular stage) are seen to be bevelled and rough, and to begin to over- 
lap. The inferior processes are seen in deep shading beyond, that of 
the 3rd of great size on the left side, that of the 6th complete on the 
left side, and partly deficient on the right. On the laminae of the 
five posterior vertebrae are seen the very rudimentary spines ; the 
more developed anapophysial processes, serial with the crests of the 
axis; and the partially cribriform condition of some of the laminae 
from their extreme thinness near their anterior margins, in this ma- 
ture or aged animal, 


Fig. 4. Front view of fifth cervical vertebra of the Stornoway 
Razorback, On the body, externally, is seen the streaked ring where 
the capsular part of the fibro-cartilage is attached, and within this 
the figure-of-8 surface where the pulp lies, somewhat raised at the 


CERVICAL VERTEBRAE IN FIN-WHALES. 55 


middle. The laminae show, about the middle, a non-symmetrical 
anapophysial process ; the spinal canal is intermediate between the 
triangular form presented by the Peterhead specimen, and the ellip- 
tical form presented by the Wick specimen. Spine also intermediate 
in length. On the lower transverse process (a) is the root stage; 
(6) the tubercular stage, with its outer and inner angular promi- 
nences; (c) the nerve-groove stage, the groove seen to be directed 
obliquely outwards and downwards. On the upper processes, (d) is 
the nerve-groove stage. At the inner part of this a small tubercle is 
seen (as it so happens, unusually developed on the left side of this 
vertebra) on the upper edge, the grooving being on the surfaces; (e) is 
the tubercular stage, marked off from the nerve-groove stage by an 
angular elevation, but the tubercular character of this stage, owing to 
the bevelling, is visible only in an upper view (see Fig. 3); (7) is the 
third stage of the superior process, the terminal plate, developed in 
the mature animal. It is seen to be less developed on the right side, 
the upper angle not having been yet formed. The rings in this spe- 
cimen present the semi-oval form, 


Fig. 5. Hinder aspect of the atlas of the Wick Razorback, 
The transverse ligament (a), flattened in the opposite direction to 
that of man, is seen dividing the canal into an upper or neural di- 
vision, and a lower, odontoid or ligamentous division. On each side 
of the lower division is seen the crescentic surface where the atlo- 
odontoid check ligament is attached, unsymmetrical in this specimen. 
The articular facet, by which it articulates with the right crest of the 
axis, is seen on the right lamina above the groove for the nerve- 
escape. The internal inter-transverse tubercle, at the upper edge of 
the root of the transverse process, is seen to be more developed on 
the left side, and the external tubercle, behind the tip of the process, 
to be also unsymmetrical. A median groove is seen dividing the 
articular surface into two, but the two are naturally continuous, the 
presence of the groove being exceptional in this specimen. 


Fig. 6. Front aspect of atlas of young (144 feet long) Pike 
Whale (B. rostrata). Transverse ligament (a) not flattened as in the 
Great Finner, but prismatic. Both upper and odontoid divisions of 
canal are proportionately wider than in the Great Finner, Median 
groove seen between condyloid cups, where ligamentous septum is 
attached. Transverse foramen for atlantal nerve already roofed over, 
Transverse processes incompletely ossified, but the twist is seen. 


NOTICE OF QUADRUPLE MAMMA, — THE LOWER 
TWO RUDIMENTARY,—IN TWO ADULT BRO- 
THERS. By P. D. Hanpysipe, M.D., F.RS.E., Teacher 
of Anatomy, Edinburgh School of Medicine. (Plate 3.) 


ONE of my pupils,—say A. B.—20 years of age, 6 feet 4 inch 
in height, of active habits, lean but muscular, well-formed and 
healthy, presented himself to me in February, 1872, having four 
mamme on his chest, the two lower of these being rudiment- 
ary (Fig. 1). He is the eldest of a family consisting of five 
males, and was a forceps-infant in the hands of Dr Hewit of 
Lauder. The Mamme Proper are normally situated, are exactly 
four inches distant from the mesial line, and are more fully 
developed than usual. The right mamma is seven-eighths of 
an inch in its long axis, which runs downwards and outwards, 
and six-eighths of an inch vertically; around its prominent and 
rose-tinted mammilla two concentric rugee appear, and on the 
inner and upper half of the periphery of a dark areola are seven 
prominent papillae (Fig. 2). The left mamma is also seven- 
eighths of an inch in its long axis, which likewise runs down- 
wards and outwards, but it is only five-eighths of an inch in 
extent vertically ; it also presents two nearly concentric ruge, 
and on the upper and outer two-thirds of its pale areola are 
one or two less prominent papilla (Fig. 3). The Lower Mamme 
are situated exactly three inches from the mesial line. The 
right, however, is 23 inches below the right mamma proper, and 
‘81 inches from the umbilicus, while the left is placed 3 inches 
below the left mamma proper, and 8! inches distant from the 
umbilicus. Again, the right lower mamma is ovate in form, 
with its base towards the umbilicus; its long axis is, therefore, 
downwards and inwards, and in length is one-quarter of an 
inch; its vertical axis being one-eighth of an inch. Its mam- 
milla is one line in diameter, round in form, and bilobed; the 
septum between the adherent elliptical lobes or nipples running 
in the long axis of the areola. This areola is of a light pink 
colour, and consists of thin delicate skin (Fig. 4). The left lower 
mamma is elliptical in shape, with its long axis placed trans- 


_ 


QUADRUPLE MAMM#Z IN TWO ADULT BROTHERS. 57 


versely. Its long axis is a quarter of an inch, while its vertical 
or short axis is one-eighth of an inch, in extent. The mammilla 
consists of two distinct elliptical elevations, which he parallel to 
each other, and in the long axis of its areola, and these eleva- 
tions -or nipples are each of one line in length and half-a-line 
in breadth (Fig. 5). In A. B.’s figure the distance from the 
serobiculus cordis to the umbilicus is 7} inches, while that 
from the umbilicus to the root of the penis is seven inches 
The umbilical cicatrice is elongated transversely, and unequally 
bilobed, and there is a trace of double linea alba below the pre- 
cordia. His genital organs are fully developed and natural. At 
puberty his mamme proper enlarged to an unusual size. 

In the second son of this family, 18 years of age, the mam- 
mee at puberty were so much developed, and discharged the 
usual milky fluid so freely, that “Dr Turnbull of Dunbar,” as I 
am informed by letter, “had to employ means to reduce them 
sufficiently to prevent their forming an impediment in his exa- 
mination on entering the navy.” 

The third son of the same family,—say C. D.,—who is 
17 years of age, and five feet ten inches in height, has also 
QUADRUPLE MAMM#. The mamme proper are placed as. usual, 
but are more fully developed. They are situated 84 inches 
apart. The lower or rudimentary mamme are 74 inches apart. 
The right lower ‘is placed 24 inches below the upper right 
mamma, and 8} inches from the umbilicus; while the left 
lower is 3 inches below the upper left mamma and 8 inches 
distant from the umbilicus. This left supernumerary mamma 
is distinctly marked, but the opposite right one is merely 
indicated by a white puckered spot of skin. 

No similar abnormality is known to have existed in their 
parents’ family on either side. The mother of these young 
men is 5 feet 4 inches in height, of a delicate habit of body, 
and all her five infants were nursed on cows’ milk alone. Their 
father, aged 62, is in height 6 feet 1 inch, and belongs to a 
long-lived family of from 70 to 80 years. He writes to me 
that all his “ five sons have so far proved themselves to be big, 
strong, muscular and masculine, far beyond their years, that 
there is the very opposite of any evidence of the blending of 
the sexes in their case, and that on the contrary they are mas- 


58 DR HANDYSIDE. 


culine to a degree.” He goes on to say—what may not be 
irrelevant to the matter in hand—“my grandfather used to 
narrate to me a tradition of a notable ancestor, a man of un- 
usual strength of body and an armourer by trade, who resided 
near Glammis in Forfarshire, and was said to be double-jointed 
in all his members, and to have forged Wallace’s sword. He is 
understood to have formed the type of ‘Hall o the Wynd’ in 
Scott's Fair Maid of Perth.’ Farther, in morphological con- 
nection with the question, it may be added, firstly, that vari- 
cocele exists in the second son, in his father, and also in his 
paternal uncle, and existed in his grandfather and his 
great-grandfather ; and secondly, that no twin births are known 
to have occurred in this family on either side. 


REMARKS. 


(1) Are these elevated spots mamme? Truly such a doubt 
no unprofessional eye has even suggested, and a mere pro- 
fessional glance, or that with a lens of low power, reveals dis- 
tinctly characteristic areolas, well-marked cutaneous glands, 
and tubercles, and nipples. Did opportunity offer, a mercurial 
injection would probably flow along as in the male mamma 
generally. These- observations apply, possibly with greater 
force, to the well-defined structures and parts so carefully 
marked by Dr Arthur Mitchell in a case of quadruple mamme, 
which I now proceed to notice. 

(2) Parallel case. I cannot trace on record an instance of 
a supernumerary Mamma, Areola, Tubercle, or Nipple in the 
male; but in notes that I took on Sept. 18, 1872, of a conver- 
sation that I then held with Dr Arthur Mitchell on the case of 
A. B., he stated that some years previously, while in Glen- 
Urquhart, Inverness-shire, he came upon a farm-servant, a male, 
about 27 years of age, who had just sustained contusions from 
a fall; in examining whose chest, Dr M. observed four mam- 
me, the two lower less developed than the upper, situated about 
a hand’s-breadth from them, and equally distant with them from 
the mesial line. Zhe wpper mamme were normal. The lower 
were less prominent; they presented faint areolas, and the 


QUADRUPLE MAMM IN TWO ADULT BROTHERS. 59 


usual tubercles were as faintly marked; but they presented 
well-marked nipples. The man was stalwart, handsome, and of 
a muscular frame; he was well bearded, had testes and a manly 
voice. He was well-conditioned mentally. No similar ab- 
normality, and no supernumerary digits or other malformation, 
had existed in his family, so far as could be learnt. Dr Mitchell 
communicated these particulars to me from memory, but was 
satisfied as to their substantial accuracy. Very shortly after 
the case came under his observation, he spoke of it to Professor 
Turner, through whom I was led to apply to him. 

(3) Blending of the sexual features. There is a slight 
approximation to the female proportions in the position of the 
umbilicus in A. B.’s figure; for, whereas the usual propor- 
tional distance in the male sex between the preecordia and the 
umbilicus is one-fifth longer than that between the umbilicus 
and the root of the penis, and in the female the space between 
the precordia and the umbilicus is one-fourth shorter than it 1s 
between the umbilicus and the base of the mons,—we have 
seen that in the case before us, the distance between the pre- 
eordia and the umbilicus is only one-fifteenth greater than is 
the distance between the umbilicus and the root of the penis. 

(4) Since it is admitted, as a teratological law, that like 
parts of two unequal bodies, the autosite and the parasite, are 
always attached near one another, it may be well, in examining 
the question of arrested twin development, not to exclude from 
consideration such cases of supernumerary mamme ; and the 
' peculiarity in the form of the umbilicus in A. B. may not be 
overlooked in connection with this remark. 


CONTRIBUTIONS TO THE ANATOMY OF THE INDIAN 
ELEPHANT (ELEPHAS INDICUS), PART II. URI- 
NARY AND GENERATIVE ORGANS’. By M. Watson, 
M.D., Demonstrator of Anatomy in the University of Edin- 
burgh. (Plate +.) 


ALTHOUGH the urinary and generative organs of the Indian 
elephant have been described by various authors who have 
examined them in whole or in part, yet these descriptions differ 
so much from one another that it may not be altogether super- 
fluous to put on record the results of my own observations on 
these parts of the animal, more especially as there is no system- 
atic account of them to which the exclusively English reader 
can refer. 

I shall in the first place consider the urinary and in the 
second the generative organs, comparing the observations of 
different authors as we proceed. 

For the opportunity of dissecting these parts I have again to 
express my thanks to Prof. Turner. 


URINARY ORGANS. 


Kidney. This viscus measures one foot in length and seven 
inches in greatest breadth, thus differing materially in size 
from that examined by Stukeley’, which measured three feet in 
length: his measurement, however, I cannot avoid thinking, 
has been somewhat exaggerated. It is triangular in form, 
tapering toward the anterior, but thick and rounded at the 
posterior extremity. Its outer border is uniformly convex, 
except where it is interrupted by lines indicating the subdivision 
of the organ into lobes. The inner border is also convex, but 
presents about its centre a deep concavity—the hilus—for the 
entrance of the renal vessels and duct. These occupy the usual 
relative positions, the ureter being situated dorsally, the vein 
ventrally, and the artery between the two. The capsule, which 
is strong and composed of dense fibrous tissue, adheres so closely 


1 Part I. On the Thoracic Viscera, appeared in this Journal, November, 1871. 
2 Hssay towards the Anatomy of the Elephant, Lond. 1723, 


ANATOMY OF THE INDIAN ELEPHANT. 61 


to the kidney as to allow of its subdivision into lobes being seen 
externally ; it is however readily separable from the contained 
organ. On separating the capsule many vessels of large size are 
to be seen passing from the substance of the kidney into and 
through the capsule, demonstrating in this animal perforating 
arteries which in all probability communicated with the parietal 
branches of the abdominal aorta in a manner similar to that 
described by Prof. Turner* in the human subject. With refer- 
ence to the number of lobes of the kidney, Camper’ states that 
there are eight or nine, Cuvier* reduces the number to four, 
whilst Mayer* observes that it is composed of only two principal 
lobes. Do6nitz® ascertained the number to be ten in that of 
the African elephant. In my own specimen the number of 
lobes in the left kidney was five, and these could be readily 
separated from one another without any laceration of the renal 
tissue. The number in the right kidney was unfortunately not 
observed in the same satisfactory manner, but, judging from the 
primary divisions of the ureter, which in the left kidney cor- 
responded in number to the lobes, there would be four. On the 
surfaces of both kidneys, moreover, indications of a farther 
subdivision into smaller lobes were observed, but these were not 
traceable to any depth without rupture of the renal substance. 
It is however probable, I think, that these lines of separation 
indicate the subdivision of the kidney into lobes in the young 
animal, and that they become less and less distinct as the 
animal grows, and may finally be obliterated altogether. That 
- the lobes are originally distinct, as in many animals, is proved 
by Camper’s® dissection of a young specimen, in which he found 
that the lobes were only beginning to unite toward the exterior 
of the organ, their inner or apical extremities being altogether 
free. If this view be correct, it will explain the diversity of 
statement of different authors with regard to the number of the 
lobes. 


Each of these lobes is to be regarded as a renal organ 


1 Brit. and For. Med.-Chir. Rev. July, 1863. 

2 Description anatomique dun éléphant male. 

3 Lecons @anatomie comparée, Paris, 1805. 

* Nova acta acad. Ces. Leo. Car. xxi. 

5 Reichert und Du Bois-Reymond’s Archiv, 1872, p. 85. 
5 Description anatomique d’un éléphant male. 


62 DR WATSON. 


complete in itself, possessing as it does a perfect system of 
tubes which do not intermingle with those of the neighbouring 
lobes. The kidney of the elephant thus presents an approach 
in structure to that of the cetacea and other aquatic mammals, 
differing however in this, that whilst in the latter the lobes are 
permanently distinct, in the former they are distinct during 
youth, but become more intimately blended as the animal 
attains maturity. With reference to the more minute structure, 
Cuvier states that there is no distinct line of separation between 
the cortical and medullary substances, while on the other hand 
Donitz found the distinction between them as well marked as 
in the majority of animals. My own observations agree with 
those of Cuvier, at the same time it must be borne in mind 
that these kidneys had been subjected to the action of spirit for 
some time, which may have rendered the distinction between 
the substances less apparent to the naked eye than would 
otherwise have been the case. The tubes of Bellini do not 
terminate on papillae as asserted by Cuvier, Mayer, Hunter’ 
and Owen’, but upon the flattened truncated extremities of the 
calyces in a manner which will be more particularly referred to 
along with the ureter. 

The renal artery divides into three main trunks, each of 
which again subdivides into numerous branches, which enter the 
substance of the kidney. Before dividing, the trunk of the 
renal artery, as observed by Camper, gives off the spermatic 
artery to the testicle—an arrangement by no means uncommon 
in the human subject. The veins passing from the kidney are 
five in number, but whether they terminate in a common trunk 
or open separately into the posterior cava, could not be deter- 
mined by reason of the organs having been removed from the 
abdomen. The calyces differ in number in the two kidneys, ten 
in the right, and thirteen in the left. It would thus appear 
that the number of calyces bears no constant ratio to that of 
the lobes, some of these being provided with two, and others 
with three calyces. The calyces are in the form of flattened 
tubes, terminating in a truncated flattened extremity, in the 
centre of which is to be observed a single aperture of large size. 


1 Essays and Observations, by Owen. 
2 Anatomy of Vertebrates, 111. 


ANATOMY OF THE INDIAN ELEPHANT. 63 


On slitting open this aperture, it is seen to lead into a cribriform 
vault, the cribriform appearance of which is due to the openings 
of the tubules of Bellini. According to Dénitz, in the kidney 
of the African elephant, the aperture just described leads into 
an elongated central canal—the tubus maximus of Hyrtl’, along 
the course of which the tubules of Bellini open: but in that of 
the Indian elephant this is not the case, as the central tubules 
of each calyx are prolonged downwards so far as to be little 
shorter than those of the periphery, which gives rise to the 
appearance of a shallow vault rather than to that of an elongated 
central canal such as is figured by Dénitz. 

With the exception of Camper, as already stated, all the 
older writers have been deceived as to the existence of papille 
in the kidney of the elephant. 

According to Cuvier and Mayer the ureter is formed by 
the union of three principal tubes. In the present specimen 
the ureter of the right side is formed by the junction of four 
and that of the left by five. In both kidneys these tubes 
emerge separately from the hilus. In the case of the right 
kidney the two anterior tubes unite to form one half of the 
ureter, the second half being formed by the junction of the two 
posterior tubes ; whilst in the left kidney the one half is formed 
by the junction of the three anterior, and the second half by 
the junction of the two posterior. As regards the number of 
calyces opening into each of these tubes, enumerating them 
from before backwards, we find that in the right kidney the 
first receives two, the second two, the third three, and the 
fourth three, in all ten; in the left kidney the first receives two, 
the second two, the third three, the fourth three, and the fifth 
three, in all thirteen. The want of symmetry as regards the 
number of these tubes in the kidneys of opposite sides seems to 
indicate that this is not constant, which would account for the 
difference between my own observations and those of the 
authors already quoted. 

There is, as stated by Cuvier’, no pelvis properly so called, 
the tubes simply uniting without marked dilatation to form the 
ureter. This tube passes backwards so as to reach the posterior 


1 Denk. der Acad. der Wissenschf. Wien xxxi. 1872. 
2 Lecons d’anatomie comparée. 


O64 DR WATSON. 


wall of the bladder, being invested on its under surface by peri- 
toneum ; as it passes between the bladder and rectum, it lies 
directly above the corresponding vas deferens. Having reached 
the back of the bladder, the two ureters are separated from one 
another by a distance of 34 inches. They then pass obliquely 
through the wall of this viscus for a distance of 2} inches, and 
open close together near the neck of the bladder. The very 
oblique and lengthened course of these tubes through the wall 
must form a thoroughly effective valve against the backward 
passage of the urine. - 
Bladder. This viscus is by no means so large as one would 
expect in an animal of such size. It is regularly oval in 
form, and occupies the usual position. Above it is the rectum, 
the vesicule seminales, and vasa deferentia intervening; 
whilst at the neck are to be observed the small prostatic 
glands, two in number on each side. The whole of the 
bladder, with the exception of the triangular interval, in- 
dicated by the points of contact of the ureters with the ex- 
terior of the bladder and the neck of the viscus, is completely 
invested by peritoneum, this investment on the lower aspect 
reaching as far back as the commencement of the urethra. 
The peritoneum covering the bladder presents moreover three 
well-marked folds or ligaments. Of these, one passes off from 
the inferior aspect of the viscus, and seems to correspond to 
that described by Camper’ as attaching the bladder to the 


pubis. The author just mentioned observed in it the urachus, - 


but no remnant of that structure could be discovered in the 
specimen under description, its absence being in all probability 
due to the age of the animal. The other two folds pass off 
from the lateral aspects of the bladder, and like the lower fold, 
each is composed of a double layer of peritoneum, and encloses, 
moreover, an artery which was still pervious in certain parts 
of its course. This artery is probably the hypogastric, but 
neither this nor the points of attachment of the peritoneal 
folds to the abdominal wall could be accurately ascertained, 
the parts having been removed from the pelvis. 

On slitting open the bladder, the apparent thickness of its 
wall is seen to be due rather to the peritoneum, and sub- 


1 Description anatomique dun éléphant male. 


; 
4 


| 


page” 


ae eee ee 


Se eee ee ee ee ee 


ANATOMY OF THE INDIAN ELEPHANT. 65 


peritoneal connective tissue, than to the proper muscular coat. 
The mucous membrane is uniformly smooth, and is not thrown 
into folds, except at the neck of the viscus, where it forms a 
single median fold of large size, which, commencing between 
the openings of the ureters, passes forwards to terminate on 
the floor of the urethra close to the base of the veru-mon- 
tanum, 


GENERATIVE ORGANS. 


Testicle. This organ, which is almost globular in form, lies 
as figured by Camper’ inferior to the posterior extremity of the 
kidney. It is entirely invested by peritoneum, with the 
exception of its upper and external margin where the vessels 
enter. The peritoneum, when traced outward, is seen to be 
reflected from the surface of the testicle, and to pass between 
it and the epididymis, the lower surface of which it also covers. 
The manner in which the peritoneum attaches the testicle to 
the posterior extremity of the kidney, notwithstanding that it 
permits of a slight degree of mobility of the former, altogether 
negatives the suggestion of Mayer’, that this organ descends to 
the perineum during the period of rut. The epididymis les 
along the outer side of the testicle and not the inner, as stated 
by some authors. 

The spermatic artery, as before said, is given off from 
the trunk of the renal, it passes backward, and after a short 
course divides into four or five branches, which supply the 
organ. It gives, moreover, several branches of small size to 
the epididymis. The testicle also receives some branches of 
small size from arteries situated in front of it, but the exact 
origins of these could not be ascertained. 

The veins leaving the testicle are remarkable for their 
number and large size. They are seven or eight in number, 
and communicate freely with one another, as also with neigh- 
bouring veins, so as to form a plexus close to the testicle. They 
finally unite to form two large trunks which open into the vena 
cava on the right side. The valves are very numerous in the 
veins composing this plexus. 

1 Description anatomique dun éléphant male. 
? Nova acta Acad. Ces. Leo. Car, xxu. 


VOL. VII. 5 


66 ; DR WATSON. 


Several nerves of large size pass to the testicle along with 
the vessels. 

The excretory ducts of the testicle (vasa efferentia) are ten 
or twelve in number, they pass off from the anterior extremity 
of the hilus of the testicle, and diverging as they pass outwards, 
enter the epididymis. With regard to the extent of this struc- 
ture, it is impossible to say where it terminates, or where the 
vas deferens begins, as the vas does not form a flexure upon 
the epididymis as in those animals in which the testicle de- 
scends into a scrotum, but is simply continuous without inter- 
ruption with the epididymis. The anterior extrenity of the 
epididymis (Globus major) is the widest part, and on tracing 
it back we find that the size of its loops gradually decreases, 
so as to become continuous with those of the vas. 

As regards this tube, with the exception of 5 inches previous 
to its termination, it is seen to be convoluted in the whole of 
its course. Its central portion. is less convoluted than either 
of its extremities, and the anterior less so than the posterior; 
the latter extending from the peritoneal fold which unites the 
vasa of opposite sides down to the point where it becomes quite 
straight, being in fact so extremely convoluted as to resemble 
rather a second vesicula seminalis, than a portion of the vas 
deferens. Throughout the whole of this part of its course each 
vas is attached to the superior abdominal wall by a double fold 
of peritoneum, which forms as it were a ligament for it. The 
ligaments of opposite sides become continuous with one another 
between the bladder and rectum, and thus form a fold corre- 
sponding in position to the broad ligament of the uterus in the 
female. The vesicula prostatica, however, does not extend 
into this fold, as it does in the goat, ass, &e. With reference to 
the straight or terminal part of its course, each vas lies between 
the corresponding vesicula and the upper surface of the bladder, 
and before opening into the urethra dilates abruptly into an 
ampulla two inches in length, which is closely connected to its 
fellow of the opposite side. Finally, the vas unites with the 
efferent duct of the corresponding vesicula, the common ejacu- 
latory duct thus formed opening into the urethra. 

Whilst Cuvier’s observations on these parts agree with my 
own, Owen, on the other hand, states that after dilating into 


ANATOMY OF THE INDIAN ELEPHANT, 67 


the ampullee, the vasa open “into the urethra distinctly from 
the vesicular glands.” 

Vesicule seminales.—These have been figured by Camper, 
and described by Cuvier. By the other writers on the ele- 
phant they are omitted. Each vesicula is an elongated sac six 
inches in length and one inch and a half in diameter, and occu- 
pies the interval between the bladder and rectum, being sepa- 
rated from the former by the ampulla of the corresponding vas 
deferens. Its inner surface is in contact throughout with that 
of the opposite side, whilst its base comes into relation with the 
peritoneal fold connecting the vasa deferentia. Each is in- 
vested by a thick layer of muscular fibre, which is continuous 
with that surrounding the membranous part of the urethra, the 
fibres diverging from their urethral extremities, so as to enclose 
each vesicula in a complete capsule. On slitting open the vesi- 
cula it is seen to be lined by a thick membrane, which is 
thrown into decussating folds throughout the greater part of its 
interior, so as to give rise to an appearance resembling the in- 
terior of the ventricles of the human heart. Towards the 
urethral extremity of the sac, however, these folds become pa- 
rallel and uniformly longitudinal in direction. A transverse 
fold of large size separates the base of each from the body of the 
sac, and so gives rise to an appearance of two compartments, 
-as described by Cuvier. Each vesicula terminates in a duct, 
which unites with the lower end of the corresponding vas to 
form the common ejaculatory duct, and finally opens on the 
side of the veru-montanum, and outside of the vesicula pro- 
statica, 

Prostate Glands. These little glands seem to have been 
observed by Duvernoi’, although he was not aware of their 
nature, for he appears to have been of opinion that the prostate 
in the elephant is represented by what is now known to be 
Cowper’s glands. Camper states that this gland in the ele- 
phant is the same as in other animals, but his figure is quite 
incorrect. Cuvier, however, describes them with tolerable 
accuracy. 

They are four in number, two on each side, and of small 
size. They are placed below, and somewhat to the outer side 


1 Comm. Acad. scient. Petropol. tom. 11. 


5—2 


68 DR WATSON. 


of the urethral extremities of the seminal vesicles, those of each 
side being closely applied to one another. In form they are 
oval, and the external one is the larger. It measures two 
inches in length, and one in greatest breadth, whilst the smaller 
or internal one is 1} inch in length, and half an inch in 
greatest breadth. They are invested with a layer of muscular 
fibre continuous with that which surrounds the seminal vesicles. 
On slitting them open each is seen to contain a central cavity 
of an oval form, lined with a membrane, which is thrown into 
well marked longitudinal folds, which converge toward the 
urethral extremity of the gland. From this extremity the duct 
of each, which is single, passes off to open into a depression 
on the floor of the urethra, on either side of the veru-monta- 
num, the openings of the ducts of each side being close to- 
gether. In consequence of the glands themselves being situated 
at some distance behind the point where the ducts open into 
the urethra, each of these runs in the wall of the urethra for a 
distance of an inch and a half. 

Cowper's Glands. Cuvier is the only author who describes 
these with any degree of accuracy. Camper states that they 
are present, but gives neither description nor figure, whilst Du- 
vernoi mentions a single gland of the size of a large apple, 
which, so far as one can make out from his description, evi- 
dently corresponds to one of Cowper's glands, although he him- 
self is inclined to regard it as the prostate. 

The glands, as usual, are two in number. They lie one on 
either side of the middle line of the perineum, and under cover 
of one of the perineal muscles, to be subsequently described. 
Each is oval in form, somewhat flattened, and measures 24 
inches in length, and 2 in greatest breadth. They are therefore 
of large size as compared with the prostates. As regards their 
structure each is composed of a number of cells or lacunze com- 
municating freely with one another; but no separation of the 
gland into two distinct portions with corresponding cavities, as 
described by Cuvier, could be made out. At the same time it 
is possible that the length of time the parts had been subjected 
to the action of spirit may have tended to obliterate these cavi- 
ties. From the anterior extremity of the gland a single duct 
measuring 3 inches in length, and sufficiently large to admit of 


| 


ne 


ANATOMY OF THE INDIAN ELEPHANT. 69 


the passage of a crow-quill, passes off, and, running for the 
anterior two-thirds of its extent through the spongy substance 
of the urethral bulbs, opens finally into the floor of the bulbous 
portion of the urethra by a valvular orifice. The orifices of 
opposite sides are separated by a distance of half an inch. 

Cuvier states that each duct is formed by the junction of 
two smaller ones coming from the two portions of the gland 
above mentioned, but this arrangement cannot be traced in the 
present dissection. In addition to the perineal muscle con- 
cealing this gland and corresponding to the transverse muscle of 
Duvernoi, this author mentions another as being divisible into 
three distinct portions, and forming a capsule to the gland. 
This I failed to perceive. 

Urethra. ‘The membranous portion of this tube measures 
8 inches in length from the neck of the bladder to the bulb 
of the urethra. It is invested by a continuous layer one 
quarter of an inch in thickness of transversely arranged mus- 
cular fibres. Passing backward toward the neck of the bladder, 
these fibres are seen to become oblique in direction, and con- 
tinuous with’those which invest the prostate glands and the semi- 
nal vesicles. In addition to this layer of muscular fibres this 
portion of the urethra is surrounded by an investment of cellular 
erectile tissue, continuous in front with the spengy tissue of the 
bulb of the urethra, and measuring an inch in thickness in 
transverse section in front, but thinning off gradually toward 
the neck of the bladder. 
' On opening this part of the urethra a median elevation is 
observed on the floor, projecting from the point where the fold 
of mucous membrane described in connection with the neck of 
the bladder subsides, and to slope obliquely forward and upward, 
terminating in the margin of the vesicula prostatica. This 
margin is circular in form, and the vesicula itself forms a cul- 
de-sac extending to the depth of a quarter of 4n inch in the 
substance of the veru-montanum. It will be observed, there- 
fore, that it does not extend into the peritoneal fold connecting 
the posterior extremities of the vasa deferentia, as in many 
animals, but forms a mere shallow cul-de-sac, as in the cetacea. 
On each side of the veru is the slit-like orifice of the common 
ejaculatory duct. A well-marked fold of mucous membrane 


70 < DR WATSON. 


extends from either side of the veru-montanum obliquely for- 
ward and outward, and in the angle between this fold and the 
veru are to be observed the openings of the prostatic ducts— 
two in number on each side. On the floor of the urethra, in 
front of the veru, are two small orifices, resembling the open- 
ings of small glands, but no such structures could be discovered 
in connection with them. ‘The spongy portion of the urethra 
presents nothing worthy of note. 

The muscles of the penis are four in number on each side, 
three of these being situated on the lower aspect of the organ, 
and one, the levator, on the upper: Ist, The Levatores penis 
have been described by Duvernoi and Cuvier, and figured by 
Camper. They do not arise, as stated by the two latter au- 
thors, from the pubis, but from the upper and lateral aspect of 
each corpus cavernosum, close to its attachment to the ischium, 
as well as, and principally from, the tuberosity of that bone. 
Each is a powerful muscle measuring 4 inches in breadth at its 
origin, where it rests upon the dorsal vessels and nerves of the 
penis, but narrows rapidly to its extremity, where it ends on a 
thick rounded tendon. This tendon unites with the corre- 
sponding structure of the opposite side at the junction of the 
posterior and middle thirds of the penis, the two together 
forming a single median tendon common to the two muscles, 
which is inserted, according to Camper, into the glans penis. 
As regards this point, however, I could not satisfy myself, as 
that portion of the organ was reserved as a Museum preparation. 
The common tendon as it passes along the dorsum is confined 
within a strong aponeurotic sheath, which is continuous with, 
and formed by, the tunica albuginea of the corpora cavernosa. 
As it lies in the sheath it is connected to the walls by a very 
lax connective tissue, which evidently permits of a consider- 
able amount of motion of the tendon within its canal. 

Of the muscles met with on the lower surface of the penis, 
the ischio-cavernosus lies to the outer side, the bulbo-caverno- 
sus to the inner, and the compressor of Cowper’s gland between 
these two. In order to an accurate description of these muscles, 
it may be as well to refer briefly in the first place to the 
perineal fascia. On removing the skin and superficial fascia 
from the region of the perineum the deep or proper perineal 


ANATOMY OF THE INDIAN ELEPHANT. 71 


fascia is seen to be of great strength, and forms a general cover- 
ing to all the muscles of this region. It is attached on each 
side to the external margin of the corresponding crus penis, and 
is prolonged forward so as to form a distinct covering to the 
corpora cavernosa. From the deeper aspect of this portion of 
the fascia on each side of the middle line two processes of great 
strength dip down to be attached to the floor of the perineum. 
Of these, one intervenes between the ischio-cavernosus and 
compressor of Cowper’s gland on the outer, and the bulbo- 
cavernosus on the inner side; whilst the other separates the 
ischio-cavernosus and compressor from one another. From 
this it will be seen that each of the perineal muscles is enclosed 
within a distinct fibrous capsule formed by the perineal fascia, 
and from certain of the aponeurotic septa just mentioned differ- 
ent muscles take their rise. 

The zschio-cavernosus muscle is described by Cuvier as 
consisting of four distinct portions, but these I failed to recog- 
nise. It is a muscle of no great size, and possesses two distinct 
origins. The posterior of these having been removed from its 
attachment could not be seen, but in all probability it corre- 
sponded to the usual origin of this muscle from the ischial tuber. 
Tbe anterior portion of the muscle, which is however quite con- 
tinuous with the posterior, takes origin from the outer side of 
the dilated extremity of the corresponding crus penis. The 
fibres all pass obliquely forward and inward to be inserted into 
the inner aspect of the crus, and thus furnishes a‘muscular in- 
‘vestment to the dilated portion of the corpus cavernosum. This 
muscle is separated from the others in this region by the septal 
processes already described. 

Compressor of Cowper's gland. This muscle, which is 
mentioned but not described by Cuvier, is also referred to 
by Duvernoi under the name of the transverse muscle of the 
perineum. It would seem, moreover, that this is the muscle 
figured by Camper in his drawing under the name of the short 
accelerator urine. The muscle has a fascial origin and inser- 
tion. It arises from the outer side of the aponeurotic’ septum, 
which intervenes between it and the bulbo-cavernosus, as also 
from the inner side of that which separates it from the ischio- 
cavernosus. ‘The fibres form an elliptical belly which embraces 


ps DR WATSON. 


and conceals the perineal aspect of Cowper’s gland. At the 
anterior extremity of this gland the fibres terminate on an 
aponeurotic septum, which is formed by the union of the two 
pieces of fascia which separate the muscle from its neighbours, 
and through the medium of this are inserted into the root of the 
corpus cavernosum. ‘This muscle is to be regarded as connected 
physiologically with Cowper’s gland, the secretion of which it is 
adapted to expel. 

The bulbo-cavernosus muscle conceals the bulbous portion of 
the spongy body. It arises principally, along with the muscle 
of the opposite side, from a median tendinous raphe which rests 
upon the bulb. It has however an additio al origin by means 
of an elongated fleshy slip from the aponeurotic structures which 
form the floor of the perineum, which slip arises, along with that 
of the opposite side, as far back as the origins of the compressors 
of Cowper’s glands, between which muscles it lies. From these 
origins the fibres pass obliquely forward and outward, so as to 
embrace the bulb, and are inserted into that portion of the 
fibrous envelope of the penis which intervenes between the 
corpus cavernosum and spongiosum. 

Transverse muscle of the perineum. Muscles distinct from 
those already described bearing this name, are figured by 
Camper, but such are not present in my dissection, nor are they 
mentioned by any other author. 

Penis. This organ, which in Duvernoi’s specimen measured 
6 feet in length and weighed 80 pounds, in the present case 
measures 2} feet from the attachment of the crus to the point. 
Each corpus cavernosum commences by a slightly dilated ex- 
tremity where it is attached to the ischium, and coalesces at 
once with that of the opposite side, so as together to form two- 
thirds of the body of the penis, the remaining third being 
formed by the corpus spongiosum, each cavernous body dimin- 
ishes gradually in transverse section from the root to the point 
of the penis, so that at a distance of 4 inches behind the glans 
each is diminished to one-third or one-fourth of its original 
diameter. The dilated extremity of each is covered by the is- 
chio-cavernosus muscle. ‘The corpora cavernosa are surrounded 
by a very strong fibrous envelope measuring one quarter of an 
iuch in thickness at the root, but diminishing to one-half of this 


— Fe ee —— 


ANATOMY OF THE INDIAN ELEPHANT. 73 


toward the point of the organ. From the middle of the dorsal 
portion of this investment a strong fibrous pectiniform septum 
dips down to separate the two cavernous bodies. This septum 
is very incomplete, and permits of the continuity of the cavern- 
ous tissue across the middle line. This tissue is disposed in 
the usual manner, being denser toward the circumference than 
at the centre of each cavernous body. The large septa described 
by Camper as subdividing each corpus cavernosum are readily 
seen in different sections of the penis, but they are quite irregu- 
lar, and’ are nothing but trabecule of larger size than those 
forming the mass of the cavernous tissue. 

The corpus spongiosum commences by an elongated bulbous 
extremity at the root, and tapers gradually to the point of the 
penis, so that in form, as remarked by Duvernoi, it may not 
inaptly be compared to a large carrot. At the anterior extremity 
of the dorsal aspect of the penis is an elongated body closely 
resembling the backward prolongation of the glans in the 
horse. It measures 3 inches in length, and 24 in breadth, and 
is, I think, to be regarded as the glans. At the same time, 
it is to be observed that this body does not reach the point of 
the penis as in the horse, but is separated from it by a distance 
of 2 inches. Four inches behind the glans the spongy body 
does not measure in transverse section more than one-sixth of 
that of the bulbous portion. As regards its structure, the 
corpus spongiosum is similar to that of the corpus cavernosum, 
except in this respect, that the investing tunic of the former is 
much thinner than that of the latter, and, in fact, is little more 
than membranous. Through the upper part of the spongy 
tissue passes the canal of the urethra, and an imperfect median 
septum rising up from the lower part of the fibrous tunic is 
attached to the floor of that canal. This septum is distinct 
enough posteriorly, but disappears entirely toward the point 
of the penis. The bulbous portion is invested by the bulbo- 
cavernosi muscles, and into this portion of the urethra open the 
ducts of Cowper. 

Vessels and nerves of the penis. Duvernoi describes an 
elaborate series of nervous and venous plexuses in connection 
with the penis, but these I failed to identify. The dorsal 
arteries lie one on either side of the middle line under 


74 DR WATSON. ANATOMY OF THE INDIAN ELEPHANT. 


cover of the corresponding levator penis. Each runs forward 
as far as the extremity of the organ, accompanied by the vein 
and nerve, the former lying to its inner, the latter to its outer 
side. In this course it gives off many branches for the supply 
of the organ, one of which, larger than the others, given off about 
the middle in length of the penis, runs obliquely downward 
and forward, to supply the lateral and inferior aspects of the 
organ. 

The dorsal veins accompany the arteries lying to their inner 
side. Close to the root of the penis the veins of opposite sides 
communicate by a number of branches, so as to form a plexus 
on the dorsum of the organ, and this plexus communicates 


freely with the interior of the cavernous bodies. It will be ™ 


observed, therefore, that there is not a single vein as stated by 
Owen, but that these correspond in number with that of the 
arteries. 

The dorsal nerves accompany the vessels, and run as far as 
the extremity of the penis. In this course they give off many 
branches for the supply of the skin and other parts of the organ. 
Of the third, or median dorsal nerve described by Duvernoi as 
forming a remarkable plexus in this region, nothing could be 
seen. Neither was there any fat present. 

Prepuce.—tThe skin is reflected from the penis just behind 
the glans, to form a well-marked prepuce. 

The orifice of the urethra is, as stated by Camper, Y-shaped, 
the two limbs being directed upward, the stem downward. 


DESCRIPTION OF PLATE IV. 


Fig. 1. The injected ureter and calyces of the right kidney. 

Fig. 2. The inferior surface of @ the membranous part of the 
urethra. 6. neck of the bladder. cc. vesicule seminales. dd. vasa 
deferentia. ee. prostate glands. The right vesicula and prostate 
have been opened. 

Fig. 83> The canal of the membranous part of the urethra has 
been opened to display a. the vesicula prostatica; 6. the bristles 
introduced into the ejaculatory ducts ; c. those introduced into the 
prostatic ducts; dd. the openings of the ureters. 

Fig. 4. Perineal muscles and fascia. aa. ischio-cavernosi, 
bb. bulbo-cavernosi. cc. compressors of Cowper’s glands. dd. in- 
ternal fibres of sphincter ani. 


SOME OBSERVATIONS ON THE DENTITION OF THE 
NARWHAL (MONODON MONOCEROS’). 


By PRoressorn TURNER. 


It is the current belief of naturalists that, in the Narwhal, two 
teeth only are produced, both of which are situated in the 
upper jaw. In the female, as a rule, these teeth remain in 
arudimentary state concealed within their sockets. In the 
male, on the other hand, the rule is for the right tooth only 
to remain rudimentary and concealed within its alveolus, 
but the left protrudes, grows in the adult to the length of 
several feet, and forms the well-known tusk or horn of this 
animal. Occasionally, however, the right tooth grows equally 
with the left, and like it projects for several feet beyond the 
mouth. In an interesting paper recently published Mr J. W. 
Clark of Cambridge’ states that at least eleven bidental skulls 
may now be found in the various museums in Europe’. 

With regard to the position of these teeth in the upper Jaw 
a difference of opinion has been expressed by anatomists. The 
illustrious Cuvier stated* that the teeth were implanted in 
the intermaxillary bones, or in an alveolus common to the 
maxilla and intermaxilla; and this view has been adopted by 
various subsequent writers, by some of whom the tusk has been 
regarded as a peculiarly modified incisor tooth. But in a paper 
read, some forty years ago, to the Royal Society of Kdinburgh’, 
-Robert Knox pointed out that the tusks were carried in the 
maxillary bones; and this view of their position has recently 
been supported by Van Beneden and Gervais®, J. W. Clark, and 
W. H. Flower’. 

1 Read before the Royal Society of Edinburgh, May 20th, 1872. 

2 Proc. Zool. Soc. Jan. 17, 1871. 

3 In the Anatomical Museum of the University of Edinburgh is the skull of 
a male Narwhal, from the left supr. maxilla of which a tusk 33 inches long 
projects. The right maxilla has been in part removed, and a eanal has been 
exposed, which extends as far back as the base of the beak, and is nearly 1 inch 
in diameter. From its length and the size of its bore it is not improbable that 
in this skull the right tusk had been developed and had protruded from the jaw, 
but the tusk and loose piece of the jaw have been lost. 

4 Ossemens fossiles, v. Part 1. 321, 322. 

5 Transactions, xt. p. 410. 1830. 


6 Ostéographie des Cétacés, p. 13. 
? British Medical Journal, May 20, 1871. 


76 PROFESSOR TURNER. 


Being desirous of satisfying myself on this point I have 
examined ten crania of this animal, all of which, with one 
exception, are in the Edinburgh Museums, and from what I 
have seen in them I entirely agree with the statement of the 
last-named anatomists. Five of these crania were males, three 
adult, and two from younger animals; two were females and 
three were well grown foetuses, and in all the specimens the 
maxillo-premaxillary suture was placed to the inner side of 
the alveolus for the tusk. In the adult male skulls this suture 
was situated in the inner wall of the alveolus, and it is pro- 
bably from this circumstance that Cuvier considered that the 
socket was hollowed out of both the premaxillary and maxillary 
bones. But in the female and fcetal crania the maxillary 
nature of the tooth was shown in so decided a manner that 
there could be no doubt as to its true position. In the female 
crania the teeth were situated close to the palatal surface of the 
bone, a little to the inner side of the outer edge of the maxilla, 
and in one specimen the rudimentary tooth lay concealed in its 
socket nearly two inches to the outer side of the maxillo-pre- 
maxillary suture. In the feetal crania the maxillary and pre- 
maxillary bones were readily separable from each other, and 
the socket of each young tusk was entirely situated within 
the superior maxilla. 

The apparent participation of the premaxilla, in the for- 
mation of the socket of the developed tusk in the adult male, is 
undoubtedly due to a partial absorption, during the growth of 
the tusk, of the thin inner wall of the alveolus next the maxillo- 
premaxillary suture, in consequence of which the premaxillary 
bone forms a small proportion of the inner wall of the socket. 

Owing to the maxillary position of the tusk it can no longer, 
therefore, be regarded as a peculiarly modified incisor tooth, 
but as it lies immediately to the outer side of the maxillo- 
premaxillary suture it should be regarded as representing a 
canine tooth. 

In the course of my observations on the foetal crania, and 
on that of a young male, I observed an appearance which led 
me to think that, in addition to the well-known pair of teeth 
in the upper jaw, each superior maxillary bone had at one time 
contained another tooth. Tor situated close to the outer border 


OBSERVATIONS ON THE DENTITION OF THE NARWHAL. "77 


of the palatal surface of this bone was a canal, which passed 
backwards, parallel and inferior to the tusk-socket. In the 
young male this canal was two inches long, and opened in 
front one and a half inch behind the mouth of the socket for 
the tusk. It had the appearance of an alveolus, and on illumi- 
nating its interior, by reflecting light from the surface of a 
mirror, I perceived a minute denticle at the bottom of the 
socket. 

Being so fortunate as to possess, through the kindness of 
Mr. C. W. Peach, a young male Narwhal (seven and a quarter 
inches long) with all the soft parts uninjured, I thought that I 
might perhaps be able to determine in it whether the Narwhal 
had originally more than two teeth developed in connection 
with its upper jaw. The surface of its palate was smooth and 
covered by mucous membrane, continuous with the tegument- 
ary covering of the upper lip, but there was no appearance of 
teeth on the surface. When the more superficial part of the 
gum was however carefully cut off, two well-defined dental 
papille, each contained in its own tooth-sac, were exposed, 
imbedded in and completely enclosed by the gum which 
covered the outer edge of each half of the upper jaw, so that 
I can now state definitely that the Narwhal, at this early stage 
of development, possesses four teeth in the upper jaw. The 
more anterior of the two papille was two-tenths of an inch 
behind the tip of the jaw, and the more posterior lay about 
one-tenth of an inch behind the anterior. 

Each dental papilla was so small as to be barely visible to 
the naked eye, and required the microscope to be employed for 
its further examination. Each papilla was continuous at its 
base with the connective tissue of the mucous membrane of the 
gum, from which it projected into the cavity of the closed-in 
tooth-sac. It was somewhat clavate in form, and was separated 
from the inner surface of the tooth-sac by a slight interval. 
When examined with high powers of the microscope, the pa- 
pilla was seen to consist of small, pale, nucleated corpuscles, 
imbedded in a delicate and apparently homogeneous matrix. 
Some of these corpuscles were rounded, others oval, whilst 
others again were distinctly caudate. Corpuscles, similar in 
form, were collected in considerable numbers at the base, and 


76 PROFESSOR TURNER. 


in the connective tissue immediately adjacent to the base of the 
papilla, whilst throughout the connective tissue of the gum 
numerous characteristic connective tissue-corpuscles were seen. 
There was no trace of calcification of the dental papilla. The 
free surface of the papilla was limited by a sharp, definite 
outline, as if a membrana limitans invested it, but no distinct 
separable membrane was demonstrated, so that the sharpness 
of definition was probably due to the tissue of the papilla bemg 
more condensed near the surface. 

The wall of the tooth-sac was entirely surrounded by the 
connective tissue of the mucous membrane. Its relation to the 
dental papilla showed it to be homologous with the enamel 
organ in man and those animals where enamel enters into the 
structure of the teeth. All connection was severed between it 
and the epithelium of the mouth, of which it had been, in all 
probability, originally an involution. It measured about +,th of 
an inch in thickness, and was not homogeneous, for its outer and 
inner portions were denser than a more delicate intermediate 
portion. It was composed of pale nucleated corpuscles, about 
equal in size to those which entered into the formation of the 
dental papilla. These corpuscles were ovoid in form, and I 
failed to recognise any elongation of the cells which formed 
the inner portion of the tooth-sac into columnar epithelium, by 
the calcification of which the rods or prisms of the enamel are 
produced in man and in those animals in which enamel forms 
a part of the structure of the tooth. No membrana limitans 
was seen on either the inner or outer surfaces of the wall of the 
tooth-sae. 

It is customary to state that the tusk of the Narwhal is 
destitute of enamel, and consists of dentine with an external 
covering of cement. As the examination of this young foetus 
revealed the existence of a structure homologous with an enamel 
organ, though at a stage too early to exhibit its characteristic 
epithelium, I thought it advisable to examine anew the micro- 
scopic character of the tusk to see if there might not be, espe- 
cially at the tip, some trace of an enamel covering. I accord- 
ingly removed a thin slice from the tip of one of the unpro- 
truded, and therefore unworn, tusks of a well-grown foetus. 
The general substance of the tusk consisted of well-formed 


| 
| 
| 


OBSERVATIONS ON THE DENTITION OF THE NARWHAL. 79 


dentine, but at the tip a depression extended for some. distance 
into this tissue. This depression was filled up with crusta 
petrosa, continuous with that which formed the external in- 
vestment of the tusk. The crusta in the immediate neigh- 
bourhood of this depression contained not only lacunz with 
canaliculi proceeding from them, but groups of fine canals, 
which resembled in size and appearance dentine tubes. Be- 
tween the dentine and the crusta petrosa was a thin ill-defined 
layer into which the dentine tubes penetrated, and which ob- 
viously corresponded to the so-called granular layer of the 
dentine in a human tooth. No trace of enamel rods could be 
seen. 

The inner surface of the wall of the tooth-sac was not per- 
fectly smooth, but possessed one or more ridge-like projections, 
which fitted into corresponding depressions on the outer surface 
of the dental papilla. It is, without doubt, to this arrange- 
ment, that the depression in the dentine at the tip of the tusk 
of the well-grown foetus owes its origin. 

There is no reason to think that the more anterior of the 
two teeth seen on each side of the upper jaw of this foetus had 
to the more posterior the relation of a milk-tooth to a perma- 
nent tooth. For they were both almost precisely equal in size 
and in comparative development, which would not have been 
the case if the latter had had to act as the successor to the 
former. In all probability the more anterior would have 
developed into the maxillary tusk, and the posterior either 
have disappeared altogether, or formed one of those irregular 
non-protruding teeth, such as Berthold described some years 
ago in the skull of a young Narwhal which he examined’. 

No rudimentary teeth were found in the lower jaw, although 
it was carefully examined. 

The ossification of the fibrous basis of tite maxillary bones 
was so imperfect that it was not possible to distinguish the 
maxillary from the premaxillary segment. But in the lower 
jaw, the ossification of the fibrous membrane, which invested 
Meckel’s cartilage, had advanced to a considerable extent. 


1 Miiller’s Archiv, 1850, p. 386. 


TISSUE METABOLISM, OR THE ARTIFICIAL INDUC- 
TION OF STRUCTURAL CHANGES IN LIVING 
ORGANISMS. By W. Arnsiie Hous, M.D. Cantab. 
Part II. Actinie continued. (Pl. vi.) — 


I HAVE elsewhere’ given an account of some experiments in- 
stituted by me with a view to ascertain the structural changes 
which can be induced artificially in the tissues of these lowly 
organised beings. I intend here giving a short résumé of some 
further researches on the same subject. 

Experiment 6. The application of acetic acid. August 3, 
1.45 p.m. <A greenish brown anemone (A. Mesembrianthemum) 
was touched with strong acetic acid, whilst withdrawn from the 
tank, it was next washed with sea-water and replaced in the 
vessel. A minute after the operation the imjured part ap- 
peared swollen and paler than the surrounding tissues, it was 
also incapable of reaction to slight stimulus (such as that made 
by scratching it with the point of a steel style); the animal 
immediately contracted itself after the operation. 3 P.M. In- 
tegument of wound of a brownish red hue, and covered with a 
pultaceous muciform mass and swollen. This material was 
examined microscopically, and was found to contain a large 
number of elongated corpuscles with nuclei (Fig. 3). It was 
very tenacious. The animal was collapsed and had partially 
relaxed its hold of the vessel it was placed in. August 4th, 
11 A.M. The injured portion (a) elevated and surrounded by 
contracting healthy tissues. The anemone partially expanded. 
On touching (a) with a style it was readily removed, and it left 
a whitish constricted scar composed of the subintegumentary 
tissues. (a) examined microscopically. Reddish brown colour 
and diffluent. It contained many spheroidal corpuscles, gra- 
nules, and oat-shaped bodies, with a few ciliated animalcules. 
Pulsating movements were observable in some of the corpus- 
cles. A small quantity of the exudation from the surface of 


1 Journ. of Anat. v1. p. 381. 


ae 7 


TISSUE METABOLISM. SL 


the newly-formed scar was carefully removed by a capillary 
glass tube and examined microscopically (Fig. 8). Several 
hyaline spheroidal corpuscles of various sizes were seen. No 
movements apparent in any of them. Several of the smaller 
corpuscles were double. August 5th, 2 p.m. Wound considerably 
puckered. Surface whitish. The healthy tissues elevated 
around it. Sept. 4th. Anemone healthy. The injured part 
indistinguishable from the other tissues except by its colour, 
which is an uniform pale green and unstriated (as is the rest 
of the body) with brown. 

The two following experiments were made in consequence 
of a suggestion of Mr Savory, whom I have to thank for several 
valuable hints in the prosecution of these researches. 

Experiment 7. The injection of fwtid sea-water into an 
actinia. July 26th, 1872, 8 p.m. Made an incision in the base 
of an anemone (A. Mesembrianthemum) and injected feetid 
sea-water in which some decomposed crab-flesh had been 
steeped for several days. The injected water was very offen- 
sive to the smell, but under the microscope (x 250) gave no 
signs of organic life; the water was, however, full of dead 
amorphous particles of matter. Immediately after the opera- 
tion the animal contracted itself. At 3.10 PM. the point at 
which the incision took place was depressed, and the surround- 
ing tissues were thrown into longitudinal and transverse folds. 
July 28, 10.30 am. Actinia contracted since last note. Scar 
still depressed. August 3rd, morning. Anemone healthy; no 
apparent result. 

Thinking that the negative results of this experiment might 
have been due to the difference between the natures of the 
decomposing tissues and the tissues of the anemone, I made the 
following experiment. 

Experiment 8. _ The injection of a portion of the slough (a) 
of Experiment 6 into a healthy anemone. August 4th, 11.45 a.m, 
Injected a portion of the slough (a) of Experiment 6 by means 
of a common hypodermic syringe into a brownish-green ane- 
mone. The anemone immediately contracted. Wound de- 
pressed. August 5th, noon. Anemone partly expanded, tenta- 
cles lax and flaccid. 5.15 p.m. The animal was closed, wound 
still depressed. August 14th. The anemone has been, con- 


VOL. VII. 6 


82 DR HOLLIS. 


tracted for the past few days. Wounds inconspicuous, no 
apparent result’. 

The following experiment shews that the evolution of 
plasmic bodies may take place in a part of these creatures 
after its removal from their body. 

Experiment 9. The absctsion of a tentacle and the subse- 
quent application of acetic acid to the abscised portion. August 8, 
noon. Tentacle abscised?. 12.15p.m. A drop of acetic acid 
placed on it for an instant, and the tentacle was then washed 
with sea-water. It was next placed on a glass slide for micro- 
scopical examination, and so covered with a thin glass that no 
pressure was exerted upon it®. 12.30 p.m. The surface was 
contracted into deep rugee and surrounded by a web of deli- 
cate thread-like bodies, hyaline, spheroidal and spindle-shaped 
corpuscles and a few granules. The tentacle was removed from 
the glass slide and placed in a vessel of sea-water contaiming 
other actiniz. August 9th, 1 p.m. Tentacle dead and textures 
soft and undergoing disintegration. Numerous ciliated ani- 
malcules* in various stages of growth were amongst the de- 
caying textures. 


1 Since performing these two last experiments I have injected some decaying 
anemone-tissue swarming with ciliated animalcules into a healthy actinia, but 
with no definite result. 

2 Tt may be as well to state that a tentacle, when placed in sea-water after 
abscission, will continue to shew signs of vitality for some hours. 

3 I found this precaution necessary, as I was previously led astray somewhat 
in several similar experiments by the rapid evolution of spheroidal corpuscles, 
after the application of the thin glass. This evolution was induced to some 
considerable extent by the pressure thus exerted upon the tentacle. 

4 Tam pleased to find that Prof. Haeckel of Jena, in his recent researches, 
has arrived at the same conclusion with regard to the constitution and the 
nature of the movements of cilia in such animals as I have done (Journ. of 
Anat. vi. p. 381). He now considers cilia (Biolog. Studien, noticed in Arch. de 
Zoolog. Expérim. et Générale, 1. p. xut. 1872) as ‘prolongations of the proto- 
plasma of the mass ealled, according to the cellular theory, by the name of 
cellule, and that their movements are amceboid. Recent researches into the 
evolution of the lowest forms of animal life would lead me to conjecture that 
possibly the locomotive corpuscles, ciliated animaleules and vermiform animals 
(Journ. of Anat. v1. pp. 384—5), described by me as the parasites of actinie, 
are different forms of the same creature, and are allied to the Euglenia, figured 
in Dr Bastian’s work (Beginnings of Life, 1. p. 526). Dr Cobbold curiously enough 
gives the figure of a nematode (Brit. Med. Journ. 1872, 1. 89), very similar in 
shape to the vermiform animals found in actiniw. This nematode was found by 
him in the urine of a patient afflicted with the ciliated Bilharsia Hamatobia. 
I have, however, been unable to trace any connection between the ciliated ani- 
malcules and the vermiform animals in my researches. That the former origi- 
nate in locomotive corpuscles I think exceedingly probable. I cannot under- 
stand why Prof. Rolleston considers that ‘ the spermatozoa of parasitic nematoids 
are spherical or oyoidal cells’ (Forms of Animal Life, p. cxxxy. 1870), ‘and move 


TISSUE METABOLISM. 83 


I give the details of the following experiment with blistering 
fluid (although similar experiments have been noticed pre- 
viously), as it illustrates better than the others the way by 
which the healing processes after irritation are carried on in 
these animals. 

Experiment 10. The application of blistering fluid to the 
tentacles of an Actina Mesembrianthemum. Aug. 12th, 1872, 
3.40 P.M. Touched a few tentacles of a red anemone (A. Mesem.) 
with blistering fluid. 3.50 p.m. The touched tentacles lax, 
swollen and semitransparent. On removing from them a little 
muciform débris on a style for microscopical examination, there 
were found in it numerous spindle-shaped corpuscles, and a few 
spheroidal ones with granules (Fig. 4). 5 p.m. The blistering 
fluid had run over the pedicle as well as the tentacles. The 
injured parts were paler and brighter coloured than the rest of 
the body, and were covered with a diffluent muciform reddish 
matter. On microscopical examination this diffluent material 
was found to contain strings and agglomerations of hyalined 
spheroidal, and spindle-shaped corpuscles (Fig. 5), the former 
were far more numerous than they were in the muciform fluid 
at the first examination ten minutes after the injury. August 
13th,2 pM. The anemone partly expanded. The tentacular 
ring on the side of the injury contracted. The extremities of 
some of the injured tentacles had sloughed off. The subinte- 
gumentary tissue (after removal of a pultaceous débris) was 
laid bare in many places. On microscopical examination the 
débris was found to contain a large quantity of hyaline corpus- 
cles, many of which were spheroidal in shape and had depressed 
centres (Fig. 6). August 14th, 11 Am. Anemone contracted 
and the injured parts covered with a reddish muciform débris, 
the consistence of which was greater than that of yesterday. 
On its removal the creature was found in the parts corre- 
sponding to the scoring of the integument to be covered with 
small nodules of tubules. The débris consisted mostly of 
spheroidal-shaped corpuscles arranged in rows and laterally 
compressed (Fig. 7). Aug. 16th, 8.25 AM. Anemone con- 


only by the protrusion of pseudopodial protoplasmic processes.’ To call such 
creatures spermatozoa appears to be asserting more than the facts of the case 
justify. 


—_ 


bo 


84 DR HOLLIS. 


tracted on the side of the injury, elsewhere partially expanded. 
Débris scanty. 

No further examination of the débris was made in this case. 
The next experiment illustrates the operation of nitric acid on 
the tissues of these animals. 

Experiment 11. The application of nitric acid to an Actinia 
Mesembrianthemum. July 16th, 4 p.m. Touched a greenish- 
brown anemone with strong nitric acid. Afterwards carefully 
washed it in sea-water, until the washings had no acid reaction. 
Immediately after the operation the animal forsook its hold of 
the vessel in which it was placed, and became strongly con- 
tracted. The part touched assumed a reddish-brown hue, and 
the surrounding tissues were thrown into fine corrugations. 
July 17th, 10 am. The animal was found attached to the 
vessel with the wounded part dependent. The tentacles on 
the upper side were expanded. The creature ate food when it 
was offered to it. The wound was covered with.a pultaceous 
reddish-brown mass; the rest of the imtegument was of a 
greenish-brown colour. On microscopical examination the 
débris was found to consist of a few granular corpuscles in 
a delicate nearly homogeneous mass. July 18th, 4PM. The 
animal still expanded. The reddish-brown slough covering the 
injured parts easily separated from the animal. It was found 
to consist of muscular fibres, granules and locomotive corpus- 
cles. The scar white and depressed. Administration of Chlo- 
roform. July 24th, 1.30 p.m. Animal contracted, the cicatrix 
puckered and depressed. Stomachal orifice drawn towards the 
scar. Chloroform vapour was applied to the creature at 2.30 P.M. 
The immediate effect of the vapour was the production of an 
cedema of the integument except at the scar, and a total loss of 
the power of reaction to stimuli. The effects of the drug lasted 
for about half-an-hour.. July 28th. Scar still puckered and 
depressed. Anemone healthy. September 5th. The cicatrix 
only to be distinguished from the surrounding tissues by the 
colour, which is greenish, and unstriated, as is the rest of the 
body, with brown. 

Recapitulation and Summary of the experiments upon Ac- 
tinie. 1. The application of an irritant to these animals 
produces (as before noticed) swelling of the tissues. This 


TISSUE METABOLISM. 85 


swelling consists essentially in a diminution of the tonic con- 
tractile force residing in the healthy tissues, and is accompanied 
by a loss (more or less considerable) of the power of reaction of 
injured tissues to stimuli. 

2. The swelling of the tissues is usually accompanied by 
an immediate production of a muciform fluid, but in such cases 
different irritants produce different results. This fluid is at 
first nearly colourless, and contains numerous spindle-shaped, 
with few spheroidal, corpuscles and granules. The above phe- 
nomena (i.e. the swelling of the tissues and the production of 
the muciform fluid) may be excited by the application of an 
irritant to a still-living part after its removal from the body. 

3. If the cause of the irritation be prolonged and violent, 
disintegration of the tissues takes place—and one of the fol- 
lowing events occurs: a. A slough is formed of the decayed 
tissues and is cast off, leaving a contracting cicatrix of healthy 
tissues bathed in a little muciform fluid similar to that de- 
scribed above (§ 2). 8. The disintegrating tissues become 
gradually eroded, and the clear muciform fluid surrounding the 
injured part becomes more tenacious and opaque’, and contains 
a large number of spheroidal and other corpuscular elements 
elsewhere described (Journ. of Anat. VI. p. 398). 

4. The injection of a solution of decomposing tissues into 
the healthy animal does not (as far as my experience goes) in- 
duce any disintegrating effect upon the tissues with which such 
a solution is brought into contact. 


ANNULATA. 


The next series of experiments I shall describe are some 
illustrating the tissue-changes which take place after the appli- 
cation of an irritant to the integument of the common earth- 
worm (Lumbricus Terrestris). It will not be necessary for me 
to recapitulate the minute anatomy of this creature, as that 
has already been most ably done by Mr E. Ray Lankester’; 

1 In my previous paper (v1. p. 393) I described this fluid as greenish-yellow. 
I find, however, from some experiments since undertaken with red-coloured 
actinie that the fluid simulates more or less closely the colour of the healthy 


tissues. 
2 Quart. Journ. of Microsc. Sc. New Series, tv. 259, and y. 7 and 99. 


86 DR HOLLIS. 


and, as far as the nervous system is concerned, by Mr (now Dr) 
Lockhart Clarke’. There are, however, ove or two particulars 
in the anatomy of the earthworm in which my own investiga- 
tions have led me to differ somewhat from the generally ac- 


cepted statements concerning it, these I shall here shortly — 


notice. 

The vascular and perivisceral fluids. The tissues of Lum- 
bricus terrestris are very generally intersected by numerous 
vessels containing a red fluid, coagulable on the addition of 
nitric acid. ‘This vascular fluid is usually described as non-cor- 
pusculated, and as thus not truly representing the blood of 
vertebrates. I have, however, after numerous carefully con- 
ducted experiments, arrived at the conclusion that this appa- 
rently homogeneous fluid in reality contains hyaline corpuscles 
and granules in variable quantity. Similar corpuscles, only in 
greater numbers, and occasionally with amceboid-like processes, 
are seen in the clear perivisceral fluid which bathes the splan- 
chnic cavity between the coats of the alimentary canal and 
the inner wall of the integument. The corpuscular elements 
upon the addition of acetic acid separate into a delicate external 
envelope and internal granular contents. The granules are 
either free or in groups, and, when seen in conglomerate masses, 
are of a greenish tint’. They are unaffected by benzine (Fig. 1). 
The perivisceral fluid varies greatly in quantity, at times barely 
sufficing to bathe the external walls of the gastro-intestinal 
canal. 

Their parasites. In the muscular layers of the tegumentary 
walls numerous vermiform animals are found (the Anguillula 
LIumbrici of Dujardin, the small nematoid of Lankester). They 
are in external respects and in size similar to the vermiform 
parasites of actiniz. Like them the anterior or oral extremity 
is pointed, and the alimentary canal passes without convolutions 
to within a short distance of their somewhat obtuse caudal 
end. On either side of the canal can be seen small granular 
particles, similar to those found floating freely in the blood of 
their host. These granules, when seen in the mass, have a 


1 Proc. Royal Soc. 1857, pp. 344—352. 
2 These bodies are similar in appearance to those observed by me in actinia 
(Opus jam cit. p. 387). 


ee 


TISSUE METABOLISM. 87 


greenish tint. These parasites do not die immediately they 
are immersed in sea-water, although their salt-water congeners 
rapidly expire when placed in fresh water. When coiled up 
amongst the muscular fibres of their host, they present an ap- 
pearance not dissimilar to encysted Trichine. 

I shall now proceed to detail some experiments I have un- 
dertaken, with a view to observe the tissue-changes that can be 
induced in these animals by the application of irritants to their 
integument. 

Experiment 1. The application of blistering fluid to an 
earthworm. July 30, 3Pp.M. Applied Liquor Epispasticus to 
sixteen segments of the tail extremity of a young healthy Lum- 
bricus, leaving seventeen caudal segments posterior to those 
injured untouched. 8.10 Pp.M. The injured part pinkish white 
and swollen. No vermicular motion in it, although the healthy 
segments behind it moved freely. 3.20P.M. The tail behind 
the seat of injury of a pink hue from congestion of the capil- 
laries, no contraction of dorsal vessel in or behind the wound. 

July 31,11 Am. The last thirty-five segments had sepa- 
rated from the rest of the worm. The terminal segment of 
worm somewhat obtuse, but the cicatrix at this poimt scarcely 
distinguishable from the healthy caudal end: worm active and 
lively. The detached end (a) was found to be almost entirely 
destitute of vitality, a slight retraction of the anal orifice was, 
however, induced upon irritating this extremity. The part (a) 
where the blistering fluid was applied was of a deep red colour 
above, and the epidermis was removed—the colour gradually 
faded off below where the integument was persistent. The re- 
mainder of ‘a’ was of a pale straw colour. The dorsal vessel 
was much congested along the denuded part. On examining 
microscopically a portion of the abraded skin several nematode 
worms were seen. The corium and muscular layers next ex- 
amined. Capillaries intensely congested (Fig. 2). Hxtravasa- 
tions of blood in parts. Corpuscles with shrunken centres, 
resembling the red corpuscles of mammals, were also seen. It 
is possible that this form is assumed by these bodies after im- 
mersion in water, the medium in which they were examined. 
The dorsal vessel of ‘a’ was next carefully pricked, and a drop 
of the dark-red-blood exuding introduced into a clean capillary 


88 DR HOLLIS. 


glass tube, and afterwards examined microscopically. It was 
found to contain numerous corpuscles of various sizes, the larger 
with a distinct red hue. The blood was coagulable with nitric 
acid; and the corpuscular elements in it were without motion. 
August 3rd, worm healthy. 

Experiment 2. Lhe application of acetic acid to an earth- 
worm. August 3rd, 1872, 1.30PM. Touched seventeen seg- 
ments of the same worm as that previously experimented on 
with acetic acid. The injured portion extended to within ten 
segments of the freshly cicatrised caudal extremity. Imme- 
diately after the operation the integument became white and 
swollen, and the injured part apparently lost the power of mo- 
tion, although the ten healthy segments behind it still retained 
that power. Upon examining the injured part ten minutes 
later with a magnifying glass, several subcuticular segmental 
blood-vessels and their capillaries were seen to be congested. 
The part had now assumed a distinct pink hue ; there was pul- 
sation of the abdominal vessel behind the injury, although no 
distinct movement was detected at the seat of the operation. 
4p.M. The last twenty-nine segments (a) partially separated 
from the rest of the body by a fissure on the left side extending 
into the abdominal cavity. The fissure, apparently caused by 
the violent tonic (hour-glass) contractions of a segmental set of 
muscles. The injured portion somewhat redder and more swol- 
len than when last observed. The last ten segments still re- 
tained the power of vermicular contraction. August 4th, 
12.15p.Mm. The part ‘a’ separated at the fissure from the rest 
of the worm. It still retained some power of independent 
motion in the ten terminal segments. A capillary tube was 
applied to the caudal extremity of the living worm, and a smal] 
quantity of fluid taken up thence and examined microscopically 
(Fig. 9). It contained hyaline corpuscles and oblong bodies of 
various sizes. Whilst I was examining the worm, and holding 
its caudal segments between my fingers, about ten segments (8) 
at the part were suddenly thrown off. Some slight move- 
ments in the dorsal vessel of ‘8’ after its separation. 

The application of irritants to the segments separated from 
the living worm. 12.30 P.M. Blistering fluid was applied to 
the still moving extremity of ‘a’; to ‘8’ a small quantity of 


TISSUE METABOLISM. 89 


acetic acid. Five minutes after the application ‘a’ was white 
at the part and swollen. A blebs appeared to be produced. 
It still retained some independent motion. ‘8’ swollen and of 
a pinkish-white colour, it appeared to have lost all contractile 
power. The bulla of ‘a’ pricked and a little fluid withdrawn. 
On microscopical examination there were seen several sphe- 
roidal and oblong corpuscles (as im Fig. 9, only somewhat 
smaller as a rule), granules, and nematode parasites. August 5. 
Both extremities (a and 8) dead. Worm healthy. 

Experiment 3. Zhe injection of decayed animal matter into 
an earthworm. August 13, 1872. 3 p.m. Injected into the 
fiftieth segment from the head of a young lumbricus a drop 
of fresh water, containing corpuscles from the slough of an 
actinia (obtained in Experiment 10 on those animals). Im- 
mediately after the experiment the segments at the seat of 
injury became slightly congested. August 14th, 11.50 A.M. 
Worm lively. At the point of puncture a small elevation of 
the skin with a thick whitish fluid beneath it. This fluid on 
microscopical examination was seen to consist of plasmic ele- 
ments of various shapes, mostly spheroidal, some granules, 
pyriform and staff-shaped corpuscles (bacillz) floating in a little 
hyaline fluid (Fig. 11). August 16,1210 p.m. The puncture 
still covered with a flap of skin and surrounded by a dark line 
(2 congested capillaries). Unfortunately absence from town pre- 
vented me from completing my observations on this animal. 
It is now (Sept. 28th) alive and healthy and the wound incon- 
- spicuous. 

As several experiments in which I had attempted to induce 
a higher stage of tissue-irritation than the mere elevation of 
the epidermis, and the production of a partially filled bulla, 
resulted in the separation of the worm at the point of injury, 
and the rapid death of the discarded segments, I undertook the 
following experiment, in which by limiting the size of the 
injury to a small local wound, and by repeated applications 
of an irritant to this spot, I hoped to produce some of the 
further changes in tissue formation, which only take place after 
the lapse of some days from the first injury. 

Experiment 4. The repeated application of irritants to 
certain segments of an earthworm. Sept. 25th, 4p.M. Applied 


90 DR HOLLIS. 


blistering fluid to a large specimen of Lumbricus Terrestris, 
forty segments from the end. This, as usual, induced a rapid 
wriggling movement of the worm. Sept. 26th, 2.45 p.m. Injured 
part an oval in shape, on the side of the worm, and extending 
over three segments. Epidermis at this point elevated and 
wrinkled, whitish. Segmental divisions, fully separated at the 
side of the injury, were there destitute of contractile power 
and sensation apparently; they were on the opposite side 
closely contracted, thus bending the body into an angle, of 
which the wound formed the apex. No capillary congestion 
was seen, but this may have been partly due to the thickness 
of the integument in this specimen, the dorsal vessel being 
scarcely visible through the skin. Placed a small quantity 
of acetic acid on the injured part, the epidermis immediately 
became elevated, semi-lucent. and tense. No wriggling of the 
worm took place on the application of the acetic acid. Worm 
active and lively. Sept. 27th, 2.10 p.m. Injured segments 
elevated, tense, and yellowish. Second application of acetic 
acid to the part. No movement of the worm at first after the 
operation; the acid, however, subsequently ‘ran over’ a fresh 
portion of the integument, when the wriggling immediately 
commenced. Sept. 28th, 10.30 am. Injured segments, now 
eight in number, the wound of an oval shape, prominent, 
whitish at the sides, and reddish-brown in the centre. Epider- 
mis removed for about a line over most prominent part, where 
the integument constantly exuded moisture. A small incision 
was made at this point and a little milky fluid withdrawn by a 
clean capillary tube and examined microscopically. It con- 
sisted of a very large number of hyaline corpuscles, mostly 
spheroidal, some, however, oblong and pyriform in shape. There 
were also numerous granules, in conglomerate masses generally, 
and of a greenish tint; again, long stringy masses of plasma 
(fibrillz). These fibrillee appeared to contain spheroidal cor- 
puscles in their substance occasionally (Fig. 12). The promi- 
nent part was again touched with acetic acid and the worm 
replaced in the case by itself. Sept. 29th, 10 Am. Worm 
active and lively. The wound of a yellower tint than it was 
yesterday. Acetic acid again applied to the most prominent 
portion. Sept. 80th, morning. Wound yellow and depressed. 


TISSUE METABOLISM. 91 


The lips of incision everted and elevated. A deep sulcus, ante- 
riorly, separated healthy from diseased tissues. No actual 
fissure. Touched the wound with blistering fluid. 

Oct. Ist, 3.15 pM. The worm was divided at the segment 
anterior to the wound and healthy. Posterior (discarded) seg- 
ments exhibited signs of vitality. On examining microscopically 
the fluid escaping from the surface of the wound, numerous 
pyriform and a few spheroidal corpuscles were observed. The 
edges of the wound retracted, colour brownish-yellow in the 
centre, greyish-white at the sides where the epidermis was 
elevated. On aclean section being made through the diseased 
tissues, they were found to be softer, less coherent, and moister 
than natural. The injury extended inwards as far as the 
intestinal walls. The diaphragmatic blood-vessels near the seat 
of injury greatly congested. The subsequent microscopic ex- 
amination shewed the natural tissues loose, shreddy, fibrils 
broken, and a large quantity of amorphous plasma appeared 
to have been poured out at one part between the muscular 
constituents of the integument. There were also numerous 
granules and plastic fibrils, few corpuscular elements; the 
fibrils were in parts granular. At one point, however, I ob- 
served a considerable number of pyriform and spheroidal cor- 
puscles. The absence of the corpuscular element to such a 
considerable extent in the metabolic processes observable in 
these creatures surprised me considerably. The evolution of 
plasma in amorphic or fibrillar masses appears to take the place 
_ of the spheroidal element in these animals, when the tissue- 
changes are far advanced. The pyriform corpuscles seen in the 
earlier stages of irritation seem to me to be only an imperfect 
evolution of the spheroidal form, due possibly to an increased 
rapidity in the production of such elements. Oct. 2nd. Worm 
healthy. Discarded terminal segments dead. 

Summary. From the above experiments it is shewn that in 
the common earthworm, irritants, when applied to the integu- 
ment, produce :—1. Swelling of the implicated tissues and loss 
of contractile power. 2. Elevation of the epidermis. from the 
true cutis. 3. The moderate production of corpuscular (pyriform, 
spheroidal, and granular) elements beneath the raised epidermal 
layer. 4 The congestion of the capillaries in proximity to the 


92 DR HOLLIS. 


injured parts. If the irritation be prolonged, besides the above 
named phenomena, we also find a softening and disintregation 
of the tissues implicated, together with the production of amor- 
phous, fibrillar and other plasmic elements. 

Conclusion. I conclude that most (if not all) the above 
phenomena denote a loss of the tonic vital contractility natur- 
ally inherent to the healthy tissues, and their consequent inca- 
pacity to react under stimulation. 


Connotation of the metabolic phenomena observable in Actinie 
and Lumbrict. 


The application of an irritant to the integument induces 


In Actiniz :— 

1. Swelling and loss of con- 
tractility in the injured tis- 
sues. 

2. The production of a mu- 
ciform fluid containing spindle- 
shaped and spheroidal corpus- 
cles with granular matter. 


3. Softening and disinte- 
gration of the tissues with 
either their subsequent remo- 
val from the body or the evo- 
lution of a large quantity of 
spheroidal and other elements. 


4, The phenomena describ- 
ed in the first two sections can 
be induced in the tissues of a 
part freshly removed from the 
body. 


In Lumbrici :— 


1. Swelling and loss of con- 
tractility in the injured tis- 
sues. 

2. Elevation of the epider- 
mis and the production of 
pyriform, spheroidal, and gra- 
nular elements held in a little 
hyaline fluid. 

3. Softening and disinte- 
gration of the tissues affected, 
with either their removal from 
the body, or their subsequent 
infiltration with amorphous, fi- 
brillar, spheroidal, and other 
plasmic elements. 

4. The phenomena describ- 
ed in the first two sections can 
be induced in the tissues of a 
part freshly removed from the 
body. 


, 


re ee ee ee ee ee 


TISSUE METABOLISM. 93 


DESCRIPTION OF THE FIGURES. 


Fig. 1. Corpuscles from the healthy blood of an earth-worm, 
a, a, amceboid-like corpuscles (x 250). 


Fig. 2. Congested blood-vessels from around ventral nerve-cord 
after irritation of integument of an earth-worm (x 90). 


Fig. 3. Tissue débris (75 minutes) after the application of acetic 
acid to an actinia mesembrianthemum (x 250). 


Fig. 4. Tissue proliferation (10 minutes) after the application 
of blistering fluid to an anemone (x 250, p. 83). 


Fig. 5. Tissue proliferation from the same specimen, as pre- 
ceding figure, about an hour and a quarter later (x 250, p. 83). 


Fig. 6. From the same specimen as the two preceding figures 
about 21 hours after the injury (x 250, p. 83). 


Fig. 7. From the same specimen as the three preceding figures, 
43 hours after injury (x 250, p. 83). 


Fig. 8. Corpuscles from a healing surface immediately after the 
removal of a slough from an anemone ( x 250). 


Fig. 9. The same from an earth-worm (x 250). 


Fig. 10. The same as preceding figure after the addition of acetic 
acid, shewing the contraction of the corpuscular contents (x 250). 

Fig. 11. Corpuscles from beneath epidermis of L. Terrestris 
about 20 hours after injection of fcetid matter into its body (x 250, 
p. 89). 


Fig. 12. Fibrille and corpuscles from tissue-proliferation after 
prolonged irritation. lL. Terrestris (x 250, p. 90). 


N.B. Examined in water. 


OBSERVATIONS ON THE DISTRIBUTION OF SOME. 


OF THE NERVES OF THE HEAD AND NECK. 
By D. J. Cunnincuam, Student of Medicine im the 
University of Edinburgh. E 


AT the suggestion of Professor Turner, and in conjunction with 
my fellow-student Mr G. Kirkwood, I have recently made a care- 
ful dissection of the Nerves of the Head and Neck, in order, if 
possible, to throw some additional light upon the nervous 
system in this region. And, though I cannot claim to have 
-discovered anything altogether unknown, I think that I have 
been able to trace the connections and distribution of some 
nerves, which had not so been traced before, and this may 
perhaps lead to the rectification of some old theories and the 
confirmation of others. The points which I have chiefly noticed 
as being less or more new are as follows :— 

Superior Cervical Ganglion of the Sympathetic.—T wo branches 
of small size proceeded from this ganglion, passed through the 
prevertebral muscles, and finally terminated in the ligaments 
and vertebre of the upper cervical region. These branches are 
not mentioned in any of the books I have consulted. The 
connection which existed between the sub-occipital nerve and 
the ganglion was very intimate. I found a whole series of 
branches—4 or 5 in number—passing from one to the 
other. Before the sub-occipital nerve disappeared from view 
over the transverse process of the atlas, it gave off a few twigs 
which passed to the articulation between the occipital bone 
and the atlas, and acted doubtless as the nerves of supply to this 
important joint. These branches have (according to Quain) 
been described by Valentin. I may mention that I could find 
no branch of communication passing between the superior 
cervical ganglion and the spinal accessory nerve, and no such 
branch has been described so far as I am aware; but it is strange 
that this ganglion, which is so liberal in giving off branches to 
the other two divisions of the eighth cranial nerve, should fail 
entirely in bestowing a similar favour upon the spinal accessory. 

With regard to the Middle Cervical Ganglion I was able to 
make out the following points:—As the phrenic nerve lay 


F 
. 
} 
j 
| 


i 


NERVES OF THE HEAD AND NECK. 95 


between the fourth and fifth cervical nerves, a slender branch 
of communication passed between it and the middle cervical 
ganglion. This communicating nerve gave off a slender but 
strong branch to the coat of the transversalis colli artery, to 
which it doubtless acted as the vaso-motor nerve. <A well- 
marked sympathetic plexus surrounded the inferior thyroid 
artery, and communicated with the fourth and fifth cervical 
nerves. From the fifth cervical nerve an accessory phrenic 
arose, and passed down on the surface of the scalenus anticus 
external to the phrenic itself and to the outer side of the 
internal mammary artery, to enter the thorax and join the 
phrenic nerve in the upper part of that cavity. This is an 
abnormality which, according to Krause, rarely occurs; but two 
cases of an almost similar nature have been described by Prof. 
Turner in this Journal, vi. p. 102. In connection with the 
Inferior Cervical Ganglion, I found a slender branch of com- 
munication passing between it and the phrenic nerve. Both 
this branch and that from the middle cervical ganglion to the 
phrenic nerve are mentioned by Krause and Telgmann*, Mr 
Kirkwood was able to make out, on the opposite side of the 
body, a connection between the inferior ganglion and the 
pharyngeal plexus. This branch passed up behind the sheath 
of the common carotid artery and, resting upon the prevertebral 
muscles, joined the lower part of the plexus. 

Long Buccal Nerve-—An additional proof that its function 
is sensory and not motor as is often represented. 

On two occasions Prof. Turner has seen this nerve coming 
off from the superior maxillary division of the fifth cranial 
nerve’, a trunk which has never been doubted to be sensory— 
and it is known that when the muscles of expression are para- 
lysed the buccinator muscle (of which the long buccal is often 
represented as the motor nerve) is paralysed with them, and 
that when the muscles of mastication are paralysed the bucci- 
nator muscle is unaffected. These two facts are almost suffi- 
cient of themselves to prove the sensory function of this nerve, 
but, as additional evidence might be obtained by dissection, 
I followed out the branches of distribution of this nerve very 


1 Die Varietéiten der Nerven, Leipzig, 1868. 
2 This Journal, November, 1866, and Proc. Roy. Soc, London, 1868. 


96 MR CUNNINGHAM. 


carefully with the following result. Of the numerous branches, 
which the long buccal gives off, as it emerges from under 
cover of the ramus of the jaw, I succeeded in finding only two 
which did not in any way communicate with the facial nerve. 
These I traced through the muscular fibres of the buccinator 
muscle by dissecting it with needles under water, and found 
that they broke up into a great number of very minute branches, 
which entered the sub-mucous coat of the mucous membrane of 
the cheek. In my dissection I failed to find a pure facial branch 
entering the buccinator muscle, and this muscle was too much 
torn up to allow of my tracing one of the’ mixed branches. 
Posterior Auricular Nerve—There appears to be some dif- 
ference of opinion in regard to the distribution of this nerve. 
Quain* and Ellis* describe it as supplying the posterior belly of 
the occipito-frontalis muscle and the retrahent muscle of the 
auricle, whilst by Flourens the posterior belly of the oecipito- 
frontalis is represented as being supplied by the small occipital 
nerve. ‘To settle this point Mr Kirkwood made a careful dis- 
section of the nerve. After leaving the facial nerve close to the 
stylo-mastoid foramen it divided into two branches in front of 
the mastoid process. One of these proceeded upwards for a 
short distance, then passed beneath the posterior belly of the 
occipito-frontalis and broke up into branches, which entered 
the deep surface of this muscle. The second division of the 
nerve ran backwards under cover of the retrahens aurem, and, 
giving a branch to the deep surface of this muscle, curved 
round the pinna, and gave branches to the deep surface of the 
attolens and attrahens aurem. It is thus seen that the pos- 
terior auricular nerve is to be looked upon as the motor nerve 
to all three muscles of the auricle and also to the posterior 
belly of the occipito-frontalis. The small occipital nerve rami- 
fied over the surface of this last muscle supplying the integu- 
ment in this region, but giving no branches to muscle. It is 
well known that nerves of supply to muscles situated super- 
ficially enter their deep surfaces, and this fact also tends to 
show that the small occipital cannot be the nerve of supply to 
the occipito-frontalis. The small occipital nerve, therefore, 


1 7th edition, by Sharpey, Allen Thomson, and Cleland, p. 612. 
2 Demonstrations, 6th edition, p. 8. 


i 
7 


NERVES OF THE HEAD AND NECK. 97 


(along with all the other branches of the superficial cervical 
plexus) is to be looked upon as a nerve purely sensory in its 
function and distributed to integument. 

Great Auricular Nerve.—The facial or anterior branches of 
this nerve appear to have a more extensive distribution than is 
generally ascribed to them, viz. the supply of the integument 
over the parotid gland. In the dissection I made of this nerve 
these branches were five in number and had the following 
distribution: (a) the most anterior and inferior pursued a 
course almost parallel with and about three-fourths of an inch 
above the inferior margin of the horizontal ramus of the lower 
jaw, and reached a spot about half-an-inch from the mental 
foramen. (b) The most posterior branch proceeded almost ver- 
tically upwards to a point about one inch from the outer angle 
of the eye, and was there lost. (c) The intermediate branches 
passed upwards and forwards towards the angle of the mouth 
and malar bone and ended about one inch from these. In 
their course these branches gave off numerous twigs to the 
skin in this region. 

Practically speaking, therefore, these branches supplied the 
integument over the parotid, masseteric and buccal regions of 
the face. Mr Kirkwood confirmed this dissection on the oppo- 
site side of the body. 


VOT. VEL. 


“J 


ON SPHYGMOGRAPHY. Part II. By A. H. Garrop, B.A., 
St John's College, Cambridge. 


In endeavouring to form a correct estimate of the significance 
of the various details of the sphygmograph trace, it will be 
necessary to enter somewhat minutely into the consideration of 
each of the several mutually related forces which, by their com- 
bined action, produce the resulting curve. As some of these 
forces are but little understood, it is clear that any attempt 
to explain the pulse movements by arguments deduced from 
tracings obtained from a ‘schema’ of the circulation, can only 
be of value, as far as they relate to forces acting on the 
‘schema’ which are strictly comparable with similar forces at 
work in the human body. This consideration has prevented 
my basing any deductions on facts derived otherwise than from 
the animal body itself. 

To commence with, it will be necessary to describe the 
tracings obtained by means of the ‘Heemadromograph’ of 
Chauveau. Dr Lortet, of Lyons, has given an excellent figure 
and description of this instrument’, which, from its simplicity 
and efficiency, is a perfect masterpiece of mechanical design. 
By means of two levers it indicates on a revolving drum both 
the modifications in the diameter of an artery during a pulse- 
beat, and also the changes in the velocity of the blood-current 
at the same spot, simultaneously. The former of these results 
is obtained by the use of an ordinary sphygmoscope, an instru- 
ment which measures indirectly the modifications in the area 
of an artery; and the latter by a long lever which, at its at- 
tached end, after passing through a slit in the side of the tube, 
projects a short distance into the vessel, which allows of its 
being moved backwards and forwards by the blood-current. 

The accompanying trace (Fig. 1), taken with this instrument 
from the carotid of the horse, is copied from Dr Marey’s work’. 


1 Recherches sur la Vitesse du Cours du Sang dans les Artéres du Cheval. 
Par M. L. Lortet, M.D. Annales des Sciences Nat., Tome vu. Zoologie, 1867. 
An earlier but very similar instrument, by M. Chauveau, is described in Marey’s 
Physiologie Médicale de la Circulation du Sang. 18638, p. 156. 

2 Loc. cit. p. 273. . 


SPHYGMOGRAPHY. 99 


Fig. 1. Hemadromograph trace from the carotid of the horse. 

a. The current trace, in which the horizontal dotted line is the zero, indi- 
cating no current in either direction. 

B. The sphygmoscope trace. 

In both, the vertical line 1 cuts the curve at the commencement of systole, 
the line 3 at the moment of closure of the aortic valve, and the line 4 when the 
centrifugal current reeommences. 

The figure reads from left to right. 


The upper curve (a) is that of the blood-current; in it the 
horizontal dotted line indicates the zero, or line of no :current 
in either direction ; all above shews a centrifugal current, and 
all below an heartward stream. The velocity of the blood’s 
movement is measured by the distance of the point which is 
under consideration from the zero line in the trace. The lower 
curve is that obtained by the sphygmoscope, and is in all re- 
spects strictly comparable with an ordinary sphygmograph 
tracing. The slightly curved vertical lines which intersect both 
tracings are drawn by the levers when the watchwork is at 
rest, and the points where similarly situated lines intersect the 
pulse traces are exactly simultaneous, to indicate which they 
are drawn. With regard to these figures, Marey makes the 
following observations :—“I. The commencement of the pulsa- 
tion coincides with the production of a rapid centrifugal current. 
II. The summit of the pulsation is not reached before the cen- 
trifugal current has already ceased. III. At the instant of 
closure of the sigmoid valves, a retrograde current is Howing in 
the carotid, as indicated by the position of the curve, which at 
that moment is below zero. IV. After the closure of the sig- 
moid valves a fresh centrifugal current originates, which consti- 
tutes the secondary pulsation—(dicrotism).” Dr Lortet explains 
the portion of the upper or current-trace which is below the 


~ 7 
(—2 


100 MR GARROD. 


zero line, by supposing that the shock of closure of the sigmoid 
valves is sufficiently great to produce a stretching of the proxi- 
mal end of the aorta and of the valves themselves, which results 
in a retrograde current for a short time in the large arteries 
after the valves have closed. Marey’s explanation, as far as it 
goes, seems much more satisfactory, and it will be necessary to 
discuss it more fully. Before doing so the precise definition of 
some of the terms employed will not be out of place. The 
main rise in the sphygmograph trace may be termed the pri- 
mary rise, and that which occurs just after the small rise and 
fall which is nearly simultaneous with the sinking of the current- 
trace below the zero, and which therefore commences with the 
secondary centrifugal current, may be termed the secondary 
rise; it is almost always very clearly indicated in sphygmograph 
tracings (c. a Fig. 2). The interval between these two rises may 
be called the sphygmosystole, for it 1s the time during which 


Fig. 2. Sphygmograph iracings, all taken from the same individual in 
health, under different conditions, to shew the effect of difference in pulse-rate 
on the trace. 

Pulse-rate 44 in a minute. 
63 
72 
103 
137 
172 


2 272 DE 


~ 


They are all drawn on one scale and read from left to right. 


SPHYGMOGAPHY. 101 


the systole at the heart influences the pulse-beat; it corresponds 
to the first part of the pulse-beat in my former communica- 
tions, and it must be remembered that it is not synchronous 
with the cardiac systole, as I have endeavoured to shew 
elsewhere’. 

From a study of tracings obtained by the use of the cardio- 
sphygmograph, I have demonstrated, in the paper just referred 
to, that in the sphygmograph trace of slow pulses, the closure 
of the aortic valve occurs when the lever is at the lowest point 
of the notch that is nearly always present in the sphygmo- 
systole, which is also clearly seen in the lower of the two 
tracings in Fig. 1, the curved line 3 cutting it exactly at the 
place which represents the moment when the valve shuts. 
Consequently, in the upper trace, the line 3 must also cut it at 
the same time, and this occurs when the retrograde current is 
at its maximum, as would be expected, the valve being closed 
by the regurgitating blood. But it may be asked, how is it 
that the backward current between the lines 3 and 4 in a Fig. 1 
is associated with a rise in the sphygmoscope trace? and to 
explain this clearly, 4t will be necessary to refer to some of the 
elementary principles which operate in the transmission of cur- 
rents through elastic tubes. First, it can be demonstrated 
that increase in the diameter of an artery may originate from 
two quite independent sets of causes, one being the simple re- 
sult of the heart sending more blood into it in a given time 
than it can dispose of ; the other being a shock-expansion, com- 
parable with the waves of condensation and rarefaction in the 
air, which constitute sound. Most of the important elements of 
the pulse-trace are referable to the former of these causes, as is 
proved by the general similarity between the two curves a and 
8, in Fig. 1. But when there are rises in the second, which are 
quite independent of changes in the first, they must evidently 
be the result of some sudden movement in the circulating sys- 
tem, originating a shock, which is not of a character to affect 
the current in any way. It is but reasonable to suppose that 
such should occur as a result of the closure of the semilunar 
valves, and the rise and fall between the vertical lines 3 and 4 


1 Proce. Roy. Soc. 1871, p. 318 et seq., and Journal of Anat. and Phys. 
May, 1871. ; 


102 MR GARROD. 


in the lower trace (8) of Fig. 1, commencing immediately on 
their closure, and being unconnected with any current changes, 
can be nothing else than a shock-wave. 

At first sight it would seem probable that when the semi- 
lunar valves had closed, the retrograde current in the aorta 
would immediately cease; but that such is not the case is 
clearly proved by the continuance below the zero line of the 
current curve, for a short period after that event (between 
the lines 3 and 4 in the upper trace). It is by the combined 
operation of two causes in the same direction that this result is 
produced. First, the orifices of the coronary arteries being 
quite close to the semilunar valves, it is evidently necessary 
that the blood which enters them during diastole must be 
derived from the aorta, and so tends to produce in it a retro- 
grade current. Secondly, the equilibrium of the arterial system 
is disturbed during the closure of the aortic valve, for imme- 
diately the systole has terminated, the only force tending to 
prevent the blood from regurgitating into the heart, is the 
statical resistance of the ventricular walls, which at that moment 
are closely approximated, causing their cavities to be completely 
‘obliterated. This resistance is clearly much less considerable 
than that offered by the heart-walls during the systole, one 
being a statical and the other a dynamical condition ; conse- 
quently the arterial blood rushes back, pushing asunder the 
ventricular walls, and in so doing, developing a sufficiently 
rapid retrograde current to close the semilunar valves. The 
interval thus arising, namely the time between the end of 
systole and the complete closure of the aortic valve, I have 
called the diaspasis, in a pamphlet on “the Law which regu- 
lates the frequency of the Pulse’,” and there is considerable 
evidence to shew that when the pulse-rate does not vary it is 
constant; also that it varies very slightly with different frequen- 
cies of heart-action, occupying in slow pulses about 0:002 and 
in quick ones 00018 of a minute*. The diaspasis is so short in 


1 Published by H. K. Lewis, 139, Gower Street, London. 

2 In my pamphlet on The Law which regulates the Frequency of the Pulse, 
from a mistake on my own part, a statement is made as to the length of the 
diaspasis, which is incorrect. Mathematical friends soon informed me of my 
error, as, when precisely stated, the problem is a manifest contradiction. In 
p. 22 the diaspasis is discussed, and in the summary of the results arrived at, 


SPHYGMOGRAPHY. 103 


duration that it has terminated before the loss of blood from the 
proximal aorta into the heart is made good by any retarda- 
tion of the onward current, or a reverse stream in the more 
distant vessels originates ; consequently this restoration of equi- 
librium has to be in great measure effected after the valves 
have closed, encroaching on the diastole to form the second 
cardio-arterial interval of the cardio-sphygmograph trace, which 
does not cease until all the complications attending the closure 
of the semilunar valves have come to an end, whereupon the 
again augmenting centrifugal current, or in very low-tensioned 
pulses, such as that figured in Fig. 1, the diminishing centri- 
petal one, originates the secondary rise in the sphygmoscope 
trace. 

In both the primary and secondary rises of the pulse-beat 
(Fig. 1), it is found on inspection that the summits of the pulsa- 
tions (8) are delayed upon, or are not reached so soon as the 
centrifugal current maxima (a); and that such should be the 
case is essential, in a circulation maintained by an intermittent 


in p. 27, the following are given as the relations of the lengths of the various 
elements of the pulse-beat to one another. 


1. The systole together with the diaspasis, or, in other words, the first car- 
diac interval varies as the square root of the whole revolution. 

2. The systole varies as the square root of the diastole. 

3. The diaspasis is constant, 


The incompatibility of these three statements it is not difficult to prove, 
for if the first and second are true, the third cannot be so, and, as a fact, the 
diaspasis is not constant. There is every reason to believe that for a given 
pulse-rate the diaspasis does not vary; and, on subtracting its length, as ob- 
tained by measurement in quick cardiograph tracings, from the first part of slow 
ones, I found that the remainder, the systolic interval, varied very nearly as the 
square root of the diastolic. On repeating the operation on the pulse of inter- 
mediate rapidity a similar result was obtained, and the error, being extremely 
small, I attributed it to my not having extracted the necessary square roots to 
a sufficient number of decimal places, and thus felt justified in making the 
generalization given above. Since the mistake has been pointed out to me I 
have repeated the arithmetical computations more carefully, and find that what 
I had first supposed were errors on my part, are constant variations, which prove 
that, the other statement standing as above, the diaspasis is slightly longer in 
slow pulses, occupying approximately 0°002 of a minute in a pulse of 61, and 
0:0018 of a minute in a pulse of 152 in a minute. This fact, therefore, leads to 
the conclusion that the rapidity of the fall of blood-tension has an influence on 
the length of the diaspasis, lengthening it slightly when the tension-fall is re- 
tarded, probably because the previous systole is then more powerful and gradual. 
It is to be noted that the second eardio-arterial interval of cardio-sphygmograph 
tracings is almost of the same length as the diaspasis, and varies in the same 
or in a very similar manner, which may be the cause of the somewhat undecided 
nature of the notch in the sphygmosystole of slow pulses. From these remarks 
it is necessary to substitute for statement 3, as given above, the following: 
3. The diaspasis varies, being slightly longer in slow pulses. 


104 MR GARROD. 


motor organ, which, like the heart, rests between its pulsations ; 
for after the rush of blood into the arteries immediately the 
sigmoid valves open, during the rest of the systole the blood 
which leaves the heart is employed in retaining the higher 
pressure in the vessels, as will be explained more fully fur- 
ther on. 


We are now in a position to consider the human sphygmo- 
graph, tracing from the wrist, and on looking at Fig 2 (a), 
which is from a slow pulse, it is evident that in all respects 
it closely resembles that taken with the sphygmoscope from the 
carotid of the horse, which has been discussed above, and there 
is every reason to believe that the details originate from similar 
causes. The primary rise (a) is followed by a gentle fall; this 
is soon broken by the shock-wave (+) consequent on the closure 
of the aortic valve, and is followed by the secondary rise (ce), 
which commences when the centrifugal current is augmented 
by the recommencing onward current in the aorta. The sphyg- 
modiastole is remarkably uniform and uninterrupted. 

Between the tracings of slow and quick pulsings there is at 
first sight not much resemblance, but it is not difficult to obtain 
a series between them exhibiting every mtermediate condi- 
tion (Fig. 2, a, 8,7). The most important cause of the varia- 
tions exhibited by pulses of different rapidities, is that the ratio 
between the length of the sphygmosystole and the sphygmo- 
diastole is not constant. For instance, when the pulse is ll4a 
minute, the sphygmosystole occupies just one half the beat ; but 
when it is 40 a minute, it only occcupies one quarter of the 
whole revolution. It is evident that this must influence the 
general appearance of the trace, and as the length of the sphyg- 
mosystole never varies in health for any given pulse-rate’, a 
knowledge of the ratio of the length of sphygmosystole to 
that of the whole beat is sufficient datum for determining the 
pulse-rate. In the paper just referred to I have given several 
measurements of these ratios, and have shewn that the length 
of the sphygmosystole maintains a very definite relation to the 
length of the beat, varying as its cube root, consequently when 


1 Proc. Roy. Soc. No. 120, 1870. 


SPHYGMOGRAPHY. 105 


the length of the revolution increases in the series 1, 8, 27, 64, 
&e., the sphygmosystole only does so as 1, 2, 3, 4, &e., so that 
if we call the rapidity of the pulse a, and the number of times 
that the sphygmosystole is contained in the beat y’, the length 
of the sphygmosystole can be found from the equation 

ay =47 J/x. 

Further, the cardio-sphygmograph’ shews that the interval 
between the closure of the aortic valve and the commencement 
of the secondary radial rise (the second cardio-arterial interval) 
varies but little with different pulse-rates, while that between 
the primary radial rise and the closure of the aortic valve (the con- 
jugate cardio-arterial interval) does so much more rapidly, both 
being longer in slow pulses. This also greatly influences the 
appearance of the pulse-trace, for, as previously shewn, the 
small rise and fall at the end of the sphygmosystole results 
from the shock of closure of the aortic valve, and as this occurs 
in slow pulses an appreciable time after the primary rise has 
reached its maximum, it is clearly seen as a separate element of 
the curve. But in quick pulses, the second cardio-arterial 
interval is nearly as long as in the slow ones, while the conju- 
gate cardio-arterial interval is much shorter, consequently the 
shock-rise and fall following the aortic valve closure is thrown 
back, as it may be termed, on the primary rise, and being blended 
with it, is not separately distinguishable. This is the cause of 

the simplicity in the sphygmosystole of quick pulses. 


1 Journ. of Anat. and Phys. May, 1871. 


THE EFFECT OF EXERCISE ON THE BODILY TEM- 
PERATURE. By T. Ciirrorp ALLBuTT, M.A., M.D., 
Cantab., F.LS. 


Reprinted from Zhe Archives of the Royal Society. 


For some years past I have been engaged upon the interesting 
study of the temperature of the human body in health and 
disease, and I have summed up certain results of my experi- 
ence in articles in thé Medical Chirurgical Review and else- 
where. In estimating these results, it is found to be of the 
greatest importance to the enquirer, to have a satisfactory basis 
for comparison, and such a basis or standard, we think, we have 
in the temperature of the healthy body. It becomes, therefore, 
a primary duty to settle as firmly as possible, the limits of 
fluctuation in health, and the effects of various external condi- 
tions upon these fluctuations. Many observers, among whom 
I may name the late Dr Davy aud Dr William Ogle, have 
laboured hard in this matter, and we have had the satisfaction 
of seeing that the regulating power of the organism is so per- 
fect that external conditions have but little effect in disturbing 
its ordinary periods. In summing up, however, I found that 
the effects of hard and prolonged exercise upon the body had 
not been tested, and this seemed a serious omission. 

When we consider the great mechanical work performed by 
the body in muscular exertion, and the molecular change corre- 
lative with it, we might well expect to find that force diverted 
in this way might leave a great deficiency of heat. The mar- 
vellous regulating power which the human body has been 
shown to possess, may, or may not, be able to make up for 
this outgoing by increased combustion within. AIL our present 
experience, however, would lead us to look for this. When we 
see that plungmg into cold water, by driving more blood 
through the lungs, and other viscera, makes up for the ab- 
straction of heat; when we find, as I and others have found, 
that exposure to heat of 300’ F., on the other hand is neutra- 
lized by evaporation from the surface and lungs, the body presery- 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 107 


ing in both instances its normal temperature; when we consider 
its resistance, I say, to cold and heat, to any interference with 
function which is not actually mischievous, and to many like 
serious disturbances, we gain so great a faith in this capacity 
for self-regulation, that we expect it to meet the strain of hard 
exercise with like success, unless the strain be beyond the 
powers of the individual. When we initiate a state of disease, 
we shall of course look for perturbation of the temperature 
curves, but knowing what we do, we should scarcely look for 
perturbation of the curves to any marked extent so long as the 
muscular exertion used did not exceed that quantity which the 
body seems built for. Such were the opinions I held in starting 
for Switzerland this summer’, but I felt I could not take the 
uniformity of the curves for granted until I had submitted 
them to the test of repeated experiments, so that I made these 
experiments one of the main objects of my tour. Unfortu- 
nately, the weather was so unpropitious, that we were pre- 
vented from making high excursions. I think, however, that 
we were able to do work enough to test fairly the effects of it 
upon the body. When I began my observations I was un- 
aware that experiments of the same kind had been carried out. 
However, Mr Ray Lankester, whom I met by chance near 
Martigny, told me that M. Lortet of Lyons had made two 
ascents of Mont Blanc for this purpose, and that he reported 
great depression of temperature as his results. 

I had been walking for a few days when I heard of this, 
and had made complete observations for two days. Two days 
afterwards I met a Swiss gentleman at Chamouni, who placed 
in my hands M. Lortet’s Deux Ascensions du Mont Blanc en 
1869: recherches physiologiques sur le mal des montagnes, par 
M. L. Lortet, 1869. I shall refer again to M. Lortet’s results 
after reading my own. Dr William Ogle makes some allusion 
to the effects of exercise upon the body, and states, in opposi- 


1 This refers to the summer of 1870. In the summer of 1871 the weather 
enabled me to take some higher excursions, but I only made a few isolated 
thermometrical observations, which presented nothing contradictory. As regards 
the observations now published, I have to say that although these probably re- 
present all that were carefully made, yet, on the other hand, the thermometer 
was rarely out of my mouth in the intervals during the day-time, so that the 
movements of the mercury were constantly controlled. Of course this added no 
little to the exertion of walking. 


108 DR ALLBUTT. 


tion to M. Lortet, that it has the effect of elevating the bodily 
temperature: but Dr Ogle does not appear to have investi- 
gated the matter on a large scale. Before setting off on my 
journey I had been engaged for some months in studying my 
own temperatures under all sorts of changing circumstances, 
so that I had become thoroughly familiar with the variations of 
my own body. I here append an average scheme of my own 
bodily temperature, which represents the average of the curves 
under ordinary circumstances, taken hourly on twenty-five 
separate days. 


Average of 25 days at various dates noted during the 
earlier summer of 1870 for the purpose of the subsequent im- 
vestigations. My ordinary habits give me but little physical 
exercise. I breakfast at 8.30, lunch at 1.30, and dine at sia. 
No temperatures were taken in bed. The gradual curve con- 
trasts strongly with the sharper curves taken in Switzerland. 


=8 8 oP oP oP CE oP oP e 
Set Pe Oe oo A) SIS EE Ste SA 
in Wee Bacal Sag SNS aes : 3 28 q 
— Pe ora) a2 ip bo — — 2 
_EcD (q2} @® oO iq) (q°) a) t¢*} : 


‘TUOLVUadI WE Y, 
AUVNIGUOG JO ANVUGAW 


“‘SaUALVAAMNAT, 


August 7, 1870. Bex to Les Pans. 


Hours. CoNDITIONS AS REGARDS E}xERCISE. 


TEMPERA- 
TURES. 


9.30 a.m.| At Bex.. After breakfast .......c0000c0000 97.8 
11.30 ... | An hour on the walk to Les Plans ...... 98.2 
ee ae Still walking, Sun very hot.................. BOL 

5.0... |) Walking on slopes. Continuous eo 99.3 

TAO. } all day with but little food............... 99.1 

8.30 ... | After supper: at Chalets .................. 98.8 

9.15 97.8 


On going to OG: zac casenantantsocmocmtcmeesaeee 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 109 


August 8, 1870. 


Dent DE MorRctes. 


Hoors. CoNDITIONS AS REGARDS EXERCISE. esa 
emOmaem. #1) En, bedi. swan. Sesh doe ak iak. dees 97.5 
4.45... Upt: Beforesbreakfast lesen. sian stu sacked 97.5 
5.30... | After breakfast. Before walking ......... 97.7 
6.30... | After aa hour’s quick walking ............ 98.6 
Sree tees Ie WV GUTS Sooo 35 ds a dinsicinlai SRO ea oe tak be eae 98.9 
9.0 After sitting without coat in cold breeze 98.5 

fonshd ani wtesinckpate acete-pacsaaeseet 
TOS... Sitting on Col, after some food ............ 99.2 
12.30p.m. | Standing about. Exercise moderate...... 99.0 
3.0 Walking home quickly. Near Les Plans 99.2 
6.0 At Les Plans. After rest. Before dinner 99.2 
8.30. After an hour’s evening walk............... 98.1 
August 9, 1870. Les PLrans To Martieny. 
8.15 a.m. | Walking about (20m. or so) before breakfast 98.4 
9.15 After Ibreakdast .ps.sa1 ce: sis sesas 05) ee acene 98.6 
120.9 Walking down to Bex from Les Plaus ... 93.8 
Z30-P.M.-|s Near Bex. Walking... 20... e202 2708 6 Sey 
‘ Near Martigny. After 4hrs. sharp walk )| 99 9 
50 ing. Wet throvch FAG hs Bao ks.) ica 
10.0 In bed. (Dined at Martigny 6.30) ...... 97.8 
August 10, 1870. Marticny to CHAMOUNT. 

aA.) in bed! at. Martigity [0 isemgsces tea Jes aoe 97.5 
7.30 After breakfast; * 605 2B x2 os. cocds éo0 cee 97.8 
9.30 Walkine® Day Nob? <....00<2tesencceete 98.8 
<) aes 
ae } Ascending steep slopes under hot sun = He 
2.30 p.m. | After rest and lunch at Barbarine......... 99.0 
4.30 Walking for two hours .........5........05. 99.2 
5.30 Walking through Argentiére ............... 99.0 
8.30 After dinner, before retiring at Chamouni 97.8 


August 11, 1870. 


PLAN DES AIGUILLES. 


9.0 A.M. 


Before starting for Plan des Aiguilles . 


Walking briskly up steep ascent. 98.5 


though ..: 74. [sees Gee eet J 
In Chalet. After food and rest............ 98.7 
Pralees down to Chamouni. Nearly a 98.8 
IONGOHT 26. eee RR ae NR COe Sole : 
98.6 


Dinner at 6.0 ... 


‘This day was of. cold drizzling rain. 


bat rest in Chamouni. 97.9 


110 DR ALLBUTT. 


August 12, 1870. CHAMOUNI TO TRELATETE. 
Hovrs. CoNDITIONS AS REGARDS EXERCISE. | Tee 
8.30 a.m. | Chamouni. After breakfast .. ............ 97.8 
10.30 {After walking nearly two hours. St Ger- 98.8 
: a | ly «wats: Road: is 2 seek. oi Aoi oes ee ; 
12.50 =... | Walking. to St: Gervais) 02). Usain ivacot SE) | 
2.45 P.M. | At Mont JoliInn ............ sone ERE 99.2 
4.30 ....-| Walking to:Contaminesy.c & sxietcecoss et 99.2 
70 Ascending steep slopes above Contamines. 100.0 
ee No sense of feverishness or of much thirst i 
8.30/23. | Before!bediatiChalets? 1... alee naestesd eee 97.8 
This day was very hot. Knapsack carried. 
August 13, 1870, Pass oF Mont Tonpv. 
3:00 amo |2 On-rising from bed-s.cace.c0rds scsvorees eee 97.5 
5.30 Ascending towards Col Trélatéte. On 97.9 
ea GIACIOR”  ciosec cists caves ence. teehee ee : 
7.0... | Thermometer removed on reaching Col... 98.2 
9.0 {Approaching Motet. Here breakfasted | 98.6 
AN Shaihd rested 1.5 shouts... :ccpetsek. tot | , 
12.0... | Reaching summit of Col de la Seigne...... 99.1 
3-0: wim,-} In A lléerBlanche@ sc. sce pcsts ices stemtscts oer 
5.0" 2... | Butering Cormayeur 2.50 sascessseceocs 99.1 
8 30 {In Cormayeur. After a cold bath and 97.8 
Vee ES Nt QUIN OR as cactevionscseee ciseeedpe puns siceesaenaen : 
August 14, 1870. CorRMAYEUR. 
9.30 a.m. |(Idle all day at Cormayeur until 3.30, 98.2 
11 ORs. when we started for the pavilion on 98.4 
2.0: Getae. ||. Montalnetin cc. se cdswseid Soepanuceae ceetad. 98.8 
5.30 ... | Taken while on ascent. Cool evening ... 98.9 
Taken just when retired to bed. Supper), " 
2 Se | with wine about 6.30 ................0006- } 973 
August 15, 1870. Cot pu GEANT. 
BO hat. re esata Before food. At Ae 97.3 
5D 2. POn top ot(Coltdn Geant). 2-2, 2 eekeeeen exe 98.0 
12.0 Noon. fOn Mer de Glace. After difficult descent) 98.6 
fA OR BERRIES: 2s Ueciisent wit, tlm adam «jee salaiem ete 
Seay icra ee: after long rest, food a 98.8 
6.0.%%.. | Muterme Chamoun, | oo 0c c.. sec canconsssses 98.6 


OiQir Votininp Woled::.id2195 le. kaos asaek eeeaee se 


97.3 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 111 


August 16, 1870. CHAMOUNI. 
Hours. | CONDITIONS AS REGARDS EX®=RCISE. eee 
9.0 a.m. | Coffee at 7.30. Taken while at rest...... 97.8 
11.0... | After breakfast with omelette. No wine 98.2 
Se Meo Atber SUCHE TONCH: oo us .cwccjoans- eters clipe ck 98.4 
Oe hee oe in lounging about the a 98.7 
0) Se Dinner, with wine; ab StXK_-6.5.schptenasases 98.5 
Uo anc, | before retiring to-bed ......2 00.0 s0ceeseesee'ne 98.0 
August 17, 1870. - Granps MUuLETs. 
aie J 
oe Just before leaving Chamouni. After)! 98.4 
PoKeaiasts Cane ots tee Mes. eth: 
100% =. Approaching Pierre Pointue. Walking 97.4 
Blea Wye sor chi kasi sapece eect eens hee 
11.0... | After breakfast with meat, wine, and rest 99.2 
Pe... ||) On way tor Grands Mulets) .5... 4-2-5. 98.0 
1.30 P.M. ms bree A cease Gin, e Seeeenae 96.3 
3.0 ... | Just reaching the Grands Mulets ......... 95.5 
3.15 ... | Seated for ten minutes in cabin ............ 98.5 
4.15 ... | Grands Mulets. After rest and food...... 99.0 
6.15 > is Just before bed ......... Jo 
The temperatures all through this day very 
curious and disorderly. 
August 18, 1870. Mont Branc. 
1.30 a.m. | At Grands Mulets. Before rising from bed | 97.5 
~ 3.30... | Walking on Mont Blane, since 3 o’clock SG 
5.0 3) 99 bP) 98.0 
G32 .. Descent commenced soon after 7 o’clock . 98.5 
8.30 . Descent to Mules, 5 min. from destination 98.5 
10.30 .... | Walking to Pierre Pointue (just starting) 98.6 
19230 Bar. {Approaching P. P., arrived as instrument) 98.0 
POMOVEG, diene teres rae eee. eee 
S50": At P. P., and after a meal with meat, &c. 98.7 
2.30: Ou way to\Chamouniis2 i)... 2 hee 98.9 
3.30 . . ee. Seren eer Ot REET Re 98.9 
5.0 At Coutet’s Hotel and after cold bath ... 98.9 
9.15 - 55 and turning into bed.. 97.6 


This was a day of very severe weather, 
and great exertion, owing to the state 
of the ice and of the snow. The tem- 
peratures were disorderly about noon. 


112 DR ALLBUTT. 


August 19, 1870. CHAMOUNI. 


Hoors. CoNDITIONS AS REGARDS EXERCISE. oe 
9.0 a.m. | After breakfast—without wine ............ 98.5 
Ot aera 98.8 
Punched at VSO ees aaacces face os seeeiereles are 
3.0 P.M. 99.2 
Wined Abi O72 Sec e cence oes «see natasseees re 
D0 Ses (On rete to bed: 2... eee mise ee one 97.5 
Poured with rain all day ; we were kept 
to the house entirely, or nearly so. : 
August 20, 1870. CHamounrt To GreNeva—By Carriage. 
5.30 a.m. |. Before rising from bed **}i23:2/:2.2.2.-5--25: 90.4 
| After breakfast—without wine. Afte) 
D230" 35. breakfast sharp diarrhoea. ‘Two evacua- 95.4 
| LIONS Hoe ee heer aa cee te ae eee See oe 
1030... 97.8 
12.30 p.m. | Tn carriage going to Geneva, and walking 98.5 
Susie bOWs Geneva. cacrcn: se eee eee acne 98.8 
a aha 98.8 
9.0 ... | Bed. Dinner with wine at seven......... Osa 


The schemes of temperature which I shall now exhibit are 
those taken while walking. My friend, T. 5. Kennedy, and 
myself, met Frangois Devouassoud at Bex, on August 6th, 
1870, with the intention of walking on the high Alps. The 
weather was very unsettled. On the 7th we commenced our 
walk. I walked generally without a knapsack, and weighted 
only with a heavy pair of boots and an axe. The thermometer 
I used was obtained from Messrs Harvey and Reynolds before 
starting: it was specially selected and tested. The instrument 
had an index, and in the majority of cases the observation was 
made by placing the instrument under the tongue, with the 
usual precautions, leaving it in situ for 15 or 20 minutes, and 
then noting the position of the index, the hour of its removal 
being noted at the same time. The observation was always 
taken while we were in motion, except in cases when the 
contrary is stated. On many occasions, however, I watched 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 118 


the thermometer while in place, and this I was able to do 
very easily with a small pocket mirror. I did not find any 
reason, however, to distrust the results of the index. As to 
the length of time of insertion, I have to say here the contrary 
of what I say in common clinical observations. In these a few 
tenths, more or less, make no matter, and we have much more 
cause to fear that medical men will shrink from the delay of 
long insertions, than that an insertion of four minutes will 
deceive them. In physiological research, however, one or two 
tenths is of great importance, and I have often found, both 
abroad and at home, when watching the instrument in place, 
that ten minutes was an insufficient time for the completion 
of arise. I have often seen the mercury rise two-tenths after 
ten minutes of insertion. On the 7th of August we walked up 
from Bex to the Chalets of Les Plans; we started about ten 
A.M., and as we extended our walk considerably by collateral 
excursions upon the slopes, we did not reach the chalets till 
late afternoon. The day was fine and very hot. On reference 
to the table it will be seen that my temperature rose to 99.4 
at 3.30 P.M. the highest temperature but one that I recorded. 
On the 8th we started at 3.0 AM, from the chalets, with the 
intention of ascending the Dent de Morcles. The weather was 
misty and chilly all day. On rising at 3.0 I found the temper- 
ature at 97.5, and this temperature I found to be my invariable 
night-temperature throughout. At 40 the temperature was 
the same and at that hour we breakfasted. At 4.45, after 
breakfast, the temperature had risen to 98.2. At 5.0 we started, 
and at 6.30 the exercise had brought up the temperature to 
98.8, an unusual height for so early an hour. At 8.30 I regis- 
tered the same, and then being hot I purposely removed my 
coat and sat on an exposed stone in the chill air for 15 minutes, 
during which time the instrument was replaced. On removal 
it registered 98.5, but I am not certain whether the drop was 
due to the coolness or to the cessation of exercise or both. At 
10.30 we had crossed the glacier and reached a depression just 
under the summit. My temperature on reaching it was 99.2, 
and I may say 99.2 was a constantly recurring number, and 
seemed, exceptions apart, to represent the regular temperature 
during the noon and afternoon when in exercise. This tem- 


VOL. VII. 8 


114 DR ALLBUTT. 


perature was often reached at 11 or 11.30, and persisted till 
about 5.30 or 6.0 P.M., when the fall set in. Under ordinary 
circumstances my temperature rarely exceeds 98.6 at any time 
of the day and seldom attains that until the early afternoon. 
On this day at 6.0, just before dinner, the evening fall had com- 
menced, the note being 98.6. 

We did not reach the summit of the peak, as the mists 
were around us, and we were unable to discover the way up 
the very ugly looking rocks at the last. I suppose we reached 
a height of about 9000 ft., the rise being a somewhat rapid one. 
The badness of the weather now broke up our plans of ascend- 
ing the Diablerets and running down the Sansfleuron Glacier 
to Lauenen. We were driven down the valley and walked to 
Martigny. On this day, the walking being easy and down 
hill, my temperature did not reach 99.2 till 4.30, the day 
however was cold and wet. 

August 10 was a hot day with a good deal of sun, We set 
off intending to walk by the new route to Chamouni, but as 
some mining operations were going forward, we passed up some 
very steep slopes as high as Salvent and dropped down upon 
the new route opposite to the Téte Noire Tunnel. The hot 
day and quick ascent brought me up to 99.2 at 10.30 in the 
forenoon. At 6.30, between Argentiére and Chamouni, the even- 
ing being cool, I registered 98.8. [Not given in curve.] 

August 11. Day chilly and foggy with light and heavy 
rains. Walked in rain up to the base of the Aiguilles, coming 
home by a détour. Although the ascent was somewhat rapid 
my temperature on this day never exceeded 98.7. I was wet 
through most of the day. 

Aug. 12. Day very different to yesterday, not very clear on 
heights but soon bright and hot. The difference between this 
day and the day before in temperature is remarkable, the 
highest range being exactly 100°; we left Chamouni at 8.30 
and walked to the Mont Joli inn at St Gervais. From thence 
we walked to Contamines, and it was in ascending some rather 
steep slopes from here to the Chalets of Trélatéte, that I found 
my temperature 100°. 

Aug. 13. A moderately fine day, not very hot. We tra- 
versed the Mont Tondu pass and the Col de la Seigne. Temper- 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 115 


ature at starting (4.0 A.M.) was 97.5 and rose steadily till 2.30, 
when it was 99.2. At this height it remained till 7 P.M. when 
it fell gradually to 98.5 on retiring to bed at 9 o'clock. 

Aug 14, Idle all the morning at Cormayeur. At three 
o'clock went up to the pavilion on the M, Frety, The nse, 
which is tolerably steep and takes about two hours, had no 
effect upon the temperature. At 9.0 it had fallen to 97.5 after 
about two hours of rest and food. 

Aug. 15. Crossed the Col du Géant. Day fine but cool. 
Temperature steadily rose all day, reaching its height at 3 P.M, 
which was 98.8, we had then been long at rest at the Mont- 
auvert. On the top of the Col at 5.15 a.m. it was 98.0. 

Aug. 16. Idle all day in and about Chamouni. 

Aug. 17. Set off for the Grands Mulets with intention of 
ascending M. Blanc, On this day I noted two very curious 
falls of temperature, like those noted by M. Lortet, and I 
noticed them on this day and the next only. The first was at 
10.0 A.M., on approaching Pierre Pointue, when the temperature 
fell to 97.4 from 98.4. It rose to 99.2, at the pavilion, after an 
_ hour's rest and breakfast with wine, &c. At 12.25, when about 
halfway to the Mules, it had again fallen to 98,0, and when 
reaching the Mules the mercury had fallen to 95.5, two degrees 
lower than I had before noted. On sitting down at the Mules 
I again inserted the instrument for 15 minutes and found it to 
be at 98.5. At 4.15 it registered 99,0, and at 6,15, 99.1. Shortly 
after this we went to bed. 

On Aug. 18 the day was unpromising. At 1.30 on starting 
my temperature was 97.15. I took continuous observations 
upon the mountain, the temperature slowly rising. At the 
Grand Plateau it was 98.0 exactly. The weather now became 
more and more threatening, the wind rose, snow began to fall, 
and we had to contend with that kind of snow underfoot which 
has a crust just too weak for support and which suddenly lets 
the voyager 18 inches or more down at each step, Reluctantly 
we felt obliged to return, and at 10.30, on leaying the Mules for 
Pierre Pointue, I registered 98.6, which was above the average 
of that hour on ordinary days at home’. At 11.30 the tem- 


1 This represented a very hard day’s work, probably much harder than a 
complete ascent in fine weather, 


§—2 


116 DR ALLBUTT. 


perature had fallen to 98.3, and just before reaching the Pierre 
Pointue (12.30), it was down to 98.0. After rest and refresh- 
ment, I found it at 1.30 to be at 98.7, and at 2.30, when 
walking down to Chamouni, it was 98.9. This temperature 
remained steady for the rest of the day. The weather was 
cool, rather chilly, and some rain. I was not particularly 
fatigued. : 

Aug. 19. Rained steadily all day. Temperature at 3.0 P.M. 
99.2, and at 9.0 on retiring to bed was 97.5, shewing the effect 
of the exercise the day before in enlarging the curve. 

Aug. 20. The weather being still bad I determined to 
leave for England. Before rising (at 5.380) I was surprised to 
find the temperature down to 95.4, and there it remained after 
breakfast (at 6.30 o'clock). I then found myself suffering from 
a sharp diarrhea. It passed off quickly, and at 12.30, when in 
the diligence for Geneva, I had risen to 98.5. 

Such are the observations I made, and from them I draw 
the following conclusions : 

1. ‘That setting aside the exceptional results of Aug. 17 
and 18, the tendency of exercise was to raise my temperature. 
This was seen not only at the time of climax but also in the 
earlier commencement of the forenoon rise. ‘This conclusion is 
strengthened by the four observations which I made upon 
Mr Kennedy during steady walking. Each observation was 
made about noon or soon after, and 99.2 was registered on each 
occasion. 

2. That this rise was compensated by the earlier occurrence 
of the evening fall, which had often reached the night level of 
97.5 at 9 o'clock in the evening. ‘his also occurred Aug. 19, 
at rest all day. I found this to be the register of all the night 
hours I took, save on the one night preceding the diarrhea, 
when it fell to 95.4. 

3. Weather had some influence upon the temperatures. On 
hot days, when walking, I reached 99.2 almost invariably, on 
cool or chilly days I commonly fell short of 99.0. 

4. To turn to the exceptional day, Aug. 17. On this day 
and the next my temperature fell quickly and twice over during 
exercise, as though the mechanical work done were more than 
combustion could compensate. ‘This brings me to the considera- 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 117 


tion of M. Lortet’s observations, with whom the two exceptional 
events of mine seem to have been a constant occurrence. M. 
Lortet seems to have had a trustworthy instrument and to have 
used it very carefully. He says, “While walking, the decrease 
of the internal temperature of the body is very remarkable, a 
is almost proportional (the italics are his) to the height at which 
one is.” From the table given “one may state that during the 
muscular effort of ascent the bodily temperature may fall, when 
one is climbing from 3500 ft. to 15,000 ft., four and even six 
degrees centigrade, an enormous fall for mammals whose tem- 
perature is said to be constant.” “The moment one stops for a 
few minutes the temperature quickly reascends almost to the 
normal figure.” M. Lortet goes on to eliminate any possible 
accidents, such as cold breezes, &e. &c., then says that his ob- 
servations are equally true for small altitudes, for he was able 
to obtain the same results in a less degree by ascending small 
elevations near Lyons. M. Lortet adds that these results are 
not true for the hour which follows a meal, and partly explains 
this by supposing that the food serves as extra fuel to meet 
the excessive demand. The rest of his chapter is occupied by 
the reasoning which attributes the loss of temperature to the 
transference of a given quantity of heat to a given quantity of 
muscular effort, &e., estimating the value of a man’s position at 
so many feet above the sea level, and so on. Now I have to 
say, 1n the first place, that except on the day stated, I never 
observed this diminution of temperature, but rather the reverse. 
_ On the following day, Aug. 18, there was a recurrence of a 
like phenomenon, but it will also be seen that at that time I 
was actually descending not ascending. I was just completing 
the easy descent to Pierre Pointue, a distance of 3500 ft., or 
nearly so. On Aug. 17, however, when the diminutions were 
certainly remarkable, I may further point out that one of 
them (12.0 noon) took place an hour after the ingestion of a 
good meal with meat and wine. 

It is difficult to see how the mere presence of food in the 
stomach can help heat developement; one reason given by M. 
Lortet is, that it probably “accelerates the capillary or general 
circulation.” Unfortunately, a page before he adduces the accele- 
ration of the circulation through the lungs in ascending as a cause 


118 DR ALLBUTT. 


of imperfect lung-combustion and consequent cooling. Food in 
the stomach may perhaps act as a cordial, and so strengthen 
the impulses of the heart, but with this we should have a less 
frequency of action. Now how it is that M. Lortet’s temper- 
ature behaved so irregularly on both his ascents, and mine also 
during one very moderate climb by a mule path to the Pierre 
Pointue, is hard to say. It is touching upon delicate ground 
to compare the relative powers of our organisms, but I must 
say that there are one or two points in M. Lortet’s experience 
which suggest some difference of this kind. M. Lortet took a 
sphygmograph up M. Blanc, though I cannot conceive how he 
could use it on the slopes with any accuracy. However he 
gives several tracings, and these seem to shew a very serious 
interference with his circulation. The pulse seems to have 
been extremely rapid and small, and the arteries to be almost 
empty. The pulse curves to me seem indeed to be pulses of 
collapse. Even on the following day at Chamouni, M. Lortet’s 
pulse shewed a very irregular trace, a trace which would make 
me uneasy if it occurred in my own person. Now I had no 
sphygmograph, and here M. Lortet has the advantage of me, 
but I am perfectly satisfied that my pulse was never irregular, 
that it was never very small, and that it never exceeded 128, 
seldom indeed 120. Certainly I did not reach the summit of 
M. Blanc, but I am now comparing my own state on the Grand 
Plateau with that of M. Lortet at the Mules. It seems to me 
that M. Lortet’s temperature behaved in some exceptional way 
when he ascended M. Blanc, in a way which is unexplained, but 
which I also noticed on one day when walking up to the Pierre 
Pointue and the Grands Mulets, and which seemed to have some 
little tendency to remain with me on the followimg day, when 
not ascending but descending from the Grand Plateau to the 
Mulets. On the Dent de Morcles, on the Mont Tondu, on the 
Col du Géant, and in my lower walks, I never noticed the same 
thing in any degree whatever. I believe, therefore, that the 
normal effect of exercise is slightly to increase temperature 
during the day, and to favour the early occurrence of the even- 
ing fall when the day’s work is done. When the day’s work 
continues till 6.30 or 7.0 the fall is postponed until the time of 
rest, when it quickly sets in. 


EFFECT OF EXERCISE ON THE BODILY TEMPERATURE. 119 


While, therefore, I agree with M. Lortet in admiring the 
grand law set forth by the beautiful labours of Meyer, Joule, 
and Tyndall, and while I admit that “it is probably equally 
true both for living bodies and for machines constructed by 
the hand of man,” yet I must say that its manifestation in the 
way claimed by M. Lortet would go far to destroy our present 
faith in that other property of moving equilibriums, namely, 
the power of self-regulation under unusual but not excessive 
strain, which is but one expression of the existence of a highly 
complex organism*. I ought, before concluding, to say a word 
upon diet, though all who know what Alpine walking is, are 
also aware how irregular diet must be. On walking days I 
take a little coffee and milk with bread and butter before start- 
ing, and breakfast on bread and meat with wine and water on 
the glacier two hours later, or thereabouts. I generally drink 
very largely of milk during the day and but little of water. 
Milk is often the staple of food until dinner in the evening. I 
never take spirits at all, and light wine only in very small quan- 
tities. I feel a strong desire for such hydrocarbons as oils, butter 
and honey, and eat them freely. These materials, no doubt, are 
most valuable in supporting the excessive combustion. That 
but little nitrogenous food should pass off as urea may be 
partially explained by the increase in size and hardness of all 
the muscles during training. This is probably due to exuda- 
tion from the intramuscular capillaries under the pressure of 
contraction, and during the first few days I am generally 
aware of a tenderness and fulness in the muscles, due to con- 
gestion. I believe, accordingly, that no investigations into the 
excretion of nitrogen can be made with any valuable result, 
until training, with the rapid increase of muscular nutrition, 
has been fully established by a month’s regular walking. 


1 Mental activity, which deals with the construction of more complex mole- 
cules than physical, does not lower my own temperature, as very numerous 
experiments tell me. 


OBSERVATIONS ON THE STRUCTURE OF THE 
HUMAN PLACENTA. By Proressor Turner. (PI. v.) 


THE Human Placenta from its great importance as the 
organ of union between the mother and the fetus in utero has 
long been a favourite object of investigation, and many distin- 
guished anatomists have from time to time directed their atten- 
tion to the elucidation of its structure. But, notwithstanding 
all the time and labour which have been devoted to the sub- 
ject, there still remained various points respecting which 
differences of opinion prevailed, and to determine the true 
meaning of which further observations were required. With 
the view of reaching a definite conclusion on these disputed 
questions, I have for some time past availed myself of many 
opportunities of investigating this organ, and have examined 
it not only after expulsion from the uterus in the ordinary 
course of labour, but whilst still connected to the wall of the 
gravid uterus. 
I shall arrange my observations in the following sections : 


Ist. The relations of the maternal blood-vessels to the 
placenta. 

2nd. The arrangement and structure of the decidua serotina. 

3rd. The minute structure of the villi of the chorion. 


Ist. The relations of the maternal blood-vessels to the 
placenta’. 

No point in placental structure has been more the subject 
of discussion amongst anatomists than the arrangement of the 
maternal blood-vessels and their relations to the foetal capil- 
laries within the villi of the chorion. All indeed now agree 
that there is no direct flow of blood from the maternal into the 
foetal vessels or vice versd, and that mother and child each 
possess an independent circulation. It is also admitted that 
in the gravid uterus, when the placenta is fully formed, the 


1 Many of the facts contained in this section were communicated to the 
Royal Society of Edinburgh, May 20th, 1872, and an Abstract of these is in- 
serted in the Proceedings of that Society. 


STRUCTURE OF THE HUMAN PLACENTA. g (4! 


curling arteries do not communicate with the utero-placental 
veins through an intermediate plexus of capillaries. 

By several obstetricians, viz. Robert Lee’, Velpeau’, Seiler’, 
Ramsbotham‘, and Adams’, it has been asserted that the ma- 
ternal blood does not pass into the interior of the placenta, 
and this view has been revived by Braxton Hicks in a paper 
published in the May number of this Journal. 

About the middle of the last century John Hunter’, from 
the dissection of a gravid uterus, where the placenta was filled 
with wax injected through the curling arteries and uterine 
veins by Dr Mackenzie, and subsequently from injections made 
in conjunction with his brother William, concluded that he 
had established the circulation of the maternal blood through 
the placenta. This important conclusion has been supported 
and corroborated by the observations of many eminent investi- 
gators, more especially E. H. Weber’, Owen§, J. Reid’, J. 
Goodsir™, Van der Kolk”, Virchow™, Kolliker’, Arthur Farre*,, 
and Ercolani®. But though there is a common understanding 
amongst these observers that the maternal blood enters the 
placenta by the curling arteries and flows through intercom- 
municating spaces—which have been variously termed the cells 
of the placenta, the cavernous spaces, the placental sinuses or 
lacune, the inter-villal spaces, or the placental bag or sac— 
prior to leaving the organ by the utero-placental veins, yet 
opinions are divided whether the blood is simply diffused 
through or extravasated into these spaces, without being con- 
fined in definite channels, or whether the spaces are limited by 


soe 


Philosophical Transactions, 1832, p. 57. 
Embryologie Humaine, Paris, 1833. : 
Die Gebdrmutter und das Ei des Menschen, 1832. 
London Medical Gazette, 1834, and Obstetric Medicine and Surgery. 
London Medical Gazette, 1845, 1. p. 758. 
Animal Economy and Collected Works, tv. p. 60. 

7 Quoted in Miiller’s Physiology, Baly’s Translation. 

8 Notes to Hunter’s Works, Palmer’s Ed. ty. p. 69. 

9 Ed. Med. and Surg. Journal, Jan. 1841, and Phys. and Path. researches, 
p. 316. 

10 Anat. and Path. Obs. 1845, and Anatomical Memoirs, 1868, 11. p. 445. 

11 Quoted by Priestley in Lectures on the Gravid Uterus. ; 

12 Wiirzburg Sitzungsberichte, 1853, and Gesammelte Abhand. 1856. 

13 Entwickelungs-geschichte, 1861. 

1 Uterus in Todd’s Cyclopedia, 

15 Mémoire sur les glandes utriculaires de l'uterus, Bologna, 1868, and 
Algiers, 1869. 


an & Ww 


199 PROFESSOR TURNER. 


a membrane continuous with and belonging to the maternal 
vascular system, or derived from the decidua. 

By the Hunters, Owen, Kolliker and Arthur Farre the 
uterine arteries and veins are supposed to open into the intra- 
placental spaces, through which the blood is transmitted so as 
to bathe directly the fostal villi, without the intermediation 
of any maternal structure, and by Virchow it is considered 
that in at least the later stages of placental formation the villi 
grow through not merely the decidua, but the coats of the 
mother’s blood-vessels, so as to bang free and naked in her 
blood. E. H. Weber, again, thought that the uterine vessels 
formed a network of wide canals within the placenta, the ex- 
tremely thin parietes of which invested the foetal vilh. John 
Reid held that the intra-placental part of the maternal vascu- 
lar system was dilated into a large sac, the lming membrane 
of which ensheathed the foetal villi. Like Weber he believed 
that the maternal blood was retained within her own system 
of vessels. John Goodsir stated that not only did the mater- 
nal vessels form sinuses within the placenta—the vascular coat 
of which ensheathed the foetal villi—but a layer of nucleated 
decidual cells was also prolonged over the villi, the latter of 
which were separated from the maternal blood by both these 
structures. Schroeder van der Kolk, again, described the inte- 
rior of the placenta as subdivided by processes of the decidua 
into compartments lined by an epithelium derived from the 
decidua, which was also prolonged over the villi contained in 
these compartments so as to separate them from the maternal 
blood as it flowed through: and Ercolani also maintained that 
the uterine vessels did not subdivide within the placenta, 
but that the blood was enclosed in lacune or sinuses circum- 
scribed by the decidua, the cells of which were prolonged on to 
and ensheathed the villi of the chorion. 

My observations have been made on uninjected mature 
placentz, and on injected specimens. The latter were as follows: 
a. normally separated placentz into the substance of which a 
pipe was inserted, and a transparent gelatine injection coloured 
with carmine was gently passed: b. an attached placenta at 
about the 6th month of gestation, the pipe being inserted into 
a uterine vein in the broad ligament and a transparent coloured 


STRUCTURE OF THE HUMAN PLACENTA. bs) 


injection passed gently into the placenta: ¢. an attached speci- 
men at the full time, the pipe being introduced into a uterine 
artery, and a transparent injection gently passed into the organ : 
d. an attached specimen at about the full time ; a section was 
made through the placenta and uterine wall and a pipe was in- 
serted into an opening on the cut face of the placenta, which 
was believed to be a divided sinus situated in a decidual dis- 
sepiment between two adjacent cotyledons, and the injection 
gently passed along it: in this preparation the fcetal vessels 
were also injected from the umbilical artery. In all the speci- 
mens the injection flowed with great ease into the placenta, 
which swelled up and became tense, as if it passed into a sys- 
tem of pre-existing freely inter-communicating spaces readily 
and naturally admitting of its diffusion, and not as if it were 
extravasated inte spaces artificially produced by the gentle 
pressure to which the injection was subjected’. In specimen 
b. the coloured gelatine was traced along the sinuses situated 
in the muscular wall of the uterus into the so-called circular 
venous sinus and into the utero-placental sinuses in the de- 
cidua serotina. When the free edge of the placenta was raised, 
and the organ carefully separated from the uterus, the injected 
utero-placental sinuses, with distinct though delicate walls, 
were torn across, and the injection which they contained was 
seen to be continuous with that within the intra-placental 
spaces: at the margin of the placenta also a direct continuity 
was traced between the injection in the so-called circular sinus, 
~ and that within the placenta. The arrangement of these 
sinuses closely corresponded to the description written many 
years ago by Prof. Owen, of a specimen he examined. Veins 
of considerable size in the decidua vera, and extending for 
about an inch into the decidua reflexa, were also injected. In 
c. not only was the placenta injected from the uterine artery, 
but the coloured gelatine had penetrated into the venous sinuses, 
in the decidua serotina, and in the muscular wall of the uterus: 
large venous sinuses, lined by a distinct smooth membrane, were 
traced for ;4,ths of an inch, into the placental substance along 
the inter-cotyledonary decidual dissepiments. In d. the injec- 


1 In preparing these injections I have been aided by the skill of my Museum- 
Assistant, Mr A. B. Stirling. 


124 PROFESSOR TURNER. 


tion was limited to those portions of the placenta adjacent to 
the spot where the pipe was inserted, but the neighbouring 
utero-placental sinuses and so-called circular sinus were also dis- 
tended. | 

Portions of the different placentee were hardened in spirit 
and thin slices were then made with Stirling’s section cutter’, 
~ and subjected to microscopic investigation. The transparency 
of the injection enabled me to study its relations to the villi and 
decidua with an accuracy such as it is not possible to attain 
with wax or other opaque injections. The drawing therefore 
(Plate v.) with which this paper is illustrated, is a picture of 
the villi and intra-placental maternal blood spaces as seen 
under the microscope, and not a mere diagrammatic conception 
of what the mode of arrangement might be considered to be, 
such as has generally been the case in the representations of 
the human placenta given by previous anatomists. 

In these sections the villi may be seen to lie in the in- 
terval between the chorion and the decidua serotina. The 
trunks of many villi may be observed to arise from the chorion, 
though in other instances they are cut across transversely or 
obliquely. Deeper in the substance of the placenta also villi 
of large size are met with, divided either longitudinally, ob- 
liquely or transversely, some of which penetrate close to the 
decidual surface, and not unfrequently are attached to it. In 
all the larger villi, the umbilical vessels may be readily seen, 
even with low magnifying powers. Numerous secondary villi 
arise from these primary stems, which branch in the tree-like 
manner so often described. But by far the greater number of 
the smaller villi with their bud-like terminal offshoots have 
been cut off from their parent stems in the act of making the 
section. The villi are not so crowded together as to touch each 
other, but are separated by intermediate spaces filled with 
coloured gelatine. These spaces are circumscribed distally by 
the surface of the chorion from which the villi spring, proxi- 
mally by the decidua serotina on the uterine aspect of the 
placenta, and laterally by the inter-cotyledonary or primary 
decidual dissepiments. The spaces within a given cotyledon 
freely communicate with each other, but the inter-cotyledonary 

1 Described in this Journal, May, 1870. 


STRUCTURE OF THE HUMAN PLACENTA. 125 


dissepiments interfere with a ready communication between the 
injected spaces of adjacent cotyledons. 

The injection was diffused with great regularity through 
each cotyledon, so as to surround not only the stems of the 
villi, but the multitudes of bud-like offshoots which branched 
off from them. Each villus was as it were immersed in a bath 
of coloured gelatine, which occupied, I believe, the place of the 
mother’s blood, and gave one a vivid conception of the mode in 
which that fluid during life laves the vil of the chorion in 
which the foetal capillaries are contained. The regularity of 
distribution of the injection is an important argument against 
its having been produced by extravasation, for if its presence in 
the placenta had been due to that cause, the gelatine would 
undoubtedly have been collected in masses in some localities 
and absent from others, and the villi would have been packed 
together and compressed. From the ease with which these 
intra-placental spaces are injected either through the artery or 
vein, or by passing the gelatine directly into the substance of 
the placenta, and from the uniformity and regularity of the 
pattern produced, few anatomists would be inclined to doubt 
that these spaces pre-exist in the placenta, that they freely 
communicate not only with each other, but with the curling 
arteries and utero-placental sinuses, and that during life they 
are distended with the mother’s blood. 

But this conclusion is supported by another important piece 
of evidence. For when thin sections through the injected pla- 
~ centee—both in those still adherent to the uterus, as in those 
normally separated—were examined with the higher powers of 
the microscope, crowds of red blood-corpuscles were seen to 
occupy the intervillal spaces imbedded in the coloured gela- 
tine. Scattered amidst these red corpuscles a proportion of 
white corpuscles was also readily recognised. Ocular demon- 
stration was, therefore, afforded of the presence of blood in 
these spaces, and as there was no appearance of rupture of the 
foetal vessels, although large numbers, not only of sections, but 
of teazed-out preparations, were examined, I have no reason to 
think that the blood could have been extravasated from them 
into the substance of the placenta. These corpuscles could, 
I believe, have been derived only from the maternal blood- 


126 PROFESSOR TURNER. 


vessels, with which the intervillal spaces were anatomically con- 
tinuous. 

I am unable, therefore, to agree with the opinion expressed 
by Dr Braxton Hicks, in the paper printed in the last number 
of this Journal, that the maternal blood does not exist in the 
intervillal spaces, and to the inference which he draws that 
there is no intra-placental maternal sinus system. For not 
only do I join issue with him on the question of fact, but 
I doubt if the specimens which he adduces in support of his 
views are of a kind to permit satisfactory conclusions on this 
matter to be drawn from them. I have experienced no diffi- 
culty in seeing blood-corpuscles in the intervillal spaces, both 
in attached and separated placente, in such numbers, and so 
generally diffused, that I cannot regard their presence, as he 
would infer, to be due to rupture of the vessels within the foetal 
villi, or to imbibition of the blood at the decidual surface of the 
placenta after detachment of the placenta from the uterus. 

Of the eight specimens which Dr Hicks adduces in support 
of his conclusions, the 5th, 6th, 7th, and 8th are admittedly 
diseased; the 5th beimg an aborted ovum “about 1 inch in 
diameter,” the others being placente “in the state called fatty.” 
The aborted ovum was in so early a stage that it is very 
questionable if the maternal blood-vessels had begun to exhibit 
the changes induced by gestation. He makes no mention even 
of blood-vessels within the fcetal villi, and until these are — 
formed we have no reason to believe that the system of ma- 
ternal sinuses is developed. The fatty placenta were so dis- 
eased that the child in each case had died before delivery, and 
not only did he find no blood in the intervillal spaces, but, as he 
very candidly observes, the villi themselves were either alto- 
gether or almost entirely bloodless. The diseased state of the 
placenta, its bloodless condition, and the death of the feetus, all 
prove that the placenta was no longer fulfilling its function as a 
great organ of circulation and nutrition, and it is not to be 
wondered at, therefore, that no blood was found in the inter- 
villal spaces. But I would submit this is no evidence that 
they were not filled with blood when the organ was healthy 
and the child was alive. For it would be just as logical, or 
rather as illogical, to say that no blood circulated through the 


STRUCTURE OF THE HUMAN PLACENTA. 127 


foetal villi, because they were bloodless in these specimens ; 
surely Dr Hicks is not prepared to support that inference ! and 
yet the one conclusion would be quite as legitimate as the 
other! Further, in the 2nd specimen, about the 3rd month, 
the foetal villi were also bloodless, a circumstance which indi- 
cates that here also a disturbance in the placental circulation 
had taken place. In his 3rd and 4th cases, about the 4th 
month of gestation, he admits that some blood was found in 
the intervillal spaces, and though his theory constrains him to 
suppose that its presence there was accidental, yet he con- 
fesses that he could not satisfactorily ascertain if it occurred 
from. rupture of the villi, or came from the maternal side. His 
Ist case, an attached placenta in the 6th month, is the only 
example adduced of a well-grown healthy specimen in which 
he found no blood in the intervillal space; but as he gives no 
account how he determined its absence in this placenta, this 
solitary example cannot outweigh the mass of evidence on 
the other side of the question which has been advanced by the 
eminent anatomists referred to in the early part of this paper, 
and by the new observations which I have myself made. 

Further, he alludes generally to differences in the amount 
of blood met with in the examination of placentae when natur- 
ally expelled during labour. In regard to this I would merely 
say that when we reflect on the great pressure the detached 
placenta is subjected to by the uterine contractions which cause 
its expulsion; that this pressure is exerted on an organ, the 
uterine face of which possesses the numerous orifices of the 
torn-across utero-placental vessels, through which the blood 
can be squeezed, and that, moreover, the pressure can be as- 
sisted by the contraction of the muscular layer’ of the caducous 
part of the decidua serotina, the wonder is not that no maternal 
blood, or only a small quantity, should be found in the inter- 
villal spaces of a separated placenta, but that any of that fluid 
should be found there at all. 

Lastly, I may state that no satisfactory explanation can be 
given of the passage of the blood from the curling arteries of 


1 In the succeeding section I shall detail the reasons in support of the state- 
ment that a layer possessing the anatomical characters of smooth muscular 
fibre exists in the decidua serotina,. 


128 PROFESSOR TURNER. 


the uterus through the utero-placental sinuses into the sinuses 
within the muscular wall, unless an intra-placental circulation 
be accepted: for the continuity of the one set of vessels with 
the other in the fully formed human placenta, either directly 
or through the intermediation of a capillary plexus, although 
the latter has been assumed, has not only not been proved, but 
is opposed to recorded observations. 

I shall now proceed to consider the theory supported by 
Weber, Reid and Goodsir, that the maternal blood-vessels are 
prolonged into the placenta, and that their lining membrane is 
reflected on to the chorionic villi, so as not only to ensheath 
and separate them from the mother’s blood, but to form a 
limiting membrane for the spaces or passages through which 
the blood circulates. 

I have been unable to satisfy myself of the accuracy of the 
special modification of this general theory, which has been 
advocated by E. H. Weber (antea, p. 122). There is indeed no 
difficulty in seeing the curling arteries pierce the decidua, or in 
tracing sinuses continuous with the utero-placental sinuses into 
the primary dissepiments of decidua which le between the 
cotyledons, and in so far the mother’s blood undoubtedly hes 
in definite canals, limited by the maternal vascular membrane. 
But I have as yet seen no evidence that the uterine vessels 
form a definite network within the cotyledons. 

The view supported by Dr John Reid, that “the inner coat 
of the vascular system of the mother, or at least a membrane 
continuous with it,” is prolonged over each individual tuft of 
villi, was obviously based, not on what he saw within the 
placenta itself, but on observations made as to the relations of 
certain villi situated at the uterine aspect of the placenta to 
the utero-placental veins, or the “sinuses placed next the inner 
surface of the uterus.” For after pointing out how these villi 
protrude into the open mouths of certain of the sinuses only, 
and that “though they were so far loose and could be floated 
about, yet they were bound down firmly at various points by 
reflections of the inner coat of the venous system of the mother 
upon their outer surface;” he then goes on to say, “the outer 
surface of the placental vessels (villi?) has a smooth appear- 
ance, and they are, we may suppose (the italics are his own), 


STRUCTURE OF THE HUMAN PLACENTA. 129 


everywhere enveloped in the inner coat of the vascular system 
of the mother, which, as we have seen above, is reflected upon 
them.” 

Dr Reid’s observations, though detailed with great precision, 
and confirmed in many particulars by Professors Goodsir, Shar- 
pey, Allen Thomson, Alison and Simpson’, have been of late 
years so ignored by many writers, that I thought it advisable to 
re-investigate this part of the subject. In an attached pla- 
centa in the 9th month of gestation, where the fcetal capillaries 
had been injected from the umbilical artery with gelatine and 
carmine, I observed, on carefully drawing the placenta away 
from the inner face of the uterus, the dilated, cavernous, ana- 
stomosing arrangement of the utero-placental veins, with their 
valve-like semilunar edges, which has been so well described 
both by Owen and Goodsir. In various parts of the uterine sur- 
face of the placenta, but as it seemed to me especially near the 
outer edges of the cotyledons, where the primary decidual dis- 
sepiments dipped into the substance of the organ, tufts of villi 
could be distinctly seen projecting into the adjacent utero- 
placental sinuses. Their capillaries filled with a red injection, 
permitted me readily to distinguish them both from the coats of 
the maternal blood-vessels and from the decidua. These tufts 
occurred in the form of patches, oval, circular, or irregular in 
outline, and varied in diameter from about an inch to some 
fractional part thereof. When examined either with a pocket 
lens, or with higher powers of the microscope, the larger patches 
exhibited a somewhat cribriform aspect, owing to interlacing 
bars of membrane passing across them in an irregular manner. 
This membrane was continuous with and obviously formed a 
part of the coat of the utero-placental sinus, and was at times so 
thick that the injected villi were with difficulty seen through 
it, but at others so thin, smooth and delicate, as to represent 
only the inner coat of the sinus. It was in part merely in 
apposition with the villi, and could be readily raised from them, 
though in part it was attached by delicate processes, and 
seemed to bear out the expression used by Reid, of being 
reflected on to them. But in other portions of the patches no 

1 See Note at conclusion of Reid’s Memoir, as reprinted in his Physiological 
and Pathological Researches. 


VOL. VII. 9 


130 PROFESSOR TURNER. , 


membrane could be seen, and the villi projected between the 
membranous bars free and naked into the canal of the sinus. 

But the demonstration of this arrangement was by no 
means limited to the uterine surface of the placenta. For 
when the so-called circular sinus, lying along the outer edge of 
the marginal cotyledons, was carefully slit up, confirmatory 
views were obtained. This it is important to note, because 
whilst attached placente are difficult to get, and but seldom 
examined, the placente separated in the ordinary course of 
labour have the marginal utero-placental sinus not unfrequently 
so little injured that a ready demonstration of the relations of 
the villi to the maternal vessels can at any time almost be 
procured. I have sometimes seen the aperture in the wall 
of the marginal sinus so small that but a single tuft of branch- 
ing villi projected through it, and the wall of the vei formed a 
distinet circular collar around the neck of the tuft. At other 
times I have seen a patch of villi upwards of an inch in length 
appear at the surface of the sinus lying next the cotyledon ; 
the wall of the sinus presenting at the same spot the cribriform 
or reticulated aspect already described. Between these two 
extremes various intermediate forms have been recognised, and 
in this locality as on the uterine face of the placenta itself, 
though the thin wall of the sinus was sometimes reflected on to 
the villi, at others the tufts were freely moveable and projected 
into the canal of the sinus. 

It is evident therefore from these observations that the 
decidua serotina does not form that complete and continuous 
membranous septum between the uterus and placenta which 
has been described by Robert Lee and many other obstetri- 
cians, but that its continuity is in various localities interrupted, 
so as to allow of the penetration of many of the placental vill 
into the utero-placental vessels. And I may further state, in 
confirmation of the above, that I have invariably found in 
examining placente separated during the ordinary course of 
labour, patches of villi, projecting through the decidua, espe- 
cially near the margins of the cotyledons. These patches were 
surrounded by a sharply defined border of decidua, which from 
its precise form was apparently a normal arrangement, and had 
not been produced by a tear or violent detachment of the 


STRUCTURE OF THE HUMAN PLACENTA. iil 


decidua from the surface of the exposed villi during the separa- 
tion of the placenta. These exposed tufts obviously corre- 
sponded to those which I have described as projecting into the 
utero-placental sinuses on the uterine surface of the attached 
placenta. 

My observations therefore are in part confirmatory of those 
of Reid; but though I have seen the wall of a sinus reflected 
on to some of the villous tufts, which project into its canal, yet 
I have not found that these villi are by any means all so en- 
sheathed; still less am I prepared to say with him, that the 
tufts throughout the entire substance of the placenta are every- 
where enveloped by the inner coat of the vascular system of 
the mother. My observations also are in part confirmatory of 
those of Virchow, as I have found numerous villous tufts unen- 
sheathed by any such membrane, but projecting free and naked 
into the canal of the sinus. The consideration, however, of the 
precise relations of the lining membrane of the maternal vascu- 
lar system to the tufts within the placenta—if, in short, each 
tuft possesses an investing membrane, such as Goodsir named 
the external membrane of the villus—must be postponed until 
the minute structure of the villi is described in a subsequent 
section. 

It is evident, in all those localities where the placental tufts 
project into the utero-placental system of sinuses, that the 
maternal blood is brought into such close relation with the feetal 
system of capillaries, that important nutritive and respiratory 
interchanges can be effected between the two kinds of blood. 
The intimate relation, therefore, which can be so easily demon- 
strated in these localities, naturally leads one to infer that similar 
relations subsist throughout the entire substance of the pla- 
centa, and gives additional weight to the arguments advanced 
in an earlier part of this essay, in favour of the intra-placental 
circulation of the maternal blood. 

Whilst reflecting on these relations, it occurred to me, that 
by a very simple experiment the direct continuity of the inter- 
villal spaces with the canal of the sinuses, supposing such to exist, 
might be easily shown, and a demonstration afforded of the 
mode in which the mother’s blood was returned from the in- 
terior of the placenta into her venous system. I accordingly 


9—2 


132 PROFESSOR TURNER. 


in a separated placenta, where the marginal sinus was uninjured, 
introduced an injecting pipe into one of the cotyledons situated 
near the edge of the organ, and very gently propelled coloured 
gelatine into the intervillal spaces. The cotyledon swelled up 
and the injection flowed both into the marginal sinus at the 
edge of the placenta, and into the sinuses situated in the 
decidual dissepiments which divided the injected cotyledon 
from those adjacent to it. But as it might be objected that 
some natural barrier between these sinuses and the interior of 
the placenta had been broken down by the force employed in 
injecting, I modified the experiment in another specimen. In 
this I opened the marginal sinus before the injection was begun, 
and, as my assistant slowly and gently depressed the piston of 
the syringe, I carefully watched one of the cribriform surfaces 
on the inner wall of the sinus, where a Jarge patch of villi was 
exposed, and saw the coloured fluid ooze quietly from between 
the villi out of the intervillal spaces of the cotyledon imto the 
canal of the sinus, and observed that no structure was torn 
down before this oozing began. No doubt then remained in 
my mind, that a natural communication existed between the 
intervillal spaces in the interior of the placenta and the utero- 
placental sinus-system of veins. 

These sinuses are so related to the placenta, as to communicate 
with the interior of the cotyledons either at the outer edge of 
the placenta, where the so-called circular sinus lies, or where 
the sinuses le within the inter-cotyledonary decidual disse- 
piments, or where they come into contact with the uterine 
face of the placenta close to the plane of entrance of the 
primary decidual dissepiments into its substance. The com- 
munication is not as if the smuses terminated abruptly by open 
mouths, as has usually been described ; but rather by possessing 
cribriform apertures in their walls as they he in contact with 
the placenta. From the relation of the sinuses to the margins of 
the cotyledons, whilst the curling arteries penetrate their uterine 
surface near their centre, the stream of maternal blood passes 
through each cotyledon from its centre to its circumference, and 
is effectually brought into contact with the whole of the feetal 
vill. 

The observations detailed in this section seem to me to put 


STRUCTURE OF THE HUMAN PLACENTA. 133 


the question of the existence of an intra-placental circulation of 
maternal blood into the category of established and demon- 
strated facts, and to deprive it of that inferential or even theo- 
retical character which it has hitherto held, and owing to which 
its accuracy has from time to time been assailed by various 


writers’. 


(To be continued.) 


EXPLANATION OF PLATE V. 


Vertical section through the entire thickness of a marginal coty- 
ledon of the human placenta and the adjacent part of the uterine wall. 
Feebly magnified. 

A. Marginal cotyledon. 

B. Portion of an adjacent cotyledon. 

C. Muscular wall of the uterus with its contained sinuses. 

ch, Chorion containing the umbilical vessels. ds, decidua sero- 
tina containing the utero-placental sinuses. pd, primary or inter- 
cotyledonary decidual dissepiments. sd, secondary, or intra-cotyle- 
donary decidual dissepiments. sv, stems of the villi arising from the 
chorion: multitudes of the smaller villi may be seen occupying the 
space between the chorion and decidua. At a, the primary decidual 
dissepiment is represented as reaching the chorion, and at 6, the 
secondary dissepiments are connected with the secondary villi. ms, 
transverse section through the marginal sinus with villi projecting 
into it through the cribriform openings in its wall. wp, utero-pla- 
cental sinus with villi projecting into its canal. up’, sinus, anatomi- 
cally continuous with the utero-placental system of sinuses, and 
situated in a primary decidual dissepiment. 


1 When kindly revising the proof-sheets of these pages, Dr J. Matthews 
Duncan directed my attention to an experiment recorded by Dr Dalton (Treatise 
on Human Physiology, 1867, p. 615), who inflated the intervillal spaces in the 
attached placenta by blowing air into one of the vessels situated in the uterine 
wall. Dr Duncan told me that he had himself not unfrequently seen in the 
newly detached placenta, bubbles of air freely move in the intervillal spaces 
from the decidual to the chorionic surface. 


ACTION OF DIGITALIS ON THE BLOOD-VESSELS. 
By T. LaupEeR Brunton, M.D., D. Sc, AND ADOLPH 
BERNHARD MeEyer, M.D. 


INDEPENDENTLY of each other, and in different ways, we both 
arrived at the conclusion that digitalin causes contraction 
of the small blood-vessels*, Wishing to support our views 
by still more conclusive proofs, we took advantage of the 
opportunities afforded to us in the physiological laboratory 
of the Berlin University to perform together, in February, 1868, 
some experiments on the subject. We are perfectly aware of 
their incompleteness, but circumstances having prevented us 
from continuing them, and the departure of one of us for a 
distant land rendering it improbable that we shall be able to 
resume them together, we now publish their results. 

We believed that by a comparison of the form of the curves 
indicating the blood-pressure before and after the injection of 
digitalin into the circulation, we should be able to determine 
exactly whether it caused contraction of the arterioles or not. 
The kymographion we employed was that of Ludwig, as modi- 
fied by Traube, and the experiments were conducted on dogs 
in the following manner. The animal being narcotized by 
hydrochlorate of morphia, a canula was inserted into the 
crural artery, and a curve (Fig. 1) showing the normal blood- 
pressure was described. Digitalin, suspended in a small quantity 
of distilled water, was then injected into the carotid artery, 
and pressure-curves again described. Injection into the artery 
was employed because Blake® found that digitalin produced a 
much greater effect on the blood-pressure when introduced into 
the circulation in this way than if injected into a vein, A 
comparison of the tracings thus obtained, after the injection, 
with that of the normal pressure and pulse (Fig. 1), showed a 
slowing of the pulse, accompanied by an increase in the mean 


1 T. Lauder Brunton On Digitalis: with some Observations on the Urine, 
London, 1868, p. 52, and A. Bernhard Meyer, Zur Lehre von den Herzgiften in 
Untersuchungen aus dem physiologischen 19 ‘aboratoriwm der Ziivicher Hochschule. 
herausgegeben von Professor Fick. Wien, 1869, p. 71. 

2 Ed. Med. Journ. 1839. 


ACTION OF DIGITALIS ON THE BLOOD-VESSELS, 135 


blood-pressure, while the height of the wave occasioned by 
each cardiac pulsation remained much the same (Fig. 2). 
The pressure continued gradually to rise although the pulse 
not only became slower and slower, but the oscillations of 
the mercurial column at each pulsation diminished in ex- 
tent (Fig.3). This rise could be due either to the heart propel- 
ling a greater quantity of blood into the aorta at each pul- 


sation, or to the arteries, contracting so as to hinder it from 
escaping from the arterial into the venous system. The di- 
minished height of the pulse-wave seems suflicient of itself 
to negative the former idea, and to show that the increased 
pressure can only be due to contraction of the arterioles, but 


136 DR BRUNTON AND DR MEYER. 


we think that a still clearer proof is afforded by the form of the 
wave. The time occupied in the ascent of the pressure-wave 
(indicated by the horizontal distance between the lowest and 
highest parts of the ascending limb) is nearly the same in 
Figs. 1 and 3, but the descending limb of the latter sinks very 


FIC.3. 


gradually indeed, while in the former it falls almost as quickly 
as it rises. What then is the explanation of this phenomenon ? 
During the diastole of the heart, the sigmoid valves when 
healthy, as they were in this case, completely close the cardiac 
end of the aorta. The whole arterial system may then be com- 
pared to an elongated elastic vessel, from which fluid is issuing 
by a narrow opening. The greater the pressure of fluid in the 
vessel the more rapidly will it escape by the opening, the more 
quickly will the pressure consequently fall, and the more abrupt 
will be the descent of the pressure-curve. Now the mean 
blood-pressure in the normal tracing is somewhat over 70 milli- 
metres’, and the maximum height of the wave 44, while in that 
taken when the action of the digitalin was greatest, the mean 
pressure is somewhat over 90 mm., and the maximum 104, The 
fall of pressure ought, therefore, to be more abrupt, but instead 
of this it is more gradual. This alteration cannot, we think, 
be explained by any oscillations of the mercurial column inde- 
pendently of the blood-pressure, and can only be due to con- 
traction of the arterioles retarding the flow of blood from the 
arterial into the venous system during the cardiac diastole. 

1 The true heights are of course nearly double these, but for convenient 


comparison with the tracings we have taken the numbers as they stand in the 
figures, 


ACTION OF DIGITALIS ON THE BLOOD-VESSELS. 137 


In a recent paper, Boehm’ considers that the rise in blood- 
pressure produced by digitalis, is chiefly due to the increased 
action of the heart, and that the condition of the arterioles has 
little or nothing to do with it. Heseems, however, to interpret 
tracings of the blood-pressure in the arteries of mammals in the 
same way as those obtained from the excised heart of the frog, 
and apparently forgets that while in the latter the form of the 
diastolic as well as of the systolic curve depends on the heart 
alone, in the former the heart can have but little or no in- 
fluence on the pressure in the arterial system during the 
diastole, since all communication between them is prevented 
by the closure of the sigmoid valves. The curves which he 
gives confirm our views, for they show the same gradual fall 
in the pulse-wave, after the injection of digitalis, that ours 
do, and being traced with Fick’s spring-kymographion, are free 
from any fallacies due to oscillations of the mercurial column. 
The continued high pressure he observed during prolonged 
stoppage of the heart, and which he attributes to continuous 
cardiac systole, we would ascribe to contraction of the vessels so 
far as it is not due to changes in the respiration. If the arte- 
rioles were not contracted the pressure would fall, as e.g. in 
the experiments of Ludwig and Hafiz”. 

We next attempted to ascertain whether the slowing of the 
pulse is due to a direct specific influence of the drug on the 
roots of the vagus as supposed by one of us*, or to the stimula- 
tion of these roots by the increased pressure of blood in the 
cranium produced by the contraction of the arterioles, as sup- 
posed by the other*. In order to do this we diminished the 
blood-pressure by the inhalation of nitrite of amyl after it had 
become high, and the pulse slow from the injection of digitalin. 
If the slowing of the pulse were due to a specific action of the 
digitalin on the vagus roots, it ought to continue although the 
pressure falls, but if due to stimulation of these roots by the 
high blood-pressure, it should disappear whenever the pressure 
is reduced. Our experiments showed that whenever the pressure 
fell after the inhalation of the nitrite of amyl the pulse became 


1 Phiiger’s Archiv, vy. 190. 
2 Ludwig’s Arbeiten, 1870. 
3 Bruuton, Op. cit. 

4 Meyer, Op. cit. 


138 DR BRUNTON AND DR MEYER. ACTION OF DIGITALIS, &c. 


quick. It might thus appear that the slowing is due in part at 
least to the high pressure, and not altogether to a direct in- 
fluence of the digitalin on the vagus; but this must be decided 
by farther experiment. 

Lastly, we tried to discover whether digitalis causes con- 
traction of the vessels by acting directly on their walls or 
on the vasomotor centre. This we sought to do by observing 
whether the injection of digitalin into the circulation caused 
any alteration in the calibre of the vessels of the rabbit's 
ear after the sympathetic nerve of the same side as well as 
both vagi had been divided in the neck. The vagi were 
divided in order to prevent the digitalin from slowing the heart, 
and thus disturbing the circulation, and the sympathetic to 
prevent any influence being transmitted to the vessels of the 
ear from the vasomotor centre. The results of these experi- 
ments were not constant, and we are unable to draw any 
definite conclusions from them; but the fact that the vessels 
of the ears were occasionally seen to empty themselves more 
quickly after the injection of digitalin than before, seems to 
us to indicate an action upon the walls of the vessels them- 
selves. 

The conclusions to which we have arrived are shortly, 
Ist, that digitalin causes contraction of the arterioles. This 
is proved by the small height of the pulse-wave, and by its 
descent becoming more gradual after the injection notwith- 
standing the increased blood-pressure. 2nd, that the slowing 
of the pulse is probably due in part to the increased blood- 
pressure which results from the contraction of the arterioles. 
We gladly take this opportunity of expressing our obligations 
to Professor Rosenthal for the assistance and advice which he 
so constantly and kindly afforded us, and to Herr Merck of 
Darmstadt, to whose kindness we owe the digitalin we em- 
ployed, 


ON THE KOMBE ARROW-POISON (STROPHANTHUS 
HISPIDUS, D. C.) OF AFRICA. By Dr THomAs 
R. FRASER. 


[THE author communicated the results of some experiments 
with this poison to the Royal Society of Edinburgh, on the 
21st of February, 1870; and an abstract of this communica- 
tion has been published in the Proceedings of the Society 
(Vol. vir. No. 81, 1869—70, p. 99). As the circulation of 
these proceedings is, however, in great part limited to the 
Fellows of the Society, the author has thought it proper to 
reprint the abstract of his communication in a Journal where 
it will have the advantage of a wider and more general cir- 
culation. 

In the following paper the abstract referred to will be re- 
produced verbatim, but a number of interpolations, consisting 
chiefly of details of experiments, will be introduced, in order to 
supply various omissions, many of which were rendered neces- 
sary by the original form of publication. 

These interpolations will be included within brackets, so 
that they may be distinguished from the original abstract. 
The author has made no experiments with this substance since 
that abstract was published ; ‘he, however, entertains the hope 
of continuing the investigation. ] 

In nearly every narrative of exploration in uncivilised tro- 
pical regions, accounts are given, often no doubt somewhat 
fanciful, of poisonous substances which are said to possess the 
most remarkable properties. Usually these poisons are of 
vegetable origin; and the great majority may be included in 
the two divisions of ordeal and of arrow poisons, according as 
they are applied to one or other of these purposes. Among the 
most remarkable of the ordeal-poisons are the Tanghinia veni- 
fera of Madagascar, the Physostigma venenosum of Old Calabar, 
and the Akazga poison of the Gaboon; and of the arrow-por- 
sons, the famous Curara or Wourali of South America, and the 
Antiaris toxicaria of Java. 


140 DR FRASER. 


The examination of these substances has not only proved of 
great value to physiology, but practical medicine has likewise 
been berefited—one of them, at least, being now an important 
medicinal agent. 

In bringing before the Society a few of the results of a 
recent examination of a new arrow-poison, the author has to 
express his gratitude to the President, who very kindly gave 
him the specimens with which the experiments have been 
made. These specimens, consisting of a number of ripe folli- 
cles, were sent to Dr Christison by Mr Walker, and were col- 
lected in the expedition of the late Bishop Mackenzie. 

Several specimens of the poison have likewise been sent to 
Professor Sharpey by Dr Kirk, H.M. consul at Zanzibar. Dr 
Kirk states “that the plant is a woody climber, growing in the 
forest, both of the valley and hills, and found at various places 
between the coast and the centre of the continent, above the 
Victoria Falls of the Zambesi. The stem is several inches in 
diameter, and rough outside. The plant climbs up the highest 
trees, and hangs from one to the other like a bush-vine. The 
flowers are of a pale yellow, and last for but a short time 
during the months preceding the first rains of the season 
(October and November). The fruit is ripe in June, and 
collected by the natives, who separate the rough outer coat 
before drying it, preserving the more leathery inner covering 
and the seeds’.” 

Dr Livingstone gives some interesting information regard- 
ing the poison in his Narrative of an Expedition to the Zambest 
and its Tributaries. He mentions that arrows poisoned with it 
are used for killing wild animals only; arrows destined for the 
more noble object of killing men being poisoned with the 
entrails of a small caterpillar. Dr Livingstone says that in 
hunting, the natives follow the game with great perseverance 
and cunning: “The arrow, making no noise, the herd is fol- 
lowed until the poison takes effect, and the wounded animal 
falls out; it is then patiently watched till it drops; a por- 
tion of meat round the wound is cut away, and all the rest 


eaten ”’ (p. 465). 
} 


1 Extract from letter to Professor Sharpey, dated January 1, 1864. 


THE KOMBE ARROW-POISON OF AFRICA. 141 


Dr Livingstone also says that the poisoned arrows are made 
in two pieces. “An iron barb is firmly fastened to one end of 
a small wand of wood, ten inches or a foot long, the other end 
of which, fined down to a long point, is nicely fitted, though 
not otherwise secured, in the hollow of the reed which forms 
the arrow-shaft. The wood immediately below the iron head 
is smeared with the poison. When the arrow is shot into an 
animal, the reed either falls to the ground at once, or is very 
soon brushed off by the bushes; but the iron barb and _poi- 
soned upper part of the wood remain in the wound. If made 
in one piece, the arrow would often be torn out, head and all, 
by the long shaft catching in the underwood, and striking 
against trees” (p. 466)*. 

[It would appear that this arrow-poison is widely distri- 
buted over Africa, for it has been found not only at Kombé, 
on the west coast near the equator, and in the Manganja 
country, near the Zambesi at the south-east of Africa, but 
also in the Gaboon district*, in Guinea’, and in Senegambia‘, 
In the Gaboon district it seems to be called Inée, Onaye, or 
Onage’.] 

The follicles examined by the author vary in length from 
about nine and three-fourths to about twelve and one-fourth 
inches, and in greatest thickness from about one inch to three- 
fourths of an inch, and they vary in weight from about 130 to 
330 grains. They contain from 100 to 200 seeds, each of 
which weighs about half-a-grain, and has attached to it a 
beautiful comose appendix, placed on an extremely brittle stalk. 
For the identification of the plant the author is indebted to 
Professor Oliver of Kew, who writes, in a letter dated 10th Dee. 
1869: “LI reopen your note to say that I have just dissected 
a flower, and conclude to name the Kombé plant Strophanthus 
hispidus, D.C” This plant belongs to the natural order Apo- 
cynacece. 


1 Specimens of these arrows, which had been presented to Professor Mac- 
lagan by Dr Kirk, were exhibited to the Society. 

2 Pélikan, Archives Générales de Médecine, Juillet, 1865, p. 115. 

eee Hasselt, Archives Néerlandaises des Sciences, T. vit. 2me. Liv. 1872, 
p. 161. 

4 Baillon, quoted by Polaillon and Carville, Archives de Physiologie, No. 5, 
1872, p. 526. 

> Baillon, loc. cit. 

6 [Since this letter was received, Professor Oliver has been led, by a further 


142 DR FRASER. 


When the seeds contained in these follicles are bruised and 
treated in a percolator with rectified spirit, a greenish-yellow 
tincture is obtained. By distilling off the greater part of the 
spirit, and drying the residue on a water-bath and in the 
exhausted receiver of an air-pump, an extract is procured which 
weighs about 25 per cent. of the seeds employed, has an in- 
tensely bitter taste, and contains about one-half of its weight 
of an inert fixed oil", From this extract the author has suc- 
ceeded in separating a very powerful active principle. [He 
proposes that this active principle should be named stro- 
phanthin. | 

As, however, the greater number of the experiments have 
been made with the extract, the results of these experiments 
only will be described in the following brief account of the 
physiological action of the Kombé arrow-poison, it being un- 
derstood that the action of the active principle is of the same 
character. 

When a small dose (one-twentieth of a grain) of this extract 
is mixed with a few minims of water, and injected under the 
skin of a frog, no distinct symptom is seen until about half-an- 
hour, when the animal’s movements become somewhat sluggish. 
Soon afterwards the respirations cease, some stiffness occurs in 
the thoracic extremities, reflex sensibility diminishes, some stiff- 
ness appears in the pelvic extremities, and in about two hours 
after the administration voluntary movements entirely cease, 
and strong galvanic irritation produces no effect, even when 
applied to exposed muscles and nerves. An examination of the 
heart shows that it is completely paralysed, the ventricle being 
pale and contracted, while the auricles are dark and distended. 

[To illustrate more fully the general symptoms that appear 
in frogs, the following experiment may be described :— 


22nd January, 1870, 1.28 p.m. One-tenth of a grain of extract of 
strophanthus, suspended in 3 min. of distilled water, was injected 
under the skin at the left flank of a frog, weighing 287 grains. <A 


examination of the botanical characters of the Kombé-poison plant, to doubt its 
identity with S. hispidus; and, accordingly, he has described it in the Icones 
Plantarum, No, 4, 1870, under the name of S. Kombé.] 

1 [Microscopic examination shows that this extract contains a large number 
of acicular crystals; and when the fatty matters are removed from it by ether, 
a hygroscopic substance is obtained, which consists in great part of crystals.] 


Si Tia” Se Fi a. ge Le Pn ee ae a a se ans ee 


THE KOMBE ARROW-POISON OF AFRICA. 143 


small quantity was lost, owing to the adhesion of the fatty matter to 
the syringe:—1.36. The frog is less active in moving and jumping 
about; but the limbs retain a normal position :—1.43. When irri- 
tated, it jumps feebly :—1.47. The thoracic extremities are weak, and 
the thorax often rests on table:—1.56. The frog is on the abdomen 
and thorax; when irritated, pretty energetic movements occur of the 
four extremities, but the frog does not jump, although it obviously 
intends to do so. The respiratory movements of the throat are 
shallow, and those of the flanks are very feeble and occur only 
rarely :—2.8. A good deal of frothy mucus has been exuded; there 
is no exaggeration of the reflex excitability :—2.12. Almost inces- 
santly, fibrillary twitches are occurring in the muscles of the abdomen. 
Both throat and chest respirations have ceased :—2.25. When placed 
on the back the frog attempts to turn, but cannot do so. No cardiac 
impulse can be detected. The fibrillary twitches continue, and are 
best seen at the lumbar region and behind the eyeballs. When the 
skin is irritated, pretty energetic movements occur in the two pelvic 
extremities and in the abdominal walls, and only feeble movements 
in the thoracic extremities:—2.45. Trritation of the skin causes 
feeble reflex movements in the two pelvic extremities; but none in 
the thoracic, which are now stiffly extended at right angles to the 
body. The fibrillary twitches no longer occur spontaneously ; when, 
however, the skin at the coccyx or upper part of the thigh is irritated, 
a series of fibrillary twitches follow at the lumbar and gluteal regions: 
—2.55, Galvanic stimulation of the muzzle excites feeble reflex move- 
ments in the pelvic extremities, but none elsewhere :—3.10. Weak 
galvanic stimulation excites no movement; a very strong galvanic 
current, however, still excites extremely feeble movements in the 
pelvic extremities. The four extremities are now stiff and extended :— 
3.12. The heart was exposed, and it was found to be motionless, with 
the ventricle contracted and white, and the auricles distended and 
dark, Even the most powerful galvanic current from a Daniel’s cell 
and Du Bois Reymond’s induction apparatus, applied directly to its 


~ surface, produced no movement whatever :—3.30. Galvanic stimula- 


tion of the exposed right sciatic nerve caused only some very feeble 
and sluggish movements of the right foot. Strong galvanic stimula- 
tion of the exposed muscles of the right thigh failed to produce any 
contraction :—3,.50. Galvanic stimulation of the right sciatic nerve 
no longer produces any effect. When the muscles of various parts of 
the body are stimulated, no movement occurs. A certain degree of 
general stiffness is present, and the four limbs continue extended :— 
23rd January, 12.50 P.M. Strong general rigor. The ventricle of the 
heart is contracted and white; the auricles are distended with 
blood. | 


It was obviously suggested by these phenomena that this 
substance acts as a cardiac poison; and, accordingly, some ex- 
periments were made in which the heart was exposed before 
the administration, of which the following is an example :-— 


144 DR FRASER. 


One-tenth of a grain of extract was injected under the skin of 
a frog. Five minutes thereafter, it was observed that the ventricular 
systole was somewhat prolonged; in six minutes, the ventricular 
diastole was imperfect, so that only portions of the ventricle dilated 
to admit blood from the auricles; in six minutes and thirty seconds, 
the greater portion of the ventricle was continuously pale and con- 
tracted, each auricular systole propelling merely a small drop of blood 
into the ventricle, where it produced a dark, pouch-like projection, 
which at times disappeared, and at other times only changed its 
position during the imperfect systole of the ventricle; in seven 
minutes, the ventricle altogether ceased to contract, while the move- 
ments of the auricles continued at nearly the normal rate; and in 
eighteen minutes, the auricles in their turn became motionless, but, 
in place of being contracted and empty like the ventricle, they were 
distended and full of dark blood. Notwithstanding this absolute 
paralysis of the heart, respiratory movements occurred for twenty-five 
minutes after the ventricle had ceased to contract, and the frog 
jumped about actively for some time after this. 


[In many other experiments, the heart was exposed before 
stropbanthus was administered, but only the two following will 
be here described. In the first, the poison was administered 
by the stomach, and in the second, by the rectum. 


15th January, 1870. A frog, weighing 310 grains, was secured 
in the usual way. 11.54a.m. The heart was exposed by removing 
the sternum:—11.56. Heart’s contractions, 13 per 30 sec.:—11.59. 
Heart’s contractions, 13 per 30 sec.:-—12. A narrow gum elastic tube 
was passed into the stomach :—12.3. 0.2 grain of extract of strophan- 
thus, suspended in 8 min. of distilled water, was injected down 
the tube into the stomach. Almost immediately violent efforts to 
vomit occurred, and a portion of the fluid was ejected :—12.6. Heart's 
contraction, 15 per 30 sec.:—12.7. ‘“ Pouching” observed at the 
heart’s apex:—12.9. Heart’s contractions, 14 per 30 sec.:—12.10, 
Heart’s contractions, 14 per 30 sec.:—12.13. Heart’s contractions, 
15 per 30 sec.:—12.18. Heart’s contractions, 15 per 30 sec.:—12.25. 
Heart’s contractions, 15 per 30 sec.:—12.32. Imperfect ventricular 
diastole; a portion only of the ventricle dilating during diastole, 
and an appearance of ‘“pouching” being thereby produced :—12.33. 
No diastole of ventricle, but a movement of that cavity occurs at 
times, although systole is continuously maintained :—12.35, A num- 
ber of ventricular contractions, with diastolic dilatations in proper 
rhythm during 20 seconds, and, then, only movements of the 
ventricle without diastole:—12.36. No movements of ventricle, 
systolic contraction being continuously maintained, The auricles 
contract regularly 13 times per 30 sec.:—12.50. Auricular contrac- 
tions, 6 per 30 sec. No movement of ventricle :—12.50, Auricular 
contractions have ceased: the auricles are dark and distended, and 
slight movements may be occasionally excited in them by direct irri- 


THE KOMBE ARROW-POISON OF AFRICA. 145 


tation; the ventricle is pale and contracted, and no movement occurs 
when the surface is irritated:—12.58, The frog was unfastened, 
and it jumped about with considerable activity :—1.30. The frog is 
jumping and moving about, but less actively, and a little stiffness is 
present in the thoracic extremities. The heart is perfectly motion- 
less :—1.50. Ditto:—-2.45. The frog no longer moves about, and it 
is lying on the abdomen and chest with the lower jaw resting on the 
table. Reflex movements can be readily excited, but they are slug- 
gish. The limbs are somewhat stiff:—16th January. 3 p.m. General 
rigor. No contractions can be excited by stimulating the nerves or 
muscles. 

20th January, 1870. 2.25 p.m. The heart of a frog, weighing 
350 grains, was exposed in the usual way, the frog being secured on 
the back:—2.28. Heart’s contractions, 12 per 30 sec., regular and 
rhythmical :—2.30. Heart’s contractions, 12 per 30 sec.:—2.31. 
Heart’s contractions, 12 per 30 sec.:—2.34.30. Injected 0.1 grain of 
extract of strophanthus, suspended in 2 min. of distilled water, into 
the rectum, by means of a gum elastic tube attached to the orifice 
of the hypodermic syringe; and, afterwards, injected 1 min. of dis- 
tilled water. A little of the fluid esecaped:—2.40. Heart’s contrac- 
tions, 12 per 30 sec.:—2.45. Heart’s contractions, 12 per 30 sec.:— 
2.48. Heart’s contractions, 12 per 30 sec.; imperfect, the diastole being 
only partial at the apex:—3.10. Heart’s contractions, 12 per 30 sec. 
The respirations continue :—3.17. Heart’s contractions irregular ; there 
being 5 ventricular contractions per 30 sec., and 3 auricular contrac- 
tions to each ventricular contraction. The ventricle remains contracted 
during two auricular beats:—3.20. Ventricular contractions, 7 per 30 
sec.; and 2auricular contractions occur for each ventricular contraction: 
—3.23, Ditto. The respirations continue, and the frog often struggles: 
—3.24. Ditto. The ventricular diastole is imperfect, the upper half 
of the ventricle remaining contracted continuously, while diastole 
occurs at the apex only :—3.26. 6 imperfect ventricular and 12 per- 
fect auricular contractions per 30 sec. The upper two-thirds of the 
ventricle now remains continuously contracted (in systole), and only 
the lowest third dilates during diastole. The diastole of the ventricle 
consists, therefore, of a mere “pouching” of the lowest third; and 
it is very brief in duration, being almost immediately followed by 
contraction (systole) of that third, which continues during two auri- 
cular contractions :—3.29. 6 imperfect ventricular contractions per 
30 sec.; the appearance of a small dark “pouch” at the apex, and 
the occasional and rare appearance of a small “pouch” at the left 
base constituting its diastole. Auvicular contractions, 12 per 39 sec., 
regular. Respirations continue, and, occasionally, there are general 
struggling movements:—3.32. 4 imperfect ventricular contractions 
per 30 sec., limited to the formation of a minute dark “pouch” at 
the apex. Auricular contractions, 12 per 30 sec., regular:—3.34. 
Only at long intervals a feeble movement oceurs in the ventricle; 
which is now altogether permanently contracted and pale. Auricular 
contractions, 10 per 30 sec.:—3.35. No movement of ventricle. Au- 
ricular contractions, 6 per 30 sec.; at times, there is a long pause :— 


VOL. VII. 10 


146 DR FRASER. 


3.37. On two occasions since last note, one imperfect ventricular 
movement occurred, following the formation of a minute dark 
“pouch” at the apex. Auricular contractions, 9 per 30 sec.:—3.39. 
Auricular contractions, 8 per 30 sec. One imperfect ventricular 
contraction during every three auricular contractions; a pouch form- 
ing at the apex ‘at the secord of the oe auricular —— and 
being cae immediately after the ar con- 
tractions, 7 per 30 see. No movement of Fonts wie is con- 
tracted and pale:—3.45. Ditto. Occasional gasping respirations :— 
3.50. Auricular contractions, 6 per 30 sec., feeble and irregular :— 

3.91.30, Auricular contractions have ceased ; the auricles are dilated 
and dark, and the ventricle is contracted and white. Occasionally 
gasping respirations occur:—3.54, Ditto. Irritation of the surface 
of the ventricle causes a slight, almost doubtful, movement; while 
irritation of the surface of the auricles causes a series of contractions 
limited to the auricles and continuing for one minute, when they 
ceased after some struggles on the part ‘of the frog:—3.57. Irritation 
of the surface of the ventricle produces no effect; and of the surface 
of the auricles, a single contraction followed by complete rest:— 
4.7. Galvanic irritation, even when powerful, applied to various 
portions of the ventricle and auricles produces no movement of the 
heart, but it excites violent struggles:—4.9. The frog was set free. It 
assumed a nearly normal posture, excepting that the thoracie extre- 
mities were extended and somewhat stiff; and, although able to move 
about pretty actively, it cannot jump:—4.30. Irritation of the skin - 
caused pretty active reflex movements of the four limbs, but after- 
wards, one pelvic extremity remained extended in a rather stiff con- 
dition :—21st Janwury, 11 a.m. General vigor: museles hard and pale; 
veins full of blood, arteries empty; ventricle pale with some red 
patches, auricles dark and dilated. 


In the last two experiments the heart was affected much 
more gradually than when strophanthus is subcutaneously in- 
jected; and the changes that take place in its action were, 
therefore, exhibited in a very distinct manner. | 

The experiments that have been performed with birds and 
mammals have likewise shown that this poison acts primarily 
on the heart. 

[By way of illustration, a few particulars may be given of 
an experiment on a pigeon, and of one on a rabbit. 


9th February, 1870. 4.5 p.m. One-tenth of a grain of extract of 
strophanthus was suspended in 4 min, of distilled water, and injected 
under the skin at the right side of a healthy pigeon, weighing ten 
ounces :—4.12. The pigeon vomited a large number of entire wheat 
grains :—4.13, Ditto:—4.15. Ditto. Some mucous substance is also 
vomited, There is decided feebleness of the limbs :—4.20. Frequently 


THE KOMBE ARROW-POISON OF AFRICA. 147 


sales since last note, and some liquid excrement is now passed :— 
4.22. The pigeon is lying on the abdomen, the wings being used to 
ede the body. The pupils are dilated: :—4,23. Some spasms oc- 
curred, which have an opisthotonie character :— 4.23.30. Respirations 
have ceased. The pigeon is dead:—4,24.30. The heart was exposed 
and found to be motionless :—4.28. Direct galvanic stimulation of a 
sciatic nerve caused active muscular contractions :—4.30. Galvanic 
stimulation, even when powerful, applied to the surface of the heart 
produced no movement. Galvanic stimulation, applied to the muscles 
of the limbs and body, produced active contractions. The pupils are 
still dilated :—5.5, The muscles of the body no longer contract under 
galvanic stimulation ; and they have become somewhat hard. 

5th February, 1870. In an active rabbit, weighing 4 ]bs, 2 oz., it 
was found that at 4.45 p.m. the respirations occurred 42 times, and 
the ae impulse 49 times, per 10 sec.; while the pupils measured 
£5 x 35 of an inch:—4.50. Two-tenths of a grain of extract of stro- 
phanthus, suspended in 1@ min. of distilled water, was injected under 
the skin at the left flank :—4.53. Respir. 43 per 10 sec. The animal 
is restless:—4.97. Card. imp. 41 per 10 sec. Right pupil, $$ x 3 of 
an inch ;—4.58. Respir. 47 per 10 sec.:—4.58.30. Card. imp. 35 per 
10 sec.:—4.59. Respir. 39 per 10 sec.:—4.59.30. Card. imp. 39 per 
10 sec.:—5.1. Card. imp. 39 per 10 sec. Grinding movements of the 
tecth :—5.2. Respir. laboured, and a sharp sound (like a “smack”) 
occurs with them:—5.3. Card. imp. 35 per 10 sec.; the impulse is 
ee y reduced in strength:—5.4, Respir. 9 per 10 sec. Right pupil, 
$3 x23 of an inch 5.5, Card. imp. 35 per 10 sec. The sounds 
continue to occur with ae ee movements:—5.6. Respir. 10 
per 10 sec. Right pupil, 4§ x $$ of an inch:—5.7. Card. imp. cannot 
be counted accurately, jaa it ies to occur about 38 times per 
10 sec. :—5.8. Respir. 13 per 10 sec., no longer accompanied with 
any sound:—5.10. Respir. 23 per 10 see.:—5.11. Card. imp. 41 per 
10 sec., again distinct:—5.14. Card. imp. 38 per 10 see. The rabbit 
is ina somewhat crouching posture:—5.14, Respir. 23 per 10 sec., 
jerking in character :-—5.16. Card. imp. 39 per 10 see.:—5.18. Respir. 
26 per 10 sec.:—5.18.30. Card. imp. 38 per 10 sec. The lips and 
mouth are frequently opened and shut. Usually the eyelids are 
nearly 5.22. Respir. 24 per 10 sec., jerking. The rabbit is 
unsteady, and tends to fall over :—5.23.30. Card. imp. 40 per 10 see.: 
—95.25. The rabbit is now sitting, and the head often falls slowly 
and is raised sharply, with nodding movements :—5.26. Respir. 13 per 
IO’ sec., HbLe and jerking:—95.26.30. Card. imp. 33 per 10 sec. 
Right pupil, £5 x33 of an inch:—5.30. Respir. 8 per 10 see. Fre- 
quent shaking movewents of the head:—5.52. Card. imp. cannot be 
counted, because of the jerking respiratory movements, and of fre- 
quent tremors; but it seems to occur about 24 times per 10 see.:— 
6.34. Respir. 14 per 10 see. Incessant tremors, chiefly of the head: 
—95.38. The rabbit is lying flaccidly, with the lower jaw resting on 
the table. Tremors oceur frequently ; and occasionally somewhat 
spasmodic movements take place, during which the body is tossed 
about :—5.41. Respir. 7 per 10 see. The rabbit lies quietly on the 


10—2 


148 DR FRASER. 


side :—5.42. Card. sounds (as heard by the stethoscope) are very feeble 
and irregular. Right pupil, }§ x ¢5 of an inch :—9.42.30. Ocecasion- 
ally a gasping respiration occurs. Right pupil, 35 x $5 of an inch:— 
5.43, No respiratory movements; conjunctiva and cornea are insen- 
sible. The rabbit is dead:—5.44. No card. imp. can be felt, nor 
sound heard with the stethoscope :—5.46. The right sciatic nerve was 
exposed: weak galvanic stimulation produced no effect when applied 
to it, or to the surfaces of the exposed muscles in the gluteal region ; 
but very strong stimulation when applied to the nerve excited a faint 
twitch of the foot, and when applied to the exposed muscles, a slow 
and feeble contraction. These conditions were present also at 5.50: 


—5.48. Right pupil, #5 x so of an inch:—5.52. The heart was ex- 
posed, and found to be motionless; while stimulation, even when 
powerful, produced no effect upon it:—5.55. Galvanic stimulation of 
nerves and muscles no longer produces any effect. The intestinal 
peristalsis is observed to be very slight:—6.30. General rigor, not 
very strong, except in the pelvic extremities. Right pupil, 59 x so of 
an inch, 


Experiments were also made with dogs and cats, and re- 
sults similar to the above were obtained.] 

An endeavour was made to ascertain by what mode of 
action these very peculiar cardiac effects are produced. With 
this object experiments were made, in which the cerebro-spinal 
axis was completely destroyed, in which the yagi nerves were 
divided, and in which the peripheral terminations of the vagi 
were paralysed by atropia, previously to the exhibition of the 
Kombé poison; but no important modifications were thereby 
caused, and it is therefore obvious that the action on the heart 
is not exerted through the cerebro-spinal nerves. In other 
experiments, after complete cardiac paralysis, the surface of the 
heart was irritated by galvanic and other stimulants, but no 
effect was thereby caused. 

[It is, therefore, perfectly obvious that strophanthus exerts 
an action upon the heart, which is independent of any change 
that it may produce in the physiological condition of the 
cerebro-spinal nervous system and of its connections with this 
organ. It is also obvious that this substance acts in a powerful 
and direct manner upon the cardiac muscular fibre; greatly 
prolonging, in the first place, the contractions of those fibres, 
and, ultimately, rendering it continuous, and only to be over- 
come when relaxation occurs as a natural consequence of post- 
mortem decomposition. Whether the intra-cardiac nerves are 


EE 


THE KOMBE ARROW-POISON OF AFRICA, 149 


likewise affected, and whether a modification of their physiolo- 
‘gical activity is concerned in the production of these remark- 
able effects, it is extremely difficult to determine, seeing that 
their condition cannot readily be discovered in presence of so 
powerful and direct an action upon the muscular fibres that 
are controlled by them. The experiments that have as yet been 
made tend to show that the final stoppage of the heart and the 
production of the virtually permanent state of ventricular 
systole are independent of any modification that may be ef- 
fected in the physiological condition of the intra-cardiac nerves. 
An example of these experiments may be here given. 


19th February, 1870. 3.5 p.m. In a frog, weighing 361 grains, 
the heart was exposed:—3.9, Heart’s contractions, 10 per 30 sec.:— 
3.14. Heart’s contractions, 12 per 30 sec.:—3.16. Heart’s contrac 
tions, 12 per 30 sec.:—3.17. One-twentieth of a grain of extract of 
strophanthus, suspended in a very little distilled water, was injected 
under the skin at each thigh (0.1 gr. in all):—3,20. Heart’s con- 
tractions, 13 per 30 sec.:—3.21. Heart’s contractions, 13 per 30 sec.: 
—3.23. Heart’s contractions, 12 per 30 sec.:—3,.24. Heart’s contrac- 
tions, 13 per 30 sec.; the ventricular systole is rather prolonged :— 
3.26. The ventricle no longer dilates; it is in a state of continuous 
systole :—3.28. Auricular contractions, 8 per 30 sec.; irregular :— 
3.29. Auricular contractions, 6 per 30 sec.; regular, but the diastole 
is only momentary:—3.32. Auricular contractions have ceased. 
Respir. 7 per 30 sec.:—3.34. The heart was removed from the 
thorax, by cutting the vessels at the base with a pair of scissors, 
and it was placed on a porcelain slab moistened with serum :—3.36. 
Galvanic stimulation, applied to various parts of the removed heart, 
produced no movement. The lower two-thirds of the ventricle was 
-cut off with a pair of sharp scissors, and stimulated with galvanismn, 
but without any effect—this separated portion remaining hard and 
contracted. 


The data that have been acquired are not, however, suffi- 
cient to permit the assertion that no action is exerted upon 
the intra-cardiace nerves. It is by no means improbable that 
these nerves are directly acted upon; and that a modification 
in their physiological condition aids in producing the irregu- 
larities in the rhythm and the intermissions in the contractions 
of the heart, which occur during the earlier stages of the 
poisoning, and before the functional activity of the cardiac 
muscle has been destroyed. | 

Another very prominent action of this poison is that exerted 


150 DR FRASER. 


on the voluntary muscles, by which their activity is gradually 
impaired, and finally completely destroyed, so that the muscles 


are quickly in a condition of true rigor 
[The nature of this action has to a 


mortis. 
certain extent been de- 


scribed in the preceding experiments; it will, therefore, be 
sufficient to give a brief account of some of the experiments 


in which its mode of production was 


investigated. 


Tt is shown by the following experiment that the effects 
on the striped muscles of the body do not result from the 
paralysis of the heart which strophanthus produces :— 


12th January, 1870. 


A: 

1.15. A frog, weighing 266 grains, was 
placed on its back, and seeured. The 
heart was then exposed :—1.20. Heart’s 
contractions, 12 per 30 see. 


1.21. 


1.51. One-tenth of a grain of extract 
of strophanthus was injected under the 
skin of the two thighs:—1.39. Ventricu- 
Jar contractions have ceased, and the ven- 
tricle is pale and contracted. Auricular 
contractions, 11 per 30 sec. and feeble :— 
1.45. Auricular contractions have ceased: 
—1.46. The frog was set free: it jumped 
about acti:ely:—1.49. The thoracic ex- 
tremities are a little stiff, though still 
mobile. The frog can no longer jump, but 
it pushes itself about by vigorous move- 
iments of the pelvic extremities. 


50; 


. A ligature was 


B. 


The heart of a frog weigh- 
ing 268 grains was ex- 
posed in the same way as 
in 4. 


Heart’s contractions, 12 
per 30 see. 


drawn 
tightly round the base of 
the heart, so as completely 
to stop the circulation ; 
pulsation continued, how- 
ever, in the large vessels 
close to the heart. 


The frog is set free, and 
jumps about actively. 


EEE OEE ey nee 


THE KOMBE ARROW-POISON OF AFRICA. 


2. The thoracic extremities are very 
stiff Irritation produces pretty active re- 
flex movements of the pelvic extremities. 
Twitches occur at the lower part of the 
abdominal wall. Respir. has ceased. 


2.10. 


2.25. Irritation of the skin produces 
no movement, unless it be powerful, when 
slow but strong reflex movements occur in 
the pelvic extremities, but only at a con- 
siderable interval after the application of 
the irritant. 

DENN .- Bae a eae = 


2.34. The pelvic extremities are now 
somewhat stiff, and when the skin is irri- 
tated very sluggish and feeble reflex move- 
ments occur in them. The twitches of 
the muscles have ceased. 


2.37. Strong galvanic stimulation ap- 
plied to any part of the skin produces no 
effect, except when it is applied to one of 
the feet, when feeble movements occur 
slowing, which are strictly localised to the 
parts through which the galvanic current 
is passed. Galvanic stimulation applied 
to the exposed left sciatic nerve produces 

- feeble movements below the left knee, but 
no reflex contractions :—2.52. Strong gal- 
vanic stimulation applied to the exposed 
sciatic nerve produces no contraction ; 
when applied to exposed muscles in the 
left thigh and calf no movement whatever 
occurs. 


4,15, 


The frog continues to jump 
about actively. 


The frog jumps only feebly 
when uritated. Active 
spontaneous movements, 
however, still occur. 


The frog jumps sponta- 
neously. 


The frog cannot now jump, 
but it pushes itself about 
by vigorously moving the 
four extremities. 

The frog has for some time 
been lying on the abdo- 
men, thorax, and lower 
jaw. Irritation causes well- 
marked reflex movements. 


152 DR FRASER. 


13th January, 1PM. ... Ae ... Irritation no longer ex- 
cites reflex movements, 
Galvanic stimulation of an 
exposed sciatic nerve causes 
ouly feeble movements of 
the limb supplied by the 
nerve; but it causes active 
contractions in the mus- 
cles, when directly applied 
to their surface. The body 
and limbs are perfectly 
flaccid. 
1.5. General rigor is present every- 

where; and the muscles are hard and pale. 

The ventricle is pale and rigid:—16th 

January, 12.10e.mM. Ditto:—17th January, 

lpm. Ditto:—18th January, 11 a.m. The 

muscles are now softer, The ventricle is 

less rigid, and at several places dark 

patches are seen. 


That the action of strophanthus upon the striped muscles of 
the body is not the result of any influence conveyed to them 
by the spinal motor nerves is rendered apparent by the fol- 
lowing experiment :— 


28th January, 1870. 3.58 p.m. One-twentieth of a grain of sul- 
phate of methyl-strychnium was dissolved in 4 min. of distilled water, 
and injected under the skin at the left flank of a frog, weighing 
290 grains:—4.25, Galvanic stimulation of the skin does not pro- 
duce any reflex movement:—4.35, Galvanic stimulation of the ex- 
posed left sciatic nerve does not produce any muscular contraction 
in the left pelvic extremity or elsewhere:—4.40. A ligature was 
passed below the left sciatic nerve, and tied tightly round the middle 
of the left thigh; the nerve, therefore, not being included within the 
ligature:—4.44, The frog was secured on the back, and the heart 
was exposed :—4.45, Cardiac contractions, 9 per 30 sec., regular and 
rhythmical: —4.47, One-tenth of a grain of extract of strophanthus, 
suspended in 3 min. of distilled water, was injected under the skin 
at the right flank:—5.44. Heart’s contractions are irregular and 
non-rhythmical: the greater part of the ventricle being continuously 
contracted and white, and only a small dark pouch being formed 
during diastole, which is emptied after each alternate auricular con- 
traction. witches occur in the muscles of the abdomen and upper 
part of right (non-ligatured) thigh:—5.52. The ventricle is now al- 
together motionless, and it is white and contracted, Occasionally 
some feeble movements occur in the auricles. The twitches of the 
muscles have ceased; and the thoracic extremities are stitf:—29th 
January, 3e.M. The ventricle is contracted, its anterior surface is 


—_— eS 


THE KOMBE ARROW-POISON OF AFRICA. 153 


pale, but its posterior is dark; and the auricles are dark and not 
distinctly contracted. Galvanic stimulation, applied to the heart, 
produces no effect. The thoracic extremities are stiff and hard; the 
right (non-ligatural) pelvic extremity is stiff, and the muscles at the 
thigh are hard, while those below the knee are less so. Galvanic 
stimulation of the muscles of the right pelvic extremity, or elsewhere 
in the poisoned region, produces no effect; but galvanic stimulation 
of the muscles (through the skin) of the left (ligatured) pelvic extre- 
mity produces active contractions in that extren.ity :—30th January, 
4pm. Ditto:—3lst January, 4 P.M. Contractions can no longer be 
produced in the left (ligatured) pelvic extremity by galvanic stimu- 
lation. 


That the action of strophanthus upon the striped muscles 
of the body is the result of direct contact with these muscles is 
rendered apparent by the following experiment :— 


24th January, 1870. 1.55~.m. In a frog, weighing 350 grains, 
a ligature was tied tightly round the structures of the right thigh 
excepting the trunk of the right sciatic nerve:—2.3. The frog was 
secured on the back, and the heart was exposed:—2.4. Heari’s con- 
tractions, 13 per 10 sec., regular and rhythmical:—2.9, One-tenth of 
a grain of extract of strophanthus, suspended in 2 min. of distilled 
water, was injected under the skin at the left flank:—2.19. Ventricu- 
lar contractions, 13 per 30 sec., and imperfect; the middle zone of the 
ventricle remaining always in systole, and only the base and apex 
dilating in diastole:—2.21. Imperfect diastole of ventricle, 7 per 
30 sec.; different limited portions become at different times pouched 
to constitute the diastole, and at times a drop of blood is distinctly 
seen to be conveyed from one part to another of the ventricle :— 
2.24. Veutricular contractions have altogether ceased, and the ven- 
tricle is contracted and white. Auricular contractions, 11 per 30 
sec. The respirations continue:—2.34. Auricular contractious have 
ceased. Respiratory movements continue:—2.36. The frog was set 
free; and it moved about pretty actively :—2.50. Spontaneous jumps 
occur, but they are rather stiff. Both pelvic extremities are moved 
actively; but the left (non-ligatured) more so than the right (liga- 
tured), the ligature round the right thigh having caused that limb to 
become a little extended:—3. Voluntary movements are feeble. 
Twitches occur in the left thigh. Respiration is represented by infre- 
quent gasping movements :—3.14. Nearly constant slight twitches in 
the left (non-ligatured) thigh. The tone of the left pelvic extremity 
is impaired, as it now usually remains partially extended :—3.25, 
Slight irritation, applied anywhere, causes a series of movements in 
the body and four extremities, but these movements are now more 
active in the right (ligatured) pelvie extremity than in the left:— 
4.44. Galvanic stimulation, applied to the muzzle, causes reflex move- 
ments in the right (ligatured) pelvic extremity, but nowhere else :— 
5, Ditto. Gulvanie stimulation applied to the sciatic nerve of the 


154 DR FRASER. 


left (non-ligatured) pelvic extremity causes movements of that ex- 
tremity:—5.30. Galvanic stimulation of the sciatic nerve of the left 
(non-ligatured) pelvic extremity no longer causes any movement: 
but when applied to the sciatic nerve of the right (ligatured) pelvic 
extremity, above the ligature, it causes active movements of the 
right pelvic extremity ; and when passed through the spinal cord it 
is followed by energetic movements of this extremity, and of no other 
part of the body. Galvanic stimulation, even when strong, of the 
muscles of the left pelvic extremity causes no contraction. It is 
found that a cut section of one of these muscles has an acid reaction. 
The muscles of the thoracic extremities are also non-contractile and 
acid in reaction:—6. Ditto. While the thoracic extremities are very 
rigid, and the left (non-ligatured) pelvic extremity slightly so, the 
right (ligatured) pelvic extremity is flaccid and its muscles are con- 
tractile and alkaline. } 


Regarding the other physiological effects, it is sufficient 
briefly to mention that the sensory and motor spinal nerves, 
the abdominal and cervical sympathetics, and the muscular 
walls of the stomach, intestines, bladder, and uterus, are 
paralysed at an early stage, although not until the blood-heart 
had ceased to contract; while the lymph-hearts of the frog 
retain a normal rate, long after paralysis of the blood-heart’. 

From these results it is apparent, that the primary action of 
the Kombé arrow-poison is isolated in the heart, and that it may 
therefore be included in the class of the cardiac poisons,—a 
class of poisons whose action has been most accurately defined 
by the researches of Kolliker, Vulpian, Pélikan, Hammond and 
Weir Mitchell, Hilton Fagge and Stevenson, Holme, Dib- 
kowsky, and others. 

(The author believes that this preliminary investigation en- 
ables him to make the following statements :—1l. Strophanthus 
acts primarily upon the heart, and produces, as the final result 
of this action, paralysis of that organ with permanence of the 
ventricular systole. 2, Pulmonary respiration continues in 
cold-blooded animals for several minutes after the heart is 
paralysed. 3. The striped muscles of the body are acted upon: 
twitches occur in them; their tonicity is exaggerated; and, 
finally, their functional activity is destroyed, the muscles being 

1 The author is indebted to Professor Sharpey of London for an account of 
some unpublished experiments made with this poison in 1862. The results 


mentioned in the above abstract harmonise in the most satisfactory manner 
with those obtained by Professor Sharpey. 


SE Ee eee Eee ee 


THE KOMBE ARROW-POISON OF AFRICA, 155 


then hard, and, soon afterwards, acid in reaction. These 
changes are accomplished subsequently to the final effect on 
the heart. They are the result of a direct contact of the sub- 
stance with the muscles themselves, and are independent of the 
action on the heart, as well as of any changes that occur in the 
physiological condition of the cerebro-spinal nervous system. 
4. The reflex function of the spinal-cord is suspended soon after 
the heart is paralysed;. but the motor conductivity of the 
spinal-cord and of the nerve-trunks continues after the striped 
muscles of the body are paralysed. 5. The lymph-hearts of the 
frog continue to contract for many minutes after the blood- 
heart has been paralysed. } 


PECULIAR MALFORMATION OF THE LEG AND FOOT. 
By W. W. Waesrarre, B.A., F.R.C.S., Joint-Lecturer on 
Anatomy at St Thomas's Hospital. (Pl. VIL.) 


THE following notes of a curious malformation are the result of an 
examination made by Mr Stewart and myself of the leg and foot of a 
man aged about 45, who had died of pyemia. The specimen is now 
in the museum of St Thomas’s Hospital, and Mr Stewart has kindly 
made the drawing of it which accompanies this paper. The existence 
of the malformation had not been known during the man’s stay in 
the hospital, so that the amount of movement or usefulness could 
not be accurately determined. 

Left leg extremely short; the Tibia measuring only nine inches, 
and the foot only six and a half. The foot had only three toes, 
and was very narrow, and was fitted to the leg in such a manner that 
its dorsum lay against the outer side of the leg; and, in conse- 
quence of the Tibia being considerably twisted outwards, the sole 
looked forwards as well as outwards. 

The fibula was absent, except in its lower end; and this portion 
was displaced upwards. The muscular irregularities were numerous, 
and due in great measure to the peculiar arrangement of the bones, 
although there weie some peculiarities which seem to have been the 
result of excesses of development. 

Boners.—/emur.— Lower extremity small, and the margins of the 
two condyles more parallel than usual. The anterior surface flat and 
expanded below; so that the articular surface was broader than normal 
in front, while it was narrower than usual behind. The inter-condy- 
loid notch not more than half its ordinary breadth, and peculiar in 
being encroached upon by articular cartilage continuous principally 
with that on the outer condyle. No groove for the popliteus on the 
outer condyle. Ligaments normal with the exception of the crucial, of 
which only a trace of the posterior could be found, while the anterior 
was deficient. 

Tibia.—Much bowed, with the convexity forwards end inwards, 
and twisted outwards below, so that the fibula lay directly behind it 
at the ankle-joint. The relation of the upper end of the tibia to the 
knee-joint was peculiar; for the cavity of the joint dipped down 
deeply on the outer side, so that more than half an inch of the outer 
surface of the head of the tibia entered into the formation of the joint, 
being overlaid at this part by a much-thickened external lateral 
ligament: the ligament was so dense at this point, in fact, as to 
represent the head of the fibula which was wanting. 

Fibula.—Entirely deficient in its upper seven-eighths. The lower 
end was present for about an inch and a half, representing the 
external malleolus, but placed an inch above the lower edge of the 
Tibia, 


ee ee ee 


PECULIAR MALFORMATION OF THE LEG AND FOOT. 157 


Foot.—The tarsal bones fused into one flat expanded irregular 

mass. The metatarsals were only three; and the phalangeals ar- 
ranged as belonging to three toes, two of which were fused together 
at their bases. The metatarsus was articulated to the end of the 
tarsus, and projected both outwards and upwards, so as to form a 
concavity on the dorsum and at the side, making an angle of about 
135° in each direction. The whole tarsus was solid, but evidences 
still remained here and there of original separation between the bones. 
Posteriorly the os calcis and astragalus were fused together, and the 
two lay nearly upon the same level, the astragalus being nearly alto- 
gether upon ihe inner side of the os calcis, The bone on the outer 
Side of the tarsus continued forward, and evidence of a cuboid 
could be traced in a well-marked ridge on the plantar surface ; 
the extremity of the bone articulated with the outermost and 
part of the next metatarsal bone. In the middle of the tarsal mass 
was a large hole which represented the normal interosseous groove ; 
its existence as a foramen was explained by the lateral displacement 
of the astragalus; it was continuous anteriorly with a cleft between 
the representatives of the cuboid and external cuneiform. On the 
inner side the distinction between original bones was not so readily 
traced as on the outer side, for the parts were more completely fused 
together, and the prominences did not correspond so visibly with 
those in the normal foot; moreover, it appeared as if considerable 
modification of shape had possibly occurred as the result of the 
malposition of the tibia and fibula, or as a concomitant of it. There 
was, however, some indication of the existence of all the bones of this 
region, but the middle cuneiform appeared most deficient, while that 
portion which represented the external was much shortened. The 
upper surface of the astragalus-region of the bone was hollowed out 
into a cup which received the lower end of the Tibia, and in such a 
manuer that the dorsum of the foot was thrown almost against the 
outer surface of the leg. There was a well-marked tuberosity on the 
portion which represented the heel. 

The metatarsus consisted of three bones which from their charac 
ter and from their relation to the tarsus seemed to be representatives 
of the first, third and fifth. 

The phalanges which were continuous with the innermost of 
these three metatarsal bones were evidently those of a great toe. 
Those continuous with the outer two were peculiar. The proximal 
phalanges were united at their bases so that the outer two metatarsal 
bones articulated with a solid bifid phalanx, upon each horn of which 
was a second phalanx, which was again surmounted by an ungual 
phalanx. 

InTEROSSEOUS Memprane.—This was well marked and its outer 
margin took the place of the absent portion of fibula. The fibres ran 
obliquely downwards and outwards, except along the outer edge, where 
they passed downwards from the head of the tibia to the remaining 
piece of fibula as a connecting cord, not, however, forming a well- 
marked structure. The membrane was perforated above by a fora- 
men through which the anterior tibial vessels and nerve passed. 


158 MR WAGSTAFFE. 


Front oF THE LEG.—Tibialis anticus: origin and insertion as 
usual; an accessory slip, sent off from it in the lower part of the leg, 
became connected with the fibrous tissues above and behind the 
proper insertion into the internal cuneiform.—Fatensor proprius 
pollicis arose from the representative of the head of the fibula and 
the upper three fourths of the interosseous membrane. In_posi- 
tion and appearance it resembled the normal extensor longus digit- 
orum. It was inserted as usual into the great toe.—Hzxtensor longus 
digitorum arose below the ext. prop. poll. from the lower fourth of 
the interosseous membrane, and occupied therefore nearly the position 
ofa normal peroneus tertius: it was, however, mserted into the dorsum 
of the representative of the outer toes in the ordinary manner.—<Acces- 
sory muscle: a large unusual muscle was deeply placed between the 7%. 
anticus and the extensors. It arose with the former, but distinct from 
it, on the inner side of the anterior tibial vessels and nerves, and was 
inserted deeply on the dorsum of the foot all round the central hole 
between the representatives of the os calcis and astragalus.— Peroneus 
longus and brevis as usual.—Peroneus tertius abseut.—Adnterior tibial 
vessels and nerve only differed from the normal in their relations, 
by having to their inner side the large, accessory muscle just men- 
tioned, 

Back oF THE Lec.—Gastrocnemius, besides arising from the femur 
as usual, was slightly connected with the Tibia on the inner side, and 
was inserted into the os calcis without receiving the tendon of the 
soleus. Its inner head was perforated by the posterior tibial vessels 
and nerves.—Solews arose from the interosseous membrane occupy- 
ing the position of the upper fourth of the fibula, but was connected 
with the tibia only by means of a muscular slip, which ran from the 
oblique line. It was muscular throughout, and was inserted into the 
inner side of the os calcis distinct from the gastrocnemius.—Plantaris 
absent.—Popliteus absent.—Tibialis posticus as usual.—llexor longus 
digitorum arose from fascia over the Tibialis posticus, but was uncon- 
nected with bone at its origin. It passed obliquely across the 7ibialis 
posticus to its outer side, where it joined with the Flex. long. pollicis. 
—Flexor longus pollicis was small, and arose from the remaining 
portion of the fibula and from the interosseous membrane above it 
to about the middle of the leg. It passed downwards and was joined 
by the tendon of the flew. long. digitorum, and was inserted into the 
great toe at the ungual phalanx.—The posterior tibial vessels and 
nerves, after passing through the inner head of the gastrocnemius, 
reached the deep part «f the leg and passed to the outer side of the 
flexor longus pollicis before arriving at the back of the ankle. 

Sore or THE Foor.—Abductor pollicis was arranged as usual ; but 
muscular slips passed from the flexor brevis pollicis into its tendon otf 
insertion.—- Flexor brevis digitorum abseut.—Abductor minimi digitt 
hada very broad origin from the os calcis, and a smaller one from the 
base of the last metatarsal bone; the insertion was into each side of 
the first phalanx of the outermost toe. The muscle gave off a large 
tendinous slip which passed across the foot to the base of the first 
phalanx of the great toe and was slightly attached to the base of the 


~~ ee 


PECULIAR MALFORMATION OF THE LEG AND FOOT. 159 


bifid phalanx. This slip appeared to represent the absent /eaor brevis 
digitorum. —Accessorius arose as usual.— Flexor brevis pollicis was very 
large; some muscular slips, however, passed from it to the insertion of 
the abdnetor. Adductor pollicis as usual.—Transversus pedis absent. 
Flexor brevis minini digiti arose as usual, but was inserted into the 
base of the bifid phalanx.—J/nterossei absent.—Lumbricales absent.— 
Plantar vessels and nerves did not present any further deviation from 
their usual arrangement than that the internal plantar nerve supplied 
the contiguous sides of the great toe and next. 

Dorsum or Foot.—fxtensor brevis digitorum as usual, but inserted 
into the two outer toes, 


Case II. A second similar case has come under my notice since 
the above notes were written, and may with advantage be recorded 
here, forasmuch as the dissection of the previous case helps to clear 
up some of the obscurities in this. 

The patient (Eliza Ann B.) is 14 years old. She is rather slow 
and dull; but the rest of the family appear not to be very intelligent. 
None of her relations are known to have had any kind of deformity : 
and I cannot discover that her mother traces the peculiarity to fright 
during pregnancy. 

She is well developed in other respects, but she walks very lame 
owing to the deformity. The right limb is from 3 to 4 inches shorter 
than the left, owing to defective development and bowing of the tibia, 
together with a peculiar displacement and deficiency of the foot, 
closely resembling that described in the first case; and this shortening 
has naturally produced a compensating tilt of the pelvis and lateral 
curvature ot the spine. 

The measurements are as follows: 

Length of right tibia along its axis (a, 6, c) 11 inches. 


* vertically (ae) LOS, 
- fet tibia along its axis 122 ,, 
Length of foot on outside, right = 8} inches. Left 94 inches. 
oy) inner ” ” gt ” ” 103 ” 


The two thigh-bones were of equal length. The tibia could be 
felt throughout its whole length, and was bowed outwards in its 
lower half, and defective on its outer side, so that the lower articular 
surface was placed at about an angle of 60° to the horizontal. The 
fibula could not be felt by manipulation, but there appeared to be a 
small mass of bone immediately under the outer condyle of the tibia 
which represented the head of the fibula. No evidence of shaft or 
lower end could be detected. If a similar arrangemeit to that in 
Case I. existed, the lower piece of fibula re presenting the base could 
not have been easily recognized. In consequence of the bowing of 
the tibia the leg was a good deal expanded and flattened above the 
ankle. 

The foot was small, and presented only 3 toes, 3 metatarsal bones, 
and a fused mass of tarsal bones. It was placed almost horizon- 
tally, and appeared to be rather displaced backwards and outwards 
from the base of the tibia, so that the inner malleolus projected con- 


160 MR WAGSTAFFE. MALFORMATION OF THE LEG AND FOOT. 


stantly below the tarsus, while the representative of the heel lay 
outside and above the lower end of the tibia. In walking, the girl 
rested upon the surface of the inner malleolus, and the inner side of 
the first or great toe. There was not, however, the great displace- 
ment outwards and upwards which appeared to have existed in the 
first case. 

The tarsus consisted evidently of a fused mass of bones, the lines 
of distinction between which could not be made out, but the heel 
projected in a very similar manner to that found in the other case. 

The metatarsus was composed of 3 distinct bones, the inuer of 
which resembled in shape an ordinary so-called first metatarsal bone, 
but was smaller than the corresponding one on the left foot. 

The phalanges were 3 to each of the 3 toes, and each toe was sepa- 
rate from its neighbour, so that they could act independently. The 
second toe was less developed than the others, but the first was evi- 
dently from its shape a fairly developed great toe. 


Case III. Mr Le Gros Clark, has informed me of a third case, 
which has been under his notice for some years. In this there is no 
evidence of the existence of a fibula, and, just as in the two foregoing 
cases, there is associated with this peculiarity the same deficiency in 
the number of toes; only three toes exist, one of which is evidently 
a great toe. 

Mechanical appliances have been resorted to to counteract the ten- 
dency of the tibia to bow more and more outwards, and the treatment 
has been so far successful, that the boy, who is now about 9, was able 
to skate last winter. 

These three cases are curiously similar. In each, a deficiency or 
absence of the fibula is accompanied by the absence of two toes: but 
I do not feel there is enough to allow one to determine which toes 
are absent, though in the dissected case (I.) the second and fourth 
appeared to be so. 

The abse: ce of the fibula is associated with a bowing outwards 
of the tibia; and this seems to point to one important function of 
the fibula—that of acting as a bony chord to the slight are which the 
tibia forms. With the absence of the outer malleolus, too, the foot 
is everted, and to such an extent, in Case I. as to allow it to be 
almost flattened against the outer side of the leg. Cases IT. and III. 
are important, also in their surgical bearing, for both have been 
treated successfully by mechanical appliances. Division of the tendo- 
achillis was performed in Case ILI. to obviate the tilting of the heel 
which existed. 


* .* During this session I haye found another instance in which muscular and 
tendinous fibres were arranged vertically on the outer surface of the external 
pterygoid, and gave origin by their deep surface to some of the horizontal fibres 
of the muscle. See Vol. y. of this Journal, p. 2381. It existed upon the right 
side; but I am unable to state whether the peculiarity was present on the left 
side. The spheno-maxillary fossa did not appear unusually large. I did not 
notice any other peculiarities in muscular development in the subject.—W. W. W. 


ee eee eee eee 


EFFECT OF STIMULI ON THE SECRETION OF THE 
PAROTID GLAND, By P. Butter Sroney, late Lesident 
Physician in St Bartholomew's Hospital. 


A HEALTHY young woman having been admitted to St Bartholo- 
mew's Hospital for a parotid fistula of long standing, I took ad- 
vantage of the opportunity to make some experiments on the effect 
of stimuli of different kinds on the secretion of the parotid gland. 
The fistula was occasioned by a cut aeross the cheek received in 
childhood. The orifice of the duct was about two inches in front 
of the right ear. It was sensitive, but not particularly tender, 
and a fine probe could be readily passed up for some distance. 
There was 10 communication whatever with the mouth, so that the 
whole of the saliva secreted was discharged by the fistula and ran 
down the cheek. The power of various stimuli to increase se- 
cretion was estimated by the rapidity with which a drop formed 
at the fistular opening before and after their application. The method 
of procedure was as follows :—The duct was emptied by passing a 
soft napkin gently but firmly along its course from the gland to 
the orifice which was then completely dried, and the time which 
elapsed before a drop formed sufficiently large to run down the 
cheek carefully noted. The duct was then emptied and the fistula 
wiped a second time, the stimulus applied, and the time required 
for the formation of a drop again observed. The effect of mastica- 
tion alone was first tested by giving the patient a glass stopper to 
chew. The experiments numbered [. II. and III. were all made at 
one time with a short interval between. The time required for the 
formation of a drop was in— 


Expt. I. II. PEE, Ve 
Natural seeretion 
without a stimulus — 3’ ohh 4’ 
While chewing 115” i 50” 1/207 
After chewing 940” 5’ 6’ or 


The effect of taste alone was tried by placing sugar or tartaric 
acid first on the tip and next on the base of the tongue, 
Expt. I. Dk III. IV. 


A little sugar placed 
on the tip of the 


tongue 3 5’ 13% 
Sugar on the base of 

tongue £15" 5! 12’ 4! 
Tartaric acid on the tip ie 20" 30” L104 
Tartaric acid on the 

base 10” 30” 60” 30” 


VOL. VII. Wh 


162 MR STONEY. SECRETION OF THE PAROTID GLAND. 


The effect of mastication and taste together was observed by 
allowing the patient to chew a piece of meat. 


Expt: 1. UU. This hfe 
No stimulus Ded oA 5’ 
While chewing 15” BU. 30” 15” 


The effect of mental stimuli was first tested by desiring the 
patient to think of something nice. This had no effect on the secre- 
tion whatever, and food was then placed before her. 


Expt, > if. Te, ENE 
Food placed before patient 1/13” 4’ 6 12’ 


These experiments show (1) that mastication alone stimulates 
the flow of saliva from the parotid to a considerable extent. (2) 
That the effects of taste vary with the sapid substance, sugar having 
no effect, while tartarie acid acts most powerfully. (3) That sapid 
substances act equally when applied to the tip and base of the 
tongue. (4) That the effect of mastication and taste together is 
much greater than that of mastication alone. (5) That mental 
stimuli had a considerable effect in one experiment, but in others 
none at all. These results accord in some respects with those ob- 
tained by Schiff in his experiments on dogs, though differing from 
them in others. This physiologist observed that mastication alone 
had little or no stimulating action on the parotid secretion in dogs, 
and Dr Brunton informs me that he has found this to be the ease 
also in rabbits, while in the experiments above described the action 
was very distinct. The effect of the application of sugar and tartaric 
acid to the tongue of dogs was the same as that observed by me. 
The slight effect of purely mental stimuli in this case is remarkable, 
as the parotid is stated by Kiihne to be readily affected by them, 
but this may have been due in great measure to the character of 
the patient, who seemed to be dull and unimaginative. An experi- 
ment was also made for the purpose of determining the time re- 
quired for the absorption of drugs and their excretion by the saliva. 
Kor this purpose iodide of potassium was administered, and the 
saliva constantly tested till it appeared. The time which elapsed 
between its administration by the mouth, and its appearance in the 
saliva from the parotid duct, was found in one experiment to be 
29 minutes 30 seconds. 


VALVES IN THE RENAL VEINS. By Watter Rrvineron, 
M.S. Lond., &e.; Surgeon to the London Hospital, and Lec- 
turer on Anatomy at the London Hospital Medical College. 


Four or five years ago I was asked by Mr Curling to examine the 
Valves at the orifices of the Spermatic ‘Veins, and while doing so met 
with many instances of Valves in the Renal Veins. At the time 
when I found them I was not aware that they had been previously 
described by Dr Edward Crisp in an Essay sent to the College of 
Surgeons in competition for the Triennial Prize for 1861. As these 
Valves are ignored in the ordinary Anatomical text-books, perhaps 
I may add here the results at which I arrived, 


Specimen I. Male. Right Spermatic Vein opened into Vena 
Cava; Double Valve situated in a fossa. Left Spermatic Vein 
opened into renal; double valve. Right Renal Vein. No Valves 
inside, but at the lower part of the opening into Vena Cava was a 
small semilunar fold. Left Renal Vein. No valves; semilunar fold, 
as in the right, at the opening into Vena Cava. 


Specimen IT. Male. Right Spermatic. Double Valve; one 
fold, not very distinct, at opening into Vena Cava, Left Spermatic. 
Two fine valves at the orifice. Right Renal. No valves. Left 
Renal. No valves. 


Specimen IIT. Male. Right Renal. Two veins opening into 
Vena Cava. At the orifice of the upper and larger no valve: at the 
orifice of the lower and smaller a beautiful double valve. Right Sper- 
matic. Opening near the lower renal, a double valve in a fossa. 
Left Renal. No valves. Semilunar fold at the orifice. Left Sper- 
matic, double valve at orifice. 


Specimen ITV. Male. Right Spermatic opened into Vena Cava. 
At the orifice is a valve, like the Eustachian Valve in the Heart, a 
semilunar fold directing the blood upwards. Inside, near the orifice, 
were two folds constituting a double valve. Left Spermatic. A 
single fold at the opening into the Renal Vein on the side nearer to 
the Vena Cava narrowing the orifice into the Renal. Right Renal. 
Fold near the orifice. Left Renal. Fold at the junction of Renal 
with Vena Cava below, apparently, to prevent blood coming up the 
Vena Cava from flowing into the Renal. 


Specimen V. Male. Right Spermatic opened into Renal, but 
being broken off the orifice could not be satisfactorily examined. 
Left Spermatic. Double Valve. Right Renal. Two large valves at 
orifice, one below and one above, the lower being the larger. Left 
Renal. Semilunar fold as already described. 


Specimen VI. Male. Right Spermatic. Opened into Vena 
Cava. A Double Valve, the ieee lip being much larger. Right 
Renal. Fold across the vein a few lines from he orifice. Left Sper- 


11—2 


164 MR RIVINGTON. VALVES IN THE RENAL VEINS. 


matic opened with the lowest tributary of the Renal into left Renal, 
and the two had a double valve. Left Renal. No valves at the 
orifice—but usual triangular fold. A single valve in Kenal vein near 
the Spermatic. 


Specimen I. Female, Ovarian Veins. Right opened into Vena 
Cava—large valve single, and at the lower side close to the orifice. 
Left. Two branches opening into Renal vein without valves. Two 


valves in the Renal vein about the middle of its course, the smaller 


one behind the other, both on the posterior wal}. 


Specimen EI. Female. Ovarian Veins. Right. Opening into 
Renal near junction with Veria Cava—a fold on the surface of the 
lining membrane, between the opening of the Ovarian Vein and the 
Vena Cava. Left. No valve. Fold across the Renal vein nearer 
to the Vena Cava. Renal Veins. Right. No Valve. Left. Small 
Valve. 


Specimen I{I. Female. Ovarian Veins. Right. Opening into 
Vena Cava—large lateral Valve. Left. No Valve. Renal. Right. 
No Valve. Left. Small Valve near orifice. There was a fold in the 
Vena Caya about + in. in length near the left Renal. 


Specimen IV, Female. Right Ovarian, opening into Vena Cava; 
single Valve. Left Ovarian, opening into Renal; double Valve. 
Right Renal. Chief branch, no Valve. Smaller and lower branch, 
double Valve. Left Renal. Semilunar fold at orifice into Vena 
Cava. Suprarenal Vein. Small Valve 3th of ineh from orifice. 
At the junction of some of the Renal tributaries Valves existed. 
Valves also existed at the orifices of the lumbar veins. 


From the examination of these and some other Specimens, I am 
led to believe that a more extended investigation would be likely to 
establish the following points :—1l, The existence of valves at the 
orifices of both the right and left spermatic veins, with a few excep- 
tions. 2. These valves are, as a general rule, double, being formed 
of two crescentic felds of lining membrane, which leave a slit-like 
aperture between them. 3. When no valves exist at the opening of 
the left spermatic into. the left renal vein, valves are generally present 
in the renal vein within a quarter of an inch from the orifice of the 
spermatic. One of the specimens here described exhibits two large 
valves at the orifice of the right renal vein. In this case the right 
spermatic vein opened into the renal, but its termination was muti- 
lated, and could not be satisfactorily examined. 

Tn another specimen, not given above, the left spermatic divided 
into two parts, opening into the renal an inch from each other. The 
part which opened nearer to the Vena Cava was furnished with two 
valves—the other had none; but two large valves existed in the 
renal vein, 2 quarter of an inch from the opening of this second 
branch into the renal. 

The Ovarian veins yield similar results; but the valves would 
appear to be more often single than in the male. I have found both 
singleand double valves in the renal veins of the female, 


LL ee ”CGUCLCLCL Ll eee errr eer err 


EXPERIMENTS AS TO THE CAUSES OF THE PRESENCE 
OF BILE PIGMENT IN THE URINE. By J. WickHau 


Lrec, M.D. 


‘I wisH to place on record the results of a few experiments made 
during the last fortnight. 


_ Naunyn’ states that the injection of the bile-acids into the 
eirculation is not followed by the appearance of bile-pigment, but 
only of hemoglobin, in the urine. But if bile, or hemoglobin, or 
zther be injected into the small intestine, the bile-pigments can be 
discovered the next morning in the urine. I can support Naunyn’s 
first statement by experiments of my own; but on these I do not at 
“present prepose to dwell. The second statement is not, however, 
confirmed by those which are now published. 

In all the experiments, Naunyn’s directions for the operation 
were closely followed; the rabbits were under the influence of 
chloroform during the operation. 

Sept. 24, 1872. A rabbit weighing 3 lbs. 6 cc. of a 12 per cent 
solution of Plattner’s crystallized bile were injected into the small 
intestine about 4 p.m. Next morning, the urine was clear, dark 
coloured like jaundiced urine, but very careful and repeated examina- 
tions gave no trace of Gmelin’s reaction with nitric acid. 


Sept. 25. <A rabbit weighing 3hlbs. 5¢.¢. of hemoglobin, pre- 
pared by alternate freezing and thawing of calve’s blood, were injected 
‘into the small intestine at 5p.m. Next morning, urine was straw- 
coloured, turbid ; shewing no trace of reaction with nitric acid’. 


Sept. 27. A rabbit. lc.c. of ether was injected into large 
intestine about 2p... Next morning, the urine was yellow, turbid, 
and gave no trace of a reaction with nitric acid. 


Oct. 4. The same rabbit as that used on Sept. 24. l5e.c. of 
zether injected into the small intestine at noon. The morning 
following, urine turbid, straw-coloured, and giving no trace of a 
reaction with nitric acid. 


I wish to express my best thanks to my friend and colleague, 
Dr Brunton, for his courtesy in placing at my disposal his pharmaco- 
logical laboratory at St Bartholomew's Hospital, in which all the. 
foregoing experiments were made, 


1 Naunyn, Arch. f. Anat. u. Phys. 1868, p. 401. 

2 This experiment has been repeated, with the same result as in the text, by_ 
F. A. Wolff (Zur Pathologie des Ikterus, Diss. Kénigsberg, 1869. Quoted in 
Henle’s Bericht f. 1869, p. 176). 


REVIEWS AND NOTICES OF BOOKS. 


Diagrams of the Nerves of the Human Body. By W. H. Fiower, 
F.R.S. 2nd Edition. London, 1872. 

Professor Flower has brought out a new edition of his very 
useful diagrams exhibiting the origin, divisions, connections and modes 
of distribution both to skin and muscles of the nerves of the human 
body. By reducing the size of the diagrams, compared with those 
in the first edition, he has put them into a more convenient and 
portable form, which will render them more acceptable both to 
students and practitioners. 


Lecons sur la Physiologie et ? Anatomie Comparée de VHomme et des 

Animaux. By H. Mitne Epwarps, Vol. x. Part 1. Paris, 1872. 

In this part of bis great work on Comparative Anatomy and 

Physiology, Milne Edwards describes the tegumentary system in the 

various groups of animals. The Part is marked by the same clear- 

ness of style and fulvess of reference to authorities which have cha- 
racterised all its predecessors. 


Untersuchungen zur vergleichenden Anatomie der Wirbelthiere, von Dr 
Cart GEGENBAUR, drittes Heft. Das Kopfskelet der Selachier, 
als Grundlage zur Beurtheilung der Genese des Kopfskeletes der 
Wirbelthiere, Leipzig, 1872. 

This is a continuation of the author’s well-known researches in 
comparative anatomy ; it is a very elaborate introduction to the study 
of the vertebrate skull, and is illustrated by twenty-two plates. 


Observations in Myology. By Proressor Humpeury. 8vo. Mae- 
millan and Co. 


This is a collection of the papers on the Myology of the Crypto- 
branch, Lepidosiren, Dog-fish, Ceratodus, and Pseudopus Pallasii, 
with the nerves of Cryptobranch and Lepidosiren, and the disposition 
of the muscles in Vertebrate animals, which have been recently pub- 
lished in this Journal. 


Roser’s Manual of Surgical Anatomy, translated from the fourth 
German edition by Joun C. Gatton, M.A. 


This is a useful book. ‘The descriptions are clear, and it is well 
translated. 


Anatomische Studien, herausgegeben von Dr C. Hasse, Prosector in 
Wiirzburg. Zweites Heft, Leipzig, 1871, drittes Heft, 1872. 

Both parts are occupied chiefly by articles on the organ of hear- 

* ing in Fishes and Reptiles, by Drs Hasse, Clason, and Ketel. There 

is also an article on the development of the atlas and the axis, by 


Dr Hasse, and one on the development of the vertebral column in 
Birds, by W. Schwarck, 


=) 


REPORT ON THE PROGRESS OF ANATOMY. 
BY PROFESSOR TURNER’. 


Ossrous System.—Albert-Kolliker communicates further obser- 
vations (Verh. der Wiirzburg Phys. Med. Gesellsch. July 19, 1872) 
on the OCCURRENCE AND DISTRIBUTION OF TYPICAL ABSORPTION SUR- 
FACES IN THE BONES. (See Leport, May 1872.) His object in this 
memoir is to furnish more precise topographical information of the 
position of these surfaces; and he conducts his enquiry partly by 
a microscopic examination of the bones of calves, which he tests 
with some other animals, and with infants in the Ist year; and 
partly by a series of madder- feeding experiments in young pigs 
and dogs. He then gives a detailed ‘description of his microscopic 
researches. With reference to the madder-feeding experiments, he 
has arrived at the following facts: Ist, the madder unites only with 
the newly deposited bone and tooth substance, and leaves the parts 
already formed quite unaltered: 2nd, the bone-substance which has 
once been coloured by madder appears to preserve its colour for a 
long time, and loses its red bone substance only in consequence of 
the absorption which takes place in the coloured localities. V. Feltz 
relates (ftobin’s Journal, 1872, 417) an experimental research into 
the power of ABSORPTION possessed by the MEDULLARY TISSUE OF THE 
RONES. Wenzel Gruber in Lull. de P Acad. Imp. de St. Petersb. 
1872, 473) records further observations on SUPERNUMERARY CARPAL 
BONES (/?eports, tv. 150, v. 375, vi. 433); an ossiculum intermedium 
in the left hand between the scaphoid, magnum and trapezoid: an 
ossicle in the distal row representing the absent process of the third 
metacarpal. Dr Gillette communicates (Ltobin’s Journ. Sept. 1872) 
an essay ON SESAMOID BONES IN MAN. He divides them into peri- 
articular and intra-tendinous. The peri-articular are true bones 
arising in pre-existing cartilage, and are developed on the flexor 
aspect of the joints. The intra-tendinous are special to certain ten- 
dons, principally those of the lower limb which are on the flexor 
aspect: not unfrequently they remain cartilaginous during life. 
Th. Simon relates a new case (Virchow’s Archiv, LY. 536) of ABNORMAL 
wiptH of the PARIETAL FORAMINA. See feporé, 1. 1082. Paul 
Broca (Bull. de la Soc. d’ Anthropologie, 1872) describes a DEFORMITY 
OF THE SKULL met with in the people of Toulouse and the vicinity, 
which he calls Ja deformation Toulousaine. It was originally care- 
fully studied by M. Gosse, who shewed how it was produced by ban- 
dages applied to the head in infancy. Broca’s observations are of 
value in showing the modifications which take place in the shape of 


1 To assist in making this Report more complete Professor Turner will be 
glad to receive separate copies of original memoirs and other contributions to 
Anatomy, 


168 PROFESSOR TURNER. 


the brain in these deformed skulls. Karl Langer communicates 
(Denk. der Akad. der Wiss. Wien, xxxt. 1870) an elaborate memoir 
on the GROWTH OF THE SKELETON, with the view of ascertaining if the 
increased growth in giants is due merely to normal processes of 
growth extending beyond what is customary, or to special modifiea- 
tions of formation.—-P. Bouland enquires into the normal cuRvA- 
TURES OF THE SPINE in man and animals (Lobin’s Journal, 1872, 
359). He states, contrary to the opmion generally entertained, that 
the human spine does possess at the time of birth an anterior con- 
vexity in the cervical 1egion, an anterior dorsal concavity, and some- 
times an anterior lumbar convexity. 


Muscvunar System.— Alex. Macalister has compiled a descriptive 
CaTALOGUE oF Muscunar ANOMALIES IN Human Anatomy (77rans. 
Roy. Irish Acad, xxv.), which will be of great service to all engaged 
in the study of this branch of anatomy. He has made a eareful 
search through the extensive literature of the subject, has classified 
the variations which have been described, and has incorporated with 
them a number of examples which have come under his own observa- 
tion. —— Wenzel Gruber continues his RECORD OF MUSCULAR VARIA- 
tions (Bull. del Acad. Imp. de St. Petersb. 1872, pp. 437 to 505): a 
m. tensor fuscie suralis, which arose from the right semi-tendinosus 
and was inserted into the fascia of the calf of the leg; it almost 
exactly corresponded with the specimen described by the Reporter in 
the last number of this Journal (Report, v1. 442): additional varia- 
tions in the m. palmaris longus (Report, 111. 197): a m. biceps brachit 
with a coracoid origin and with an origin from the humerus close 
to the insertion of the tendon of the pectoralis major, but with no 
glenoid origin: a m. radialis internus brevis with two heads of origin, 
one the Jarger from the border of the radius, between the anterior 
and lateral surfaces; the other from the aponeurosis of the fure-arm ; 
in the same arm (the left) a m. cubito carpeus was present, and in 
both arms the abd. digitt minimi had a supernumerary head of origin 
from the fascia of the fore-arm: a case in which the palmaris longus 
was absent, but where the flexor carpi radtalis was subdivided into 
two bellies, one inserted normally, the other into the anterior annular 
ligament: a case of m. costo-coracoiders supernumerarius situated 
below the pectoralis minor, which arose from the 5th and 8th ribs, 
and was inserted into the tip of the coracoid process. Gruber also 
has described (Leichert u. du Bois Leymond’s Archiv, xi. 663) 
cases of m. tibio-astragaleus anticus extending from the tibia to be 
inserted into the neck of the astragalus: and on p. 669 enters into 
an elaborate account of the m. ilio-costo-cervicalis, and of a new bursa 
mucosa on the tubercle of the Ist rib, and some accidental bursie 
on the back.——S. Pozzi notes (Robin's Journal, 1872, 269) as a 
frequent variation in the m, peroneus brevis the passage of a slip 
from its tendon of insertion forwards to be inserted into the poste- 
rior end of the Ist phalanx of the little toe.——S. Haughton con- 
tinues (P. 2.8. Lond. May 2, 1872) his researches into some ele- 
mentary principles in ANIMAL Mecuanics; No. vy, refers to the most 


ee ee OO eee 


REPORT ON THE PROGRESS OF ANATOMY. 169 


perfect form of a plane quadrilateral muscle connecting two bones, 
and No. vi. enquires into the theory of skew muscles, and investigates 
the conditions necessary for maximum work.—-—-An elaborate expe- 
rimental research into Locomotion 1n Maw has been conducted by 
G. Carlet. (Ann. des Se. Nat. 1872, 1). It has been carried on in the 
laboratory of M. Marey by the aid of his graphic method. After 
giving a brief historical sketch of the subject he describes the appa- 
ratus employed, and then proceeds to consider what takes place when 
a step is made, and the part played by the lower limbs in walking. 
The movements of oscillation, inclination, rotation and torsion of the 
trunk are then considered, and the movements of the upper limbs. 
The essay concludes with a brief statement of the principal conclu- 
sions arrived at, and of the theory of walking which he considers 
that he has established.— —M. Schlagdenhauffen communicates (fo- 
bin’s Journal, Sept. 1872) the Ist part of an essay on the MECHANISM 
or THE Musc es and treats of the action of the extensors.——G. M. 
Humphry has republished (Cambridge and London, 1872), with addi- 
tions, the OpservaTioNs IN Myonoegy which appeared in the 6th 
Vol. of this Journal, The volume contains a description of the 
myology of Crypto-branchus, of Lepidosiven, of the Dog-fish, of 
Uromastix spinipes, of Ceratodus and of Pseudopus pallisti, with an 
account of the nerves of Crypto-branchus and Lepidosiren, and an 
essay on the disposition of muscles in vertebrate animals, 


Neryous Sysrem.—With the object of facilitating and making 
more exact the method of determining the DEPTH OF THE GREY 
MATTER OF THE CEREBRAL Conyouvtions, H. C. Major (West Riding 
Lunatic Asylum Reports, u. 1872) has invented an instrument which 
he calls a Vephrylometer. It is made on the principle of a cork-borer, 
or cheese-taster, and consists of a glass tube 5 inches long, graduated 
in fiftieths of an inch, The bore of the tube is uniform throughout, 
the largest size he has employed is ;sths of an inch, the smallest, 
goths, The wall of the tube is about equal ta a goose-quill in thick- 
ness, but is made to taper off at one end so as to facilitate its passage 
into a convolution. In using the tube its tapering end is carefully 
inserted into the mddle of the free surface of a gyrus, from which 
the pia mater has been removed; the middle of the gyrus is se- 
lected so as to escape the grey matter which clothes its lateral as- 
pects, About half the length of the tube is passed into the brain, 
and the forefinger is then pressed closely upon the free end and 
the tube is withdrawn. The thickness of the grey matter may 
be read off on the graduated scale, His observations have as yet 
been made only on morbid brains in which he has found that the 
depth of the cortical layer in the frontal and parietal lobes, and the 
annectent gyri is not subject to much variation, though in the two 
former it is as a rule somewhat greater than in the latter; in the 
occipital gyri there is a marked decrease in thickness, whilst the 
temporo-sphenoidal convolutions have thicker grey matter than the 
occipital. In the orbital convolutions the grey matter is diminished 
in thickness, in the island of Reil it is deep ; the conyolutions of the 


170 PROFESSOR TURNER. 


inner aspect of the hemisphere correspond closely with the temporo- 
sphenoidal, whilst the cuneate and uncinate gyri invariably show a 
falling off. H. C, Major also in the same feports, p. 41, records 
an examination of the MINUTE STRUCTURE of the CORTICAL SUBSTANCE 
of the Bray in a case of chronic brain-wasting, and compares it with 
the appearances seen in a healthy brain. In a second memoir on 
the HISTOLOGY AND PHYSIOLOGY OF THE Nerves (Archives de Phys. 
1872, 427) Louis Ranvier describes the connective tissue of the 
nerves. He arranges it as follows: the ensheathing tissue forming 
layers, the peri-fascicular and the intra-fascicular. The surfaces of 
the Jayers of the ensheathing tissue are occupied by endothelial cells. 
The intra-fascicular tissue is extremely delicate, the fibres run longi- 
tudinally and the plate-like cells which it contains are situated 
partly on the surface of the nerve-fibres, partly on the vessels, and 
partly on the groups of small bundles of the connective tissue. He 
recognises both blood and lymph-vessels in the trunk of a nerve, but 
he has not yet determined any communication between the latter and 
the intra-fascicular connective tissue-———M. Lavdowsky records 
(Reichert u. du Bois Reymond’s Archiv, 1872, 55) observations on 
the minute STRUCTURE AND MODE OF TERMINATION OF THE NERVES in 
the urinary bladder of the frog. 


Ear.—Urban Pritchard communicates (P. 2. S. Lond. May 30, 
1872) an account of the SrrucTuRE AND Functions of the Rons of the 
‘CocHLEA in man and mammals and in Quart. Journ. Mie. Se. Oct. 
1872, a description of the methods of PREPARING THE COCHLEA FOR 
MICROSCOPIC INVESTIGATION, ‘These papers are in amplificatiun of his 
graduation thesis submitted to the medical faculty of the University 
of Edinburgh in 1871. He points out that the rods of Corti vary m 
length according to their position in the cochlea; at the base the 
outer are as nearly as possible equal to the inner, but proceeding 
upwards both rows increase in length with great regularity, although 
not in the same ratio, the outer increasing with much greater ra- 
pidity, so that near the apex they are twice uae length of the inner, 
The rods vary in length from about sjgth to s}oth inch, and there are 
about 3 rods in the inner to 2 in the outer row, He believes that 
not only for each tone or semitone, but even for much more minute 
subdivisions of the same, a particular rod is caused to vibrate, 
H. N. Moseley also in the same number of the Micros. Jowrn, 374, 
describes nethods of preparing the OnGAN or Corti for microscopical 
investigation, 


Eyr.—R. J. Lee in a paper on the sense of sight in birds (P. 
Rh. S. Lond., May 16, 1872) describes the CILIARY MUSCLE in three 
species of the order Rapaces; the Eagle Owl, the Egyptian Vulture, 
and the Buzzard; and in a supplementary notice the eye in ‘hea 
Americana, Phanicopterus antiquorum and Aptenodytes Humboldtii. 
H, N. Mose ley gives (Quart. Journ. Mic. Se. Oct. 1872) some in- 
structions as to the mode of. making microscopic preparations of 


ta) 
the Eyxrs or Inszcrs, Robinski commences an Essay (Letchert wu 


REPORT ON THE PROGRESS OF ANATOMY. 171 


‘du Bois Reymond’ Archiv, 1872, 178) on the anatomy, physiology, 
and pathology of the CRYSTALLINE LENS in man and mammals, 


Vascutar System.—Wenzel Gruber communicates (Virchow’s 
Archiv, uv. 445) an elaborate memoir on the ARTERIA THYROIDEA 
IMA and on variations in the origin of the INTERNAL MAMMARY and 
THYREO-CERVICAL ARTERIES. He not only reviews the literature of 
the subject, but communicates a series of new observations.—Paul 
Berger investigates the INTERNAL STRUCTURE OF THE UMBILICAL 
VESSELS (Ar chives de Phy ys. 1872, 551), and confirms the observations 
of Hyrtl on the presence of semi- “Tunar folds, connected with the wall 
of the arteries. In the umbilical vein also semi-lunar folds can be 
demonstrated. They are analogous to those feund in the arteries, 
but larger. Often however they are rudimentary. He does not, 
‘however, with Hyrtl regard these folds as valves, but conceives 
them to be formed by the entire thickness of the wall of the vessel. 


Tretu. —C, 8. Tomes supports (Quart. Journ. Mie, Sc. Oct. 
1872) the view that Nasmyru’s Memprane on the enamel of a tooth 
is the homologue of the thick coronal cement of the herbivora. He 
believes it to be continuous with the outermost layer of the cement, 
which accounts for its imperfect calcification. 


Kipyey.—J. Hyrtl contributes a beautifully illustrated memoir 
(Denk. der Akad. der Wiss. Wien, xxx. 1870) on the PELVIS OF THE 
KIDNEY IN MAN AND MAMMALS. The various modifications in form 
which this structure may exhibit are described and in most instances 
figured. W. Donitz describes (Reichert und du Bois Reymond’s 
Archiv, 1872, 85) the sTRUCTURE OF THE KIDNEY of the African 
Elephant.——C. J. Eberth gives an account (Centralblaté, 1872, 225) 
of a network of SMOOTH MUSCULAR FIBRES which he states that he has 
found on the surface of the HUMAN KIDNEY. These fibres are not in 
connection with the muscular coat of the blood-vessels. He has not 
found them in the stroma or in the capsule, but immediately beneath 
the latter. He has never seen this museular layer in the kidney of 
the ox, sheep or pig, 


CoMPARATIVE ANATOMY. 


MamMariA.—Rt. Hartmann contributes the first part of a memoir 
(Reichert u. du Bois Reymond’s Archiv, 1872, 107) on the ANTHRO- 
pop Aprs, in which he treats of the Chimpanzee, and figures and 
gives a description of some crania of this animal. C. J. F. Major 
notes (Ann. Nat. Hist. Sept. 1872) the occurrence of some Fossit 
Monkeys in Italy, and precedes his description by a review of fossil 
quadrumana in general. Alex. Macalister gives a short notice 
(Ann. Nat. Hist. July, 1872) of the anatomy of CynocepHatus Ha-° 
MADRYAS. Alex. Macalister communicates (P. 2. 8S. Lond. Jan. 11, 
1872) an abstract of a memoir on the MyoLogy OF THE CHEIROPTERA. 


172 . PROFESSOR TURNER. - 


From the study of the brains of some, and of casts of the cranial 
cavity of other specimens, Paul Gervais prepares an elaborate memoir 
(Nouvelles Archives du Muséum, 1870) on the forms of the Brain in 
THE CARNIVORA. J, Chatin communicates (Ann, des Sc. Nat. xv. 
1872) observations on the Myonocy or Hya@moscuus, viz. on the 
muscles of the fore and hind limbs. His researches lead to the con- 
clusion that this animal has many points of affinity with the pigs, 
A. Sanson, in a memoir on the Hyprip OF THE HARE AND THE 
Rapsit (Ann. des Sc. Nat. xv. 1872), gives an account of the crania, 
and concludes that of the two kinds of hybrids obtained by M. Gayot 
one is absolutely identical with the rabbit, the other approaches more 
to the hare, less however by the shape of its skul] than by its ex- 
ternal characters. Alph. Milne Edwards communicates his obser- 
vations on the PLACENTA IN THE TAMANDUA (Ann. des Sc. Nat. xv. 
1872). The specimen was 7’, tetradactyla, from New Grenada, 
Placenta unilobed and circular occupied a considerable portion of the 
surface of the oyum, and was very arched in its form. He applies to 
it the name placenta discoidal envahissant, The villi are not simple, 
but with very serrated vascular vegetations amongst them, and pre- 
senting a considerable thickness in the central portion, so as to give 
‘to this part of the organ a spongy appearance; the border is well- 
defined, and beyond it the chorion is smooth. There are appearances 
of a decidua, but the condition of the specimen does not permit him 
to speak positively on this point, He found no trace of the allan- 
tois, but owing to the hardened state of the specimen, from its pro- 
longed immersion in spirit, he was unable to separate the various 
membranes, and to isolate the constituent parts of the umbilical 
cord. G. Pouchet records (/obin’s Journal, 1872, 539) observa- 
tions on the composition of the VERTEBRAL COLUMN OF MyrkMrcoPHAGA 
JUBATA. Paul Gervais, from the study, more especially of casts 
of the cranial cavity, draws up a memoir (Nowvelles Archives du 
Muséum, 1869) on the forms of the BRAIN IN THE MARSUPIALIA. 
Rich. Owen, in continuation of his researches on the Fossil Mammals 
of Australia, communicates (P. R. S. Lond, Dec. 7, 1871, April 18, 
1872) a memoir on the genus Puascotomys.——Alex. Macalister 
communicates some additional observations (Ann. Nat. Hist. July, 
1872) on the Myonocy or SarcopHILus uRSINUS, and in the August 
number of the same volume an account of the MyoLtocy oF PHAScOLARC- 
TOS CINEREUS.——CeEtacEeA.—Thos. Dwight describes (Mem. Boston 
Soc. Nat. Hist, xt. 1871) the external characters and the skeleton ofa 
young Balenoptera musculus, 48 feet long, captured in Oct. 1870 off 
Gloucester, Massachusetts. The flipper was 5 ft. 4in. long, and the 
height of the dorsal fin along the anterior edge was 1 ft. 2in, The 
baleen was of a very light straw oolour anteriorly, whilst farther back 
dark stripes appeared on it, until the hindmost blades were of a uniform 
dark slate colour. He compares the skeleton, which is preserved in the 
Boston Museum, with the descriptions of the skeletons of B. muscu- 
lus given by previous writers, and points out the variations which he 
has observed; the most remarkable being the presence of 63 ver- 
tebre instead of 62 or G1, which is the number usually seenn—- 


REPORT ON THE PROGRESS OF ANATOMY. 173 


P. Fischer notes the ¢ases (Journ. de Zoologie, 1872) where the 
Sperm-Whale has been stranded on the sea-coasts of France. 
MM. Delfortie and Fischer make notes (Actes de la Soc. Linn. de 
Bordeaux, 1872) on some Cetacean bones from Léognan. Wm. 
Turner records for the first time the occurrence of Ziphius cavirestris 
in the British Seas (Zrans. Rk. S. Edinb. 1872). The specimen was 
captured off Hamna Voe, Shetland, in 1870. The skull is described 
~and compared with that of Sowerby’s whale, a specimen of which is 
in the Edinburgh Museum of Science and Art. Reasons are given 
for separating Sowerby’s whale from the genus Ziphius, and Gervais’ 
name of Mesoplodon Sowerbyi is adopted. The differences in the 
form of the naso-premaxillary region in the two animals are, in the 
opinion of the author, sufficient to base generic distinction on. 
Paul Gervais communicates (Vowvelles Archives du Muséum, 1871) 
a memoir on the Baleen Whales. He gives an account of the speci- 
mens in the Museum at Paris; he then enters into the characters of 
the skull as displayed in an antero-posterior median section; the 
relations of the vomer and the great mesial cartilage of the beak; 
the arrangement of the palatine bones, the characters furnished by 
the sternum and the form of the tympanic bones; he then considers 
the cerebral forms proper to the baleen whales, and concludes with 
some remarks on their classification. W. H. Flower gives an 
account (Z’rans. Zool. Soc. 1872) of the external characters and 
skeleton of Misso’s Dolphin, which he associates with the genus 
Grampus as Gr. griseus. W. H. Flower also (Ann. Nat. Hist. 
1872, 440) notes the discovery of the bones of a sub-fossil whale in 
Cornwall, which he considers to be the same as a Swedish specimen, 
described by Lilljeborg as Lschrichtius robustus. P. J. van Bene- 
den describes (Lull. de V Acad. Roy. de Sc. de Belgique, No. 7, p. 6, 
1872) various specimens of FossIL WHALES from Antwerp, which he 
refers to the following genera: Balena primigenius, Probalena du 
Buisii, Balenula balenopsis, Baleenotus insignis, Megapteropsis ro- 
busta, Plesiocetus garepit, Cetotherium hupschii, brevifrons, dubium 
and burtinii, Burtinopsis similis, and Herpetocetus scaldiensis. 
Jas. Hector gives an account (dun. Nat. Hist. 1872, 436) of the 
New Zealand Bottle-nose, Lagenorhynchus clanculus. H. Bur- 
meister states (Ann. Nat. Hist. July, 1872) that after perusing 
Gervais’s criticism of the Globio-cephalus Grayi, described by him 
in Anales del Museo Publico de Buenos Ayres, tomo 1, he has come 
to the conclusion that this cetacean is not a Globio-cephalus, but 
a true Pseudorca, which he regards as differing both from Ps. crassi- 


f=) 
dens and meridionalis, and to which he gives the name of Ps. Gray. 


.  Brrps.—M. Marey publishes a second memoir on the FLIGHT OF 
BIRDS AND INSECTS (Ann. des Se. Nat. xv. 1872). He compares these 
groups of animals in reference to their mode of flight; he attempts 
a synthetic reproduction of the phenomena of flight in the bird, 
and records a new series of experiments to ascertain the dynamical 
conditions under which flight is effected. With reference to the 
figure of 8, described by the wing during flight, he gives a different 


174 PROF. TURNER. REPORT ON THE PROGRESS OF ANATOMY, 


interpretation of the cause of its production, and of the nature of 
the movements by which it is produced, from that which has been 
advanced by J. B. Pettigrew (Reports, v. 385). Jas. Murie gives 
an account (Ibis, July, 1872) of the genus CoLtus, its structure and 
systematic place, and figures many of its bones. He considers it to 
be an aberrant form, and to afford the strongest proof of the inter- 
linking of type; and whilst it presents affinities to the sparrows, 
parrots, woodpeckers, hornbills, the hoazin, the rollers, touracoes, 
and finches, yet it cannot be said to belong to any one of these 
groups. He conceives it to be annectent between the Coccygomor- 
phe and Coraco-morphe.——J. Murie also in 7Z’rans. Zool. Soe. 
Nov. 7, 1871, makes a further note (Report v1. 447) on the PowpEr 
Downs of Rhinochetus jubatus; and in Month. Mic. Journ. Apvril, 
1872, he gives an account of the development of VEGETABLE ORGAN- 
isms within the THORAX of Livine Birps. 


Fisurs.—J. Oellacher makes a series of contributions (Stebold u. 
Killiker’s Zeitsch. 1872, 373) on the DEVELOPMENT OF THE OVA OF 
qHE River TRovr. W.K. Parker made the struCTURE and DEVE- 
LOPMENT OF THE SKULL OF THE SaLMon the subject of the Bakerian 
Lecture (P. &. S. Lond. May 30, 1872). 


REPORT ON PHYSIOLOGY. By T. Lauper Bruytoy, M.D., 
D. Sc, Joint-Lecturer on Materia Medica and Therapeutics, 
St Bartholomew's Hospital, and Davip Ferrier, M.A., M.D., 
Professor ef Forensic Medicine, King’s College, London. 


Nervous System. 


PuystoLoey or THE Corpora QuapRIGEMINA.—Knoll (Eckhard’s 
Beitriige, 1869, Centralb. 17, 1872) investigates some disputed points 
in the relation of the corpora quadrigemina to the movements of the 
iris, According to Flotrens these structures are the centres of the 
reflex action of the optic nerve on the pupil. Budge confirms this, 
but also states that lesion of the inner half of the nates causes para- 
lysis of the iris on the opposite side, while, on the other hand, Rienzi 
states that in such lesions he never observed paralysis of the iris, 
but only a transitory contraction. Knoll attributes some of these 
differences to the fact that, in all probability, in some cases the fibres 
of the optie tract, which spread themselves out on the nates, may 
have been injured. He makes experiments in rabbits, and finds, on 
cutting through the optic nerve between the eye and the commissure, 
there is complete dilatation and paralysis of the pupil on that side 
alone. If section is made between the commissure and the corpora 
quadrigemina complete paralysis ensues of the opposite iris, and of 
that alone, a fact which shows complete decussatien of the fibres of 
the optic tracts in the chiasma. The action of the oculo-motor on 
the iris is purely reflex. Sections of the optic thalami and corpora 
quadrigemina cause no affection of the iris, so long as the fibres of 
the optic tract are avoided. If these are involved then paralysis of 
the opposite pupil results. Lesions of the corpora quadrigemina do 
not seem to affect the movements of the animal. Irritation of the 
nates causes dilatation of both pupils, especially that of the same 
side, This ceases to be the case if the cervical sympathetics are cut, 
which seems to show that the action is to be attributed to downward 
irritation of the oculo-spinal centre, 


GALVANISATION OF THE Brain.—Hitzig (Reichert u. du Bois Rey- 
monds Archiv, 1871, 716—770 and 771—772) describes peculiar 
phenomena caused by passing constant currents through the head ; 
some of which had already been observed by Purkinje. Vertigo is 
produced, especially if the current is directed from one mastoid fossa 
to the other, or if one pole is placed in one fossa and the other on 
any other point. In addition to the vertiginous feeling there is an 
inclination of the head, or of the whole body, towards the anode 
when the current is formed, and towards the cathode when it is 
broken. Objects appear to sink towards the side of the cathode 
when the current is formed. Strong currents also cause involuntary 
movements of the eyes, similar to nystagmus. Besides these, other 


176 DR BRUNTON AND DR FERRIER. 


movements of the eyeballs, first to one side and then to the other, are 
observed, following each other with a certain amount of rhythm, 
beginning from the anode. When the currents are very strong both 
eyeballs are fixed in the angles towards the cathode, and oscillate 
there. Circular movements of the eyeballs are also seen. From 
various experiments Hitzig thinks that the movements are due to 
irritation of parts of the third and fourth nerves in the left eye when 
the current passes from left to right, and in the right eye to irritation 
of other parts of the third and the sixth when the current passes in 
the contrary direction. The intra-cerebral irritation appears due to 
the same causes as the electrotonic state ef nerves. He thinks that 
some of the vertiginous phenomena are in reality due to the invo- 
luntary movements of the eyeballs, and the consequent apparent 
movement of external objects. The other movements, which result 
from opening and closing the current, occur, whether the eyes are 
open or shut, and ate apparently voluntary in character, caused by 
the feeling as if the head were inclined towards the cathode, and the 
desire to restore the equilibrium of the body. If the eyes are opened 
the movements of the body itself cease, but are transferred to ex- 
ternal objects. 

Similar movements are observed in rabbits. Hitzig attributes 
the rotatory movements, which are observed in animals on lesions of 
the cerebellum, to voluntary efforts, directed with a desire to recover 
equilibrium, 


Cause oF SiEep.—Obersteiner (Archiv f. Psychiatrie, Bd. 29) 
considers the proximate cause of sleep to be the accumulation of 
acid products in the brain, just as in the case of muscular fatigue. 
The periodicity is merely a matter of custom like many other bodily 
functions, All conditions which interfere with the conveyance from 
the brain of acid combustion-products tend to induce sleep, and all 
conditions which favour their removal have a contrary effect. The 
conditions lie chiefly in the state of the circulation. Active hy- 
pereemia in this way prevents sleep. Passive hyperemia or venous 
congestion has an opposite effect. Anemia, by diminishing the ac- 
tivity of interchange between the blood and tissue, favours sleep, 
Changes in the chemical constitution of the blvod, such as by nar- 
cotics, interfere with the absorption and excretion of the acid pro- 
ducts, and hence cause sleep. He compares the cerebral vessels to 
the bronchial vessels, and destined only to nourish the tissue. The 
real initiative to cerebral action proceeds from the nervous system 
itself. Hence to allow of rest from the ordinary association links of 
cerebral action the brain must be disconnected from outward stimuli. 
When this is done a tendency to sleep ensues. So also if one im- 
pression is kept up long. The continuance of an impression is 
equivalent to none at all. The will acts as a sort of inhibitory 
centre to the reflex cerebration. In sleep this is at rest, and hence 
cerebration shapes itself into dreams. These may be conditioned by 
stimuli which are not of sufficient intensity to rise into con- 
sciousness, . 


sed 


ee 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 70 


INJECTIONS INTO THE BRAIN SUBSTANCE.—Beaunis (Gazette Med. 
Nos. 30—31, 1872) thinks that the injection of corrosive and other 
fluids into the brain by means of a fine needle would be of great 
service in investigations into the physiology of the great nerve 
centres, inasmuch as the lesions so produced should be more localised 
than by the process of vivisection. The results have generally been 
fatal convulsions after movements of various kinds when injections 
were made into the corpora striata and cornu ammonis.——H. Noth- 
nagel (Cextralblatt, No. 45, 1872) also gives a method by which the 
functions of various parts of the brain may be studied. It consists 
in injecting a concentrated solution of chromic acid through a small 
hole in the skull, by means of a subcutaneous syringe. The part 
injected becomes green and hard, surrounded by partial encephalitis. 
The function of the part is completely destroyed, while the lesion is 
exactly circumscribed. Nothnagel has made over one hundred expe- 
riments, the animals living at least eight to fourteen days. He pro- 
mises soon to publish his results. 


CoNDUCTION OF SENSORY IMPRESSIONS IN THE SpinaAL Corp.— 
Nawrocki (Zudwig’s Arbeiten, v1. 89) has continued and extended the 
research begun by Miescher under Ludwig’s direction (see this 
Journal, Noy. 1871, p. 220). Miescher found that the sensory im- 
pressions, which produced an action on the blood-pressure, were 
chiefly conducted up the cord in the lateral white columns, but he 
did not determine whether they were not partly conducted along the 
grey matter as well. Nawrocki’s experiments, however, show that 
the impressions are conducted entirely along the lateral columns, and 
not at all by the grey matter. They were conducted in the same 
way as Miescher’s, with this improvement, that the two small knives, 
used to limit the extent to which the cord was divided, were united 
into one instrument instead of being used separately. 


INSENSIBILITY OF THE SpinaL Corp.—Wolski (Pjliiger’s Archiv, 
v. 290—291) agrees with those who hold that the substance of the 
cord is itself insensible to irritation. In order to localise the irritant 
he used fine needles. No irritation was caused in frogs or dogs until 
the needle was inserted close to the roots of the nerves. 


TrritaTIon OF THE CuraNeous Nerves By SuLtpHuric AciID.— 
Baxt (Ludwig’s Arbeiten, v1. 69) sought by experiments on frogs, 
after the removal of the brain, to determine the time required by 
various dilutions of sulphuric acid to produce reflex contractions, 
While doing this he noticed that the frog is less sensitive to the acid 
at first, so that the reflex contractions occur more and more quickly 
when the same acid is applied at regular intervals; until after several 
applications have been made they require the same time at each irri- 
tation. A weak acid, applied after a stronger one, will produce 
a contraction more quickly than it would have done without the 
previous application of the stronger one. “His experiments were 
made with very dilute acids from 0.0003 to 0.003. The former never 
produced a contraction at alJ, the latter did so in about a second. 


VOL. VII. fg 


178 DR BRUNTON AND DR FERRIER. 


The sensibility varied in different frogs. No direct proportion exists 
between the strength of the acid and the time it requires, but if 
acids of different strengths are successively used a proportion is found 
between the differences of strength and the differences in time, for 
while the former diminish in an arithmetical, the latter do so in 
a geometrical ratio, The irritation must then be due not only to 
the amount of acid which reaches the nerves but to the rapidity 
with which it does so. From this it appears that it is not the con- 
dition of the nerve which exists after the acid has reached it, but 
only the process of alteration in it that causes irritation. 


Tropuic Action oF Nervous CENTRES ON MuscuLaAr TISSUE; 
ALTERATIONS IN MUSCLES AFTER LESIONS OF NERVES.— Vulpian(Compt. 
rendus, LXX1v. p. 964) finds that the grey matter of the spinal cord 
and medulla, and the corresponding parts in the Pons Varolii, exert 
a trophic influence upon muscles through their motor nerves as well 
as upon the motor nerves themselves. When the motor nerves are 
cut, the muscles, which are thus severed from connection with their 
trophic centres, become atrophied, and their contractility diminished. 
That the trophic influence is conveyed to the muscles through the 
motor nerves, and not through sympathetic fibres, is shown by the 
facial muscles becoming atrophied after division of the facial nerve 
close to its origin, when it contains only motor and no sympathetic 
fibres, as well as by the atrophy of muscles, which occurs after dege- 
neration of the anterior horns of the grey substance in the spinal: 
cord. 


Movements IN Nerve Freres From Evectric IRRITATION.— 
Engelmann (Pfliiger’s Archiv, v. 31) on irritating either the sciatic 
nerve of the frog, the nerves of the web or the small ones on the inner 
side of the leg by single induction shocks, noticed that the white 
substance of Schwann in some of the fibres became thicker, and its 
outlines jagged. Similar appearances were also produced in dead 
nerves and in pieces of white matter separated by ligatures from the 
centre parts of the nervous system. These changes he attributes to 
the rise in temperature caused by the current passage of them, as 
similar ones could be produced by warming the nerves to 33—35°C. 
He believes that in ordinary experiments on the irritation of nerves 
these changes may play a considerable part. 


An Errect oF GALvANIc CurRENTS ON MuscLE AND NERVE.— 
Hermann (Pfliiger’s Archiv, v. 223—275) finds that the conduction in 
living muscle is better in the longitudinal direction than in the 
transverse, but if the musele is brought into the state of rigor by 
heat, there is no difference in the amount of resistance offered to the 
current. The same is true of the nerves. Hermann explains these facts 
by internal polarisation in the region between the sheath and its con- 
tents. The resistance manifests itself soonafter the closure of the current, 
and is the same in interrupted constant currents as in continuous. 
With changes in the direction of current, however, the resistance in 
the transverse direction is less than when the current is constant, 


REPORT ON THE PROGRESS OF PHYSIOLOGY, 179 


though it is still greater than the longitudinal resistance. In nerve, 
little difference is observed. In order to account for these phenomena 
by polarisation, it must be supposed that in muscle and nerve, con- 
trary to other electrolytes, the signs soon become reversed with fre- 
quent reversal of the current. Investigation of the polarisation after 
opening of the current supported this idea. When the polarised 
currents of a transverse and longitudinal preparation of muscle were 
examined after opening the current it was found that the polarisation 
current of the transverse was greater than that of the longitudinal. 

¢ is otherwise in nerve. The longitudinal polarisation current ex- 
ceeds the transverse, so that the longitudinal polarisation of the dead 
nerve is greater than the transverse polarisation current of the living 
nerve. ‘This is attributed by Hermann to the fact of the transverse 
polarised current disappearing more rapidly than the longitudinal on 
account of the comparative approximation of the electrolytes. That 
the resistance in the transverse direction is due to internal polarisa- 
tion seems probable also from the fact that the polarisation increases 
to a maximum with strengthening of the current, and again dimin- 
ishes. 


ELEcTRICAL STIMULATION OF NERVE AND Muscie. — Willy 
(Pfliiger’s Archiv, v. 275—280) investigates the influence which the 
length of the intrapolar region exerts on the stimulation of nerve by 
electricity. The irritation increases the nearer the cathode is brought 
to the muscle, and the further the anode is removed. This is always 
true of a stimulation by closure of the current, but has exceptions 
in the case of stimulation by opening the current. Kulenburg 
(Berlin. Klin. Wochenschrift, No. 21, 1872) obtains the strongest 

stimulation of nerve, both from the cathode i in closure of the current, 

and from the anode in opening, when he places one pole on another 
nerve of the same plexus, or on the corresponding spinal centre. 
in this way a greater intrapolar length is obtained. Du _ Bois 
Reymond (feichert u. du Bois’s Archiv, 1871, 561—567) ‘On the 
influence of extra-conduction (neben-leitungen) on the current in the 
gastrocnemius of the frog.———‘ Directions for the use of the round 
compensator.’ (/bid. pp. 608—618). Fuchs (Zeitschrift f. Biologie, 
vu. 1, 100—123, 1872) investigates the laws of nerve-stimulation 
with unipolar currents, 


SENSATION. 


Vision witH Compounp Eyes. Boll (Reichert u. du Bois Reymond’s 
Archiv, 1872, H. 5, pp. 530—549) investigates the conditions of 
vision in animals which possess compound eyes in special reference 
to the question whether, as I. Miiller supposed, the individual facets 
each form an image of individual parts of the objects only, or 
whether, according to more recent researches, each facet forms a 
complete inverted image of the whole object. The latter view would 
necessitate the supposition that behind each facet there is a kind of 
retina, a supposition not altogether in harmony with the microscopical 


12—2 


180 DR BRUNTON AND DR FERRIER. 


researches of Max Schultze. Though nerve-fibrils may be traced to 
the central end of each crystalline cone, still they do not form a 
structure analogous to a retina. The crystalline cones of the eyes 
of anthropoda do not seem to be adapted for forming images, but they 
refract light, so that the light falling in the axis of the cone can be 
concentrated on the end of the nerve-fibril, and the light from neigh- 
bouring parts excluded. Boll agrees with the view expressed by 
Miiller. It would thus appear that the compound eyes are not to be 
regarded as an aggregation of simple eyes. The cornea and crystal- 
line cone of the anthropoda are not analogous to the cornea and 
vitreous humour of the vertebrate eye, but rather, that the cornea of 
the compound eye is analogous to the bacillary layer of the vertebrate 
retina, 


PERCEPTION OF SPACE IN THE SKIN OF THE LOWER EXTREMITY. 
Paulus (Zeitschrift f. Biol. vit. 237) in continuation of the research 
of Kollemkamp and Ullrich (see p. 210, Vol. v. of this Journal) 
examined at Vierordt’s suggestion the sensibility of the skin of the 
lower extremity. This differs from that in the arm, for in the latter 
it steadily increases from the shoulder downwards to the hand, 
while in the former, it increases from the hip downwards to the 
knee, and from the ankle onwards along the back of the feot and 
great toe. In the leg, however, the sensibility is least midway 
between the knee and ankle, and increases upwards to the knee, and 
downwards towards the ankle. The increase is more rapid in the 
latter than the former direction, The skin over the joints is very 
sensitive. 


Circulatory System. 


ACTION OF THE RIGHT AND LEFT Vaaus. Masoin (Bull. de? Acad. roy. 
de Médicine de Belgique, T. vt. 3 Serie, p. 4) and Arloin and Tripier 
(Arch. de Physiol. Norm. et Path. July, 1872), from experiments on 
rabbits and a pigeon, have found, independently of each other, that the 
right vagus has a much more powerful inhibitory action on the heart 
than the left. This action is exerted equally over every part of the 
heart. A year or two ago A. Bernhard Meyer (Das Hemmuwngs- 
nervensystem des Herzens, 1869, p. 61) observed that the left vagus 
of a tortoise (Hmys lutaria) obtained from certain districts in Ger- 
many had no inhibitory action on the heart whatever, while the 
right vagus had a powerful one. In specimens of the same species 
obtained from Italy the left vagus had a slight inhibitory action. 


INNERVATION OF THE Heart.—Schmiedeberg (Ludwig's Arbeiten, 
v1. 34) discusses the anatomy and physiology of the cardiac nerves 
in the dog. The investigation presented great difficulties from the 
numerous vessels which must be divided before the nerves are ex- 
posed. The experiments were made entirely on the nerves of the 
right side. As is well known the vagus and sympathetic are 
united in the neck of the dog into one common trunk, the vago- 
sympathetic, which descends along the carotid artery to the last 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 181 


cervical ganglion. This ganglion is the point of junction where the 
nerves not only of the heart but of other organs meet, and whence 
their fibres, after becoming mixed together in various ways, pass on 
to their several destinations. The chief nerves in relation with this 
ganglion are (1) Two filaments which connect it with the first 
dorsal. (2) The vagus trunk which passes onwards from it. (3) The 
recurrent laryngeal. (4) The superior cardiac nerve. (5) The infe- 
rior cardiac nerve. These latter three either spring from the gang- 
lion or its immediate neighbourhood. In describing the anatomical 
relations of these nerves he mentions that (1) the second or under- 
twig of the two connecting filaments sometimes joins the vagus 
instead of the lower cervical ganglion. (2) The recurrent laryngeal 
either springs from the lower end of the ganglion, or from the 
trunk of the vagus near it. At the point where it bends round 
the subclavian it gives off one or more branches which form a plexus 
with several twigs proceeding from the vagus below the ganglion. 
Its ascending part is connected with the ganglion by a tolerably 
strong filament opposite to it. (3) The superior cardiac nerve may 
either spring from the lower cervical ganglion itself, from the upper 
twig connecting the two ganglia, or from the vagus immediately 
below the ganglion, and passes along the innominate to the heart. 
(4) The name of inferior cardiac nerve is given to a branch which 
springs from the vagus near the ganglion, but below the origin of the 
recurrent instead of above it, like the superior cardiac branch. Fre- 
quently only one or other of the cardiac nerves is present. They only 
differ in their position, and their functions are sometimes exercised 
by the one, sometimes the other. Besides the nerves above men- 
tioned, the functions of the vago-sympathetic and of the filaments 
which pass from the spinal cord along the vertebral artery to the 
Ist dorsal ganglion, or roots of the ganglion, were investigated. The 
accelerating fibres which Rutherford found in the vago-sympathetic 
seem not to be contained in it above the inferior cervical gang- 
lion. It contains afferent fibres which cause increased blood-pres- 
sure, and sometimes a quickened pulse. The roots of the first dorsal 
ganglion contain efferent vaso-motor fibres which regulate a large 
vascular district. Accelerating fibres could not be shown to exist in 
them, but this might be due to their action being counteracted by 
the increased blood- -pressure. They contain dilating fibres for the 
pupil. A peculiar phenomenon was observed during “their irritation 
which renders it not improbable that they contain vaso-motor nerves 
for the pulmonary vessels. During stimulation the blood-pressure 
rose and the pulse became slow. Both effects disappeared atter the 
irritation was discontinued, but in a few seconds again re-appeared 
ina more marked manner than even during the invit: ition, and after 
lasting a few seconds again vanished. (1) The filaments: ‘connecting 
the two ganglia contain efferent fibres which accelerate the heart, 
but hardly influence the blood-pressure, and afferent fibres which 
cause acceleration of the heart, and sometimes a rise in blood-pres- 
sure. It is not certain whether this acceleration is reflex or not. 
The functions of the roots are not always distributed in the same 


182 DR BRUNTON AND DR FERRIER. 


way between the filaments, for irritation of the central end of the 
under twig at one time quickened the pulse, while irritation of the 
upper one slowed it and raised the blood-pressure; at another time 
irritation of the upper one quickened the pulse while that of the undi- 
vided under one left it unaltered but raised the blood-pressure. The 
vagus trunk below the ganglion contains sometimes inhibitory fibres 
(efferent) alone, sometimes a mixture of inhibitory and accelerating. 
The recurrent sometimes contains efferent accelerating fibres alone, 
sometimes inhibitory fibres alone, and sometimes a mixture of both. 
It contains afferent fibres which raise the blood-pressure. The superior 
and inferior cardiac nerves may either contain inhibitory fibres alone, 
or accelerating fibres alone. The inferior nerve may also contain a 
mixture of both, and afferent fibres which cause increased blood- 
pressure. Inhibitory fibres are more easily excited and more easily 
exhausted than accelerating fibres, and therefore when a nerve con- 
taining both is stimulated, the pulse is at first slowed, but gradu- 
ally becomes quicker and quicker, as the stimulation is continued, 
and the accelerating fibres begin to assert their preponderance over 
the inhibitory ones. Stronger currents are required to stimulate 
the accelerating than the inhibitory fibres, and while the latter may 
be paralyzed by numerous poisons, none is known which destroys 
the former, their action remaining perceptible on the pulsating parts 
of the frog’s heart after poisoning by digitaline or veratria. The ac- 
celerating nerves have often no influence whatever on the blood- 
pressure. During their action the pulse waves diminish in size. It 
is probable, though not certain, that this is due to imperfect dilata- 
tion of the heart during diastole. It is uncertain whether the dimi- 
nution of the pulse-wave depends on the quickness of the pulse, 
‘or vice versd, since they occur simultaneously through the heart of the 
dog. In that of the frog the wave becomes altered before the pulse-rate. 


On PeEcuLIARITIES OF THE IRRITABILITY OF THE MuscuLAR 
Fipres oF THE Heant.—H. P. Bowditch (Ludwig's Arbeiten, V1. 
p- 139) made use of the excised heart of the frog for his experiments 
on this subject. A glass cannula was passed into the ventricle from 
the auricle, and the lower two-thirds of the ventricle separated from 
the auriculo-ventricular groove, and therefore from the chief cardiac 
ganglia, by a ligature firmly tied round it. The ventricular apex was 
filled with serum, and suspended in a vessel also containing serum, 
It was irritated by shocks from an induction coil, and its contrac- 
tions registered by a small mercurial manometer. For a description 
of the apparatus we must refer to the original. When the pulsa- 
tions are equal in number to the irritations applied, he terms the 
pulse regular, when fewer, intermittent, and when more numerous, 
supernumerary. The supernumerary pulse was very rare, and gene- 
rally appeared only when the ligature round the ventricle was not 
sufficiently low. When weak shocks are used, the pulse is inter- 
mittent, but as their strength is increased it becomes regular, It can 
be also rendered regular by increasing the interval between the 
instead of increasing their strength, Although the strength 


shocks, 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 183 


of the shocks necessary to produce a regular pulse may be diminish- 
ed if the interval between them is increased, the diminution cannot 
be carried far without producing intermittence. It thus appears 
that great intervals of time are compensated by small differences in 
the strength of the irritant. The sensibility of the heart becomes 
increased after the repeated application of the irritant, so that after 
some time weaker shocks than at first suffice to produce a regular 
pulse. The amount of contraction has no relation to the mode of its 
recurrence. An irritation, which is only sufficient to produce an 
irregular pulse, suffices to excite the maximum contraction of whieh 
the ventricle is capable. An irritation, which produces a contraction 
each time, he terms infallible (unfehlbar), one which is able to produce 
the maximum contraction, but only occasionally, he calls sufficient 
(hinreichend), Exhaustion is not the reason why a sufficient irri- 
tation is not always infallible. The ligature round the ventricle has 
probably an inhibitory action, and when a second one is applied a 
stronger irritation is required to produce regular pulse. When fresh 
serum is supplied a weaker irritation is often sufficient. The amount 
of each contraction depends much on the time between it and the 
preceding one. <A succession of infallible irritations produce a series 
of contractions, each greater than its predecessor, to which he gives 
the name of a stair (Treppe). The form of the stair, the minima 
and maxima of the contractions, are independent of the direction 
and strength of the induced current. The length of the interval 
between two shocks greatly infinences the amount of contraction. 
When there is a short interval only between two shocks, but a long 
one between each pair, the contraction induced by the second shock 
of the pair is always greater than that of the first. The minimum 
and maximum height of the stair is also regulated by the interval, 
the height of the lowest step diminishing as the interval increased, 
and the greatest height being attained at an interval of between 4 
and 5 seconds, and diminishing with an interval either greater or less. 
The power of a heart exhausted by constant contraction can be 
partially restored several times by the introduction of fresh serum, 
but after each successive introduction the heart has less power, and 
is more easily exhausted. The exhausted heart differs from the fresh 
one only in the amount of its contractions and the length of time it 
can work. ‘The stair-like increase in the amount of its contraction, 
and its dependence on the interval between the irritations, are ob- 
servable in it, just as in the fresh one. When a weak solution of 
salt and gum was used, instead of serum, the stair-like appearance 
was absent, the height of the first contraction after an interval 
increasing with its length, and being greater than that of succeeding 
contractions. Muscarin diminished the height of the contractions 
and increased the stair-like appearance. In hearts poisoned by it 
the height of the curve increased as the interval diminished. Atropia 
nearly destroyed the stair-like appearance and increased the height 
of the contractions. The greatest height was obtained after a long 
interval. Delphinia in large doses destroys the irritability of the 
heart at once, small ones lessen it, so that an irritant sufficiently 


184 DR BRUNTON AND DR FERRIER. 


strong to be infallible only produces intermittent contractions. The 
stair disappears. Contractions also take place without external irri- 
tation; sometimes these occur when the most powerful shocks have 
lost all effect. Increased temperature causes diminution in the 
amount of contraction. This is not hindered by atropia. A ligature 
round the auricle increased the stair-like appearance. 


INHIBITION OF THE Heart’s Action tn Motiusca.— M. Foster 
(Pfliiger’s Archiv, v. pp. 191—195) describes a peculiar inbibitory 
action he has observed in the heart of Anodon and Helix. He 
found no nerve corresponding to the vagus. By direct irritation of 
the heart, when entirely removed from the body, by means of weak 
induced currents a condition of stoppage in the state of diastole was 
produced. When the current was increased in strength, first vermi- 
cular movements, and then tetanus resulted. During the stoppage of 
the heart electrical and mechanical stimuli have the power of ex- 
citing it. The facts are explained by the molecular condition of the 
contractile tissue of the heart itself, apart from nervous influence. 
Atropin, as in the vertebrate heart, paralyzes the inhibitory mecha- 
nism, and causes the systole to be remarkably prolonged. 


Meruop or Opraininc LympH in LArGEe Quantities. — Lesser 
(Ludwig’s Arbeiten, v1. 94) obtains lymph by inserting a cannula into 
the thoracic duct of dogs paralyzed by the injection of a small quan- 
tity of curare repeated from time to time. Lymph flowed in an almost 
unbroken stream from the duct, even when the animals on which he 
experimented had fasted for 24 hours, and the limbs were perfectly 
motionless. Variations in the rapidity of the stream occurred with 
a certain regularity. In some experiments, the flow underwent a 
gradual diminution from the beginning to the end, in others it was 
at first slow, and became gradually quicker till near the end, when it 
again grew slower. For a long time (as much as 69 minutes) after 
the cessation of the cardiac pulsations the lymph continued to flow 
almost as rapidly as before. This must have been due to the passage 
of fluid from the tissues into the lymphatic vessels, which was pro- 
bably caused by the intermittent pressure exerted by the movements 
of artificial respiration on the abdomen, producing a similar effect to 
alternate extension and flexion of the joints (see Genersich, in this 
Journal, vi, 228). In other experiments such passive movements 
were made on the posterior extremities. Like Genersich, he found 
that they increased the quantity of lymph. They also altered its 
quality, that which flowed during rest being opalescent, while during 
the continuance of passive movements the opalescence became very 
slight or disappeared altogether. The stream from the thoracic duct 
of a dog, which had been fed, was somewhat less during rest than 
from that of a fasting animal. Fasting animals, without movement, 
generally die without apparent cause, after large quantities of lymph, 
corresponding to }—} of the total quantity of blood, have been 
taken from them. When passive movements of the lower extremi- 
ties are made, a much larger quantity (as much as. 23 per cent.) can 
be withdrawn without death ensuing. ‘The blood is concentrated by 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 185 


the removal of lymph, its colour becoming more intense, and its power 
of taking up oxygen increased. Loss of lymph, therefore, acts dif- 
ferently on the blood from loss of fluid by the urine or sweat, which 
does not alter its composition. The difference is probably due to the 
.diff-rence in the forces which govern the secretion of the lymph and 
urine. The serum of the blood, after the extraction of a large quan- 
tity of lymph, was found in one instance to contain less water and 
ash, and more organic substance. The blood-pressure, instead of 
being lessened by the loss of lymph, was generally increased by it. 


INJECTION OF THE BLOOD-VESSELS IN COLEOPTERA. — Moseley 
(Ludwig's Arbeiten, vi. 60) proceeds to inject the blood-vessels in 
large beetles, such as Dytiscus marginalis, Hydrophilus piceus, Melo- 
lontha vulgaris, by cutting the elytra straight across while the insect 
is alive, and observing tb what point in ihe section the largest drops 

of blood make their appearanee. The position of the largest vessel 

being thus ascertained, he inserts into it a fine glass cannula. To 
the outer end of this a piece of indiarubber tubing is connected, 
and both it and the cannula are filled with the injecting fluid. The 
outer end of the tubing being closed, the fluid it contains is forced 
into the vessel, either by pressure with the fingers, or by a small 
weight. By inserting the cannula into the vessels of the severed 
dndiof, the elytra, or ‘the part still connected with the body, either 
the elytra alone, or the whole insect, may be injected. 


Temperature, 


AniMAL Heat.—Horvath (Centralblatt, No. 45, 1872) publishes 
an instalment of his observations on hybernating animals, concerning 
the condition of which, during their winter sleep, there: are many 
contradictory statements. He has made his observations in Russia, 
on the souslik marmot (Spermophilus Citillus). He finds that these 
-marmots do not sleep continuously the whole winter, but occasionally 
wake up, so that they would sleep from one to four days, and remain 
awake for a similar period. This irregularity in the sleep enabled 
him to make some very interesting observations regarding the tem- 
perature of these animals in their different states. The internal 
temperature of the souslik during the waking condition, as deter- 
mined by the thermometer, in the rectum, was from 35—37°C. 
During the state of hybernation the temperature is the same as that 
of the surrounding medium. Horvath observed a marmot, which a 
short time before had an internal temperature of only 2°C. awake, 
and become lively and active. He gives this as the only example on 
record of a warm-blooded animal surviving a ae ee sO near 
the freezing point. 


On ApsorPTION oF Hear purine Incupation.— Moitissier 
(Comp. Rend. LXXx1v. 54) noticed that when eggs are warmed to 41°C., 
and then allowed slowly to cool, those in process of hatching cool 
more quickly than others down to 35° or 36°, at which temperatures 


186 ._ DR BRUNTON AND DR FERRIER. 


they remain for ten or fifteen minutes, and then they become gra- 
dually colder, Just like those in which no chick was contained. This 
phenomenon is ascribed by the author to part of the heat between 
41° and 36° being absorbed, and applied to the development of the 
embryo. The specific heat of incubated eggs seems less than that of 
others. 


Digestive System. 


SECRETION OF THE SupmaxtLuaARy Guianp.—R. Heidenhain 
(Pfliiger’s Archiv, v. 40—47) describes the action of some poisons 
on the submaxillary gland, of great interest in a physiological point 
of view. When atropin is injected into the circulation, in doses 
sufficient to cause paralysis of the inhibitory action of the vagus, 
paralysis of the chorda tympani is also induced, so that irritation of 
this nerve no longer causes increased flow of saliva, although the 
the rapidity of the blood-stream in the veins remains unaffected. 
This shows that the secretion of this gland is independent of the 
circulation. The action is not due to paralysis of the secreting ele- 
ments of the gland, inasmuch as irritation of the sympathetic pro- 
duces the usual results. The cells are therefore in full functional 
activity, but their innervation by the chorda must be different from 
that by the sympathetic. Calabar bean stimulates the chorda, and 
removes the paralysis caused by atropin, but if given in large doses, 
this effect is apparently counteracted by changes induced in the cir- 
culation, from irritation of intraglandular vaso-motor centres, so 
that the stoppage of the blood-flow leads to resistance and paralysis 
of the secretion. Nicotin at first stimulates the chorda, and causes 
salivation, but ultimately causes complete paralysis of the secreting 
and vaso-inhibitory power of the chorda. Digitalin increases the 
flow of saliva by stimulating the central end of the chorda, Large 
doses do not cause paralysis. 


Formation or Pepsin in THE Stomacu.—LEbstein and Griitzner 
(Pfliiger’s Archiv, v1. p. 1) find that the pyloric part of the stomach 
contains pepsin. On separating the superficial from the deep layer 
of the pyloric mucous membrane, the latter was found to digest more 
actively than the former. As the chief cells (hauptzellen) of Heiden- 
hain are the only cells in the pyloric end of the stomach, and are more 
abundant in its deeper layers, the authors consider, in opposition to 
Friedinger, that the formation of pepsin takes place in them. 


PuystoLocy or Vomrrinc.—Hermann and pupils (Lfliiger’s Ar- 
chiv, v. 280-—282) investigate the physiology of vomiting, in special 
reference to the experiments of Grimm (see Journ. of Anat. and 
Physiol., No. 1x. p. 242). They find that tartar emetic acts 
more slowly when introduced directly into the blood than when 
introduced into the stomach. Hermann attributes the vomiting to 
direct irritation of the stomach by the drug, and not to direct action 
on a “vomiting centre.” The fact that tartar emetic may cause 
vomiting, even after the extirpation of the stomach, as in Magendie’s 


i CO 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 187 


experiment, is explained by reflex irritation from the mucous mem- 
brane of the pharynx and intestine. 


ForMAtIoN OF GLYCOGEN IN THE Liver.—Dock (Pfliiger’s Archiv, 
v. 571), in a series of experiments performed by him under Her- 
mann’s direction, found that when rabbits are deprived of food for 
several days glycogen disappeared completely, or almost so, from their 
livers. After a few injections of sugar into the stomach, glycogen 
again became abundant in the liver in all but one instance. Injec- 
tions of water or albumin did not produce glycogen. Glycogen did 
not appear in the liver, or only in traces after the injection of sugar 
into the stomach, if the floor of the fourth ventricle had been pre- 
viously punctured in three instances. In a fourth case glycogen 
appeared notwithstanding the puncture. Although no glycogen -was 
formed in those three cases, sugar did not appear in the urine after 
its injection into the stomach. In animals poisoned by curare, 
and kept alive by artificial respiration, injections of sugar into the 
stomach were not followed by the formation of glycogen in the 
liver, but sugar was found copiously in the urine. Poisoning by 
curare produced diabetes in animals which had fasted so long that all 
the glycogen must have disappeared from their livers, although no 
sugar was injected into the stomach. 


Tarrys Fistuta:— Albini (Alcune considerazioni sulla fistola 
intestinale secondo il metodo di Thiry, abstracted in Henle and 
Meissner’s Jahresbericht for 1871, p. 113) finds that the piece of 
intestine isolated in Thiry’s fistula becomes atrophied although its 
nerves and vessels are intact. 


Urinary System. 


Solowieff (Centralblatt, No. 22, 1872) describes the histological 
changes which result in the liver from gradual obliteration of the 
-portal vein. According to the length of time that the animals have 
lived changes are observed in the vessels, the hepatic cells, and the 
connective tissue. For details reference must be had to the original 
paper. 


A New Mernop or inpucinG Diaseres.—Bock and Hoffmann 
(Reichert wu. du Bois’s Archiv, 1871, 550—560) produce diabetes 
in rabbits by injecting rapidly large quantities of a one per cent. 
solution of common salt into the carotid or femoral artery. Melituria 
was always preceded by polyuria. The amount of sugar gradually 
rose to a maximum, and then gradually disappeared. At such period 
the liver gave neither sugar nor glycogen on analysis. On these 
facts the authors base a hypothesis that it is the result of nervous 
influence or change in the circulation. Further research is promised. 
Kiintzel (Jnaug. Diss. Berlin, 1872, Centralblatt, No. 44, 1872) 
obtains the same result with other solutions such as do not act as 
direct poisons, viz. various sodium salts, and also gum-arabic. 
Tiegel in a research “On a ferment-action of the blood” (Pfliiger’s 


188 DR BRUNTON AND DR FERRIER. 


Archiv, v1. pp. 249—266) makes some observations which have a 
bearing on the preceding. He finds that the blood-corpuscles when 
undergoing destruction possess a diastatic action on glycogen. This 
takes place in the liver. He thinks that diabetes may be induced in 
three ways. 

(1) When the glycogen is not increased, but when the destruc- 
tion of blood-corpuscles is more rapid than usual. To this class 
belong Harley’s experiments of injecting chloroform, &e. into the 
blood. 

(2) It may result without increase in the glycogen or increased 
destruction of the blood-corpuscles, simply if more glycogen is dis- 
solved. To this he attributes diabetes caused by injections of one 
per cent. salt solution into the arteries. 

(3) Diabetes may be produced by injecting glycogen into the 
vessels, 


Skin. 


Curaneous Resprration.—Rohrig (Deutsche Klinik, No. 23, 25, 
1872) gives the results of investigations on the cutaneous respiration. 
He finds that in the mean temperature the skin excretes 38 mgm. 
of CO, and 1:91 of water per hour. This is increased by food and 
also by increased temperature, as well as by irritants applied to the 
skin. He found also that the quantities of CO, and water given off 
&c. have very much increased during an attack of bronchial and 
nasal catarrh. He thinks that gases may be absorbed by the 
sound skin. A rabbit whose body was enclosed in a vessel of sul- 
phuretted hydrozen in such a manner that it could only gain access 
to the sys stem though the skin died with symptoms of poisoning by 
this gas. Chloroform also was found to exert a narcotising influence 
through cutancous absorption. In another paper (Archiv d. Heil- 
kunde, xu. 341—388) Rohrig gives the results of experiments on 
cutaneous absorption. Turpentine is readily absorbed, as well as 
camphor. They are readily recognised in the urine. Conia placed 
on the shaved skin of a rabbit caused death in twenty-eight minutes. 
Many other substances also are shewn to be capable of absorption by 
the unbroken skin.—Von Paalzow (Pfliiger’s Archiv, 1872, p. 492) 
finds that baths of water rich in carbonic acid do not increase the 
respiration. He finds, on the other hand, that powerful irritation of 
the skin, such as is produced by mustard, causes increase both in the 
amount of carbonic acid evolved and oxygen taken in. 


Suppression or Persprration.—Socoloft (Centralblatt, No. 44, 
1872) gives an abstract of the results which follow varnishing the 
skin and suppression of the cutaneous secretion. 

(1) A few hours before the death of the animals so treated, 
clonic and tetanic spasms appear in various groups of muscles, while 
the temperature in the rectum sinks in a marked degree. 

(2) Enveloping the animals in wadding did not serve to raise the 
temperature or arrest the fatal result. 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 189 


(3) Respiration of oxygen proved ineffectual to resuscitate the 
animals. 

(4) In the stomach ulcers were observed, the result of deep 
extravasations. 

(5) Albumen appeared in the urine very soon after the skin was 
varnished. 

(6) In all cases a diffuse parenchymatous inflammation of the 
kidneys was observed—sometimes swelling of the cells, and some- 
times fatty degeneration. This result was independent of the nature 
of the varnish used, whether turpentine varnish, or gelatin, or gum. 

Lang (Arch. d. Heilkunde, x11. pp. 277—287, 1872) investigates 
the cause of death when the skin has been varnished. In addition 
to other phenomena he found an hour or two after death ‘triple 
phosphate crystals’ in various parts of the body, and some of the 
uriniferous tubules blocked with a finely granular dark mass. He 
thinks that the triple phosphate crystals are the result of decomposi- 
tion of urea, and that the cause of death is uremia, 


Muscle. 


A full report of an important paper on muscle by H. Kronecker 
(Ludwig's Arbeiten, v1. 177) will appear in our next number. 


Miscellaneous. 


ON THE Gases oF Doc’s Lympnu.—Hammersten (Ludwig's Ar- 
beiten, vi. 121) finds that dog’s lymph contains on an average 35°38 
per cent. of carbonic acid, which is not simply absorbed, but is in 
a state either of loose or firm combination. It contains no oxygen, 
or only traces of it. Oxygen may, however, be present in it without 
being consumed. Its nitrogen amounts to 1-17 per cent. on an aver- 
age. In order to discover whether the easily oxidizable substances 
found by A. Schmidt in the blood of asphyxiated animals, were 
formed in the tissues, in which case they would be present in the 
lymph, or were formed in the blood itself, lymph and arterial blood 
were mixed together, and the gases analyzed after some time. If the 
lymph contained oxidizable substances the O of the blood would be 
diminished and the CO, increased. The experiment showed that the 
lymph from the body generally contained almost no oxidizable sub- 
stances, although it is possible that they may be present in the lymph 
of special organs. 


PERMEABILITY OF THE CorRNEA .TO Fiurms.—Laqueur (Central- 
blatt f. d. med. Wiss. 1872, p. 577) finds that fluids do not pass through 
the cornea of an excised eye, so long as it is covered with epithelium, 
but pass readily either from within outwards, or vice versd, when the 
epithelium is removed. Descemet’s membrane is permeable to fluids 
as well as the substance of the cornea. ‘The periphery of the cornea 
is more permeable than the centre. The diffusive power of the ante- 
rior epithelial layer of the cornea during life differs from that of the 


190 DR BRUNTON AND DR FERRIER. 


excised eye, since a solution of atropia passes readily through the 
former. 


FIXATION OF THE ARYTENOID CARTILAGES DURING PHONATION.— 
Jelenffy (Wiener Med. Wochenschrift, 1872, Nos. 3 and 4) has disco- 
vered, by experiments on the excised larynx, and by observations in 
three cases of paralysis of the muscles of vocalization, and one of 
spasm of the glottis, that the arytenoid cartilages are fixed in the 
first instance on the cricoid by the combined action of the arytenoidei 
postici and laterales, which draw them inwards and downwards, 
The arytenoidei laterales then turn the car tilages on their vertical 
axes by drawing the muscular processes forw ards, The vocal pro- 
cesses, Which form the other end of a two-armed lever, are conse- 
quently pulled backwards and fixed. As soon as this is effected the 
vocal cords can be rendered tense by the cricothyroid muscle. 


NEW METHOD OF ESTIMATING THE Lime AND PHospHoric ACID IN 
SeruM,—Pribram (Ludwig's Arbeiten, vi. 65) obtains serum in large 
quantities, by collecting blood directly from the vessels in long glass 
cylinders, cased in lead, which, as soon as coagulation has taken 
place, are laid in the drum of a centrifugal machine, the bottoms of 
the cylinders being directed outward. The drum being then made to 
rotate with considerable speed, the clot is driven outwards to the 
bottom of the cylinder and the serum collects at its upper end. The 
serum thus obtained is freed froth blood corpuscies by treating it 
a second time in a similar manner. In order to estimate the calcium 
it was precipitated by ammonium oxalate. As this precipitate sub- 
sides very slowly from serum, and cannot be collected by filtration, 
the fluid was again placed in the centrifugal machine, and in half 
an-hour the oxalate was obtained as a firm layer at the bottom of the 
cylinder. After washing the calcium in it was determined in the usual 
way. The remaining serum passed readily through a filter, and the 
clean filtrate was precipitated by ammonio-magnesium phosphate. 
This precipitate was also separated by the centrifugal machine, and 
then weighed as magnesium pyro-phosphate. 

Almost the whole of the calcium, but less than one-third of the 
phosphoric acid contained in the serum, is precipitated by this method. 
Even were it possible that tribasic calcium phosphate could be kept 
in solution by some organic substance present in the serum, the phos- 
phoric acid precipitated would be insuflicient to form this salt, and at 
least a part of the calcium must therefore be combined with some- 
thing else, but whether this is albumin or not is uncertain. The 
presence of lime as phosphate in the serum is also rendered impro- 
bable by the fact that the greatest part of the phosphoric acid can be 
extracted by alcohol. 


A CERTAIN sIGN oF Deara.—Magnus (Virchow’s Archiv, ty. 511 
—517) gives as a certain sign of death the complete cessation of the 
circulation, as determined in the following manner. If a finger is 
tightly ligatured even when life is at its lowest ebb there is observed 
at the point of ligature a white circle, due to arterial anemia, while 


——s -- — ——s 


——— ee ee 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 191 


the part beyond the ligature gradually becomes red, and ultimately 
livid from venous hyperemia. In the dead body these phenomena do 
not manifest themselves, M. Rosenthal (Wien. Med. Presse, Nos. 
18—19, 1872) investigates the CHANGES IN ELECTRICAL EXCITABILITY 
IN THE MUSCLES AFTER DEATH. Shortly after death—from one-and- 
a-half to three hours—the reaction of the muscles to electrical stimu- 
lation disappears. He thinks that in cases of apparent death electrical 
exploration of the muscles will be of great value. He mentions 
a case in which he in this way diagnosed apparent death lasting 
forty-four hours, and thus prevented burial. 


THE SpHincter Vesicz.—Kupressow (Pfliiger’s Archiv, v. 291 — 
294) measures the resistance offered by the sphincter vesicee by a 
column of water acting from the ureter. In rabbits the sphincter 
is forced by a pressure of from 40—50 cm. Section of the spinal 
cord between the fifth and seventh lumbar vertebra causes the re- 
sistance to sink to 12—16cm. The centre of innervation of the 
sphincter les between the fifth and sixth lumbar vertebre. The 
retention of urine in the bladder is attributed by Budge to the 
urethral muscles alone. This is not the case, though they aid the 
true sphincter. Shitting up of the urethra causes a sinking of the 
column one half im male rabbits, and one fourth in female. 


DR FRASER’S REPORT. 
Physiological Action of Medicinal and Poisonous Substances. 


Brommpe or Portasstum.—Believing that this salt forms the 
efficient agent in Bibron’s antidote, Dr E. A. Anderson administered 
it freely in two cases of snake bite (American Journal of the 
Medical Sciences, April, 1872, p. 366). Both cases recovered; and 
the opinion is expressed that death would have occurred had the 
bromide not been given. It is said, especially, to have promptly 
allayed pain and to have produced sleep. (Along with it, however, 
very considerable doses of alcohol were given; and this introduces 
an obvious fallacy in estimating the value of the antidote, even 
although it were granted, which it cannot be, that a sufficient 
quantity of the venom had been introduced to cause death in the 
absence of all treatment.) 


Mercury.—Dr H. Byasson has made some experiments (Journal 
de TAnatomie et de Physiologie, No. 5, 1872, p. 500) for the purpose 
of ascertaining within what time a soluble salt of mercury, intro- 
duced by the stomach, appears in the urine, saliva, and perspiration 
of human beings. The mercury was given in the form of perchlo- 
ride, in doses of 0.15 gr., combined in a pill with 0.015 gr. of 
hydrochlorate of morphia; and its presence in the fluids mentioned 
was detected by a delicate yet simple process of analysis described in 
the previous number of Robin’s Journal (p. 397). The experiments 


192 DR FRASER. 


led to the following conclusions :—1. Perchloride of mercury, admin- 
istered by the stomach, can be detected in the urine about two hours 
after its administration. 2. In the saliva in about four hours. 3. 
It does not seem to be discoverable in the perspiration. 4. Its 
elimination may be regarded as finished in twenty-four hours after 
administration. 


Cutorat.—Among other statements regarding the effects on 
rabbits of non-lethal and lethal doses of chloral, exhibited by subeu- 
taneous injection, Dr A. Edward M°Rae (£dinburgh Medical Journal, 
August, 1872, p. 133—149) makes the following:—Sleep was some- 
times produced by 10 grs.; sometimes, however, 30 grs. had scarcely 
any effect; death resulted, in two experiments, from 60 grs., but in 
another experiment this dose did not prove fatal’. About four 
minutes elapsed between the injection and the manifestation of 
the first symptoms. These consisted of increase of the number of 
the respirations, immediately following the injection, then slowing 
with jerkiness of character until sleep was produced, and still further 
slowing until death, which always occurred when the rate had 
diminished to 28 per min.; the production of hypereesthesia with all 
the doses that were given, followed by anesthesia when the doses 
were lethal; loss of special sensibility, vision being affected before 
hearing, hearing before smell, and smell before touch; and distinct 
elevation of temperature when small doses were given, but diminu- 
tion of temperature when the doses were large. Shivering was not 
observed in any experiment where death supervened, but it pre- 
ceded resolution in non-fatal experiments. Salivation occasionally 
occurred, and likewise relaxation of the sphincters, with increased 
peristalsis of the bowels and bladder. Rigidity set in immediately 
after the respirations ceased. The injection of the above doses almost 
always caused local inflammation, which ended in resolution, suppu- 
ration or gangrene. In the fatal experiments, the superficial blood- 
vessels were empty; while the internal vessels were full of blood; the 
walls of the heart were flaccid, and both sides were gorged with 
blood; the lungs were congested, with here and there apoplectic 
spots; the vessels of the encephalon were full of blood; and the 
odour of chloral was distinctly perceived in the substance of the brain. 


CHLORAL-SULPHYDRATE.—By submitting anhydrous chloral to the 
action of sulphuretted hydrogen, M. Henri Byasson obtained a 
white crystalline substance, having a disagreeable odour, and a pecu- 
liar taste suggestive of anhydrous chloral (Journal de (Anatomie et 
de la Physiologie, No. 3, 1872, p. 297). An analysis of this sub- 
stance has led M. Byasson to propose for it the formula C* HCl’ O°, 
2 HS. Under the influence of hydrated alkalies or ammonia, it is 
decomposed in an analogous manner to chloral, as is shown by the 
equation, 

C, HCl, O,, 2 HS+2 (KO, ee, HCl,+C, HKO,+ KS, HS+ 
2 HO. 


2 Unfortunately no information is giyen regarding the weight of the animals, 


REPORT ON THE. PROGRESS OF PHYSIOLOGY. 193 


It is decomposed also by water and alcohol, which renders difficult 
the examination of its physiological action. By dissolving it in 
ether, however, M. Byasson was able to administer it to guinea-pigs. 
He found that, in non-lethal doses, it produces diminution of tem- 
perature to about one degree Cent., muscular resolution, and sleep 
lasting about two hours, without any notable alteration of sensibility, 
but with slight acceleration of the rate of the heart’s contractions. 
At the end of the sleep the animal quickly recovers its normal con- 
dition. 


Formic Eruer.—In a research on chloral hydrate by MM. 
Byasson and Follet (see Journal of Anatomy and Physiology, Vi. 
1872, p. 493), the opinion is expressed that a certain portion of the 
action of this substance depends on the formic acid which is pro- 
duced under the influence of the alkaline salts of the blood. This 
opinion was difficult to prove because the formiates are physiologi- 
cally inert. It was important, therefore, to find a substance which 
produces formic acid as a result of its decomposition. M. Byasson 
(Comptes Rendus, 29 April, 1872, p. 1202) now publishes the result 
of some experiments with formic ether, a substance that fulfils the 
required indications, as it is readily converted by alkalies into alcohol 
and a formiate. When this substance was adininistered to animals 
by inhalation, it rapidly affected them, but not so rapidly as chloro- 
form; and the symptoms were those of asphyxia, with lowering of 
the temperature, incomplete muscular flaccidity, and diminution, 
without total suspension of sensibility. Return to a normal state 
was not accomplished until several hours. Similar symptoms were 
pronounced by subcutaneous injection. In an experiment on man, 
where the administration caused a marked tendency to sleep, formic 
acid was detected in the urine. By way of comparison, acetic ether 
was given in similar experiments, but no decided effect was observed. 
M. Byasson believes that these results confirm his previously ex- 
pressed opinion, that the physiological action of chloral hydrate is 
“not the same as that of chloroform introduced into the blood in 
small successive doses, but that it is an action special to that sub- 
stance, resulting from the combined actions of the chloroform and the 
formic acid which are produced in the economy under the influence 
of the alkaline carbonates of the blood. 


CYANATES.—Continuing his important researches on the trans- 
formations undergone by substances introduced into the system, Dr 
Rabuteau has made some experiments (Jowrnal de Anatomie et 
de la Physiologie, No. 3, 1872, p. 335) to discover if cyanates 
are poisonous, after they become reduced to cyanides; or if they 
undergo in the organism the same transformation they experience in 
contact with water. He finds that cyanates are not poisonous, and 
that in the body they produce alkaline carbonates, which may be 
detected in the urine. It is much the same thing, therefore, to 
administer an alkaline cyanate, or to administer an acetate, lactate 
or tartrate, 


VOL, VII, 13 


194 DR FRASER. 


Optum.—The action of the chief immediate principles of opium 
has formed the subject of an elaborate research by Dr Rabuteau 
(Journal de VAnatomie et de la Physiologie, No. 3, 1872, p. 302). Be- 
sides investigating the soporific, convulsant, toxic and analgesic effects 
of these principles, Dr Rabuteau has examined the influence they exert 
on the exosmosis of fluids into the intestinal canal, according to the 
method of Dr Moreau. His experiments in great measure confirm 
those of Claude Bernard respecting the soporofic, convulsant, and 
toxic power of these principles on the lower animals; but they seem 
to show that in man the relative prominence with which these actions 
are exerted is somewhat different than in the lower animals. This, 
and the other main results of his experiments, are shown by the 
classification of the opium principles according to their physiological 
action on man, which he has published :—In their order as Soporijics: 
morphia, narceia, and codeia; the other principles do not produce 
sleep. In their order as poisons: morphia, thebaia, codeia, papa- 
verin, narceia, and narcotine. In their order as analgesics (pain- 
destroyers): morphia, narceia, thebaia, papaverin, and codeia; nar- 
_cotine does not appear to lessen pain. In their order as anexosmotics: 
morphia, narceia; the other principles do not arrest diarrhea. Dr 
Rabuteau further points out that opianin, a principle that is present 
only in minute quantity in opium, has an action very similar to 
that of morphia, and that meconin and meconic acid are devoid of 
physiological activity. 


THE COMBINED Errects oF Oprum ALKALOIDS AND CHLOROFORM.— 
The statement of Claude Bernard and Nussbaum that by administer- 
ing morphia and chloroform in combination very complete anes- 
thesia may be obtained with a quantity of chloroform greatly less than 
is required when chloroform is alone given, has received confirmation 
and extension from the observations of MM. Labbe and Guyon and 
Dr Rabuteau. The former observers mention (Journal de Pharmacie 
et de Chimie, Mai, 1872, p. 398) that among their other results they 
found that this combination produces an anesthesia of long duration, 
which may be prolonged still further by successive small doses of 
chloroform. They believe, therefore, that by this combination the 
risk of fatal accidents is greatly diminished.—Dr Rabuteau, towards 
the termination of his paper on the opium active principles (oc. cit.), 
expresses his adherence to the statements of previous observers, and 
adds that nearly all the opium alkaloids, even those which are not 
soporifics, are able to continue the analgesic action of chloroform, 
because nearly all of them possess the power of lessening sensibility. 
Narcotine, however, is not able to do so; but he has found that 
narceia is nearly as efficient as morphia, while codeia and papaverin 
only feebly continue the action of chloroform. 


ApromorpuiA.—The evidence in favour of the valuable emetic 
properties of apomorphia has received an important addition from 
the observations of Dr Siebert (Archiv der Heilkunde, December, 
1871; and Gazette Hebdomadaire, No. 22, 1872, p. 366). He 
finds that this agent does not produce any effect on the digestive 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 195 


organs except the simple production of emesis. After its injection 
under the skin the pulse becomes irregular and slightly accelerated, 
and these changes are most pronounced at the moment of nausea 
and before vomiting, while after vomiting slowing takes place. In 
short, apomorphia affects the circulation in very much the same way 
as tartar emetic and ipecacuanha; a similarity that is further shown 
by the lowering of the temperature which succeeds the emesis. It 
differs from these substances, however, in being altogether destitute 
of irritant property; its injection under the skin does not produce 
inflammation, and neither diarrhea nor disturbance of digestion 
follow its administration. r Siebert found that the minimum 
quantity requisite to produce emesis by subcutaneous injection is, in 
man, from 0.09 gr. to 0.1 gr., in cats 0.3 grs., and in dogs 0.015 gr. 
to 0.03 gr. He could not discover the maximum dose that can be 
given to cats and dogs without producing injurious effects, because even 
when doses 80 or 100 times as large as the above were given, there 
were no symptoms of cerebral disturbance. He believes that in man 
the dose may, without risk, be increased to 0.2 or even 0.6 gr. 
When given by the stomach much larger doses are of course required. 
Thus, a man of 49 years, took 0.95 gr. and only nausea resulted, and 
when he took 1.5 gr. emesis oceurred without any injurious effects. 


Quinta.—Several years ago Professor Binz made the extremely 
valuable discoveries, that quinia has the power of preventing putre- 
faction, that it is an energetic poison for low vegetable and animal 
organisms, and that it is able to arrest the movements of the white 
blood-corpuscles. The last of these discoveries has lately attracted 
great attention from its obvious relationship to Cohnheim’s views 
regarding inflammation and suppyration, according to which pus is 
mainly a collection of white blood-corpuscles that have passed through 
the walls of the vessels. If quinia be able to arrest the movements of 
these corpuscles, it may be expected to exert a very beneficial influ- 
ence as a remedy in inflammation. Many observers, accordingly, have 
turned their attention to this point. Of these, Stricker threw some 
doubts on the value of Binz’s results, by suggesting that the arrest in 
the movements of the white corpuscles was simply the result of the 
avid contained in the solution of quinia that was employed. This 
criticism has induced Dr Kerner to reinvestigate the subject (Central- 
blatt, No. 9, 1872, p. 143). To avoid the objection raised by Stricker, 
he made use of perfectly neutral solutions of the hydrochlorate and 
carbonate of quinia. He mixed the blood of cats and dogs with a 
solution of one part of the salt in ten of water, in the proportion of 
one part of the solution to 4000 of blood, and found that the move- 
ments of the white corpuscles were immediately arrested, and that 
these bodies soon became rounded and granular. By way of com- 
parison, he made a number of similar experiments with solutions of 
salicin, caffein, atropia, and arsenate of potassium, and he was able to 
satisfy himself that these substances exert no decided action on the 
white blood-corpuscles. An elaborate series of researches by Dr 
Geltowsky (Practitioner, June, 1872, p. 321) confirms the more im- 


196 pus DR FRASER. 


portant of the statements of Binz, Kerner, and those who coincide 
in their views. In two points, however, Geltowsky differs from them: 
namely, in the degree of toxic strength of quinia—held by Binz, 
Kerner, &c. to be about 1: 4000 of human blood, but by Geltowsky 
only 1: 2800—; and in the possibility of obtaining the special effect 
on the white corpuscles by administering quinia to man. Both 
of these differences are criticised in a recent paper by Professor Binz 
(Practitioner, September, 1872, p. 141). In reference to the first of 
them, Binz not only reiterates the statement that the proportion of 
one part of quinia to 4000 of blood is a sufficient one to destroy the 
movements of the white corpuscles, but he likewise gives the details 
of some experiments made with the blood of a leucocythemic patient, 
which showed that the white corpuscles of human blood may be 
paralysed by quinia in the proportion of one part of quinia to 10,000 
parts of blood. In reference to the second point of difference, which 
Geltowsky has given expression to, by stating that “in the case of 
man, * * * it would be necessary, in order to obtain the special effect, 
to take almost one drachm of quinine, which would be impossible.” 
Binz quotes the experience of Professor Socin of Bale, who found it 
quite practicable to administer from six to seven grammes of hydro- 
chlorate of quinia daily (6 grammes=about 13 drachm). He also 
states that it can never be our object to kill the white blood-corpus- 
cles in the human body by quinia. They are necessary for life, and 
their death would be our death. All that is intended to be done to 
them is to decrease their number and the activity of their movements; 
and this can undoubtedly be done by quinia. 


STRYCHNIA.— Some observations by Brown-Séquard on the influence 
of carbonic acid in arresting epileptic convulsions promise to afford a 
valuable practical application in the treatment of strychnia poisoning 
(Archives de Physiologie, No. 2, 1872, p. 204). He finds that when 
epilepsy is artificially produced in guinea-pigs, the fit may be at once 
arrested by directing with considerable force a current of carbonic 
acid gas against the mucous membrane of the fauces. On account of 
the great sensibility of the mucous surface of the nares in man, he 
believes that the same result may be obtained in patients suffering 
from epilepsy by injecting this gas into their nostrils ; and he recom- 
mends this treatment in poisoning by strychnia. Brown-Séquard 
asserts that Rosenthal has fallen into an error in ascribing the ar- 
restment of respiratory movements, or of strychnia convulsions, by 
pulmonary insufflation to superoxygenation of the blood. He believes, 
on the contrary, that these results are due simply to mechanical irri- 
tation, by the forcibly inspired air, of the ramifications of the vagi 
nerves in the bronchi, and of the phrenic and other diaphragmatic 
nerves. ‘The experiments which lead him to adopt this view show 
that when the spinal cord is divided above or below the origin of the 
phrenic nerves, or when the vagi nerves are divided, pulmonary in- 
sufflation no longer causes a cessation of the respiratory movements. 
In like manner, the forcible injection of carbonic acid acts as an 
irritant to the nerves in the air-passages, and an inhibitory influence 


REPORT ON THE. PROGRESS OF PHYSIOLOGY. 197 


is thereby exerted upon respiration and upon convulsive movements. 
Carbonic acid, which is an excitant of convulsions when acting upon 
the nerve centres, is thus, also, an arrester of convulsions when it 
acts upon certain of the mucous surfaces. The action is, however, in 
both instances the same: it irritates the nerve-elements (fibres or 
cells) with which it comes into contact. If these elements possess the 
power of producing convulsive movements, these movements occur ; 
if, on the contrary, they possess the power of arresting the activity 
that causes convulsions, this activity ceases, and the convulsions no 
longer occur. 


BELLADONNA.—In some interesting communications (Practitioner, 
August and October, 1872) Dr Sydney Ringer brings forward an 
abundance of evidence to prove that belladonna and its active prin- 
ciple are able to check and prevent sweating, whether the result of 
disease or induced by exposure to an elevated temperature. In the 
former case, his observations enable him to conclude that gjoth of a 
grain of atropia, injected under the skin, is generally sufficient to 
check sweating for one night. This dose produces dryness of the 
fauces, but it does not dilate the pupils. Stramonium, it was found, 


is able to exert the same influence. 


‘ampHor.—Dr John Harley describes some observations (Practi- 
tioner, October, 1872, p. 210) which appear to shew that camphor 
acts with but little energy on man. So large a dose as thirty-five 
grains merely caused giddiness, followed in thirty-five minutes by a 
full, bounding, but not accelerated pulse, by flushing of the cheeks, 
and by a little somnolency; all of which symptoms had nearly dis- 
appeared within two hours after the dose was administered. The va- 
pour, he finds, acts with considerable energy as a poison on moths, 
and produces very decided hypnosis in mice, 


Antaconism.—(Between chloral and strychnia.) M. Oré, in a 
series of communications to the French Academy (Comptes Rendus, 
T. uxxiv. Nos. 24 and 26, T. uxxv. Nos. 1 and 4), describes a re- 
search undertaken to test the truth of Liebreich’s statement, that 
strychnia is an antidote to chloral. He, in the first place, points out 
that Liebreich has omitted to determine the toxic dose of either of 
these substances with the required accuracy, and, accordingly, that 
he has fallen into the error of concluding that the same dose is suf- 
ficient to kill animals of different weights. Oré finds that the mini- 
mum-lethal dose of chloral, given subcutaneously, is about 60 grains, 
and that of strychnia 0.03 grains, for rabbits weighing four pounds. 
In a number of experiments, where this minimum-lethal dose of 
chloral was injected at various intervals before different doses of 
strychnia, he found that death always occurred, although in some in- 
stances its production was apparently postponed for some hours, by 
the influence of the strychnia. Oré concludes, therefore, that strych- 
nia is not an antidote for chloral. At the same time, he found that 
the chloral had an obvious influence in preventing or diminishing the 
convulsive effects of the strychnia, but that this influence disappears 


198 DR FRASER. ~ 


when the dose of strychnia is increased beyond certain limits. (This 
research is not quite conclusive, as the doses of antidote (strychnia) 
were restricted to those extending from slightly below to considerably 
above the minimum-lethal).—(Between physostigma and atropia.) 
Dr Thomas R. Fraser has published (Z'ransactions Royal Society of 
Edinburgh, Vol. xxvi. 1872, p. 529 —713) a continuation of his pre- 
liminary experiments on the antagonism between physostigma and 
atropia (see this Journal, Vol. tv. 1870, p. 168). By the result of a 
large number of experiments he confirms his previous statement, that 
atropia is able to prevent the fatal effect of large lethal doses of phy- 
sostigma. He further shews that this antagonisin is successfully 
exerted only within a well-defined range in the doses of the two sub- 
stances. For the purpose of defining this range he performed three 
series of experiments: the chief objects of the first two of which were 
to ascertain the maximum dose of physostigma that can be success- 
fully antagonised by atropia, and the range of doses of atropia that 
can successfully antagonise lethal doses of physostigma ; while the 
chief object of the third series was to ascertain within what limits of 
time between the administration of the two substances successful an- 
tagonism occurs. The results obtained by these three series of expe- 
riments have been graphically represented in diagrams, in which the 
region of successful antagonism is mapped out.—(DLetween physo- 
stigma and strychnia.) An interesting case is described by Mr Ash- 
mead (Edinburgh Medical Journal, September, 1872, p. 235) of 
strychnia-poisoning treated by physostigma. One drachm of solution 
of strychnia (=0.5 gr. strychnia) had been taken, and recovery took 
place after the administration of a little more than two grains of 
extract of physostigma.—(Detween atropia and morphia.) Careful 
details are given by J. Magee Finny (Dublin Journal of Medical 
Science, July, 1872, p. 38) of a case of atropia-poisoning successfully 
treated with morphia. Although there is no evidence to show that 
the dose of atropia was as large as the minimum-lethal one, and, 
therefore, that the morphia counteracted the lethal action of atropia, 
still the account of the case appears to show that very distressing 
atropia symptoms may be relieved by morphia. 


ABSORPTION OF MEDICINAL SUBSTANCES BY THE SKIN.—By ex- 
posing the surface of the body to vapour obtained from water in 
which iodide of potassium was dissolved, M. Bremond (Comptes Ren- 
dus, 24 June, 1872, p. 1583) has been able to satisfy himself that 
this salt is absorbed by the skin. He finds that unless the skin has 
been well cleansed with soap before the vapour-bath, absorption does 
not take place below a temperature of 100.4 F.; but that if it has 
heen thus cleansed, absorption takes place at temperatures varying 
from 93,2 to 96.8 F. The absorbed salt commences to be eliminated 
with the urine at about two hours after the bath; and elimination 
after a single bath altogether ceases within twenty-four hours, what- 
ever may have been the quantity of salt in the vapour, or the tempe- 
rature and duration of the bath. In cases, however, where ten or 
twelve vapour-baths have been taken, elimination continues during 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 199 


three or four days, and where twenty-five or thirty have been taken, 
it continues during ten or twelve days after the last bath. 


ACTION OF VARIOUS SUBSTANCES ON THE Nerves or THE SUBMAXx- 
ILLARY GLANDS.—The well-known fact that dryness of the fauces 
results from the action of atropia, led Paul Keuchel (Das Atropin und 
die Hemmungsnerven) to examine the influence of this substance upon 
the secretory fibres of the chorda tympani. He discovered that these 
fibres are paralysed by atropia ; and inferred that this paralysis is the 
result of a direct action upon certain inhibitory nerves, R. Heiden- 
hain has lately repeated these experiments (Centralblatt, No. 21, 1872, 
p. 326; from Pfliiger’s Archiv, v. 40—47) on curarised dogs, by in- 
jecting into the jugular vein a dose of atropia sufficient to paralyse 
the cardiac filaments of the vagi. After this injection, excitation of 
the chorda tympani no longer cansed secretion : nevertheless, there 
occurred an acceleration in the flow of venous blood through the 
gland, which differed in no essential respect from that which resulted 
before the administration of atropia. This observation proves that 
the secretion produced by excitation of the chorda tympani is entirely 
independent of any modification in the circulation of the gland, and, 
therefore, that totally different sets of nerve-fibres are concerned with 
the secretion and circulation of this gland. Indeed, notwithstanding 
the incapacity of the chorda tympani to excite secretion when it is 
acted upon by atropia, the sympathetic nerve still excited this secre- 
tion when it was irritated. It is, therefore, the extremities of the 
secretory fibres of the chorda tympani alone which are affected by 
atropia ; and, accordingly the relation of these fibres with the secret- 
ing cells must be different from that of the fibres of the sympathetics, 
Heidenhain finds that the paralysis of the chorda tympani by atropia 
may be removed by physostigma. The latter substance has, besides, 
a special action on the gland ; for if it be given to an animal after one 
of the chorda tympani nerves has been divided, there occurs a copious 
secretion on the side of the uncut nerve, and but little on the side 
where the nerve has been cut, shewing that the action is exerted on 
the central portion of these nerves. Nicotia and digatalin influence 
the secreting apparatus in the same way as physostigma; but large 
doses of the former substance paralyse the chorda tympani. 


CHANGES UNDERGONE BY THE SECRETIONS UNDER THE INFLUENCE OF 
CERTAIN AGENTS THAT MODIFY THE BLoop-GLoBuLE.—In a thesis of 
great ability for the doctorate in Sciences of Paris (Des modifications 
que subissent les sécrétions sous Vinfluence de quelques agents qui modi- 
jient le globule sanguin, 1872) Mr Ritter, of Strasbourg, describes 
some important investigations made with oxygen, nitrous oxide, 
carbonic oxide, compounds of antimony and arsenic, phosphorus, and 
biliary acids. He specially examined the influence exerted by these 
substances upon the blood, urine, and general nutrition of the body. 
It is impossible to mention more than a few of the very interesting 
results he has obtained. Oxygen, inhaled in quantities of 25 or 30 
litres, was found to augment the acidity of the urine, to diminish the 
nitrates, urea, and uric acid, and to considerably increase the ammo- 


200 DR FRASER. REPORT ON THE PROGRESS OF PHYSIOLOGY. 


niacal salts. The urine voided after inhalation of oxygen, presented 
the singular.character of very soon undergoing acid fermentation, and_ 
of remaining acid for a long time. Ritter states that only after two 
months does it become alkaline. Nitrous oxide, apparently a some- 
what active poison for rabbits, pigeons, and frogs, always diminishes 
oxidation; and it cannot support life when mixed with an equal 
quantity of air. A saturated watery solution of this gas augments 
all the constituents of the urine—it produces, in short, true diuresis, 
Carbonic oxide likewise exerts a very decided influence on the urine, 
for it diminishes all the constituents of that fluid which are looked 
upon as the results of oxidation, and renders it albuminous. When 
tartar emetic, sulphide of antimony, arsenious acid or phosphorus is 
given to man, Ritter finds that large or toxic doses produce a more 
or less decided alteration of the blood, while small doses have a less 
energetic action. The blood-globule is deformed, and at the same 
time crystals of hemoglobin make their appearance; the blood be- 
comes anemic, the albumen and globules diminish, the fibrin in- 
creases and the proportion of gas lessens; generally, the glucose 
augments; and the fatty bodies, including cholesterine, are always 
increased, These substances also modify the composition of the 
urine, They diminish the total quantity of nitrogen and urea; 
lessen the acidity and sometimes even cause it to become alkaline; 
and always augment the uric acid. Where the blood-globule is 
decidedly altered, and especially where crystals of hemoglobin have 
appeared, the urine contains various abnormal constituents, and 
especially bile, albumen, and sometimes hemoglobin. Finally, under 
the influence of these substances the formation and the deposit of 
fat is exaggerated, but only when they are given in moderate doses, 


Hournal of Anatomp and Phpstology. 


ON THE ACTION OF INORGANIC SUBSTANCES 
WHEN INTRODUCED DIRECTLY INTO THE 
BLOOD. By James BuakE, M.D., F.R.CS., San Fran- 


cisco, California. 


In the present paper I propose to bring forward some experi- 
ments showing the action of the salts of Soda, Lithia, Czesium, 
Rubidium, Thalium, and Silver, when introduced directly into 
the blood. For the manner in which the experiments were 
performed, I would refer to my communication in the May 
Number of this Jowrnal, 1870. 

/ Exp. 1. A dog, weight about 18lbs. Pressure in the 
arteries 6 to 7 in. 15 grs. of nitrate of soda, in half an ounce 
of warm water, was injected into the jugular. In 7” the pres- 
sure in the arteries diminished 2 in., but in a few seconds re- 
gained its former level. Respiratory movements rather deeper. 
After 10’ the animal seemed about the same as before the in- 
jection. 10’, pressure 6 to 7 in.: inject 30 grs.; 8”, pressure in 
the arteries rapidly falling ; respiratory movements almost sus- 
pended. 30”, pressure 2in., although the heart is beating 
strongly. 45”, pressure in the arteries increasing. 1' 15”, 
pressure 6 to 7in., respiration again becoming regular, and 
the animal apparently not much affected. 10’, pressure 5 or 
6in.; inject 40grs. In 10’, rapid diminution of pressure. 
30”, pressure 1in., violent spasm with opisthotonos. 35”, spasm 
gradually relaxed, but no respiratory movement for more than 
two minutes ; respiration was again established, and the animal 
became conscious. 2' 30”, pressure 4in., respiration very quick. 


VOL. VII. 14 


202 DR BLAKE. 


3', frothy fluid escaping from the air-passages. 5’, animal died. 
On opening the thorax the heart was found beating, and re- 
tained its irritability for some minutes: the right cavities 
much distended with blood: left cavities contained a smaller 
quantity of partially arterialized blood. The lungs were cede- 
matous and red; and all the air-passages were filled with a 
frothy fluid. 

Injected in larger doses into the veins the salts of soda 
completely and suddenly stop the passage of the blood through 
the lungs, causing violent opisthotonos from accumulation of 
blood in the venous system, and thus producing immediate 
death’. 

Exp. 2. Dog, weight 30]bs. Pressure 5 to 54in. Inject 
60 grs. of nitrate of soda into axillary artery. Immediate ex- 
pression of pain. 10”, general-spasm with opisthotonos. 16” 
after the injeetion, pressure 14in., action of the heart unaf- 
fected; on the spasm relaxing, the animal was dead: pressure 
falling slowly; heart’s action regular. 3’, pressure 6in. 5’, 
pressure 3in., heart’s action quick and weak. On opening the 
thorax, the heart was found beating rhythmically; both cavities 
full of dark blood ; lungs slightly ceedematous ; blood coagulated. 
The nitrate is the most poisonous of the salts of soda with 
which I have experimented. Much larger quantities of the 
sulphate or the carbonate can be injected without producing 
death. 


Salts of Lithia. 


Exp. 3. A rabbit; weight about 3$lbs. Pressure 4:2 to 
4°5in. Inject 3grs. of sulphate of lithia into the jugular; no 
effect. 5’, inject 3grs.; no effect. 4’, inject 5grs. 4”, pressure 
falling. 10”, pressure 2}in., respiration quickened. 30”, pres- 
sure 4in., inject 7grs. 10”, pressure 2in., animal not suffering ; 
inject Sgrs. 4”. pressure falling. 10”, pressure 1}in. 457, 
pressure 34in., heart beating regularly, respiration quickened. 
About 4” after the last three injections there is a well marked 
single jerking inspiration; pressure 3}in., inject 12grs. 4", 


1 There can be no doubt but that this opisthotonos is the result of the 
increased pressure of the blood in the venous system, forasmuch as it is pre- 
vented or relieved by dividing the jugular. Direct experiment shows that on the 
arrest of the pulmonary circulation the pressure in the venous system becomes 
equal to an inch and a half of mercury. 


ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 203 


violent struggles. 10”, opisthotonos, which relaxed in about 1’; 
animal dead; pressure not observed. On opening the thorax, 
the heart was found beating rhythmically ; right cavities dis- 
tended with blood not very dark; left cavities contained small 
quantity of blood only partially arterialized; lungs red and 
cedematous; frothy fluid in the bronchial tubes. 

Exp. 4. <A rabbit; pressure 4 to 4°2in. Inject 7egrs. of 
sulphate of lithia in 31] water into carotid. 3”, pressure rising, 
expression of pain. 10”, pressure Gin. 20”, pressure 5in. The 
pressure gradually fell to 4in., respiratory movements quick- 
ened, no marked effect. 7’, pressure 4°3 to 45in., inject 6grs. 
3”, pressure rising. 10”, pressure Gin. 30”, pressure 4.8 to 5in. 
5’, pressure 5 to 5:2in., inject 7grs. 5”, increase in pressure. 
10", pressure Gin.; general spasm; heart’s action slow and 
irregular. 1’, pressure 4in. 1°30’, pressure 2in.; respiration 
very rapid; conjunctiva sensible, but no expression of pain or 
reflex movements on pinching the tail. 5’, pressure 2 to 24in., 
inject 8ers. 10”, pressure 3 to 34in.; general spasm; respira- 
tion arrested; animal dead. On opening the thorax, both 
cavities of the heart and the veins and arteries were full of 
blood, the blood in the left cavities and arteries partially 
arterialized. On teking out the blood, heart contracted for 
some minutes. Lungs cedematous; frothy secretion in the 
bronchial tubes and trachez. Blood coagulated firmly; the 
corpuscles much crenated; molecular movements well marked 
for many hours after death. 


Salts of Thalium. 


Exp. 5. Rabbit: pressure 54 to Gin. Inject lfegrs. of 
sulphate of Thalium, in 3i. water into the jugular. 5”, pressure 
diminishing. 12”, pressure 2in. 30”, pressure 64in., heart’s 
action quicker; respiration accelerated; animal appears not 
to suffer. 5’, pressure 54in.; inject 2ers. 6”, pressure falling. 
12”, pressure 2in.; respiration irregular, intermittent for 2° 30”, 
when the animal died, the pressure gradually falling. On 
opening the thorax the heart was found beating rhythmically: 
right cavities much distended with dark blood; left cavities 
contained a small quantity of bright scarlet blood. 


14—2 


204 DR BLAKE. 


Exp. 6. Rabbit: pressure 4°8 to 5in. Inject 1gr. sulphate 
of thalium into the carotid; no effect. Inject 2grs. 7’, pressure 
falling. 10”, 34 to 4in. 30", pressure rising. 40”, pressure 
Gin. 2’, pressure 5in.; animal apparently not affected. 5’, 
pressure 5 to 5hin., inject 3egrs.; breathing quickened. 12”, 
pressure falling. 25”, pressure 3 to 3}in. 2’, pressure 5in.; 
respiration natural. 7’, pressure 5in., inject 4$grs. 3”, respi- 
ration quickened, slight struggle. 10”, pressure falling. 20”, 
pressure 24 to 3in. 45”, pressure 4in.; respiration natural ; 
animal apparently not affected. 5’, pressure 44 to 5in., inject 
5ers.; respiration immediately quickened. 7”, pressure falling. 
20”, pressure 2in., heart beating regularly. 40”, pressure 
rising. 1’ 30”, pressure 3 to 3:2in.; animal sensible, apparently 
not much affected. 6’, pressure 4in., inject 12ers.; the im- 
mediate effect on the circulation could not be ascertained, 
owing to the struggles of the animal. 30”, pressure 2in. 45”, 
pressure 4in. 1’ 30’, respiration stopped; pressure 3in. On 
opening the thorax, the heart was found beating rhythmically ; 
right cavities distended; left cavities contained small quafi- 
tities of arterialized blood ; lungs red, but contracted. 


Salts of Rubidium. 


Exp. 7. Rabbit: pressure 5in. Inject 2grs. of chloride 
of rubidium into the jugular. 5”, pressure falling. 10”, pres- 
sure 1din. 1’ 30", pressure 4°8 to 5in.; the respiration quick- 
ened. 8’, pressure 5in. Inject 2ers. 4’, pressure falling. 
15", pressure 2}in.; spasm; respiration stopped one minute 
after the injection. On opening the thorax, heart contracting 
rhythmically: right cavities distended with dark blood: left 
contained a small quantity of scarlet blood. Lungs scarlet. 
Blood coagulated. 

Exp. 8. Rabbit: pressure 44 to 5in. Inject 2grs. of chlo- 
ride of rubidium into the left carotid. 7”, pressure less. 
15”, pressure 2}in.; slight spasm. 25”, pressure rising. 1’, 
pressure 4}in.; heart’s action regular; respiration quickened. 
5’, pressure 4in.; inject 8grs. 7”, pressure less. 12”, pressure 
2hin. 45”, pressure 4in.; heart’s action regular, but oscillations 
slight. Owing to an accident the hemodynamometer could no 


ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 205 


longer be used. 7’, inject Ggrs. 10”, tonie spasm; respiration 
suspended for 15”, then irregular for 2’; sensibility apparently 
unimpaired. 3' respiration regular, but very rapid. 5’, inject 
10grs. 10”, respiration suspended. 45”, respiration com- 
menced and continued irregularly for about 1’; animal then 
died. On opening the thorax the heart was beating rhythmi- 
cally; right .cavities much distended; left contained a small 


quantity of scarlet blood. 


Salts of Casium. 


Exp. 9. Rabbit: pressure 4 to 4:2in. Inject I}gr. of 
chloride of cesium into the jugular. 5”, pressure falling 
slightly. 15’, pressure 34 to 4in.; respiration quicker. 5’, 
pressure 3 in., inject 38grs. 5”, pressure falling. 7”, respi- 
ration quickened. 10”, pressure 2in. 45”, pressure 3in.; 
respiration more natural; animal sensible. 5’, pressure 33in. 
Inject 4¢grs. 5”, pressure less; 7”, general spasm; 10”, pres- 
sure lin., animal dead. On opening the thorax the heart was 
found contracting rhythmically, and continued contracting 12' 
after death: right cavities distended: left cavities contained a 
small quantity of scarlet blood. 

Exp. 10. Rabbit: pressure 4 to 4}in. Inject 4grs. of 
chloride of cesium into the carotid. 3”, general spasm; re- 
spiration suspended. 35”, respiration renewed, regular but 
quick ; posterior extremities and trunk stiff, as in rigor mortis. 
' 1’, cornea insensible. 4’, respiration natural; lids close about 
2" after irritating cornea ; animal lies quite still, limbs stretched 
out as in a state of rigor, no sign of consciousness, but reflex 
movements on pinching the tail. 10’, signs of returning con- 
sciousness, respiration regular, 15 in a minute. 12’, inject 
4c7s., no marked effect: respiration 15 in a minute, principally 
diaphragmatic. 5’, inject 5grs. 3”, violent spasm lasting about 
10”, animal dead. On opening the thorax the heart was beat- 
ing rhythmically, and continued contracting for 10°: both cavi- 
ties contained blood; that in the left rather brighter than in 
the right but still dark. (Note in consequence of the hamody- 
namometer becoming obstructed, the pressure could not be 


observed.) 


20G DR BLAKE. 


Salts of Silver. 


Exp. 11. Dog: weight 15lbs. Pressure 5 or 6in. Inject 
3ers. of nitrate of silver into the jugular. 
30”, pressure din. 40”, pressure rising. 1' 20", pressure 8 to 
Qin.; respiration quicker. 2’, pressure 3in. 5’, pressure 24in.; 
heart’s action slower, 58; breathing rapid, 48. 7% frothy fluid 
escaping from the mouth; pressure 2in. 12’, respiration stopped ; 
the heart still felt beating through the chest-walls, although 
causing no oscillation in the mercury. On opening the thorax 
the heart was contracting rhythmically, and continued contract- 
ing for 10’; the right auricle contracted spontaneously half an 
hour after the thorax had been opened. The blood both in the 
right and left cavities was dark and viscid; it did not coagu- 
late. The lungs were red, partially hepatised ; the air-passages 
were full of a frothy secretion. 

Exp. 12. Dog: weight 15lbs.; pressure 6 to 64in. Inject 
ler. of acetate of silver into the axillary artery. 4’, pressure 
rising. 8”, pressure 12in. 20”, pressure 10in. 1’, pressure 
Yin. 3’, pressure 74 to 8in.; respiration quickened. 5’, pres- 
sure 7 to 74in., inject 2er. The immediate effect on the pres- 
sure was not observed, owing to the tubes being obstructed. 
General spasm and arrest of respiration. At 1’ to 1'30”, pres- 
sure 10in.; heart slow, 35 in a minute. 3’, respiration again 
commenced, lasted irregularly for two minutes, and then stop- 
ped: animal dead, five minutes after the injection pressure in 
the arteries gradually fell. On opening the thorax the heart 
was beating rhythmically; its contractions continued for 8’; 
right cavities distended ; left cavities moderately full of dark 
blood which coagulated very imperfectly. Lungs hepatized, 
red, and some spots of ecchymoses: frothy secretion in bronchial 
tubes: twitchings in the voluntary muscles 8 minutes after 
death. 

The substances, the action of which has been described in 
the above experiments, form an interesting group, possessing, as 
they all do, certain isomorphous relations, and yet not so closely 
agreeing in all their chemico-molecular properties as some other 
isomorphous groups. An analogous remark will apply to their 
physiological reactions. When introduced directly into the 


at 


7’, pressure less. 
0" 


ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 207 


blood they all have certain well marked resemblances ; but still 
there are differences which serve to distinguish some of them 
in their reactions on the different organs. Amongst the most 
marked reactions they produce is the manner in which they 
act on the lungs. Without exception they are all lung-poisons. 
They all kill when injected into the veins, either by directly 
arresting the pulmonary circulation, or by causing changes in 
the lung-tissue which rapidly prevent the aération of the 
blood. They also all agree in another important respect. Some 
of them, notwithstanding the marked changes they cause in the 
blood, do not at all diminish the irritability of the heart. In fact 
I believe they all increase the irritability of the heart muscles ; 
so that these continue to contract longer without the stimulus 
of arterialized blood. There is another property which they 
all seem to possess, with the exception of cesium ; and that is, 
that they exert no direct effect on the functions of the nervous 
system. In striking contrast to their action in arresting the 
passage of the blood through the pulmonary capillaries is the 
facility with which they all, with the exception of silver, pass 
through the systemic capillaries, unless introduced mto the 
arteries in large quantities. Nothing can more strikingly show 
the want of action of these substances on the nervous system, 
and on the systemic circulation, than the fact that they can be 
introduced in much larger quantities into the arteries than into 
the veins without proving fatal. With the salts of thalium, 
for example, three and a half grains, when injected into a vein, 
will instantly kill by arresting the pulmonary circulation ; 
whilst six times the quantity can be injected into the arteries 
without producing any marked symptoms. They pass over the 
heart through the nervous tissue, and find their way through 
the systemic capillaries without apparently causing any marked 
change im these organs; and it is only when injected into the 
arteries, in quantities large enough for them to reach the lungs 
in a sufficiently concentrated state there to give rise to these 
well marked effects, that they kill. These properties are found 
in all the more strictly isomorphous members of the group. 
The only exceptions met with are in the salts of cesium and 
silver; the latter evidently an outlying member of the group. 
The former of these salts exerts a decided action on the ner- 


208 DR BLAKE. 


vous system when introduced into the arteries, and a small 
quantity of a salt of silver, in the arterial blood, causes ob- 
struction to its passage through the systemic capillaries. 

As to the mechanism of the arrest of the pulmonary cir- 
culation, two explanations are admissible. The pressure of 
these substances in the blood gives rise, either directly, or through 
reflex action, to contraction of the smaller pulmonary arteries > 
or they arrest the passage of the blood through the capillaries 
by the changes they make in the shape of the blood corpuscles. 
I have found that all these substances, so far as examined, pro- 
duce changes in the shape of the blood corpuscles. The fol- 
lowing appearances were observed in a microscopical exami- 
nation of the blood six hours after death, from a rabbit that had 
been killed by caesium (see Exp. 9). Blood from right side of 
the heart coagulated, but not much contracted : blood from left 
side perfectly fluid. In the venous blood not a natural coloured 
corpuscle was found. About one-third of the corpuscles were 
colourless, and adhered to the slide. The others had all lost 
their natural discoid shape. They were thickened, and nearly 
all crenated generally with six indentations. Where they still 
retained a circular outline, the border seemed thickened; as 
they rolled over, they were all found to be curved: the colouring 
matter was collected in points or around the thickened border. 
In the blood from the left cavities not a crenated corpuscle 
could be found. They all retained their circular outline, but 
were contracted and thickened. The size of the crenated cor- 
puscles in the venous blood was 5°5™™"", of the round corpuscles 
67™™". In this instance it would seem as if the lungs had 
acted as a filter by which the roughened corpuscles were re- 
tained; as there must have been many roughened corpuscles 
in the blood for some time before the last injection. This is 
the most marked example I have met with of the absence of 
roughened corpuscles in the blood from the left cavities of the 
heart. But they are always in a smaller proportion than in 
tlie venous blood. In none of the specimens of blood examined 
after the injection of any one of these salts have I seen the 
corpuscles agglutinated in rouleaux. The molecular movements 
are very active many hours after death. Should this mechani- 
cal explanation of the arrest of the pulmonary circulation be 


ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 209 


correct, we are still to consider how it is that many of these 
substances when injected into the arteries find their way through 
the systemic capillaries without causing obstruction, unless in 
large doses. This point however, together with a more detailed 
account of the changes caused in the lung-tissue, J must leave 
for a future communication. 


ERRATA IN PRECEDING PAPER, Vou. VI. 


Page 95, line 13, from top, for 2 or 2°2 inches, read 2 to-2°2 inches. 

Line 9 from bottom, for respiration stopped; in 40” hearts—read respiration 
stopped in 40”; hearts, 

Page 96, lines 3 and 4, for down to 3:5, respiration suspended; for 30” 
animal sensible—read down to 3:5; respiration suspended for 30”; animal 
sensible, 

Line 6, for arrested; at 2’, read arrested at 2’; 

Line 8, for suspended; 1°30”—+read suspended 1':30"; 

Page 97, line 9, from bottom, for quick; but regular respiration—read quick 
but regular; respiration. 

Page 100, line 4, for overdisturbed state, read overdistended state. 


ON THE PHYSICAL NATURE OF THE COAGULATION 
OF THE BLOOD.—By ALFrep HUTCHISON SMEE, 
F.CS., F.SS. 


Read at a Meeting of the Royal Society. 


THE cause of the coagulation of blood fibrine has long been a vexed 
question among physiologists. In bringing this subject under the 
notice of the Royal Society it is my intention, first, briefly to review 
the various theories which have been held at different times, and, 
then, to state those views on coagulation which have been enforced 
upon my mind by direct experiment, and also by the behaviour of 
colloidal substances analogons to fibrine. 

Hunter, the great upholder of vital force, thought that coagulation 
of the blood was an act of life, and was analogous, to some extent, to 
the contraction of a muscular fibre. Hewson, a contemporary, in 
opposition to Hunter’s views, noticed that blood could be kept fluid 
for months by addition of certain neutral salts. This experiment has 
been urged as conclusive evidence against Hunter’s theory of coagu- 
lation; for it is impossible to conceive that vital power could last for 
such a length of time. Gulliver found that blood remained fluid for 
one year on the addition of nitre, yet still retained its power of 
coagulation on the addition of water. 

Coagulation was held by other physiologists to depend upon the 
stasis of the blood in the vessels; and they poinied to the coagulum 
at the point of Lgature in an artery which had been tied as the result 
of the stasis. 

Again, exposure to air was stated to be necessary for the for- 
mation of a coagulum; and it was supposed that something, pro- 
bably of a gaseous nature, was given off from the blood at the time of 
coagulation. 

The evolution of carbonic acid from the blood was considered to 
be the cause. Dr Richardson considered that fibrine was held in 
solution by free ammonia which escaped from the blood on exposure, 
when the fibrine began to coagulate. 

In an experiment which I related in my paper on the artificial 
formation of fibrine from albumen, I demonstrated that. ammonia, 
in quantities greater than could ever be found in the blood of the 
living body, might be added to albumen; nevertheless when this 
albumen was submitted to the action of oxygen at temperature of 98° 
fibrine formed, although it was afterwards slowly dissolved. 

Lester pointed out in one of his experiments, that, on carefully 
neutralizing the blood with acetic acid no alteration was made in the 
power of its coagulation. Coagulation however was observed to take 
place more rapidly in vacuo, giving some justification to the views of 


PHYSICAL NATURE OF THE COAGULATION OF THE BLOOD. 211 


those who hold to the theory that the blood exhales from itself some 
volatile principle at the time of coagulation. 

Heat likewise is said to favour, and cold just above the freezing 
point to retard, coagulation. 

In asphixiated animals coagulation was found to be retarded. 
This fact was regarded as evidence that carbonic acid kept the blood 
fluid. 

Astley Cooper and Thacknot thought that the blood-vessels, 
exerted a specific influence in actively preventing the coagulation of 
the fibrine in the blood. Briicke supported this view. He found 
that the blood of a turtle injected into an empty heart remained fluid 
for many hours, whilst some of the same blood exposed in an open 
vessel coagulated in a few minutes. Lester found that, on inserting 
a tube into the circulating system, fibrine, in a short time, separated 
from the blood, the coagulum coating the internal surface of the tube ; 
but that, after a time, fibrine ceased to separate from the blood. 
This was a case of blood coagulating in a sealed vessel and disposed, to 
my mind, of the exhalation theory of coagulation. Morrant Baker, 
in the Ist Vol. of the St Bartholomew Hospital Reports, remarks that 
the blood, in blood tumours, remained fluid, and that the fibrine in 
most cases had separated from the liquor sanguinis. He further 
remarked that the internal surface of the tumour was coated with a 
coagulum which had apparently undergone organization, and was, in 
tumours of old duration, hard, and like fibrous tissue. The specific 
gravity of this blood had fallen from 1052 (normal blood) to 1020, 
Sometimes, however, a portion of blood coagulated when the tumour 
was opened. He thinks this is due to bleeding from the small vessels 
which have been punctured by the incision, the fresh blood mixing 
with the blood of the tumour, and a second coagulation takes place. 
I think this phenomena may be explained by Lester’s tube experi- 
ment, where he shews that when blood is first effused, or passed into 
or through a tube which had not previously been coated with blood 
fibrine, the fibrine separates into a coagulum, this coagulum then 
undergoes organization into fibrous tissue; but, if on a second heemor- 
rhage taking place into a blood tumour, or if more blood is passed 
through the fibrine coated tube, the fibrine will not separate from the 
liquor sanguinis. When a needle or wire is inserted into a blood- 
vessel whilst the blood is circulating it acts as a foreign body, and it 
soon gets coated with a fibrinous deposit: upon this principle is 
founded a treatment for the cure of aneurism. 

Lester has demonstrated that the blood will remain fluid in the 
jugular vein from 24 to 48 hours after death, provided the vein has 
been tied just before or just after death, but when the blood is turned 
out coagulation immediately takes place. Buchanan has shewn that 
the fluid from a hydrocele yielded a coagulum on the addition of blood 
serum although each fluid, without the addition of the other, might 
be kept separate for any length of time without coagulation taking 
place. Schmidt also held the same view, but considered that it 
required a fibro-plastic substance of the nature of globulin to combine 
with another substance which he termed fibrogen. 


212 MR SMEE. 


Lionel Beale has studied the phenomenon of coagulation under a 
magnifying power of upwards of 2000 diameters (33 inch). The first 
change he noticed was a film-like appearance in the liquor sanguinis, 
especially in the track of the red corpuscles as they slowly traversed 
the field. This film-like appearance was succeeded by delicate threads 
apparently corresponding with the track of the blood-cells, these lines 
gradually increased in density and refractive power. 

He thinks that this coagulable matter exists, in the first instance, 
as a diffused plasma probably formed from the white cells which, 
gradually separating from the serum, contracts, acquires density, and 
thus becomes visible under the microscope. 

During coagulation the red cells become stellate, refract more 
highly, lose diameter and fluid; at which time probably globuline 
escapes. 

Lastly, when blood is stirred up with twigs coagulation proceeds 
more rapidly. 


Having thus briefly reviewed the principal theories held by 
physiologists, I now venture to submit that view which, in 
my opinion, best accords with all observed facts. 

From a careful review of all the circumstances of the case 
we may fairly consider that the coagulation of the blood takes 
place in obedience to a purely physical Jaw, namely, the power 
of soluble colloid matter, whether organic or inorganic, to pec- 
tize, or in other words, spontaneously to coagulate. In order 
that I may illustrate this view of the coagulation of blood, I 
must ask the Fellows of the Society to travel with me into the 
paths of inorganic chemistry, especially calling attention to 
Graham’s experiments on colloid matter published in his paper 
“on liquid diffusion applied to analysis.” 


The act of pectization of a colloid body may be regarded as the 
equivalent of the act of crystallization of a crystalloid body. 

Take for instance a solution supersaturated with sulphate of soda, 
it will remain fluid for days. Stir it, or even drop a particle of dust 
into the fluid, it will instantly begin to crystallize. 

In this case we observe a perfect analogy between the action of a 
particle of dust which determines the act of crystallization in this 
solution and that of the wires or twigs which, when applied to blood, 
produce a rapid formation of a coagulum. No one can say that this 
crystallization of the sulphate of soda was an act of vitality. 

Graham has shewn that the essential characters of all colloids are 
to form a jelly and not to dialyse. This jelly he regards as the pec- 
tous or insoluble state; whilst the soluble state of a colloid he regards 
as the peptous. An inorganic colloid in the pectous condition is a 
vitreous mass, homogencous and perfectly structureless, in which state 


PHYSICAL NATURE OF THE COAGULATION OF THE BLOOD. 213 


it apparently remains an indefinite time, gradually losing water and 
becoming more and more dense, and probably after the lapse of years 
it is capable of undergoing transformation into a more or less crystal- 
line state. In nature the structureless flint may be found gradually 
being converted into the crystalline. When an organic colloid 
assumes the pectous condition it contracts and crushes up into fibres. 
This is the case with fibrine, albumen and gluten. 

Graham has also shewn that all organic colloid substances have 
high chemical equivalents, and are at the same time chemically inert 
in the ordinary sense, but possess a compensating activity from other 
physical properties. Crystalloid bodies appear to shut out external 
impressions, whilst colloids possess properties, to use Graham’s own 
words, which enable them to become the medium of diffusion like 
water itself. Another characteristic quality of colloids is their con- 
stant mutability. Their existence is a continual change. A fluid 
colloid may assume a pectous modification, and often passes, without 
any visible external influence, or even of internal change, from the 
first (the fluid) to the second (the pectous) condition. Colloidal sub- 
stances, such as gelatine, which gelatinize, but still retain their power, 
again become fluid by heat and are soluble in water, cannot be re- 
garded as assuming the pectous condition, but the gelatinous. 

Graham has demonstrated that when a soluble colloid has assumed 
the pectous or coagulated condition it cannot again per se become 
fluid. In the case of certain colloids, however (silicic acid, for instance), 
the addition of a small quantity of an alkali to the jelly causes it to 
again liquefy. If the fluid containing silicic acid plus the alkali is 
placed upon a dialyser, the alkaline salt is removed, leaving the 
silicic acid in a pure and soluble state, in which condition it will 
remain for some time, when it will again assume the pectous state. The 
addition of the alkali, however, appears to have had no chemical 
action on the silicic acid jelly, but only changed its physical condition 
from the pectous to the peptous or fluid state. On the other hand, 
the act of gelatinization may be repeated ad infinitum, as the gelatine 
- may be melted over and over again. 

I must now direct attention to Graham’s experiments on the 
behaviour of hydrated silicic acid, and to the analogous physical 
behaviour of film. Graham has shewn that hydrated silicic acid, in 
a state of great purity, can be obtained and held in solution, but 
cannot be preserved in that state for any length of time. It will 
remain fluid for days, and even for a longer period, if it is in a sealed 
tube; but it will ultimately spontaneously coagulate and become 
insoluble. A concentrated (14 per cent.) solution assumes the pectous 
condition in a few hours. A 5 per cent. solution may be kept for 
some days. A 2 per cent. solution will keep two or three months; 
and a 1 per cent. will remain fluid even after two years. | 

The addition of solid matter in the state of powder to liquid 
silicic acid greatly favours the act of pectization, the solid matters 
apparently acting as a nucleus, like the dust in sulphate of soda in 
solution, and set up crystallization. 

The addition of one 10,000th part of an alkaline carbonate to fluid 


214 MR SMEE. 


silicic acid causes it to immediately pectize. The silicates of the 
alkalies are themselves soluble in water; but no one would assert 
that the addition of an alkaline carbonate could have any other effect 
on silicic acid than that of a foreign body setting up a new physical 
condition, for no chemical action can possibly have taken place during 
the transition of the silicic acid from the fluid to the pectous state. 
Acids and other neutral salts likewise cause coagulation. Caustic 
ammonia, however, has no action on fluid silicic acid. Alumina like- 
wise has the power of existing in the fluid and pectous state without 
the intervention of an acid; but soluable alumina is one of the most 
difficult substances to prepare owing to its unstable nature in the fluid 
state. A vessel washed out with ordinary water (which contains a 
trace of sulphate of potash) is sufficient to cause it at once to coagu- 
late. The addition of sulphate of potash, in the one case, or of an 
alkaline carbonate, in the other, cannot be said to have effected any 
chemical change in these colloids to cause them to pectize. I believe 
that neutral salts added to the fluid silicic acid act in a similar 
manner to the handful of twigs used for stirring up blood, as they 
act simply as foreign bodies. 

Soluble peroxide of iron is more interesting to the physiologist 
than either soluble silicic or albumina, on account of iron being one 
of the constituents of the blood; and it is more than probable that 
the iron in the red blood corpuscles is in a fluid state. Graham re- 
marks that soluble peroxide of iron remains fluid for 20 days, provided 
it is in a weak solution and kept in sealed glass tubes, when it will 
suddenly spontaneously pectize without any apparent cause. Water 
containing 1 per cent. of hydrated peroxide of iron has the deep red 
colour of venous blood. This solution can be concentrated to a certain 
point, when it will suddenly pectize. The coagulum is a deep red 
coloured jelly resembling blood-clot. Graham remarks that the feeble 
circumstances which suffice to produce this change is highly sugges- 
tive of blood. 


The following experiments which I have made upon fresh- 
drawn blood illustrate the manner in which fibrine pectizes in 
animal fluids. These experiments clearly demonstrate that the 
act of coagulation takes place in accordance with purely physi- 
cal law. ‘ 

On the addition of an equal quantity of a solution of sul- 
phate of soda to fresh-drawn blood coagulation will not take 
place, and the blood will remain fluid an indefinite time. The 
blood-cells however will gradually subside, leaving the liquor 
sanguinis containing the uncoagulated fibrine bright and clear. 
If some of this liquor sanguinis is placed in a dyalyser and then 
the dyalyser placed in distilled water, in the course of a few 
hours the sulphate of soda will dyalyse out of the liquor san- 


PHYSICAL NATURE OF THE COAGULATION OF THE BLOOD. 215 


guinis, and a thin gelatinous film of fibrine will be formed at 
first, in direct contact with the parchment paper of the dyalyser. 
This film of fibrine will gradually extend and become thicker 
as the sulphate of soda slowly dyalyses out of the solution; and 
after the lapse of 10 or 12 hours the whole will have become 
one uniform structureless clot. This clot will, after some 
days, contract, squeeze out the liquor sanguinis, lose its struc- 
tureless appearance, and crush up into fibres. 

If-a second quantity of the liquor sanguinis is placed on the 
dyalyser which has been previously coated with fibrine jelly, 
the second quantity will require a greater length of time for 
coagulation to begin. This is especially the case if a cup-shaped 
depression has been made in the fibrine jelly for the second 
quantity of the liquor sanguinis. When the solution of sulphate 
of soda is added to the liquor sanguinis in excess, the time of 
coagulation is delayed in the direct ratio to the quantity of the 
sulphate of soda added to fresh-drawn blood. 

If the fluid which is placed upon the dyalyser contains not 
more than one part of the liquor sanguinis to 30 pints of the 
sulphate solution no pectization will take place, even after the 
sulphate has been removed by long-continued dyalysing; and 
probably the fibrine remains fluid upon the dyalyser sufficiently 
long to get oxydized, and converted into some other substance. 
Upon this point I purpose making further investigation. The 
fibrine jelly formed by dyalysing is dissolved slowly in caustic 
potash, leaving behind a small quantity of hexagonal crystals 
_ which are soluble in acetic acid. I have placed fibrine jelly 
which has been redissolved in potash upon a dyalyser to remove 
the alkaline salt, but I have failed in every instance to get the 
film a second time to pectize. 

I have observed a very curious property in fibrine jelly 
(which has been made from a dilute solution) of breaking up and 
becoming again fluid, and passing through the blotting paper 
filter used to separate the liquor sanguinis from the fibrine jelly. 
I can find no satisfactory explanation of this remarkable change, 
unless we accept the view that very minute causes are sufficient 
to determine the physical condition of colloid substances. 

Dr Goodman has noticed that albumen suspended in a 
ropy condition in cold water, coagulated after some time, 


216 MR SMEEF. 


became white, dense, insoluble, and finally fibrous. This change 
he regarded as evidence of the formation of fibrine from 
albumen. I have frequently repeated the experiment. I be- 
lieve that no new substance is formed out of the albumen by 
this method, which I believe has only changed its physical 
condition from having its salts removed by dyalysis, which 
causes the albumen to change from the peptous to the pectous 
state. 

Having brought under the notice of the Society some of the 
principal characteristics of colloid, I will now proceed to a 
comparison between the behaviour, physically, of the inorganic 
colloid silica with that of the organic fibrine. 

In every essential point it will at once be recognized that 
these two dissimilar substances agree, and, at the same time, 
that there is no circumstance during the act of coagulation 
which cannot be explained by physical law. 

The analogy between fibrine and silica in their physical 
behaviour will be best observed by comparing their properties 
together. Ist. Fibrine and silica colloidal substances are known 
to exist in the fluid as well asin the coagulated condition. 2nd. 
When either fibrine or silicic acid assumes the pectous state it 
is incapable of being per se redissolved, so as again to be able to 
spontaneously pectize. The existence of both these substances 
is a continual metastases: fibrine is a typical instance of this 
metastases. 38rd. All colloids in the fluid condition, whether of 
organic or inorganic origin, after an interval of time, longer or 
shorter according to their specific characters, spontaneously 
coagulate. 4th. This coagulation takes place without the in- 
tervention of any chemical agent which is capable of producing 
a change in that colloid. 5th. The condition of neutral salts to 
inorganic colloids ina fluid state (just as the falling of a speck of 
dust into the supersaturated solution of sulphate of soda favours 
crystallization) favours the coagulation of those colloids. The 
neutral salts in these cases must be regarded as foreign bodies. 
Possibly the white blood-cells, altered in their physical con- 
dition by exposure to air, become as foreign bodies to the 
blood, and have a similar influence as twigs and rods used ordi- 
narily to defibrinate blood. 

Lastly, the capacity of all colloids to remain in the fluid 


PHYSICAL NATURE OF THE COAGULATION OF THE BLOOD. 217 


condition is greatly promoted. Ist. By the weakness of the 
solution (less than 10 per cent.); 2nd. By being contained in 
sealed vessels. 

The time required by a fluid colloid to pectize apparently 
depends upon its molecular equivalent. 

The fluid condition appears to be less stable in colloids 
with high molecular equivalents, and the act of pectizing takes 
place more rapidly and with less apparent cause, and apparently 
in direct ratio to the molecular equivalent of the colloid. Con- 
sequently soluble peroxide of iron pectizes sooner than alumina, 
and alumina more rapidly than silica. Therefore fibrine, with 
a molecular equivalent vastly higher than either of these colloids, 
might be expected to coagulate almost immediately. 

Occasionally masses of natural silica containing fluid are 
found in nature, of which I myself possess a remarkably fine 
specimen. These appear to me to have a great resemblance 
in their formation to that of the blood-tumours described 
by Morrant Baker. The enclosed fluid may be analogous to 
that in Lester's experiment where fibrine ceased to coagulate 
or separate when passed through a tube which had been pre- 
viously coated with blood-fibrine. 

In the case of the natural stone what has taken place? Ist. 
Silica has been deposited. 2nd. Coagulation has taken place; 
and the coagulum has contracted and formed a cavity in its 
inner surface. 3rd. The addition of more (fluid) silica has taken 
place; and the external coagulum has been transformed, passing, 
after the lapse of time, from the vitreous to the crystallized state. 

The second quantity of the fluid silica has coagulated; but 
it has coagulated in the gelatinous form, squeezing out the 
remaining water. 

This recalls to our minds what takes place in blood-tumours, 
Ist. Blood is effused; 2nd. The fibrine is coagulated; 3rd. 
The coagulum contracts and becomes organized into the dense 
hard fibrous margin. Ifa second effusion of blood now takes 
place the blood will remain fluid, and will remain so for a con- 
siderable length of time; but at last the fibrine will coagulate 
squeezing the liquor sanguinis. 

To my mind the analogy between the formation of the 

VOL. VII. 15 


218 MR SMEE. NATURE OF THE COAGULATION OF THE BLOOD. 


fibrine in a blood-tumour and the deposition of silicic acid in 
certain natural stones is complete. 

The above considerations of the causes of the coagulation of 
blood-fibrine may be briefly summed up. Ist. That the coagu- 
lation of fibrine is a physical act, and cannot be considered to 
be in any way identified with a vital property such as the con- 
traction of muscular fibre. 2nd. The coagulation of fibrine 
depends upon and is regulated by the same laws which cause 
all soluble colloid substances, whether organic or inorganic to 
become pectorous. 38rd. That the soluble or fluid form of fibrine 
ought to be regarded as its allotropic form; and, as in the case 
of its colloidal analogue, silicic acid, its presence in the blood in 
the fluid condition depends upon the physical conditions under 
which fibrine is found in the living body. 


ON THE LAW WHICH REGULATES THE FREQUENCY 
OF THE PULSE. By A. H. Garrop, B.A. Cantab. 


THE paucity of mechanical theories to explain the frequency 
of the pulse, probably arises from the very general assumption, 
that in all cases when the rapidity of the heart’s beat is caused 
to vary, the action of nerves, with special powers of retarding 
or quickening it, is brought into play; and the relation of 
heart power to work to be performed has not been introduced 
into the problem. 

The theory of energy has of late spread so far and wide the 
necessity for finding in all cases where work is done, a sufficient 
source for the production of that work, in one form or other, 
that a vague statement to the effect that heart frequency de- 
pends solely on nerve action, is far from sufficient for the re- 
quirements of physiologists. It is now necessary to shew that 
with different amounts of work to be performed in the cir- 
culation, different supplies of nutrient substance must be pre- 
sented to the motor organ, just as in the steam-engine the 
amount of fuel must be varied according to the work required 
from the machine. 

When the microscope revealed the existence of a well- 
marked muscular coat to the smaller systemic arteries, it be- 
came evident that the different diameters of those vessels, 
consequent on the degrees of contraction of their walls, varied 
the amount of force necessary to propel the blood through 
them; and these variations have been considerably studied of 
late. Dr Marey of Paris, the introducer of the sphygmograph, 
has, in his most scientific treatise On the Circulation of the 
Blood’, strongly drawn attention to this subject, and he has 
worked out a theory respecting the law regulating the fre- 
quency of the pulse, which is based mainly on the variations in 
arterial resistance. 


1 Physiologie Médicale de la Circulation du Sang. Paris, 1863. 
15—2 


220 MR GARROD. 


This theory of Marey’s it will he necessary to recapitulate 
here, and to examine the facts on which it rests. The followmg 
is the law in the two forms in which he gives it. 

1. “The heart beats so much the more frequently, as it 
experiences less difficulty m emptying itself.” 

2. “The frequency of the pulse varies inversely as the 
arterial tension.” 

As reasons for the accuracy of this law are given :— 

Ist. The analogy of other intermittent muscular move- 
ments, as the followmg—A man can walk a certain distance 
quicker, the less he is loaded. Or this—The hand can be 
moved alternately backwards and forwards more quickly in 
air than in the more resisting fluid, water. 

2nd. The pressure can be made to change by variations in 
the amount of blood in circulation, and by modifications in the 
degree of arterial or capillary resistance, both of which vary the 
pulse-rate in the manner required by the theory. 

To prove the effects of different amounts of blood in circu- 
lation, the experiments of Hales are quoted, in which he found 
that loss of blood increased the frequency of the pulse. 

To prove the effects of varied arterial or capillary resistance 
many satisfactory and original results are referred to, among 
them, the effect of compressing the abdominal aorta, or the 
femorals, which retards the pulse; the effects of cold baths, 
according to Drs Bence Jones and Dickinson, when the pulse 
was greatly reduced in frequency ; the quickened pulse follow- 
ing successive additions of warm clothing over the body is also 
proved. 

From these latter results it is clear that Marey assumes that 
by varying the capillary resistance the blood-pressure is also 
varied at the same time, but this assumption is not necessarily 
true in a circulation that is maintained by a pulsating motor 
organ, whose rate is variable, as can be easily shewn by an 
analogy from electricity, which is a useful one im many ways to 
students of the circulation, and is quite worth bemg worked out 
by each. It is this—Suppose a battery connected, through a 
break-and-make key, to a long uniform insulated line or tele- 
graph cable, insulated at the other end, and connected with a 
static galvanometer, 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 221 


First connect the two parts by the key and thereby charge 
the line, and then break connection ; upon this the charge will 
fall in tension slowly, and this’ fall may be observed on the 
galvanometer; when the tension has fallen one half, reconnect 
and break again. It is evident that if this process be repeated 
a definite current is maintained between the cable and the 
surrounding bodies to which it leaks. If the line be now 
halved in length, whereby the resistance is doubled, and again 
instlated at the free end, it is evident that by again breaking 
and making contact as before, when the tension is halved, the 
maximum tension will not be changed. So with the circulation, 
if the resistance in the arterial peripheral vessel is varied and 
the length of the pulsation depends on the time of fall in ten- 
sion only, the pressure does not vary, if the vascular capacity is 
constant. 

It is thus seen that the blood-pressure need not depend on 
the arterial resistance, but if the pressure does not vary, the 
pulse-rate must do so. 

A desire to arrive at the genuine value of this theory of 
Marey’s led me to make experiments similar to his own, as to 
the accuracy of his fundamental facts. My observations were 
divided into two series, to find,— 

1st. Whether the pulse-rate was related to the capillary 
resistance. 

2nd. Whether the pulse-rate depended on the pressure of 
the blood in the arteries. 

These points will be considered separately. 

Ist. The relation of the pulse-rate to the arterial resistance. 

The effect of exposing the surface of the body to the in- 
fluence of different temperatures, whereby, as it has been my 
endeavour to prove elsewhere’, variations in the calibre of the 
cutaneous vessels are produced, was carefully examined, and the 
following tables embody my results, the curves being those of 
changes in pulse-rate. 

Experiment I. Temperature of the air 51°5° F. Nude at 
11°57 P.M. Lay down on floor, carpeted, on right side, at 11°58 
P.M., with head on footstool. Did not feel cold. Got up and 


* Proceedings of the Reyal Society, 1869, p. 419. 


222 MR GARROD, 


Nieut 50 55 60 65 


put on night-shirt and jumped into bed at 12°29; a skin glow 
came on at 12°29. Same position maintained in bed as when 
on carpet. 


Experiment II. Temperature of air 50° F. Nude at 11°40 
P.M. and lay down on right side. Experiment conducted exactly 


as the last. Got up at 12 night, and in bed in less than a 
minute. A glow came on at 12°6. 

Experiment III, Temperature of air 525° F. To shew 
that the change in pulse-rate did not depend on the effort of 


50 55 60 65 


11°10 p.m, 
1115 


11°30 


11°45 


12 Nieut j 


getting into bed. Experiment conducted exactly as the first. 
Nude, lying on rigbt side, at 117 P.M. Got up at 11°26 P.M., 


o 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 223 


put on night shirt, took it off again and lay down again on 
floor. Got up again at 11-45'5” and went into bed, in night- 
shirt. A glow came on at 11°54'5" P.M. 

From these observations it is apparent that the effect of 
simply altering the condition of the cutaneous vessels, by vary- 
ing their relations to external agencies, varies the pulse-rate in 
a definite manner; and thermometric results shew that on 
warming the skin, as by covering it with bad conductors, the 
vessels are increased in calibre and the arterial resistance re- 
duced. These experiments therefore shew that reducing the 
resistance quickens the pulse. 

Marey’s own observations, specially as they are recorded 
mostly by the graphic method, are of themselves sufficiently 
convincing on this point. He compressed the abdominal aorta 
of a horse, per rectum, and found the pulse thereby rendered 
much slower. The same result followed compression of the 
human femoral arteries. 

The quickened pulse produced by the Turkish bath (in one 
case reaching the extreme rapidity of 172 in a minute on my- 
self) is well known; as is the slow one following a cold bath, as 
shewn by Drs Bence Jones and Dickinson. 

From these many facts, all tending in one direction only, it 
may be stated that the rapidity of the pulse varies inversely as 
the resistance to the flow of blood from the arteries. 

2nd. The relation of the pulse-rate to the amount of blood 
ain circulation, or to the blood pressure in the arteries. 

The following experiments were made— 

Experiment IV. An old donkey which had been standing 
for more than half an hour in the room in which the experi- 
ment was conducted, had at 7°30 A.M. a pulse of 34 a minute. 
At 7:40 half an ounce of chloral hydrate was given it in 2oz. 
of water. 


At : Pulse a minute. 


7°50 6 standing unsteadily as if intoxicated. 
7°52! it fell down asleep. 
‘bo 43 
759" 40 
8:8’ 48 a tap having been put in the jugular vein but 


no bleeding having occurred. 


224 MR GARROD. 


At Pulse a minute. 


8:12’ 52 Bleeding slowly from jugular. 
S15 67 | minute after minute, the animal having lost 
& 64 | : 
62 ¢ altogether about one pint or a little more 
% 60 of blood. 


8:19’ 59 Bleeding freely. 
8:20 52 Bleeding ceased 


8:21 AD 55 55 

8:22! 48 x " 

8:22’ 30” Bleeding resumed 

8:23’ 30” 49 Bleeding. Resp. 11:5. 

8:24) 15” Bleeding ceased after loss of another pint and 
half. 


8°24’ 30” 43 Bleeding ceased. 

8:25’ 15" 42 No bleeding. Resp. 11°5 
8-26 42 i 

8:28’ 15” 42 Bleeding freely. 


8°30 30” 42 - - Resp. 13: 
8:34 30” 87 i < 
836 38 ¥ : 
840" 37 . é 
8°42’ 35 55 > Resp. 14 
S45 36 if » Resp. 16° 


and from this time until 9°10’, by which time more than half 
a pailful of blood had been lost and the carotid pulsations were 
very feeble, the pulse remained at 35°5 to 35 in a minute, with 
the respirations varying from 12 to 13 in the same time, and 
the loss of blood being continuous throughout. 

The animal did not move once through the whole experi- 
ment. 


Experiment V. <A terrier dog had 30 grains of chloral 
given it in two doses, 15 grains first and another 15 grains 
about an hour afterwards. This did not render it quite coma- 
tose, so it sniffed chloroform until insensible, when a hamady- 
namometer was connected with one of its carotids, and a pres- 
sure of 66 inches was immediately registered, which was 
steadily maintained, undulating with the respiration. 


~~ 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 225 


Pulse Pressure. 


in sf SA in a minute. 
40 186°5 6°45 inches 
In 3 minutes 42 192 6:3 = 
2 40 188 . 64 RY 
2 cms 184 64, 
2 49 182 6°6 
if 52 164 GS5) 
1 159 ! re oi 
1 i 150 c y 
15 70 138 6:9 FS 
4 142 6-7 i 
5 149 6'8 o 
6 156 58 H, 
4 168 Garis 
6 102 17 s 
10 129 ra | = 
5 138 19 Fs 
2 a8: 141 2k rs 
3 64 142 741 | re 
5 


138 1) 


In this experiment the bleeding occurred from the carotid, 
and took place between the pulse-countings, which were traced 
on a revolving drum. The great fall in pressure indicates the 
excessive bleeding in one case. In others a much less quantity 


~ of blood was lost on each occasion. 


There was no bleeding after the fall in pressure to 17 
inches and from that time the pressure and pulse became less 
and less till the animal died. 


Experiment VI. A rabbit was made comatose by 15 grains 
of chloral, and was bled to death, the whole operation taking 
half an hour. An hemadynamometer was connected with the 
carotid, and the blood was lost from the jugular of the same 
side, in drops continuously. The pressure at first, at about 
6 inches, fell at the end of the experiment to less than one 
inch, when death occurred. The following are the pulse-rates 
taken at equal intervals during the half hour. 


226 


In 10 seconds, 
42°5 
41°9 


Bleeding began 46° 


MR GARROD. 


In 10 seconds. 


42°75 
43° 
42° 
42: 
42°5 
41°5 
4.2 
42°4 
41° 
41°75 
42: 
42° 
4]° 
41° 
42° 
40°6 
42° 
42: 
39° 


Experiment VII. <A rabbit under the influence of 15 grains 


of chloral— 


When pressure 4°8 inches. 


‘5 3 43 
ss - 28 
5: 4 laf 
5 55 28 
15 £ 2°4 
. “ es) 
3 * i 
, » 5 


> be ” 9 
Death from loss of blood, 


Pulse 136 in a minute. 


Serpe: 
127 a 
12 ie 
Loa hes, 
(58s & 
PAG. 
(Caan 
het hae 
1101 eo 


From these experiments it is evident that the pulse does 
not increase in frequency with loss of blood, as it did not do 


so in any one of them. 


In Experiment IV. the pulse-rate rose on making the in- 
cision in the skin necessary to expose the jugular vein, and 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 220 


continued to do so shortly after bleeding commenced, but soon 
diminished, and after reaching 86 a minute remained perfectly 
constant, notwithstanding a continuous and considerable loss 
of blood from the vein until the animal was almost exsangul- 
nated. 

With the rabbits the difficulty in keeping them completely 
under the influence of the hypnotic, with the tendency to 
struggle, makes the results less uniform, but in all the cases 
there was a fall in pulse-rate, not a rise, accompanying the 
reduction in blood-pressure. This fall, which was not very 
great, may result from the cooling of the surface, consequent 
on the lessened circulation. 

From these observations it may be concluded that variations 
in the amount of blood in circulation do not vary the rapidity 
of the pulse, and consequently that the pulse-rate is not de- 
pendent on the blood-pressure, as Marey supposed. 

The next question was—What law as to the frequency of 
the heart’s beats would satisfy these two above proved facts, 
namely, the dependence of the pulse-length on the arterial 
resistance and its non-dependence on arterial pressure ? 

The method adopted by Mr Fleeming Jenkin for detecting 

the insulation of long cables at different times occurred to me 
as being subject to exactly similar laws, the time of fall of cable 
charge from tension to half tension, which he employs, varying 
directly as the leakage, and as that only. 
Can it be that the heart always recommences to beat when 
the tension falls a certain invariable proportion, and then only ? 
This theory it was my next object to analyse, and the different 
elements into which it resolves itself were, and will be now, 
considered separately. 


a uniform circu- 


First, as to the full meaning of the term, 
lation. A uniform circulation is one in which the quantity of 
fluid flowing through all cross sections of the circulating system 
is the same; for if the flow through one part were less than 
through another, there would be a tendency for the fluid to ac- 
cumulate in front of the obstruction, which is incompatible with 
the premises. 

As a consequence of this, the heart must always recommence 
to beat directly as much blood has left the capillaries as was 


228 MR GARROD. 


sent out from it in the previous pulsation, and therefore the 
length of the pause or diastole must depend on the relative ca- 
pacities of the heart and of the arterial system, and on the 
rapidity of the flow of blood through the capillaries. 

At this point the work of Poiseuille respecting the flow of 
fluids through capillary tubes is invaluable. He found* that, 
other things being the same, the flow of fluids through capillary 
tubes varies directly as the pressure. These results were veri- 
fied by a Committee of the Academy of Sciences; and, by an 
entirely different method, I have been enabled to do the same 
on the vessels of the animal system. 

My method was the following in a particular case-——The 
kidneys of a deer, with the aorta and renal vessels intact, were 
removed from its body and placed for some time in water at 
100 F.; the aorta was ligatured just below the origin of the 
renal arteries, and a uniform glass-tube was tied into it just 
above them. Water at 100 F. was poured into the tube and it 
distended the organs; the tube was maintained full by a con- 
tinuous supply which was suddenly stopped, and the time of fall 
of the column from tension to half tension at different initial 
pressures observed, and it was always found that it took exactly 
the same time to fall from 40 inches to 20 inches as from 20 
inches to 10 inches, thus verifying the law. 

This law being thus true, it is evident that if the capacity of 
the arterial system, including the left ventricle, varies directly 
as the pressure, then the heart must always recommence to beat 
when the arterial tension has fallen a certain proportion; for 
with double pressure, and consequently double amount of blood, 
the time of flow through the capillaries is constant, as the flow 
varies directly as the pressure ; and with double resistance and 
unvaried pressure the time of flow is double also, for the heart 
pumps again when as much has gone from the capillaries as it 
has sent into the arteries, and the relative capacities of the heart 
and arteries do not vary according to the assumption. 

But does the capacity of the arterial system vary as the 
pressure 4 

1 Recherches expérimentales sur le mouvement des liquides dans les tubes de 


tres-petits diametres. apport de VAcadémie des Sciences. Comptes Rendus. 
Tome tv. 1842. 


OO CCl 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 229 


This is a point which it is very difficult to prove. With re- 
gard to the heart the following facts bear on it. By connecting 
a syringe with’ the coronary arteries, or by tying it into the 
aorta and pumping backwards, it can be shewn that increasing 
the pressure in the coronary arteries increases the capacity of 
the ventricles. Also in many post-mortem examinations the 
heart is found with the ventricular cavities fully obliterated, and 
as they are not then in action, the capacity of the heart and the 
pressure in it are at a minimum together. This is all the direct 
evidence that it is in my power to bring on this point. 

With regard to the arterial system and its capacity, the ab- 
sence of blood in the arteries after death has been known from 
time immemorial, and if their capacity varied directly with the 
pressure, it is evident that that must be the case, both capacity 
and pressure being at a minimum. 

A direct method of determining this point having occurred 
to me, the following description will illustrate it. In a rabbit 
one of the carotids was put in communication with a kymogra- 
phion ; and during the time the recording drum was revolving, 
the chest was suddenly opened and the ventricles cut across 
transversely. The pressure fell rapidly to zero, and it is clear 
that the fall must have arisen from the escape of the blood 
through the peripheral vessels, as the aortic valves would close 
immediately. The curve of descent would take a definite form, 
which is easily expressed in mathematical language, if the ca- 
pacity diminished as the pressure. Unfortunately the time 
required to open the chest, and other difficulties connected with 
the operation, prevented my results from being of much value ; 
and Dr Michael Foster suggested to me that the same object 
would be attained if the heart were made to stop by the action 
of the interrupted current on the pneumogastric nerve. Mr 
Martin, of Christ’s College, Cambridge, kindly sent me some 
traces thus taken, and one of the two which are suitable for 
measurement entirely conforms with the law, that the capacity 
of the vessels varies directly as the blood-pressure, assuming 
Poiseuille’s law to be correct. The other curve does not exactly 
fulfil the requirements, but varies very little from them. When 
further opportunity occurs, I hope to repeat these experiments 
on a larger scale. 


230 MR GARROD. 


It can also be shewn in other ways that the arteries do not 
obey the laws of ordinary elastic tubes. They are covered by a 
dense, scarcely elastic, fibrous coat which limits their distension, 
and they are surrounded by organs and muscles which are press- 
ing on them in all directions. So it may be said at least that 
they do not vary in capacity as simple elastic tubes, and that 
the difference is towards their varying directly as the pressure. 

However, the indirect evidence proves that the capacity of 
the arterial system, the ventricle included, varies directly as the 
pressure: for the facts above considered as to the frequency of 
the pulse depending on the resistance, and not at all on the 
pressure, can only be explained on this assumption. 

If the direct evidence as to the capacity of the vessels had 
- been contradictory, it is true that it would have been necessary 
to assume some error in the method of conducting the pulse ex- 
periments ; but, as above shewn, it is quite in the right direction, 
and only lacks partial direct verification. 


So much in the verification of theories connected with Phy- 
siology must depend on the way in which collateral facts are 
explained by them, that it will be advisable now to consider 
some of them; and these considerations will be divided into two 
sections,—Ist, The explanation of the known variations in pulse- 
rate in health, and 2ndly, The explanation of the cardiograph 
laws. 

Ist. Variations in Pulse-rate in Health. With regard to 
these points, as on this theory change in pulse-rate can only 
depend on change in arterial resistance, it is evident that Ma- 
rey’s law will, upon his supposition as to the relation between 
blood-pressure and arterial resistance, explain the phenomena 
equally well. 

The following are some of the best known:— 

The effects of Respiration on the Pulse-rate. 

Physiologists, though not completely agreed as to the effects 
of respiration on the pressure of the blood in the arteries, all 
acknowledge that during inspiration the pulse quickens, and 
during expiration it gets slower, whether the pressure rises or 
falls. The theory under consideration clearly shews that this 
must be so, for during inspiration the expansion of the chest 


LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 231 


must reduce the pressure in the intra-thoracic aorta, and con- 
sequently its contained blood must fall in tension more rapidly 
than if the chest were motionless, and the more rapid tension 
fall causes increase in pulse-rate. In expiration the opposite 
occurs, diminution in chest capacity reduces the size of the 
aorta, and consequently delays the time of fall of tension, and 
therefore slows the pulse. 

If other remote effects of respiration tend to modify the 
pressure in the vessels, it is evident that they would co-exist 
with the above and influence it but slightly, explaining the 
existence of the experimental discrepancies. 

The effect of position of the body on the Pulse-rate. 

The experiments of Dr Guy led him to explain the differ- 
ences in pulse-rate following change of position as depending 
on the amount of muscular effort necessary to maintain the 
positions assumed, and his explanation, assuming that muscular 
effort of itself can change pulse-rate, is very complete. It is 
curious that the theory under consideration gives an inter- 
pretation of the same facts, though very different from that of 
Dr Guy. 

The following are the most essential facts. The pulse is 
quickest while standing erect, slowest while lying, intermediate 
while sitting, slow while standing leaning and while supported 
entirely, as by being bound to a wheel in any position. 

The following is the explanation. While standing, the only 
soft parts of the body which support the weight of the body are 
the soles of the feet, and the weight is transmitted to them 
through non-vascular and rigid tissues, cartilage and bone. Con- 
sequently the blood flows freely through almost all the vascular 
system unobstructed. But while lying, most of the weight is 
supported by highly vascular tissues, as the shoulders, arms, 
thighs and legs, and consequently much of the circulatory sys- 
tem is greatly reduced in capacity from the compression it ex- 
periences, and considerable resistance to the flow of blood is 
introduced into the system, the fall of tension is retarded, and 
the pulse therefore rendered slower. 

In sitting, an intermediate condition is the result, and an 
intermediate rate of pulse is produced. 

Leaning while standing, and entire support on a wheel, 


232 MR GARROD. THE FREQUENCY OF THE PULSE. 


both by introducing resistance from compression of soft parts, 
tend to make the pulse slow. 

Thus, according to Dr Guy’s assumption, the slow is the nor- 
mal pulse, and the quick the induced ; upon the fall of tension 
theory the reverse is the case. The occurrence of bed-sores 
and the paleness of a compressed part prove that pressure dis- 
turbs the uniformity of the circulation. 

The rapid pulse after a meal, during digestion, depends on 
the relaxation of the vessels of the alimentary canal while its 
functions are being performed. 


(To be continued.) 


A CONTRIBUTION TO THE VISCERAL ANATOMY OF 
THE GREENLAND SHARK (Lemargus borealis). 
By Proressor TURNER 


Read before the Royal Society of Edinburgh, March 17, 1873. 


NaTuRALISTS have recorded a few instances of the capture 
of the Greenland Shark in the British seas. Dr Fleming 
states’ that one was caught in 1803 in the Pentland Firth, 
and that one was found dead at Burra Firth, Uist, in 1824, 
Mr Yarrell’ refers to a specimen caught on the coast of Durham 
in 1840, which has been preserved in the Durham University 
Museum. In May 1859 a specimen, about ten feet long, was 
caught in the Firth of Forth near Inchkeith, the stuffed skin 
of which is preserved in the Edinburgh Museum of Science 
and Art. In April 1862, a specimen was caught on the Dogger 
Bank and brought into Leith. <A brief description of its 
external characters was read by Mr W. S. Young to the Royal 
Physical Society of Edinburgh*, On April 27, 1870, Dr John 
Alexander Smith read before the same Society a notice of a 
female specimen caught about 30 miles east of the Bell Rock. 
It had become entangled in one of the deep sea fishing lines, 
many of the hooks attached to which had stuck into its body. 
It measured about 15 feet in length and 3 ft. 1 in. between 
_ the tips of the tail lobes. The stuffed skin of this fish is also 
preserved in the Edinburgh Museum of Science and Art. 

In the month of February of the present year three 
specimens were caught by fishermen at sea some miles east 
of the Bell Rock and brought into Broughty Ferry. One was 
taken to Dundee for exhibition, the others were brought to 
Edinburgh for the same purpose. By permission of the pro- 
prietors I was enabled to examine the latter specimens, and to 
acquire for the Anatomical Museum the viscera and other 
parts. One was a large female 11 ft. 8 in. in length, the other 
a smaller female 84 feet long. 

1 History of British Animals, 1828. 


2 British Fishes, 1841, p. 528. 
3 Proceedings, May 7th, 1862. 


VOL. VII. 16 


234 PROFESSOR TURNER. 


The skin possessed the characteristic shagreen structure, 
and in colour was bluish-gray. The sides of the body were 
barred with faintly marked narrow stripes, about 1 inch asun- 
der, which extended transversely from the dorsal mesial line 
towards the ventral surface. These stripes were due to a 
somewhat darker colouration of the integument along the bars 
than in the intervals between them. The lateral line was dis- 
tinctly seen on the side of the body, and the mucous canal corre- 
sponding to it ramified anteriorly both on the upper and under 
surfaces of the head. Thick mucus oozed through the pores 
of the canal on to the surface of the skin*. The following 
measurements of the larger specimen were taken*. 


it, aie 
From tip of ‘snout to end of tails... ccc. menteaccoss 11 8 
to back of Ist dorsal fin ......... 6 0 
to back of 2nd dorsal fin......... WE) 
to anterior edge of ventral fin... 7 9 
to anterior edge of pectoral fin... 8 5 
Rey ALO MOS UTILS me iasts sere rane tT atc eeceee 0 5 
tO ISPIZACIC. ose cae eee 1 * 76 
OREN C reac cere 5 oes cicaeiescenener bya! 
5 doth DOLENOUGL cccnse het ceon eens seen ee Poe 
HMeiahtiot. Ustidorsal tra)... 5s seeces oneness cae deee Cra 
as BM fase f usck cert Coee sme se ae eee eee 0 54 
Dieneth ot pectoral tin co. ccc. ngee ve clorasee ceca vecaaneeer 1 3 
Breadth of wae yes erin dot eceut Oe ae pee cone aeoae be | 
Deneth-of ventral Gn 425.000. concnsennessegaceee- serio“ Ls 
Breadth of ventral fin from cloacal aperture ......... Ont 
Between tips of tail-lobes............sccccesssenssnnsseees yes 


Snout short and bluntly rounded. Nostrils situated on 
under surface of snout and near its outer edge. Spiracles on 


1 The figure of this Shark given by Seoresby (Arctic Regions) errs in making 
the snout too pointed, and in placing the gill apertures too far in front of the 
pectoral fin. Couch (British Fishes) errs in the same particulars, and moreover 
makes the colour ash-grey, instead of bluish-grey. He represents, however, 
the barred appearance of the side of the body, though he does not make the 
stripes sufficiently numerous. Yarrell’s figure gives a much better representation 
of the fish, and is correct in the position of the gill-openings. 

2 All the dimensions given in this paper, except when otherwise stated, are 
from the larger of the two sharks. 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 235 


a line with outer edge of snout, 44 inches above and behind the 
eyes, also behind a line drawn vertically from the angle of the 
mouth. 5 gill-apertures on each side lying in an oblique line 
in front of the pectoral fin, the posterior opening being so close 
to the root of this fin, that the membranous lining of the 
aperture was in contact with the surface of the scapular arch. 
Each opening was 5 inches in vertical diameter, and the distance 
between the anterior and posterior aperture was 94 inches. 
The greatest depth of the body of the fish, immediately in 
front of the 1st dorsal fin, was 3 ft. 9in., the girth at the root 
of the tail was 1 ft. 4 in. 

Attached to the cornea of one of the eyes of the smaller 
fish was a specimen of the parasitic crustacean named Lerneo- 
poda elongata. This remarkable parasite was first observed on 
the eye of the Greenland Shark by Dr Scoresby, but its true 
character was not understood until Professor Grant published a 
description of it’. Some additional particulars of its mode of 
attachment to the eye were given some years ago by Dr H.S. 
Wilson and myself, in an appendix to a memoir on Lerneopoda 
Dalmanni?. 

The transverse diameter of the mouth was 15 inches. Eight 
rows of teeth with truncated cutting edges were arranged 
transversely on the lower jaw. One row was placed on the 
free margin of the jaw, and formed the line of the cutting 
edge; one row was situated on the outer surface below this 
line, and six rows lay in successive tiers on the inner surface 
which had not yet reached the cutting line. The two deepest 
of the inner series were rudimentary both in shape and size, 
and seemed to be invested by the mucous membrane. Seven 
transverse rows of sharply pointed teeth projected from the 
free surface of the upper jaw. The mouth was lined by a 
smooth mucous membrane stained with grey pigment, which 
was prolonged into the canal of the spiracle which opened 
between the cartilage of the upper jaw and the great cartilage 
of the skull. The mouth opened directly into an cesophagus, 
equal in calibre almost to the mouth itself. 

When the abdominal cavity was opened an enormous 

1 Brewster's Edinburgh Journal of Science, 1827, p. 150. 
2 Trans. Royal Society, Edinburgh, 1862, p. 85. 
16—2 


236 PROFESSOR TURNER. 


2-lobed, pale-yellow, oil-containing liver was exposed, which 
broke away by its own weight, when it lost the support of the 
abdominal walls. 

The stomach was a capacious sac, and empty, some viscid 
yellow mucus coating its inner surface. Anteriorly it opened 
into the wide and short cesophagus. A few inches in front of 
its posterior end it had a transverse diameter of 14 inches. It 
then abruptly diminished to a breadth of only 54 mches and 
formed a short cul-de-sac, rounded posteriorly, which I shall 
name the pyloric compartment, and from which the pyloric 
tube arose 2 inches in front of its rounded end. <A shght con- 
striction externally marked the separation of the pyloric com- 
partment from the general sac of the stomach, whilst internally 
they communicated by an orifice, large enough to admit the 
fist, bounded by a fold of the mucous membrane. The mu- 
cous membrane of the pyloric compartment, like that of the 
stomach generally, was elevated into small and very tortuous 
closely arranged folds, which resembled on a small scale the 
convolutions of the cerebrum. This arrangement differed from 
that usually found in sharks, but was apparently not unlike 
what Duvernoy has described in Galeus thalassinus’, though 
in the latter the convolutions were probably of larger size. 
The pyloric tube communicated with its compartment by an 
orifice which barely admitted the little finger, and after curving 
forwards for 6 inches it opened into the duodenum by an 
orifice so small that the tip of the little finger only could be 
passed into it. The internal diameter of the pyloric tube was 
1 inch at its commencement, but it tapered so as to be not 
more than half an inch at its duodenal end. Its mucous lining 
was quite smooth, and contrasted in a marked manner with 


the corresponding membrane both in the stomach and duo- | 


denum. 

The duodenum was a cylindrical tube 3 ft. 2 in. in length, 
which at first ran forward for 14 inches, then, bending at an 
acute angle, passed straight backwards for 24 inches, where it 
was constricted, partially surrounded by a collar-hke fold of 
mesentery, and became continuous with the part of the in- 


1 Cuvier, Lecons, tv. 2nd Part, p. 166. This shark is the Thalassorhinus 
vulpecula of Valenciennes, Miiller and Henle. 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 237 


testine also straight in its direction, which contained the spiral 
valve. The spiral valvular part of the intestine was 13 inches 


DESCRIPTION OF FIGURE 1. 


a. hinder part of great sac of stomach. b. pyloric compartment. c. pyloric 
tube. d. duodenum. e. shorter and f. longer duodenal ceca. g. bile-duct. 
h. pancreas. i. great pancreatico-intestinal vein. k. spiral valvular intestine. 
l. glandular body. m. rectum. The arrow is placed in the aperture of commu- 
nication between the great sac of the stomach and the pyloric compartment. 


long, and terminated behind in the rectum, which extended for 
7 inches backwards to the anal orifice. The total length of 
the intestine was 4ft. 10in. The external circumference of 
the duodenum was 54 inches. Its mucous lining was floccu- 
lent and villous. 
Continuous with the duodenum, immediately opposite its 
pyloric orifice, was a ccecum (e), 6 inches in length, which pos- 
sessed almost the same internal diameter as the duodenum 
itself. The ccecum was not free, but lay parallel to, and so 


238 PROFESSOR TURNER. 


intimately blended with one side of the wall of the pyloric 
tube, that it required a careful examination to recognise that 
two distinct tubes were enclosed within a common peritoneal 
investment. Its ccecal end was in contact with the wall of 
the pyloric compartment of the stomach. Its canal was in line 
with the duodenum and had a similar villous mucous mem- 
brane, so that the duodenum seemed rather to be a continua- 
tion of the ccecum than of the pyloric tube. Seven inches 
beyond the mouth of the pyloric tube a second cecum (/) 
184 inches long opened into the duodenum by a wide orifice 
14 inch in diameter. It extended backwards, enclosed in a 
fold of peritoneum, parallel but distinct from the straight part 
of the duodenum, to terminate in a free rounded end. This 
ccecum was 44 inches in circumference, and lined internally by 
a flocculent, villous, mucous membrane. The pancreatic duct 
entered the wall of the duodenum in the retreating angle of its 
bend, and the hepatic duct pierced it in nearly the same locality. 

The valvular part of the intestine was dilated so as to have, 
when distended, an external circumference in its widest part 
of 12 inchés. The coats of the gut, at the constriction which 
separated the duodenum from the valvular part, were very 
thick, and the lumen of the tube barely admitted the tip 
of the index finger. The valve formed a broad thin membra- 
nous band, which made 23 turns, each of which lay almost 
transversely and extended so far into the lumen that but a 
narrow central passage was left. The mucous covering of the 
valve was smooth on the surface. The transverse arrangement 
of the folds of the valve was indicated by the circular course 
of the branches of the intestinal artery and vein, the outhne 
of which could be seen on the surface beneath the serous 
membrane. 

The terminal part of the gut, or rectum, not so dilated 
as the valvular part, was distinguished by longitudinal mark- 
ings on the outer surface owing to the fasciculated arrange- 
ment of the muscular fibres. The anus opened into the cloaca 
immediately in front of the two ureters. Four inches in front 
of the anus an ovoid body two inches in length was attached 
to the wall of the rectum. It was invested by peritoneum 
continuous with that surrounding the gut, and a prolongation 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 239 


of the same membrane passed from it to an adjacent part of 
the wall of the abdomen. A duct opening into the rectum 
passed along the centre of this body nearly to its free end. 
The inner coat of this body was formed of a softish material 
continuous with the smooth mucous coat of the rectum. It 
had the appearance of a gland, and without doubt discharged a 
secretion into the canal of the duct. A similar body probably 
exists in the Sharks and Rays generally, and to it Duméril, who 
recognises its glandular nature, applies the name of appendice 
digitiforme’. 

The pancreas was in close relation to the posterior straight 
part of the duodenum. Its body, flattened and quadrate in 
form, was 6 inches in breadth and lay across the intestine. 
From the posterior angles two elongated processes passed back- 
wards, one for 18 inches, the other for 21 inches, and the latter 
was much more attenuated, especially at its free posterior end. 
These processes lay parallel and connected to the straight part 
of the duodenum by a fold of peritoneum, and the shorter one 
occupied the interval between that part of the intestine and 
the larger of the twe duodenal cceca. The surface of the 
gland was quite smooth, invested by peritoneum, and showed 
no appearance of a lobular subdivision. The duct was con- 
cealed in the compact substance of the gland and sent a branch 
along the centre of each of the elongated processes. The duct 
emerged from the body of the pancreas at one of the antero- 
_ lateral angles, and after a course of 4 inches in a peritoneal 
fold, it pierced the wall of the duodenum, through which it 
ran obliquely, and twisted upon itself before it opened into 
the canal of the gut at the summit of a small papilla. 

The hepatic duct, 14 inches in length, also enclosed in a fold 
of peritoneum, pierced the wall of the duodenum 13 inch from 
the place of entrance of the pancreatic duct, and after an 
oblique and twisted course opened at the summit of a papilla 
situated on the opposite wall of the duodenum to the orifice 
of the pancreatic duct. 

Enclosed within the same fold of peritoneum as the bile- 

1 Hist. Nat. des Poissons, 1. 157. Monro figured it as an Appendix vermi- 
formis in the Skate. E. Home figured it in Spinax; Owen refers to it, Verte- 


brates, 1. p. 424, fig. 352; and Gegenbaur, Vergleich. Anat. fig. 267, figures it in 
a Squatina, 


240 PROFESSOR TURNER. 


duct, at about two inches from the duodenum, was a bi-lobed 
body (below g, Fig. 1), each lobe of which was the size of a 
small almond. From the more anterior of these two lobes a 
slender cord, which seemed on external examination to be solid, 
passed forwards parallel to the bile-duct. On transverse section 
a canal, so fine as only to admit a bristle, but bounded by a 
thick wall, was seen in the centre of this cord. This canal 
passed into the lobes, where it apparently formed a convoluted 
tube, for on making a section through the lobes it was repeat- 
edly cut across, sometimes transversely, at others obliquely. In 
the more posterior of the two lobes the calibre of the tube was 
less than in the anterior, or than in the cord, but whether this 
arose from a diminution in the size of the tube itself, or because 
it subdivided into smaller branches, I cannot say. A large part 
of the substance of each lobe was made up of a dense fibrous 
tissue, continuous with the thick wall of the duct, which, on 
microscopic examination, was found to be largely composed of 
pale smooth fibres, resembling involuntary muscle, mingled with 
w hich was a nucleated protoplasmic substance, in which an 
arrangement into not very well defined caudate corpuscles could 
occasionally be seen. Sections through the tubes showed an 
appearance which indicated the presence of an internal epithe- 
lium, but from the length of the interval between the death of 
the animal and the time when the microscopic examination was 
made, neither the form of the cells, nor their method of ar- 
rangement, could be satisfactorily determined. It is not very 
easy to pronounce an opinion on the function of this small 
organ. It resembled in many points a convoluted or compound 
tubular gland, the secretion of which could be expelled by the 
muscular wall of the duct. But this secretion was not poured 
into the intestine, for the duct passed away from it towards the 
liver. Whether it reached that organ, or had some other mode 
of termination, I am unable to say, as the duct had been cut 
short in eviscerating the animal, and before indeed it had been 
observed, It is possible that it may have communicated with 
the bile-duct, for, as is well known, the bile-duct has, in many 
animals, gland structures not unfrequently opening into it. 

A large mesenteric vein passed forward from the spiral 
part of the intestine, and reached the hinder end of the more 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 241 


attenuated limb of the pancreas. It ran forwards in close con- 
tact with this part of the gland, receiving in its course veins 
from the pancreas and duodenum, and at the anterior border of 
the gland it was joined by a large vein coming from the oppo- 
site limb, into which a number of small duodenal veins had 
opened. The great pancreatico-intestinal vein was almost equal 
in size to the human inferior vena cava. 

The spleen was attached by a gastro-splenic fold of peri- 
toneum to the stomach. It was 17 inches long and 6 inches 
wide anteriorly, but was much more attenuated in its middle 
and posterior two-thirds. Its surface was marked by many deep 
fissures, which mostly ran longitudinally and gave to its surface 
a corrugated appearance. 

The kidneys lay one on each side of the spine and extended 
backwards almost to the cloaca. At the hinder end of each 
gland the ureter became free, and the two ducts opened close 
together in a shallow fossa at the back of the cloaca. 

The ovaries, two in number, were situated in the anterior 
part of the abdominal cavity, and connected to its dorsal wall 
by a fold of peritoneum. Each ovary in the larger shark was 
23 inches long, and consisted of numerous parallel folds or 
laminz which lay obliquely. The folds were attenuated along 
their Jine of attachment to the mes-oarium, but were club- 
shaped and succulent at their free ends. Multitudes of ova 
were seen in the fibro-vascular stroma, the smallest of which 
"were mere specks, others again corresponded to shot of different 
sizes, and a few were as big as small bullets. Extending along 
the mes-oarium close to the attenuated ends of the folds was 
a thin-walled blood-vessel, into which an injecting pipe was 
introduced, and a carmine and gelatine injection gently passed 
into it. The vessel and its branches at once became distend- 
ed, and the injection at the same time passed into a system 
of freely anastomosing spaces both in the mes-oarium and in 
the ovarian substance. These spaces swelled up so rapidly 
and continuously as to give one the impression, not of an 
extravasation, but of the existence of a cavernous vascular 
arrangement into which the injection freely passed. When 
examined microscopically the injection was seen to be con- 
tained in a net-work of dilated vessels, which often presented 


242 PROFESSOR TURNER. 


DESCRIPTION OF FIGURE 2. 


Cloacal region of Greenland Shark. The bristles were introduced into the 
two ureters. The rectum opened into the darkly shaded hollow in front of the 
bristles. The abdominal pores may be seen one on the outer side of each bristle. 


bulgings or varicosities on their walls. The coats of these 
vessels seemed to consist merely of a nucleated membrane, so 
that the vessels themselves may be regarded as colossal capil- 
laries: though as regards size it should be stated that they 
were by no means uniform; for whilst some were but little 
larger than ordinary capillaries, the greater number exceeded 
these in diameter five or six times. 

Distinct puckerings, radiating from a central depression, 
were observed in the ovarian stroma, immediately superficial 
to that aspect of both the larger and smaller ova which lay 
next the surface, and it is probable marked the spots where the 
ovary ruptured at the period of discharge of the ova. In the 
injected part of the ovary these puckerings preserved their 
white appearance, and contrasted with the red injected stroma. 
No oviduct was seen in the abdominal cavity, and there were 
no oviducal openings in the region of the cloaca. 

Immediately posterior to the mouth of the cloaca, where the 
skin was soft, smooth and pinkish, two rounded openings pro- 
tected by fimbriated processes of the soft integument were 
seen. These openings were the abdominal pores and through 


DESCRIPTION OF FIGURE 3. 


View of the posterior end of the peritoneal cavity. R. the rectum trans- 
yersely divided. The two ureters lie immediately behind it. On each side the 
funnel-shaped prolongation of the cavity leading back to the abdominal pore is 
represented in shadow. 


them large bristles could be passed into the peritoneal cavity, 
which communicated with them by means of a funnel-shaped 
and somewhat twisted prolongation, one on each side of the 
terminal part of the rectum. In the smaller shark each ovary 
was only 14 inches long, and the ova were so small in size as to 
be scarcely visible. 

The heart was contained in a well-defined pericardium. It 
had the general form and subdivisions into auricle, ventricle 
~and bulbus (conus arteriosus) of the heart of the cartilaginous 
fish. The great venous sinus opening into the auricle was 
situated within the pericardium, and its orifice was guarded 
by a bi-segmented semilunar valve. The valve at the auriculo- 
ventricular orifice was bicuspidate, and was attached by chord 
tendine to the surface of the wall of the ventricle and not to 
projecting musculi papillares. It lay to one side of the opening 
into the bulbus (conus), and a pouch-like prolongation of the 
ventricle passed forwards in front of it. The bulbus (conus) 
was three inches long. Four transverse tiers of valves pro- 
jected from its inner wall within 13 inches from its ventricular 
origin. The first tier situated around the bulbo-ventricular 
orifice consisted of six cuspidate segments, three larger and 
three smaller. The apices of the cusps were directed forward 


244 PROFESSOR TURNER. 


away from the ventricle. The surface of each cusp lying next 
the lumen of the bulbus was smooth, whilst the opposite sur- 
face and the free edges were connected to slender chord ten- 
dine arising either from the valves in the tier immediately 
above or from the wall of the bulbus itself. The second and 
third tiers each consisted of five segments, three larger and 
two smaller. The fourth tier consisted of only three small 
segments. The segments in the three upper tiers were not so 
distinctly cuspidate as in the lowest, and attached to both their 
surfaces as well as to their free edges were chorde tendinez. 
The attachment of the chorde tendinee to the surface next 
the lumen gave it a corrugated appearance not seen in the 
cusps of the tier next the ventricle. Between the most ante- 
rior tier and the end of the bulbus, where the great artery 
arose from it, was an interval of 14 inches, and around the 
orifice of this artery was situated a valve composed of three 
semilunar segments, which in form, size and arrangement 
closely resembled the valve at the mouth of the human aorta. 
The muscular wall of the bulbus though not so thick, yet had 
the same colour as the wall of the ventricle, and consisted of 
transversely striped fibres, so that it formed, lke the ventricle 
proper, a chamber with a pulsatile wall. The aorta, on the 
other hand, had the usual yellow colour of a* large artery. 
From a consideration of its anatomical characters I am disposed 
to agree with Gegenbaur’ in regarding the bulbus in the 
Selachia as an elongated and to some extent independent 
segment of the ventricle, prolonged forward beyond it, which 
is homologous with the conus arteriosus in the higher verte- 
brates, rather than with the bulbus aorte of the osseous fishes, 
and to which the name of conus, rather than bulbus arteriosus, 
should be applied. 

So far as I can ascertain the visceral anatomy of the Green- 
land shark does not seem to have been previously examined by 
anatomists; and, as my dissections have revealed several impor- 
tant structural peculiarities in which this fish differs from the 
sharks generally, I shall now proceed to direct attention to 
them. 


1 “ Ueber den bulbus arteriosus der Fische.” Jenaische Zeitschrift, p. 360, 
yu. 1865, 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 245 


The stomach possessed the “siphonal” form met with in so 
many fish, but the pyloric tube, instead of arising directly from 
the great sac of the organ, as is the case in the Selache maxima* 
and various other sharks, proceeded from a “pyloric compart- 
ment.” Its opening into this compartment was so constricted 
that I at first regarded it as the proper pyloric orifice, and it 
was not until I had slit open both the pyloric tube and the 
duodenum that I recognised the former to be not the commence- 
ment of the intestine, but the tubular pyloric termination of 
the stomach. 

The cylindrical duodenum, its very considerable length, and 
the absence of any dilatation, such as constitutes the Bursa En- 
tana properly so called, formed most important features of 
difference between this fish and the sharks generally. The 
cylindrical form resembled indeed the duodenum of the sturgeon, 
though it did not possess that double turn which Carus has 
figured in the intestinal tube of the latter fish’. 

The co-existence of a well-developed pancreas with two large 
ceca opening into the commencement of the duodenum, is an 
anatomical arrangement which had not previously been recog- 
nised in the Plagiostomata. If we are to regard these cceca as 
homologous with the pyloric cceca in the osseous fishes—and 
their relations to the duodenum, both as regards their situation 
at its pyloric end, and the continuity and structural correspond- 
ence of their coats with the serous, muscular and mucous coats 
of the intestine point to this homology—then a most important 
piece of evidence is furnished against the view so generally 
entertained by anatomists that the pyloric cceca are the homo- 
logues of the pancreas. Many years ago Alessandrini indeed 
described in the sturgeon, in addition to the pyloric coeca which 
are united into an irregular ovoid organ, a proper glandular 
pancreas opening into the intestine by a distinct duct. But his 
interpretation of the pancreatic nature of the gland, though 
accepted by Milne Edwards’, who regards it—along with some 


1 As figured by Home, Phil. Trans. xcix. Copied in Owen’s Anatomy of 
Vertebrates, Vol. 1. p. 416, and in other works. 

2 Tafeln zur vergleichenden Anatomie, Part tv. Pl. rv. Fig. x1. 1835. 

3 Lecons sur la Physiologie et V Anatomie comparée, Vol. v1. p. 412, 1861. 
Milne Edwards also, p. 515, directs attention to the observations of Stannius 
on the co-existence of a pancreas with pyloric ceca in the salmon, and states 


246 PROFESSOR TURNER. 


experiments by Claude Bernard, which show that the secretion 
of pyloric cceca differs chemically from pancreatic juice—as 
conclusive against the morphological identity of the two organs, 
has been called in question by various anatomists’. The great 
size, however, both of the cceca and of the pancreas in the Green- 
land shark places their co-existence in this fish beyond all ques- 
tion, and should Alessandrini’s view of their presence in the 
sturgeon be substantiated, it would make another feature of re- 
semblance between these two fish to that afforded by the 
cylindrical duodenum already referred to’. 

The bile-ducts did not, as Owen has described in the Selache 
maxima*, open by separate orifices into a common pouch, which 
then communicated with the duodenum by a single orifice; but 
a long single bile-duct pierced the duodenal wall. Owing to 
the breaking down of the liver I was unfortunateiy unable to 
ascertain if a gall-bladder was present or absent. 

In connection with the number of transverse rows of valves 
in the muscular bulbus of the Plagiostomata, Johannes Miiller 
has described* two rows in Carcharias, Scyllium and Galeus; 
three in Sphyrna, Mustelus, Acanthias, Alopias, Lamna, Rhino- 
batus, Torpedo; four in Hexanchus, Heptanchus, Centrophorus 
and Trygon; four to five in Raja; five in Scymnus, Myliobatis, 
Pteroplatea and Squatina. As he merely enumerates the num- 
ber of rows without giving any description, I am unable to say 
if he includes the valve at the anterior end of the bulbus, or if 
he restricts himself to those situated on the wall in proximity 
to the ventricular cavity. In either case however it is clear 
that there is considerable variety in the number of transverse 
rows in the sharks and rays. If the semilunar arterial valve be 
included then Lemargus borealis will correspond with Scymnus, 
Myliobatis, Pteroplatea and Squatina. 


that in the herring, perch, turbot, and one or two other osseous fishes, both 


organs co-exist. 
1 Cuvier, Lecons, rv. Part u. p. 615, Ed, 1835. Salter, Cyclop. Anat. and 


Phys., Article ‘‘ Pancreas.” 

2 Since this paper was in type I have received the last published part of the 
Ann, des Sciences Naturelles, 1873, in which is an elaborate memoir by M. P. 
Legouis, ‘‘ On the Tubes of Weber and the Pancreas of Osseous Fishes.” From 
his researches M. Legouis concludes that all osseous fishes kave a well- 
developed pancreas, and that the system of tubes discovered by Weber in 1827 
forms the excretory duct of the gland. 

3 Comparative Anatomy of Vertebrates, Vol. 1. p. 426, 

4 Miiller’s Archiv, 1842, p. 484. 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 247 


In its genital apparatus the Greenland shark departed in a 
most remarkable manner from the type of structure of the 
Plagiostomata. It is well known that in the sharks and rays 
special canals or oviducts for the reception of the ova exist, and 
that the ova are impregnated in their passage along these ducts. 
In some the egg becomes invested by a case in its passage, and 
is then deposited to undergo development outside the body: in 
others the oviduct dilates into a uterus, and the development of 
the embryo takes place within the parent fish. But in the 
Greenland shark the oviducts were absent; and from the ap- 
pearance of the ovary it was clear that the ova, when ripe, 
would be discharged into the abdominal cavity, which would 
serve the office of an oviduct, and pass backwards along it to be 
discharged by the abdominal pores. In the organisation of its 
genital apparatus the female Greenland shark is much less 
advanced than the other Plagiostomata, and exhibits, in the 
mode of evacuation of its ova, arrangements corresponding to 
those which have been described in the lamprey and other 
Cyclostomatous fishes. In the number of its ovaries it corre- 
sponds to the Rays and to the genera Spinax, Scymnus, Hexan- 
chus and Heptanchus, but differs from the dog-fish, and those 
sharks which possess a nictitating membrane, in which, accord- 
ing to Johannes Miiller’, the ovary is unsymmetrical. 

By Fleming, Yarrell and other zoologists, the Greenland 
shark has been included in the Cuvierian genus Scymnus, sp. 
borealis. Miiller and Henle, in their revision of the generic 
characters of the Plagiostomata’, limited the European species 
of the genus Scymnus to the Liche or Sc. lichia, and made for 
the Greenland shark the new genus Lemargus, and this limita- 
tion has been adopted by Duméril in his Histoire Naturelle des 
Poissons, and by Dr Giinther in his Catalogue of Fishes*. The 
characters relied on for establishing this generic difference were 
derived from the shape of the teeth. In Scymnus the upper 
teeth are straight and narrow, the lower crooked, pyramidal, | 
equilateral. In Lemargus the upper teeth are narrow, conic, 


1 Untersuchungen tiber die Eingeweide der Fische. Berlin, 1845, p. 20. 

2 Wiegmann’s Archiv fiir Naturgeschichte, 1837, p. 399, and Charlesworth’s 
Magazine of Natural History, 1838, p. 89. 

® Vol. vyiir. p. 356, London, 1870. 


248 PROFESSOR TURNER. 


straight or curved outwardly, whilst the lower have a transverse 
edge, oblique and truncate. 

It may be well that we should now compare the visceral 
arrangements in Lemargus with those of Scymnus, so far as the 
latter have been recorded. Johannes Miiller, in his memoir on 
the arrangement of the viscera in fishes, has, several times, 
referred to what he saw in Scymnus lichia, in which he found 
the stomach to terminate directly by its posterior end in the 
pyloric tube, which he says was rather short. The duodenum 
also formed a dome-like Bursa Entiana, as in the sharks 
generally, and differed therefore very materially from the 
tubular duodenum of Lwmargus. He makes no reference to the 
presence of pyloric cceca in Scymnus, so that presumably they 
were absent. Further, Scymmnus possessed a uterus, the mucous 
membrane of which was studded with longitudinal rows of villi, 
and an annular gland for the secretion of an egg-shell existed, 
though, he says, it is remarkable that notwithstanding the 
presence of the gland no egg-shell is formed. 

The anatomical differences, therefore, between Scymnus and 
Lemargus are very much greater than those referable to the 
form of the teeth, on which systematic zoologists have hitherto 
relied in their classification. ‘These differences indeed, more 
especially in the form and extent of the duodenum, in the co- 
existence of a pancreas with large pyloric cceca, and in the 
absence of oviducts, affect not only the relations of Lwmargus 
to Scymnus but to the sharks generally, and call for a recon- 
sideration on the part of the zoologist of the place which the 
Greenland shark ought to occupy amongst the Plagiostomata, 
and require the establishment of a separate family for the 
reception of the genus Lemargus. This family would exhibit 
the following characters: 


LEMARGIDZ. 


No nictitating membrane; two dorsal fins; no anal fin; duo- 
denum cylindrical; both a pancreas and duodenal coeca; in the 
female no oviducts. 

Lemargus. Dorsal fins short, the second not quite so high 
as the first; lower teeth oblique, truncate. 


VISCERAL ANATOMY OF THE GREENLAND SHARK. 249 


Whether the Mediterranean species, which has hitherto 
been associated with this genus as Lemargus rostratus, pos- 
sesses the same visceral peculiarities, will now need to be 
enquired into by those who may be so fortunate as to obtain 
specimens. 


ADDENDUM. April 29th. 


Since this communication was in type I have found in Carus 
and Engelmann’s Libliotheca Zoologica a reference to a paper 
by S. Kneeland, Junr. M.D., entitled “Dissection of Scymnus 
Brevipinna (Lesueur),” published in the Boston Journal of 
Natural History, 1847, Vol. v. No. Iv. pp. 479, 485. This 
paper appears to have been overlooked by subsequent writers 
on the Plagiostomata, doubtless because it appeared in a Jour- 
nal the circulation of which is limited. The shark was a female 
7 ft. 5in. long, caught on the coast of Massachusetts. Dr Knee- 
land briefly describes the liver; the heart with the valvular 
arrangements within the bulbus; the stomach and the spleen. 
He states that the duodenum arises directly from the stomach, 
and evidently considers the pyloric tube as a part of the intes- 
tine. He gives the length from the pylorus to the anus as 
5 ft. 6in. He describes and in a rude outline wood-cut figures 
the cylindrical form of the duodenum and the two pyloric cceca. 
He also figures the spiral valve and the glandular appendage to 
‘the rectum. He states that a pancreas with two lobes existed. 
The arrangement, both from the description and figure, closely 
corresponds with what I saw in Lemargus borealis. Kneeland 
however makes no reference to the morphological importance of 
the arrangement, nor does he recognise in how far it differs 
from other Plagiostomata. He states that “the ovaries con- 
sisted of two membranous sacs, on the internal border of which 
was a light-coloured tubuliform glandular mass filled with 
innumerable granules, about half the size of a pin’s head: each 
ovary had a duct about the size of a crow-quill, extending 
nearly the whole length of the spinal column.” The ducts, 
which he describes as belonging to the ovaries, were, I believe, 
the two ureters, their length and position corresponding with 

VOL. VII. 1% 


950 PROF. TURNER. ANATOMY OF THE GREENLAND SHARK. 


that of the kidneys, and their size being precisely that of the 
ureters in ‘the Greenland Shark. He says nothing about the 
cloaca and the openings into it. 

The Scymnus brevipinna has been referred both by Duméril 
and Giinther to the genus Leamargus. In the arrangement of 
its intestinal canal and pancreas the fish described by Kneeland 
undoubtedly possessed the characters, not of Scymnus but of 
Lemargus, so that there can be no doubt that it ought to be 
referred to the latter genus. Giinther considers brevipinna to be 
merely a synonym for borealis, and refers them to the same 
species. Duméril, however, principally on account of a differ- 
ence in the relative length of the fins to that of the body, 
regards brevipinna as a distinct species of the genus Lemargus. 
Kneeland says nothing of the external characters of his speci- 
men, except that the upper lobe of the tail was not so much 
prolonged as in Sharks generally, so that we have no data 
before us to aid in determining whether or no his fish possessed 
the external configuration of the Greenland Shark, or was a 
distinct species of the same genus. It should be stated, how- 
ever, that the figure of Scymnus brevipinna which Dr Storer has 
given in his memoir “on the History of the Fishes of Massa- 
chusetts” (Memoirs of American Academy, Vol. 1x. pl. XXXVI, 
Fig. 2 and p. 235, 1867), bears a very strong resemblance to 
Lemargus borealis. 


ON THE SOURCE OF NERVE FORCE. A THEORY. 
By A. H. Garrop, B.A., Scholar of St John’s College, 
Cambridge. 


THE universally acknowledged inefficiency of any theory at 
present before the scientific world, to account for the origin of 
the force by which, through the intervention of the nervous 
system, organs at a distance from one another are placed in 
communication, makes me feel justified in publishing a theory 
which has been in my mind for some time, and which during 
that time accumulated information has not in any way shaken. 

The following is my proposition:—The species of energy 
which exhibits itself in the form of ‘nerve force,’ is electricity 
of thermo-electric origin, resulting from the fact that the sur- 
face of the living animal body is always colder than its inte- 
rior. In other words—The available energy resulting from the 
interior of the living animal body being at a higher tempera- 
ture than that of the surrounding medium, is expended wholly, 
or in great measure, in generating the force called nerve force. 
In the discussion of this problem, the following are the con- 
siderations on which it is based. 

1. The temperature of the interior of the living body is 
always greater than that of its surface, because all animal life 
is only a form of chemical degradation, and is therefore neces- 
sarily attended with evolution of heat; which, no other force 
intervening, will always keep the interior of the body hotter 
than the medium in which it lives. When the temperature of 
the air in higher animals exceeds or nearly approaches that of 
the body, which varies but little, special arrangements (perspi- 
ratory, &c.) come into play to diminish that of the surface. 

2. There is an available source of energy in the body, 
which has been but little considered by physiologists, depending 
on the temperature of the surface being lower than that of. the 
interior. The theory propounded gives employment for this 
force. 

3. In the struggle for life, the individuals that economise 
the forces at their disposal are most likely to survive; it is 


17—2 


252 MR GARROD. 


therefore improbable that this not inconsiderable source of 
energy should have escaped employment in this struggle. 

4. The actual construction of the nervous system is quite 
sufficient for the working of such a force as the one proposed. 
In addition to the already known properties of the nerves, in- 
cluding their good conducting power, it is only required that a 
thermo-electric current be capable of being generated between soft 
tissues of different composition or structure, such as the extremi- 
ties of the sentient nerves and the corium in which they are 
embedded. 

Although Magnus was not able to produce any thermo- 
electric current between fluid metals, his experiments are not 
in any way conclusive against the possible existence of such 
currents between different tissues ; the subject is in a position 
for actual demonstration, no doubt, and I much regret that as 
yet I have not had opportunities of attempting to prove or dis- 
prove it. Notwithstanding this, some of the circumstantial or 
collateral evidence is so strong, that, without any direct proofs, 
I feel justified in assuming its correctness. The following points 
strongly favour my theory. 

1. Within certain limits, which are those to which the 
body is most generally exposed, the energy of the individual 
(which must be closely related to the supply of nerve force) is 
greater as the temperature of the air is less. On a cold day, in 
a cold air, there is more will and power to work physically than 
in hot weather, in a hot air, during which languor is a promi- 
nent feature. In a paper published in this Jowrnal I have 
proved that the circulation through the skin in man varies ac- 
cording to the temperature of the atmosphere, and that when 
70° F. of the air is reached, perspiration commences. Conse- 
quently by this means a difference of about 30° PF. is always 
maintained in health between the surface and the interior ; and 
exposure to the highest temperatures of the Turkish Bath, when 
continued for some time, does not disturb this condition. But 
a hot-water bath is followed by very different consequences; if 
the body and face be immersed in one of 100° F., and breathing 
be performed through a tube projecting from the mouth to the 
surface, as I have tried, faintness, or loss of nerve power, comes 
on very rapidly, and is it not because the temperature of the 


THE SOURCE OF NERVE FORCE, 253 


surface is raised to that of the interior? At all events this ex- 
planation is as reasonable as any other. This faintness is soon 
recovered from on the reapplication of cold to the skin. 

2. The effect of muscular exercise is to raise the tempera- 
ture of the body, and so to increase the difference of temperature 
between it and the external air; and when great muscular ex- 
ertion has to be sustained, as in rowing, most of the clothes 
have to be removed in order to allow of the rapid cooling a the 
surface, the necessity for which is keenly felt. 

3. During intra-uterine life the active chemical changes 
going*on in the body of the foetus maintains its temperature at 
a slightly higher point than that of the mother; but imme- 
diately after birth the nerve force generated by the cooling of 
the surface brings the intercostals and other muscles into play, 
and -the child is otherwise much more active than. when in 
utero. 

4. The source of the body heat beimg central, and the cir- 
culation of the blood tending to render that of all the parts 
uniform, it is evident, as a simple physical deduction, that the 
temperature of the portion of the body which corresponds to 
the heated end of the thermo-electric couple (which in the 
theory now under discussion can only be the brain), must be 
lower than that of the blood, because the heat supplied to, and 
developed in it (the brain), is partly employed in generating the 
electric current which is circulating. Dr John Davy noticed 
‘that the temperature of the brain of the rabbit was peculiarly 
low, considerably below that of the abdominal viscera; his re- 
sults have been considered improbable by some, but have never 
been refuted ; they are strongly in favour of my theory, which, 
as shewn above, explains them completely. 

Upon my theory the mechanism of the nervous system may 
be thus summarised— 

The afferent nerves are the conductors to the nerve centres 
of the electric current which is generated by the contact of their 
peripheral ends with the tissues of the cooled skin, which they 
supply. The brain is the largest of the centres towards which 
the nerve current is directed, the other ganglia forming the 
smaller. Through these centres the currents, as through an 
elaborate commutator, are split up or concentrated in a manner 


254 MR GARROD. THE SOURCE OF NERVE FORCE. 


not understood as yet, to be directed along the efferent nerves, 
which are always so situated as to be beyond the reach of exter- 
nal cooling influences. Where an organ acts in any way auto- 
matically, it generally has centres of its own, of a size varying 
in degree according to its automaticity, and these minor centres 
are only to a certain extent subject to the influence of the 
brain. 

As in the working of the electric telegraph, no return or 
second special conductor is required to carry back the current 
to the point from which it started; for where an efferent nerve 
terminates in a muscle, it loses its insulating covering, and 
so is put into indirect communication with the peripheral sen- 
tient nerves through the intervention of the mass of body tissue 
generally, which, though its resistance is much greater, offers 
an incomparably larger mass to be traversed by the current. 


NOTE ON THE CARPUS OF THE SLOTHS. By 
W. H. Fiower, F.R.S. 


Ir is well known that in the existing Sloths (Bradypodide), as 
in their gigantic extinct allies, the megatherioids, the trapezium 
is not present as a distinct bone in the carpus. 

The absence is accounted for by Cuvier’, Owen’, and most 
other writers on comparative anatomy, by the assertion that it 
is united with the scaphoid. The hook-like prolongation of the 
latter bone towards the inner side of the carpus lends some 
countenance to this view, but it is nevertheless incorrect, as in 
reality the representative of the trapezium is to be found in 
what appears at first sight as an enlarged proximal end of the 
rudimentary first metacarpal. This may be demonstrated in 
the following manner. 

1. The aforesaid hook-like prolongation is common to all 
the other Edentates, which possess the trapezium as a distinct 
bone, though not developed to the same degree as in the Sloths. 
Moreover, in the youngest Sloths which I have examined there 
is no sign of its being ossified from a separate centre, as would 
be the case if it were a coalesced trapezium. 

2. The position and relations of the apparent enlarged 
head of the first metacarpal (especially in the two-toed Sloth or 
_ Unau) is exactly that of the trapezium of the Armadillos, Ant- 
eaters, and Pangolins, its proximal surface being rather above 
the level of the head of the second metacarpal, and it articu- 
lates with the last-named bone, the trapezoid and the scaphoid, 
whereas the head of the first metacarpal in all these animals is 
placed more distally than that of the second, and articulates 
only with the trapezium. 

3. The most crucial test is afforded by development. In 
the skeleton of a young two-toed Sloth (Cholopus didactylus), 
No. 2387, Mus. Roy. Coll. Surgeons, the bone which has been 


1 “Dans V'unau, le scaphoide se soude avec l’os d’en dessous ou le trapéze: 
e’est une chose qui lui est toute particuliére.” ‘Dans lai, la soudure du 
scaphoide au trapéze a toujours lieu.”—Osteologie des Paresseux. 

2 Comp. Anat. and Physiology of Vertebrates, Vol. 11. ‘* Memoirs on the My- 
lodon, Megatherium, &c.’’ 


256 MR FLOWER. NOTE ON THE CARPUS OF THE SLOTHS. 


considered the first metacarpal in the adult consists of three 
distinct ossifications. a. a rounded nodule having the usual 
relations of the trapezium, and without doubt representing that 
bone. 06. a smaller intermediate ossification, the epiphysis, and 
c. an elongated cylindrical bone, the shaft, of the metacarpal. 


PS. Since writing the above I observe that Professor 
Humphry has noted the coalescence of the trapezium with the 
first metacarpal in a two-toed Sloth, of which he described the 
muscular anatomy (Journal of Anatomy, Vol. Iv. p. 18). The 
specimen is now mounted in the Anatomical Museum of the 
University of Cambridge; and being exactly at the best age to 
show the disposition of the still partially united bones most 
clearly, Professor Humphry has kindly allowed a drawing of 
them to be added in illustration of the subject (see Fig. 1). 


Fig. 1. Inner side of carpus and metacarpus of a Two-toed Sloth (Cholopus 
didactylus) in the Cambridge University Museum, J. lunar. s. scaphoid. td. 
trapezoid, tm. trapezium. ml. first metacarpal; e. its epiphysis. mll. 
second metacarpal. mIII. third metacarpal. 


ON A CASE OF HYPERTROPHY OF THE RIGHT 
CEREBRAL HEMISPHERE WITH COEXISTENT 
ATROPHY OF THE LEFT SIDE OF THE BODY 
By J. Barty Touxes, M.D. (Edin.) F.R.C.P.E. Medical 
Superintendent Fife and Kinross District Lunatic Asylum: 
visiting Physician to the Saughton Hall Institution for the 
Insane. 


HMistory.—THE only account his mother gives of this man, is 
that at birth he was not an unusually large child, nor was her 
confinement difficult or tedious; that when he was 10 days 
old a drunken man stumbled against the child’s head (which side 
she could not remember), and that immediately afterwards epi- 
leptic convulsions set in and continued for 9 months, when they 
ceased. She states that the head began to grow very much 
after the fits commenced, and that the doctors stated to her 
that he had water on the brain. He never showed any evidence 
of intelligence as other children do, and the left side of the 
body was always smaller than the right. 

There is no history of hereditary predisposition except that 
a near relative died of hydrocephalus. Her son was placed 
in a poorhouse when 10 years of age, when he again became 
subject to fits, which continued until he died. 

He was clinically examined on Sept. 28th, 1871, and the 
following is the report. 

General Appearance.—Patient is a man of average stature. 
The whole of the left side of the body is considerably atrophied, 
whereas the right appears normal. The face is drawn to the 
left; the left side of the face seems on a lower level than that 
of the right, which is fuller. The mouth is drawn to the left. 
The left arm is hardly half the circumference of the right. 
The left forearm rests at right angles to the arm, and the hand 
to the forearm. The whole arm is kept applied to the trunk. 
No anchylosis exists in either joint, passive movements can 
be made though there is no power of voluntary movement. The 
left leg is atrophied to a half, the knee is kept in an inverted 
position, and the foot is both slightly extended and inverted. 


258 DR TUKE. 


The knee-joint is capable of slight passive movement, but is 
not so as regards voluntary action. The ankle-joint is inca- 
pable of either. Progression is solely performed by the right 
leg, the left being lifted and carried after by the pelvic muscles. 

The atrophy of the trunk is best seen in the thorax. The 
thorax is flat, its antero-posterior diameter being 6.3, inches; 
its left side is lower than the right. The circumference is 31} 
inches, the right half being 17} inches, the left 152. 

The head has the following peculiarities and measurements. 
The forehead is low and slightly retreating. The right side is 
higher than the left, the vertex being formed by the right pari- 
etal bone. The occipital region is drawn towards the right 
shoulder. Circumference measures 22 inches, antero-posterior 
diameter 7,7, inches, transverse auricular diameter 63, inches, 
from ear to ear over vertex 12} inches, and from between supra- 
ciliary ridges to occipital protuberance 143 inches. 

Nervous System.—The epileptic fits occur about once a week, 
and are of medium severity. The convulsions last about half 
a minute, the face beimg very much drawn to left shoulder. 
The coma is profound. The eyes do not act in a co-ordinate 
manner, the action of the left being impaired. 

Mental Phenomena.—Expression of face imbecile; mouth 
always half open, saliva runs from it; smile and laugh childish. 
Articulation very imperfect and deficient. He can utter a few 
words in a childish manner. A marked feature in his case is 
constant singing, his great fondness for it, and the correctness 
of his tunes, which are limited in number. His affection is 
strong for his companions, but his temper is very bad and irrita- 
ble. Ideation very limited, if any. 

Course of Case. In May, 1872, signs of Phthisis pulmo- 
nalis were discovered. This disease made rapid progress, and he 
died on Oct. 2nd of the same year, at the age of 37 years. 


Report of Post-Mortem Examination. 


General Condition—Body much emaciated, yet unilateral 
atrophy, as described in case, very marked. 

Head.—Scalp thin; calvarium unequally thin, very dense, 
and of ivory hardness in certain places; diploe present only on 
left side of frontal bone, and in one limited spot, in right. The 


A CASE OF HYPERTROPHY. 259 


thickness of left side of frontal bone above supra-orbital notch 
was ;4, of an inch, and at corresponding part of right side %. 
Other parts of the circumference were also unequal. The skull 
was large, the dome high, ridged at the summit, to the right 
of which the parietal bone was round and full, to the left it 
was flattened. The circumference was 21%, in., the lateral 
measurement from nasal eminence to occipital protuberance on 
right side being 10,%; in., on left 103 in. The antero-posterior 
measurement from same points over vertex was 138 in., the 
transverse from one meatus auditorius to the other over vertex 
was also exactly 13%in., the measurement on the right side 
being 6% in., and on left 6Z in. The depth of dome from fora- 
men magnum was 52 in., the antero-posterior diameter 7 in. and 
the transverse 518 in. The capacity was 110} cubic inches. 

The sutures were very faintly marked, ossification being very 
far advanced, especially in the sagittal. 

The facial portion presented the following abnormalities :— 
the external inferior angle of the right orbit on a lower level 
than the left, and the right infra-orbital foramen correspond- 
ingly lower, as was also the inferior border of the right nasal 
aperture ; in short, the right side of the face was on a lower level 
as a whole than the left. The molars in the right upper jaw 
were only two in number (wisdom tooth being undeveloped) ; 
in lower jaw the same condition existed. 

External base showed a very thinned state of the bones 
immediately posterior to the posterior condyloid foramina; on 
the left side this spot was a mere shell. The left condyloid 
articular surface was divided into two facets. The ridges on 
the occipital bone were well marked. 

Internal surface. The superior longitudinal sinus took a 
sharp turn to the left about an inch above the point of the 
lambdoidal suture. The sutures were indistinguishable. On 
the inner table of frontal bone, on either side of longitudinal 
sinus, were numerous small dense bony elevations to which the 
dura mater was adherent, especially on the right side. 

The internal base presented the most marked abnormalities. 
Between the anterior fosse a sharp ridge ran upwards and 
forwards for more than two inches, and where it ended the 
shallow and irregular superior longitudinal suleus began. Both 


260 DR TUKE. 


the anterior and middle fossee were very rugose. The area of 
the right middle fossa was somewhat greater than that of the 
left. The body of the sphenoid bone was out of the mesial 
line to the left. The petrous portion of the left temporal bone 
differed in shape materially from the right; the groove of the 
lateral sinus of the left side had become expanded into a con- 
siderable antrum and changed the size and form of the foramen 
Jacerum posterius. This sulcus bifurcated from the superior 
Jongitudinal markedly to the left of the mesial line, the pos- 
terior fosse being thus rendered of different areas, the left 
being about half that of the right. The twist to the left of the 
superior longitudinal sinus caused the cerebral fossee of the 
occipital bone to be unequal. The groove of the left lateral 
sinus was well marked. The right was very faintly imdicated. 
A rugged exostosis 3ths of an inch long and {th of an inch high 
grew from the upper surface of the basilar process into the 
eranial cavity’. 

Membranes.—At vertex, on left side, all the membranes 
were adherent to each other. The vessels of the pia-mater, 
especially the veins, were deeply injected. Lateral sinuses gorged 
with dark blood. A bony plate, } inch by } inch, was found on 
the first left frontal convolution about its middle, between 
dura mater and arachnoid, and attached to both. At the cor- 
responding part on right side smaller particles of bone were 
found. 

Brain—general characters.—Of large size, heavy (60 0z.), 
firm consistence, tough and resilient. No impression was made 
upon it by a column of water 5 feet in height. The left oc- 
cipital lobe was perceptibly larger than the right: the relative 
weights of the two hemispheres were, right 304, left 23$0z.: 
the difference being 6j0z. The right ascending frontal convo-_ 
lution was very tortuous, and the fissure of Rolando joined the 
great longitudinal fissure further back than usual. ‘The fissure 
of Rolando on the ieft side did not reach the great longitudinal 
fissure, being bridged across at the vertex by a knuckle which 


1 Such exostoses have been described by Virchow (Entwickelung des Schi- 
delgrundes, 1857), who ascribes them to ossification of an elevation of the car- 
tilage of the basicranial synchondrosis, and Professor Turner has noted the same 
condition in a scaphocephalic cranium. (Vide Paget’s Lectures on Surgical 
Pathology, 3rd edition, edited by Turner, p, 531.) 


A CASE OF HYPERTROPHY. 261 


joined the ascending frontal and parietal convolutions. The 
third left frontal convolution was much better defined than 
the right. On the right side, a knuckle from the second frontal 
interposed between the portion bordering the fissure of Sylvius 
and that running down to the island of Reil. 

The specific gravity of the two hemispheres as proved en masse 
was exactly 1036: that of the cerebellum slightly greater. 
Separate portions of grey and white matter taken from various 
localities on both sides were tested and found to be sp. gr. 1036. 

On section there was a marked difference between the grey 
matter of the two hemispheres as to colour and thickness. The 
left hemisphere had the grey matter of all its convolutions 
markedly darker than the opposite side; the colours were on 
the left dark grey, on the right light pink grey, in the outer 
layers; the inner ones being of a darkish brown, a white band 
separating them. The line of demarcation was obvious to the 
naked eye. The grey matter of the left side, as measured by 
Dr Major’s tephrylometer’, varied from ;{, of an inch in the 
frontal lobes to 4, in the occipital. On the mght side it was 
much thicker and much more even over the whole hemisphere. 
In the frontal and anterior convolutions of the parietal lobe 
the thickness was 4, of an inch; in the posterior convolutions 
of the parietal (more especially the annectant convolutions) 
and the occipital lobe the thickness was #13, of an inch. 

On viewing the superior surface of the cerebellum, the 
left side was found to be one-third less in bulk than the right. 
The folia of the left side were narrower than those of the ght. 
The notch or vallum was filled by an abnormal lobule, springing 
from the inferior surface. On inspection of inferior surface, 
the left hemisphere was found, although much smaller than the 
right, more normal in its arrangement, for its lobules were fairly 
defined. The vallecula was but very slightly marked. The 
notch was represented by a slight depression. There was no 
appendix vermiformis. Proceeding from above downwards on 
the right side, the lamellz, immediately below the longitudinal 
fissure, were normal for two-thirds of their extent. The biven- 
tral lobe could be traced; also the amygdala, although its 


1 See a description of this instrument in West Riding Asylum Reports, 1, 
and this Journal, vit. 169. : 


262 DR TUKE. 


appearance was quite abnormal, inasmuch as it was composed 
of several convoluted lamelle, its shape was round, and it was 
unduly elevated. The slender and posterior inferior lobes were 
represented by a confused mass of lobules of convoluted laminz 
extending right across the vallecula, and attached to the oppo- 
site side by membranes. 

No difference was observed in the size of the corpora 
striata, optic thalami, or in any other of the organs of the 
cerebrum. The pons varolii was symmetrical. The medulla 
oblongata was markedly larger on the right side above the 
point of decussation. The spinal cord was symmetrical, and 
no difference could be observed in the size of the ganglia of 
the different sides. The vessels were deeply injected, and a 
small amount of serum was found. A marked difference ex- 
isted in the calibre of the vertebral arteries, the right being 
about twice the size of the left. 

Microscopic Examination.— Cerebellum. In a specimen taken 
from the grey matter of the deformed portion of the right 
hemisphere, several well-marked cells of Purkinje were seen. 
Vessels normal. On the left side the nerve fibres were pe- 
culiarly distinct and large. Cells of Purkinje and molecules 
were seen, the former large but not very distinct. In ex- 
amining a punctum vasculosum, the vessel forming it was 
carefully extracted, and the cellular coat was seen to be 
thickened, and the muscular fibres well pronounced. <A con- 
siderable deposit of hematin was found on the outside of the 
vessel. In a specimen prepared in chromic acid the structures 
appeared nearly normal, except that the granules of the granular 
layer were not so closely packed as usual. In the white matter 
spaces of molecular matter were seen which had taken the 
place of fibres. This specimen was taken from the deformed 
left postero-inferior lobule. On the right side no abnormality 
was noticed. 

Right Frontal Lobe.—In a recent specimen taken from the 
1st frontal convolution, the nerve-cells were found in greater 
abundance than elsewhere. A very fine molecular deposit was 
observed on the vessels, and the nuclei of neuroglia and of the 
vessels were proliferated and enlarged. The external granular 
layer was productive of great numbers of irregular round bodies, 


A CASE OF HYPERTROPHY. 263 


of the size of a blood-corpuscle. Microscopic section of the 
same convolution confirmed the above. The cells of the outer 
layers were small, rounded, irregularly distributed, and nu- 
merous; those of the two inner layers were well-marked, nor- 
mal im size and regular in arrangement, the poles long and 
well-defined. The fibres, although thickened, were properly 
distributed. In some places they were rather widely separated. 
The external layer was thickened. 

Left Frontal Lobe-—The only difference between the 3rd 
frontal convolutions of either side was the definite evidence 
of increase of connective tissue in the right; in it 4, 5 or 
6 fibres could be seen pushed together into fasciculi, be- 
tween which connective tissue and nuclei of neuroglia existed 
in large quantities. In the white matter few fibres could be 
traced, although several clearing agents were employed, tur- 
pentine, glycerine, and oil of cloves. The molecular matter 
was increased and arranged in a fibroid manner. Taken alto- 
gether, the constituents of these lobes were more nearly ap- 
proaching the normal condition than the other parts of the 
brain. 

Right Parietal Lobe—Various recent specimens taken from 
the right ascending parietal convolution, near vertex, showed 
the nerve-cells to be scarce and but faintly defined. A chromic- 
acid preparation showed the large cells of the deepest layer of 
grey matter well, with the poles traceable for long distances. 
There was a brownish molecular discoloration at their bases, 
but not more than is often observed in health. The smaller 
cells were irregular in shape and distribution and without per- 
ceptible nuclei. The increase of neuroglia was well-marked, 
the nuclei were much proliferated in plasm and on vessels. 

Right Occipital Lobe.—In several specimens taken from this 
region and examined in a recent state, and by section after 
being steeped in chromic acid, there was seen a very evident 
increase of molecular matter between the nerve elements; with 
this were associated fine fibrillar structures. The fibres were 
much enlarged, and were moniliform. Both these conditions 
might have been to some extent due to pressure, but the ex- 
perience gained from many brains, examined under exactly 
similar circumstances, leads to the conclusion that it was not 


264 DR TUKE. 


altogether so. On the capillary vessels a small amount of very 
fine colourless molecular matter was sparsely scattered. The 


Portion of white matter of right occipital lobe. The lines represent the nerve 
fibres. The intermediate nucleated material is the hypertrophied neuroglia. 


grey matter was examined in successive slices from without 
inwards. Very few cells were found, and these were im no way 
actively diseased; they were atrophied to a great extent; were 
small in size, irregularly shaped and ill-defined. In some chromic 
acid sections a fine layer of yellow molecular matter was found 
between the pia-mater and the brain substance, presenting 
the appearance of pale hematin. In the same sections the 
fibrils became fainter as they proceeded inwards; when more 
than half an inch from the grey matter, the fibres disappeared, 
and the field of the microscope was filled with fine molecular 
matter arranged in a fibroid manner. 

In the Left Occipital Lobe the microscopic appearances ap- 
proached more nearly to those of health. The fibres were 
not displaced, and there was not the same evidence of increased 
connective tissue. The cells were in the same atrophied con- 
dition, and did not occur in any greater number. 

Remarks.—1 am inclined to regard this case as one of uni- 
lateral hypertrophy of the brain, primarily due to acute hydro- 
cephalus, and eventually resulting in atrophic hemiplegia. Al- 


bo 


. A CASE OF HYPERTROPHY. 265 


though no great stress need be laid on the statement of the 
mother as to the causation of the hydrocephalus and the fits, 
the general characteristics of the calvarium indicate that the 
former did exist to a certain degree, and that it was due to 
effusion into or below the arachnoid sac. As occasionally hap- 
pens the calvarium was asymmetrically dilated. The disease 
having been arrested after the skull had become ossified and the 
fluid being gradually absorbed, a compensatory substance was 
demanded, which was supplied by an increase of the neuroglia 
of the right hemisphere pari passu with the absorption. This — 
sclerosis of the packing substance of the right hemisphere im- 
pairing its trophesial influence, resulted in arrestment of develop- 
ment of the opposite side of the body. It is a matter of every- 
day experience that bilateral atrophy of the body occurs in the 
subjects of bilateral brain disease, and that deformed and 
rickety dwarfs are very frequently possessed of abnormally large 
heads. It appears to me more than probable that in the latter 
class both hemispheres of the brain have undergone the hyper- 
trophy which was demonstrated in the right hemisphere only 
of the case under consideration. 

In Van der Kolk’s’ case of atrophy of the left hemisphere 
of the brain with coexistent atrophy of the right side of the 
body, the right hemisphere of the cerebellum, and the left side 
of the spinal cord were found atrophied. In the case under 
consideration the right hemispheres of both cerebrum and cere- 
_ bellum were hypertrophied, and no difference could be detected 
in the size of the right and left sides of the cord, or of its 
ganglia; in fact, the sclerosis was symmetrical, and did not 
extend below the decussation in the medulla. 

It may here be mentioned that this is the fourth case of 
disease of the cerebellum which has come under my notice. 
In three atrophy was the abnormal condition, and it had impli- 
cated in one case five-sixths of the organ: in the other two one 
hemisphere was destroyed. In none of these cases was there 
any psychical or physical indication during life in consonance 
with theories which are held by some physiologists as to the 


1 Case of atrophy of the left hemisphere of the brain with coexistent atrophy 
of the right side of the body, by J, L. C. Schroeder Van der Kolk, translated by 
the New Sydenham Society, 1861. 


VOL. VII. 18 


266 DR TUKE. A CASE OF HYPERTROPHY. 


functions of the cerebellum. In the three cases of atrophy 
there were no symptoms at all. 

It does not appear from Van der Kolk’s monograph that he 
determined by microscopic sections which of the brain elements 
were atrophied. In my case there exists no doubt that the 
general sclerosis produced a distinct and definite change in the 
continuity and arrangement of cells and fibres, and that im- 
paired trophesis was the result. I deeply regret that the exami- 
nation was not completed by accurate measurements of bones 
and of muscular and nerve fibres, in the same exhaustive 
style as characterized the researches of this eminent observer. 
It is more than probable that in his case the chief loss of 
brain substance occurred in the neuroglia; my experience 
at least bemg that it is the first of the cerebral constituents 
to give way. Whatever may have been the condition, it is 
interesting to note that both hypertrophy and atrophy produce 
the same results. 

It is worthy of consideration to what extent the exostosis 
might have had influence on the epileptic convulsions and the 
permanent tonic spasm. [f can find no definite statement as to 
the period of life at which ossification of the basicranial synchon- 
drosis occurs. It is generally said to take place “at an early 
age,” and Henle states that the suture becomes obliterated after 
the 20th year. The faet of the return of the epilepsy at the age 
of ten, is suggestive that ossification of the cartilaginous eleva- 
tion may have been the cause, but nothmg can be founded on 
the suggestion. It is more likely that the chronic tonic spasm 
may have depended on this constant cause of irritation: but 
the most probable hypothesis is that it had no effect im the pro- 
duction or maintenance of either epilepsy or spasm. 


ON THE ORDER DINOCERATA (Marsh). By A. H. 
Garrop, B.A., Cantab. 


It is very seldom that specimens are now obtained, either fossil 
or recent, of mammalian forms that have, provisionally at least, 
to be placed in a new order by themselves, on account of the 
presence of peculiarities hitherto quite unknown and unex- 
pected. Such, however, is the case with a large series of fossils 
which have been, within the last two years, obtained from the 
Eocene deposits of Wyoming in North America, Our chief 
source of information respecting these remains consists of a 
series of papers by Prof. D. C. Marsh of Yale College, abstracts 
of which have recently appeared in The American Journal of 
Science, and in The American Naturalist. Prof. E. D. Cope has 
also published several papers on the subject, and some excel- 
lent photographs of one of the most important forms, sent by 
him to this country, it has been my good fortune to see. Prof. 
Leidy has also named one of the genera. Most of my infor- 
mation is obtained from the papers by Prof. Marsh on the Order 
Dinocerata, together with the critical remarks made by him on 
Prof. Cope’s descriptions, which I have also by me. 

Dinoceras mirabilis is the name given by Marsh to a huge 
- ungulate animal, of which there is an almost complete skeleton 
in the museum of Yale College. Its size must have been very 
nearly that of a full-grown Elephant, as the length of the skull 
of a closely allied genus was a little more than a yard. The 
skull was peculiarly long and narrow, and supported three pairs 
of horn cores in rows, on its superior surface. The anterior 
pair were situated on the anterior ends of the nasal bones, they 
were short, conical, and directed nearly straight upwards. The 
median pair were conical prolongations upwards from the maxil- 
laries, they were longer and more cylindrical than those on the 
nasals, and the fangs of the huge canine teeth entered their 
bases. The posterior pair, the largest, were very peculiar, being 
extensions upwards from near the middle of large lateral longi- 
tudinal crests, which were formed by the occipital, parietals, 

18—2 


268 MR GARROD. 


and frontal bones. The dentition was peculiar, the upper inci- 
sors were deficient, and the premaxillee consequently small. 


Skull of Dinoceras mirabilis, copied from Prof. Marsh’s figure, and kindly 
lent by the Proprietors of Nature. 


The upper canines formed huge downwardly directed tusks, 
nearly straight, but directed somewhat backwards. These, after 
a gap equal to their breadth, were followed by six small molar 
teeth. Each of these was composed of two well developed 
nearly straight, transverse ridges, which joined at the imner bor- 
der of the tooth and diverged as they ran outwards, so forming 
a simple >-shaped pattern, that continued well marked till 
the tooth was much worn, The posterior of the two ridges was 
quite transverse, and the anterior was slightly concave back- 
wards, and ran obliquely forwards and outwards. These teeth 
diminished in size from before backwards gradually, and there 
were no intervals between them. All that I can find about the 
lower jaw is, that it was slender, and its tusks small. 

The orbit was not separated from the temporal fossa, which 
latter was large, extending up the outer side of the lateral pa- 


THE ORDER DINOCERATA. — 269 


rieto-o¢cipital crest. The malar completed the anterior portion 
of the zygomatic arch; the lachrymal was large, forming the 
anterior border of the orbit, a large oval foramen perforated its 
facial surface. The squamosal sent down a large post-glenoid 
process. ‘The nasals were massive and greatly prolonged for- 
wards, at the tip carrying the anterior horn-cores; they also 
sent slight processes up the inner faces of the maxillary horn- 
cores. The premaxillaries were peculiar in that they almost 
enclosed the anterior nares; they united posteriorly with the 
maxillaries just in front of the canines, and then divided into 
two branches, one of which, the lower, corresponding to the 
premaxillary of the Rhinoceros, ran forwards free, the other 
closely uniting with the adjoining nasal went upwards to 
strengthen the support of the horn-cores. 

The extremities were tetradactylate, and in the pes the 
astragalus articulated with the cuboid as well as with the sca- 
phoid bone. The humerus was short and massive, with the 
great tuberosity not rising above the articular head, and the 
condylar ridge of the distal end not continued up the shaft. 
The radius which was free, was not so oblique as in the Ele- 
phant. The femur had no pit on its head for the ligamentum 
teres, and there was not any third trochanter. The phalanges 
were short as in the Elephant. 

The number of the vertebrae are not noted by Marsh, so 
that I cannot say how many there were in the dorso-lumbar 
region, a point of great importance. There were four in the 
sacrum, the last being small and supporting a slender tail. The 
ribs had rudimentary uncinate processes. 

The name Tinoceras has been given by Marsh to a very 
closely allied species, which differs from Dinoceras in having 
the anterior, or nasal-horn cores compressed on the top, larger 
and projecting more forward; the maxillary horn-cores are also 
proportionately longer, more cylindrical, and directed slightly 
forward. The photographs of this genus above referred to 
shew very clearly that the palate was completed opposite the 
posterior molars by the palatine bones. The differences be- 
tween Tinoceras and Dinoceras seem to be scarcely generic. 
Another closely allied genus, of which I have seen no descrip- 
tion, is the Uintatherium of Leidy. The name Zobasileus, in- 


270 MR GARROD. THE ORDER DINOCERATA. 


troduced by Prof. Cope, is a synonym of Tinoceras, and being 
of later introduction, must be sunk. 

From the facts given above, Prof. Marsh is led to placing 
these undoubtedly peculiar animalia in an order different from 
any yet established, intermediate between the Proboscidia and 
the Perissodactylata. To me it seems much more probable that 
they belong to the Ungulata proper, and that no separate order 
is necessary for their reception. In the characters I have given, 
there are none which shew them to have any true Proboscidian 
affinities, whilst there are several which seem to indicate that 
they belong to a family of the Artiodactylata, and not to the 
Perissodactylata. Among the reasons in favour of Dinoceras 
and its allies being Artiodactylate are the following: 


The astragalus has a well-developed cuboid facet. 
The palate is complete between the posterior molars, 
There is no third trochanter to the femur. 

The premaxillae are edentulous. 

The anterior premolar is not devoloped. 


A ola a OM 


ON THE SO-CALLED PRICKLE OR CLAW AT THE 
END OF THE TAIL OF THE LION AND OTHER 
FELINES. By Proressor TuRNER. 


Homer, in the 20th Book of the Ziad, when describing how 
the God-like son of Peleus rushed on the high-souled, warlike 
fEneas, compared him witb a destructive lion— 

“As he lashes 


Fiercely his sinewy flanks with his tail, each side in succession 
Rousing himself for the fight.” 


Various authors and critics have supposed that the tip of 
the lion’s tale is armed by a pointed sting or prickle, and that 
when the animal uses his tail in the manner so vividly de- 
scribed by the poet, he goads himself by its prickly sting prior 
to making a rush on his prey. Although comparative ana. 
tomists generally had taken no notice of any such structure, 
yet some support was given to the opinion formed of its 
existence by the authors referred to, by a statement made by 
Blumenbach in his work on Natural History, who said that he 
had seen something like a sting in the tail of a lion. 

Some years ago this subject was investigated by Prof. 
Leydig, who wrote a short but interesting Essay, entitled Ueber 
den Schwanz-stachel des Lowen, in Reichert und du Bois-Rey- 
- mond’s Archiv for 1860. Leydig had examined in the previous 
winter the tail of a splendid lion which had died in the Zoolo- 
gical Garden at Stuttgart, and found, on turning on one side the 
hair which covered the end of the tail, a perfectly smooth and 
hairless papilla, rundlich-kegelformig in shape, with a con- 
stricted base and an elevated apex. Leydig made a careful 
microscopic examination of this papilla, and came to the con- 
clusion that it was not a special and peculiar organ, but merely 
a papilla of the skin endowed with vessels and nerves, so as 
in all probability to be an organ of sensibility like a touch 
organ at the point of the finger. Leydig also refers to a 
monograph with the title Der Stachel des Léwen an dessen 
Schweifende, published anonymously at Darmstadt in 1855, in 
which the author described the ‘sting’ or ‘ prickle’ in the tail 


Ns 


(2 PROFESSOR TURNER. 


of a recently dead lion, and in several stuffed specimens. This 
author also stated that the Puma, Felis concolor, possessed a 
similar structure, and that in Bos urus, some species of Macro- 
pus, and various genera of monkeys a nail-like structure had 
been observed at the end of the tail. 

A few years ago Dr J. E. T. Aitchison, now British Commis- 
sioner in the Punjab, read a paper before the Royal Medical 
Society of Edinburgh, in which he stated that a claw-hke 
appendage was to be found at the tip of the tail in the large 
felines, but doubts were expressed at the time as to the ac- 
curacy of his statements. Dr Aitchison has however availed 
himself of the opportunities afforded by his residence in the 
Punjab to secure the tails of two leopards, which he forwarded 
for examination early in the present year to my colleague, the 
Professor of Natural History. Owing to the absence of Dr 
Wyville Thomson, in charge of the Deep Sea Exploring Expe- 
dition, the specimens have been handed over to me. 

On turning on one side the hairs at the tip of the tail of 
one of the specimens, I found a grey-coloured, hard, hairless 
conical structure, which projected for ;2,ths of an inch beyond 
the roots of the nearest circle of hairs. It terminated in a 
sharp, prickly point, and possessed a diameter at its base, of 
not more than jth inch. It exhibited two circular constric- 
tions, which gave me at first sight the impression that it might 
be formed by very minute terminal caudal vertebre, but this 
impression was not borne out by more careful observation. 

In the second specimen, the hard structure only projected 
zth inch beyond the roots of the nearest circle of hairs, and 
was slightly broader at the base than in the first specimen. It 
did not end in a sharp point, but had the form of a rounded, 
hairless, nipple-like projection. In neither instance could the 
structure be seen until the hairs were turned on one side, 
although in both cases their hardness and slight projection 
enabled me to feel them without difficulty. 

I soaked the latter specimen in water so as to soften it, 
to enable me to examine. its connections. I then carefully 
cut away the hairs in its immediate neighbourhood, and ex- 
perienced no difficulty in seeing that it was nothing more than, 
the hairless termination of the integument at the end of the 


CLAW AT THE END OF THE TAIL OF THE LION. Dia 


tail, which though hard and horny in the dried condition, was 
now soft and flexible, like the ordinary skin of the tail with 
which it was continuous. 

I then scraped some soft whitish material off the surface, 
and examined it microscopically. It consisted of well-defined 
nucleated, and stratified squamous epidermal cells. A thin slice 
of the sub-epidermal tissue made perpendicularly to the surface, 
was then placed under the microscope. In it I recognised a 
number of small papille, having the structural characters and 
mode of arrangement of the papille of the cutis. There could 
be no doubt, from its structural characters, that the so-called 
claw or prickle at the end of the leopard’s tail, as Leydig had 
already pointed out in the case of the lion, had the structure of 
skin, but imstead of consisting, as in the lion, of a single large 
papilla, it possessed numerous small papille. Notwithstanding 
the poetical idea of its function as a sting, it is to be regarded 
therefore, not as a specially developed organ, but merely as a 
hairless part of the integument. 


ON AN EDENTULOUS CONDITION OF THE SKULL 
OF THE GREY SEAL (Halichoerus gryphus). By 
PROFESSOR TURNER. 


Ty Vol. v. of this Journal, p. 270, I stated that specimens of 
the grey seal had recently been captured on the East coast of 
Scotland, and I referred to two young animals caught in the 
salmon nets near Montrose. Since those specimens were re- 
corded, I have received the skull of another animal of that 
species from the same locality, which exhibited a remarkable 
defect in the development of the teeth. 

The skull was from an animal about half-grown, and mea- 
sured 8$ inches in length. The basi-cranial synchondroses were 
unossified, and the sutures between the bones of the face and 
those between the bones of the cranial vault were so loose, that 
many of the bones could, without much force, be separated 


274 ~ PROF. TURNER. SKULL OF THE GREY SEAL. 


from each other. No teeth were present in the upper jaw, 
except the pair of canines, which were well developed, pro- 
jected ;4,ths of an inch beyond their sockets, and occupied their 
usual position immediately behind the maxillo-premaxillary 
sutures. Similarly no alveoli existed in the upper jaw, except 
those in which the canine teeth were lodged. 

There was a complete absence also of teeth and alveoli in 
the lower jaw except the pair of canines and the sockets in 
which they were lodged. Owing to the absence of teeth the 
dentary borders of both upper and lower Jaws were much nar- 
rower than in a well formed cranium, and the vertical diameter 
of the horizontal ramus was appreciably smaller than in a 
normal jaw of the grey seal of the same size. The dentary 
borders of both jaws were studded with small foramina, which 
opened into canals apparently for the transmission of blood- 
vessels. 

The Anatomical Museum of the University contains a skull 
of this species of seal of almost precisely the same dimensions, 
which originally formed a part of the Monro collection, but of 
which there is unfortunately no record of the locality whence it 
was obtained. In this skull the dentition is perfect; all the 
teeth project prominently from their sockets, and present the 
following formula— 

5/1,3) 3-15 

Geile? (2a 
It is clear therefore that the grey seal at this period of life has 
completed its permanent dentition, and that the peculiarity 
exhibited by the cranium I have described is due either to a 
non-development, or premature atrophy, of all the teeth of the 
permanent series, except the upper and lower canines. This 
edentulous cranium is not without interest in connection with 
the consideration of the zoological affinities of the Seals to the 
Cetacea. 


ON THE PHYSIOLOGICAL ACTION OF LIGHT. No. I. 
By James Dewar, Esq. and Jonn G. M°KeEnprRick, M.D., 
of the Unwersity of Edinburgh’. 


It is the object of this communication briefly to describe some 
of the results we have already obtained in an investigation into 
the Physiological Action of Light on which we are at present 
engaged. We have more especially directed our attention to 
the problem of the specific effect produced on the retina and 
optic nerve by the action of light. Numerous hypotheses have 
been made from time to time by physicists and physiologists ; 
but up to the present date our knowledge of the subject is 
without any experimental foundation. 

It is evident that, in accordance with the principle of the 
transference of energy now universally accepted, the action of 
light on the retina must produce an equivalent result, which 
may be expressed, for example, as heat, chemical action, or 
electro-motive power. It is well known that the electro-motive 
force of a piece of muscle is diminished when it is caused to 
contract by its normal stimulus, the nervous energy conveyed 
along the nerve supplying it; and similarly a nerve suffers a 
diminution of its normal electro-motive force during action. In 
the same manner the amount and variations of the electro- 
~ motive power of the optic nerve affected secondarily by the 
action of light on the retina are physical expressions of certain 
changes produced in the latter ; or, in other words, are functions 
of the external exciting energy, which in this case is light. 
Considerations such as these led us to form the opinion that 
the problem of what effect, if any, the action of light has on the 
electro-motive force of the retina and optic nerve would require 
for its investigation very careful and refined experiment. 

The enquiry naturally divided itself into two parts, first, to 
ascertain the electro-motive force of the retina and nerve; and 
second, to observe whether this was altered in amount by the 
action of light. With regard to the first of these questions, 


1 Abstract of paper read before the Royal Society of Edinburgh, April 21, 
1873. 


276 MR DEWAR AND DR M°KENDRICK. 


Du Bois-Reymond found, while pursuing his great investigations 
into the electro-motive properties of living tissues, that every 
point of the external surface of the eyeball of a large Tench was 
positive to the artificial transverse section of the optic nerve, 
but negative to the longitudinal section. This he accomplished 
by the use of his well-known non-polarizable electrodes formed 
of troughs of zinc carefully amalgamated, containing a solution 
of neutral sulphate of zinc, and having cushions of Swedish 
filter paper and clay-guards, on which to rest the preparation. 
These electrodes were connected by him with a galvanometer, 
and the preparation was placed so that the eyeball, carefully 
freed from muscle, rested on the one clay-guard, while the 
transverse section of the optic nerve was in contact with the 
other. By following Du Bois-Reymond’s method we have had 
no difficulty in obtaining a strong deflection from the eyes of 
various rabbits, a cat, a dog, a pigeon, a tortoise, numerous 
frogs, and a gold fish. 

In the investigation of the next question, namely the effect 
of light on the electro-motive force, we found more difficulty. 
The method followed was to place the eyeball on the cushions 
in the manner above described, to note the deflection of the 
galvanometer needle, and then to observe whether or not any 
effect was produced on the impact of a beam of light, during 
its continuance, and on its removal. In a few of our earlier 
experiments we used Du Bois-Reymond’s multiplying galva- 
nometer, but finding the amount of deflection obtained was 
so small, that the effect of light could not be readily observed, 
we have latterly used Sir W. Thomson’s exceedingly sensitive, 
reflecting galvanometer, kindly lent us by Professor Tait. We 
met also with secondary difficulties, such as the dying of the 
nerve, the impossibility of maintaining an absolutely constant 
zero, and an absolutely constant amount of polarity, the effects 
of heat, &c.; but these difficulties we have overcome as far as 
possible by the most approved methods. The changes in 
polarity of the apparatus occurred slowly, and could not be 
mistaken for the changes produced by the action of light, which 
we found occurred suddenly and lasted a short period of time, 
It is also important to state that the deflections we observed do 
not at present profess to be absolute, but only relative values. 


THE PHYSIOLOGICAL ACTION OF LIGHT. O77 


The effects of heat were carefully avoided by covering over the 
electrodes on which the eye under examination rested with a 
spherical shell of water, of at least an inch in thickness, by 
means of a properly constructed glass apparatus. 

The result of our work at present is that we have been able 
to demonstrate that the action of light on the retina alters the 
amount of the electro-motive power to the extent of from three 
to seven per cent. of the total amount of the natural current. 
This is most readily seen by using the eye of the Frog. On 
placing the eye on the cushions, so that the eyeball rests on the 
one cushion and the transverse section of the nerve on the other, 
there is no difficulty in obtaming a deflection of between 300 and 
400 degrees on the scale of our reflecting galvanometer. This 
deflection slowly decreases for a time, until it reaches a point 
where it is, for a space of half an hour or an hour, comparatively 
steady, in amount. We then find the eye to be remarkably 
sensitive to light. A flash of light, lasting the fraction of a 
second, alighted match held at a distance of 4 or 5 feet, the 
light of a small gas jet suddenly turned on, coloured light pro- 
duced by allowing light from a small gas-flame enclosed in a 
lantern, to pass through a globular glass jar (12 inches in 
diameter) filled with a solution of ammoniacal sulphate of 
copper or bichromate of potash, have all produced remarkable 
changes in the amount of the electro-motive power, 

When a diffuse light is allowed to impinge on the eye of the 

frog, after it has arrived at a tolerably stable condition, the 
natural electro-motive power is in the first place increased, then 
diminished, during the continuance of light it is still slowly dimin- 
ished to a point where it remains constant ; and on the removal 
of light, there is a sudden increase of the electro-motive power 
nearly up to its original position. The alterations above re- 
ferred to are variables depending on the quality and intensity of 
the light employed, the position of the eyeball on the cushions, 
and modifications in the vitality of the tissues. 
_ Similar experiments made with the eye of warm-blooded 
animals, under the same conditions, have never given us an 
initial positive variation, as we have above detailed in the case of 
the frog, but always a negative variation. The after mductive 
effect on the withdrawal of light occurs in the same way. 


278 MR DEWAR AND DR M°KENDRICK. 


A large number of experiments has been made with different 
portions of the spectrum, which all tend to shew that the 
greatest effect is produced by those parts of the spectrum that 
appear to consciousness to be the most luminous, namely, the 
yellow and the green, 

Similarly, experiments made with light of varying imtensity 
shew that the physical effects we have observed vary in such a 
manner as to correspond closely with the values that would 
result if the well-known law of Fechner was approximately 
true. 

The success of these preliminary experiments leads us to be- 
lieve that by the employment of proper appliances this method 
of research may be greatly extended, not only with regard to 
vision, but also to the other senses. We expect soon to be ina 
position to communicate quantitative results involving time asa 
variable element in the case of the action of light on the retina 
and optic nerve, and we also intend prosecuting similar observa- 
tions on the terminal structures and nerves of the other organs 
of special sense. 


ON THE PHYSIOLOGICAL ACTION OF LIGHT. No. I. 
By James Dewar, Esq., and JoHN G.M *Kenprick M.D.' 


In continuing our experimental enquiry on the Physiological 
action of Light on the Eye, we have found it necessary to con- 
struct a true graphical representation of the variations of the 
electro-motive force occasioned by the impact and cessation of 
light. It is clear that to register minute galvanometrical altera- 
tions, the only plan that could be employed would be to pho- 
tograph on a sensitive surface, covering a cylinder rapidly re- 
volving on a horizontal axis, the alteration of position of the 
spot of light reflected from the mirror, just as continuous mag- 
netic observations are registered. As the apparatus required 
to execute these observations is very complicated, and would 


1 Abstract of paper read before the Royal Society of Edinburgh, May 5, 1878. 


THE PHYSIOLOGICAL ACTION OF LIGHT. 279 


require much preliminary practice, we have in the mean time 
adopted a simpler method of registration. This plan is to note 
the position of the galvanometer at equal intervals of time, be- 
fore, during, and after, the impact of light on the eye. In these 
observations we have used a seconds pendulum giving a loud 
beat. One observer reads aloud the galvanometer, the other 
marks every interval of two and a half seconds, registers the 
numbers obtained, and regulates the supply of light. A little 
practice in the method above deseribed has enabled us to ob- 
tain very satisfactory results, agreeing very closely in different 
observations, and shewing in a decided way the salient poimts 
of the variation curve. Figures 1 and 2 represent two curves 
thus obtained shewing typical forms of light variation. 


The central dotted line would represent the stable position 
of the electro-motive force in the dark. The curve above 
this line represents increase of the electro-motive force, while 
that below it is the reverse. The curve shewn in Fig. 1 
represents the change observed when the cornea was in con- 
tact with the one cushion, and the transverse section of the 
nerve in contact with the other. On the other hand, the 
eurve delineated in Fig. 2 is that obtained by placing the 
sclerotic instead of the cornea on the cushion, the transverse 
section of the nerve touching the other cushion. These curves 
shew that on the impact of light there is in Fig. 1 a sudden 
increase of the electro-motive force; during the continuance 
of light it falls to a minimum value; and on the withdrawal 
of light, there is what we have formerly styled an induc- 
tive effect, that is to say, a sudden increase of the electro- 
motive force which enables the nerve to acquire its normal 
energy. In other words, the falling off of electro-motive force, 
shewn in the diagram, is the physical representative of what, 
in physiological language, is called fatigue; the inductive effect 


280 MR DEWAR AND DR M°KENDRICK. - 


exhibiting the return of the structure to its normal state. In 
the case of Fig. 2, the impact of light is followed by no in- 
crease. There is, however, a gradual rise during the continu- 
ance of light, and on the removal of light there is the inductive 
effect we have spoken of formerly. 

We have carried out, since our last communication, four 
distinct sets of observations. rst, as to the pigment-cells of 
the skin or choroid shewing any sensibility to light. It is 
well known there is no difficulty in obtaining a strong current 
from the skin of the frog, but we have not observed any sensi- 
bility to light. The same is the case with the pigment-cells 
of the choroid. Second, as to the action of different well-known 
poisonous.substances on the sensibility of the retina. We have 
found that Woorara, Santonin, Belladonna, and Calabar Bean, 
did not seem to destroy the sensibility to light. Third, as to 
the action of the anterior portion of the eye. On carefully bi- 
secting an eye of a frog, so as to remove completely the anterior 
portion including cornea, aqueous humour, iris, ciliary-muscle, 
and lens, and on bringing the retina into actual contact with 
one of the clay pads, we readily obtained a large deflection 
which was as sensitive to light as when the whole eye was em- 
ployed, thus eliminating any possibility of the contraction of 
the iris under the stimulus of light having to do with the re- 
sults previously obtained. On using the anterior portion of the 
eye so that the cornea and posterior surface of the crystalline 
lens were the poles, we obtained a large deflection, which was, 
however, insensible to light. The sclerotic and nerve without 
the retina, in the same manner, give a large natural electro- 
motive force, also not sensitive. The distribution of the electro- 
motive force between the different portions of the eye and cross 
section of the nerve may be stated as follows: The most posi- 
tive structure is the cornea, then the sclerotic, then the longi- 
tudinal surface of the nerve; the cornea is also positive to the 
posterior surface of the crystalline lens, and the retina itself 
seems to be positive to the transverse section of the nerve, 
Fourth, as to the effect of varying luminous intensity on the 
electro-motive force. Perhaps the most remarkable result ar- 
rived at during the course of our experiments is the small dif- 
ference between the amounts of alteration observed under con- 


THE PHYSIOLOGICAL ACTION OF LIGHT. 281 


ditions of varying luminous intensity. If a candle is placed at 
a distance of one foot from the eye, and then is removed ten 
feet, the amount of light received by the eye is exactly one 
hundredth part of what it got at a distance of one foot, whereas 
the electro-motive force, instead of being altered in the same 
proportion, is only reduced to one-third. Repeated experiments 
made with the eye in different positions has conclusively shown 
that a quantity of hght one hundred times in excess of another 
quantity only modifies the electro-motive force to the extent 
of increasing it three times as much, certainly not more. 

It was apparent to us that these experiments would ulti- 
mately bear upon the theory of sense-perception as connected 
with vision. It is now generally admitted that no image, as 
such, of an external object, is conveyed to the sensorium, but 
that im reality the brain receives certain mmpressions of altera- 
tions taking place in the receiving organ. The natural query 
then arises—are the physical effects we have described and 
measured really comparable in any way with our sensational 
differences in light perception when we eliminate all mental 
processes of association, &e., and leave only perception of dif- 
ference of intensity? In other words, are these changes the 
representative of what is conveyed to the sensorium? It would 
appear, at first sight, that this problem is altogether beyond 
experimental enquiry. There is, however, a way of arriving at 
very accurate measures of the variation of our sensational dif- 
ferences in the case of hght, and this has been developed theo- 
retically and experimentally by the justly renowned physiologist 
Fechner. Stating the law of Fechner’ generally, we may say, 
the difference of our sensations is proportional to the logarithm 
of the quotient of the respective luminous intensities. A recent 
series of experiments by Dalbceuf? has entirely confirmed the 
truth of this law. If, therefore, the observed differences in 
electro-motive power, registered under conditions of varying 
luminous intensity, agree with this law of Fechner, regulating 
our sensational impressions, then there can be little doubt these 
variations are the cause of, and are comparable to, our perception 
of sensational differences. Now, we have stated above, that 

1 Fechner, Elemente der Psychophysik. Helmholtz, Physiological Opties. 

2 Recent Memoir to Belgian Academy. 


VOL. VII. 19 


982 MR DEWAR AND DR M‘KENDRICK. ACTION OF LIGHT. 


with a quantity of light 100 times in excess of another quantity, 
the electro-motive force only becomes three times greater. Ac- 
cording to Fechner’s law, we may say the difference of our sen- 
sations, with that variation in the amount of luminous intensity, 
would be represented by 2, the logarithm of 100. Our experi- 
mental results being as 3 to 1, the difference is also 2, thus 
agreeing very closely. It is to be remembered, however, that 
these results have been obtained by experiment on the eye of 
the frog, but similar changes have been observed in the eyes of 
Mammals. In the latter, however, the amount of alteration is 
not so great, in all probability owing to the rapid death of the 
parts. 

In order to see how far we could trace this alteration of 
electro-motive force in the course of the optic nerve, we have 
made some experiments without removing the eye from the 
orbit, leaving the nerve intact. On placing the cross section 
of the posterior part of the optic lobe on one cushion, and the 
cornea on the other, a variation, amounting to about one per 
cent. of the electro-motive force, could still be obtained. 

Since the above results were communicated to the Royal 
Society of Edinburgh, we have made two experiments of some 
interest. In the first place, the same kind of results has been 
obtained by operating on the eye of the cat while it remained 
in the orbit of the living animal. Secondly, the eye of the frog 
has been experimented on with a beam of uncondensed moon- 
light and was found sensitive, thus eliminating all traces of 
radiant heat, as possible causes of the changes we have described. 


CAUSE OF THE RETARDATION OF *THE PULSE 
WHICH FOLLOWS ARTIFICIAL OR VOLUNTARY 
CLOSURE OF THE NOSTRILS IN THE RABBIT. 
A reply to certain criticisms. By WILLIAM RUTHERFORD. 
Professor of Physiology, King’s College, London. 


In 1868 I discovered that if chloroform, ether, amylic nitrite, 
acetic acid or ammonia be held before the nose of a rabbit, the 
heart’s action soon becomes greatly retarded. I at first fancied 
that the inhibition of the heart in this case results from the 
inhalation of the vapour, but I soon abandoned this idea; for I 
found that when I opened the trachea, and placed a cannula in 
its lower end, and permitted the vapours to be inhaled through 
the cannula the retardation of the heart did not occur. I next 
supposed that the influence of the vapour might be owing to 
reflex action: viz. to an excitement of the cardio-inhibitory 
centre in the medulla following the local stimulation of nasal 
nerves. I observed, however, that when the vapour was pre- 
sented to the nose of the animal, it closed its nostrils and 
ceased to breathe.° During this, the inhibition of the heart 
appeared. And I further noticed that this retarded cardiac 
action ensued when the nostrils were closed by the hand in the 
gentlest manner possible. From these faets I suspected that the 
cardiac retardation was not owing to reflex action, but to stimu- 
lation of the cardio-inhibitory nerves by the state of the blood 
which results from arrest of the respiration; and this idea was 
confirmed by finding that closure of the nostrils by the hand, or 
the placing of chloroform, etc., before the nose did not retard 
the heart if the trachea had been previously opened, and the 
air prevented from passing through the nose by means of a 
cannula tied in the lower end of the trachea and having no 
communication with the upper end. I never published these 
experiments, but I alluded to the results in a paper on the 
“Tnfluence of the Vagus upon the Vascular System,” Journ. 
of Anat. and Physiology, Vol. 11. I there (p. 408) stated that 
“the theory that the inhibitory influence of the vagus upon the 
heart is due to exhaustion of the cardiac ganglia produced by 


19--2 


284 PROFESSOR RUTHERFORD. 


over-stimulation, seems to me irreconcilable with the following 
fact. If any irritating vapour, such as that of chloroform, ether, 
alcohol, acetic acid, etc., be held before the nose of a rabbit, 
it instantly closes its nostrils and ceases to breathe—often for 
30 or 40 seconds. Within three seconds after the cessation of 
the respiration the heart comes almost to a standstill, and con- 
tinues to beat very slowly until respiration be re-established. 
This arrest of the heart is due to stimulation of the inferior 
cardiac branch of the vagus by the asphyxiated condition of the 
blood, for the slowing of the heart does not set inif the vagi have 
been previously divided. The perfect calmness with which a 
rabbit will often sit with its heart almost stopped, seems to 
forbid the idea that in such a case the vagi are over-stimulated.” 
By the term “inferior cardiac branch of the vagus” I simply 
meant the cardio-inhibitory fibres of the vagus, which in the 
rabbit are known to be contained in the inferior cardiac branch. 
My meaning was this—the asphyxiated condition of the blood 
arrests the heart through the agency of the vagi, because the arrest 
of the heart does not appear if these nerves have been previously 
divided. I did not intend anyone to suppose my meaning to be 
that the asphyxiated condition of the blood is the cause of the 
heart’s arrest, because the heart is not arrested if the vagi have 
been previously divided. In the paper above mentioned I 
alluded to this matter in order to bring a new argument against 
Schiff’s notion that the inhibition of the heart is due to ex- 
haustion of the vagi. 

Dr Brown-Séquard, in a paper on “The Sudden or Rapid 
Arrest of many Normal or Morbid Phenomena” (Brown-Séquard 
and Seguin’s Archives of Scientific and Practical Medicine, New 
York, January, 1873), refers to this matter (p. 90). He states, 
that “if the supposed asphyxiated condition of the blood were the 
cause of the arrest of the heart, we should see the same effect 
from other causes of asphyxia, while the reverse is what we 
generally observe.” And he further states, that the effect is 
evidently due to a reflex influence proceeding from the nasal 
mucous membrane. I trust that my distinguished friend will 
bear with me while I give the results of experiments which 
prove lis explanation to be untenable, and mine to be, notwith- 
standing his strictures, the correct one. The experiments 


RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 285 


formerly performed by me were done without the aid of a 
kymograph; I have recently repeated them with the aid of this 
instrument, and I am satisfied with the perfectly accurate 
manner in which the following facts have been ascertained. I 
trust that my friend, and also other authorities’, will repeat 
these experiments, and I shall be very glad to know whether 
after such repetition they still cling to such an explanation as 
that given by Brown-Séquard. 

In the following experiments a mercurial kymograph (heemo- 
dynamometer) was connected with an artery, and the pulsa- 
tions recorded upon a Secretan’s cylinder. At the same time a 
metronome and Neef’s hammer were used to record seconds 
upon the paper on the cylinder. The seconds are indicated by 
dots on a straight line in the tracings (Figs. 1, 2 and 3). The 
interval between any pair of dots is not always of the same 
length, because the motion of even a Secretan’s registering 
apparatus is not perfectly regular. The straight line upon 
which the dots occur is not an abscisse. It has nothing to do 
with the line indicating the zero pressure which is usually 
drawn on the paper in such experiments. The wavy line in 
the tracings indicates the heart’s pulsations. Every wavelet is 
a pulsation. The risings and fallings of the wavy line indicate 
risings and fallings of the arterial blood-pressure. All the 
tracings are to be read from left to right. 


Experiment I. &abbit. Cannula of kymograph in lower 
end of right carotid artery. 


No. of Ob- | 


1 + be 
servation. | General Notes. 


| 
Time. Pulse in 2” 


dd 
ys) 
57 
| 59 


| 
Obs, A | ie 3 es) 


CH Om or 


' See Handbook for the Physiological Laboratory, edited by Dr Burdon San- 
derson, p. 274. 


286 


PROFESSOR RUTHERFORD. 


No. of Ob- 
servation. 


Obs. A 


(continued). 


Obs. B 


Obs. C 


Obs. D | 


Time. Pulse in 2’ General Notes. 
9/4” | 
D8 : : '| During closure of the nostrils by 
the fingers the retardation of the 
5 5) heart was decided at the seventh 
7 3 second after the closure (which 
9 3 began at 5. 31’. 59’... The graphic 
11 3 | representation of this is given in 
Fig. 1. 
13 3 
15 3 
17 3 
5.34.48 8 
50 7 
52 7 
54 8 
306 8 
58 7 
oD. 8 
2 u 
4 6 
6 6 
8 5 The above was repeated. The clo- 
10 4 sure of the nostrils began at 5.35’. 
The retardation of the heart was 
12 4 decided at the fourth second after 
14 3 the arrest of respiration. 
16 3 
18 2 | 
20 3 
22 4 
24 6 
5.41.58 8 
42’, 7 
2 7 
4 7 A drop of strong ammonia held on 
6 6 paper before the nostrils. The 
8 5 animal immediately ceased to 
10 4 breathe. The retardation of the 
12 4 heart began at the jifth second. 
14 3 
5.54’, The trachea was opened and a can- 
14 8 nula was tied in its lower end. 
16 7 
18 8 
20 8 


RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 287 


No. of Ob- 


Sea ee Time. Pulse in 2” General Notes. 


Obs. D 5.54.22” 
(continued). DA 


Nostrils were again closed by com- 
pression with the fingers. The 
breathing continued (no retarda- | 
tion of heart). 


oe 
ore) 
YIWIINNDAWODBDNGSCHAMBDO 


Obs. E 


ou 
ba 
ce 
os 
bo 


Sars) 


A drop of strong ammonia placed 
on paper and held before nose. 
The breathing continued, but at the 
same time became somewhat slower. 
Observe that the heart’s action was 
slightly retarded. Compare this 
with Obs. C. 


UY 
struggle 


Or 
ie) 
ip 2) DDAHANIBANWWGAAM cow NIISTDDHODHDH 


Obs. F 6.0'.0 


288 


| No. of Ob- | 
| servation. | 
| 
Obs. F 


\(continued). 


Obs. G | 


Obs. 


Pulse in 2” 


ey) 
struggle 


WADM Os Aanananrtegnl 


UY 
struggle 


WWWOUWO DD worm nm MArntrooaon 


WD rm wWDWOO® 


aos a) 


PROFESSOR RUTHERFORD. 


General Notes. 


Obs. EK was repeated. The breath- 
ing continued, but became some- 
what slower. 


The nasal mucous membrane was 
pricked with the sharp points of a 
pair of scissors. The breathing 
was not arrested but the animal 
struggled violently. 


During the whole of Obs. H, artifi- 
cial respiration was kept up by 
means of Richardson’s bellows at- 
tached to the cannula in the tra- 
chea in order to prevent deficient 
oxygenation of the blood. 


Ammonia held before the nose as 
in Obs. Cand E. WNo retardation 
of the pulse. 


RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 289 


No. of Ob- 


: ep 
pain Time. Pulse in 2 General Notes. 


Cts I | 6e19'.52" 


Artificial respiration still maintain- 
ed during Obs. I. 


At * a drop of strong ammonia was 
applied to the mucous membrane 
of both nostrils, and it was washed 
off at **. No retardation of the 
pulse. 


Obs, J 54 


Respiration arrested by closing the 
cannula. Pulse retarded. 


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sg 
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PROFESSOR RUTHERFORD. 


290 


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RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 


Experiment II. Rabbit. 


291 


Cannula of kymograph in femoral 


artery. 
Novor hs Time. Pulse in 5’ General Notes. 
Obs. A .| 2.59'.187 21 
19 16 During arrest of respiration by clo- 
sure of the nostrils by the hand. 
Obs. B 3.0.13 20 
20 9 During arrest of respiration conse- 
quent upon holding a drop of 
strong ammonia on bibulous paper 
before the nostrils. 
Obs. C Obs. B was repeated with like re- 
sults. 
aya eys Cannula placed in lower end of 
trachea. 
Obs. D 3.20'.6 21 
9 
18 20 | During closure of nostrils by the 
21 | fingers. Breathing was not arrested. 
O12 20 
13s 
21 20 Ammonia held on paper before 
30 20) nose. Breathing not arrested. Pulse 
BO) | not retarded. 
Pulse in 1” : : 
Obs. E 3.94.9 5 See Fig. 1 (b) for a graphic repre- | 
; 1 0 5 sentation of Obs. E. 
1 4 
12 5 
13 5) 
14 4 
15 4 
16 5 Respiration arrested by closure of 
i by 4 cannula. Retardation of the heart 
18 3 decided at the fourth second. 
19 2 
Pulse in 
10” 
Obs. F | 3.30.14 45 
21 Nasal mucous membrane pricked 
36 39 and scratched by means of a 
needle. There was slight strug- 
48 40 gling. The respiration became 
49 slower. 


PROFESSOR RUTHERFORD. 


No. of Ob- : Pulse i 
Baton. Time. 10” 7 
3.37'.9” 44 
10 
20 42 
29 Struggle 
Trace 
not suffi- 
| ciently 
3.50.4 if clear to 
show the 
| pulse- 
J rate. 
8 
17 43 
29 44 
. 38 
Obs. I 3.56.9 42 
14 
34 |40 or 42? 
40 


General Notes. 


Artificial respiration had recourse 
to in order to prevent deficient 
oxygenation of the blood. 


Nasal mucous membrane again 
pricked and scratched. During 
artificial respiration, 


A drop of strong ammonia applied 
to the nasal mucous membrane. 


Ammonia washed off. Artificial 
respiration was kept up during the 
whole of Obs. H. (Although the 
pulse could not be ascertained at 
3.50’.4” it may be safely said 
that in this Observation there was 
no retardation of the pulse.) 


Ammonia applied to nasal mucous 
membrane. 

Ammonia washed off. Artificial 
respiration kept up all the while 
asin Obs. H. No retardation. 


RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 293 


Experiment III. Rabbit. Cannula of kymograph in femoral 
artery. 


No. of Ob- 
servation. 


Obs. A 


Obs. B 


Obs. C 


Obs. D 


Obs. E 


Obs. F 


Time. Pulse. | General Notes. 


in 2” | 
Suatrial le oa) 
20 8 
22 8 1) 
24 8 
26 8 
28 6 Respiration arrested by gently com- 
30 6 pressing the nostrils with the fin- 
39 5 gers. See Fig. 2 c—c’ for a graphic 
representation of Obs. A. 
34 5 
36 3 
38 
A drop of strong ammonia upon 
bibulous paper was held before the 
nose. The animal shut its nos- 
trils and the pulse was retarded as 
usual, The results are not given 
because the metronome apparatus 
failed to record the seconds on this 
oceasion. 
The trachea was opened and a can- 
nula placed in its lower end. 
in 10” 
aor 12 40 
15 Nostrils closed by fingers. Respira- 
28 41 | tion still continued through the 
31 cannula. (See Fig. 2 d d’ for a 
J graphic representation of Obs. C.) 
4.3'.8 33 
14 Ammonia held in front of nose as 
before. Respiration continued 
27 33 through the cannula unchanged. 
29 Notice that in Obs. C and D the 
J pulse was not retarded. 
4.6’.12 33 
Mucous membrane of nostrils prick- 
16 ed and scratched with a needle. 
27 25 The respiration became slow and 
| feeble. Notice the retardation of 
the pulse. 


4.7.6 30 


294 


| No. of Ob-| 
| servation. 
| Obs. F | 


(continued). 


Obs. G 


Obs. H 


Obs. I 


PROFESSOR RUTHERFORD. 


Time. ae 
7 TB ie 0 hale 
21 27 
31 24 
A10°50 | 34 
rae | 
12 35 
14 
4 18' Ad ol 40 
19 | 
34.1 9 42 
BS: | read 
59 
4.28’ .24 40 
35 | 
52 41 
54 


General Notes. 


A drop of strong ammonia applied 
to mucous membrane of nostrils. 
The respiration became feeble and 
slow, and continued to be so until 
4.7’. 35” when the ammonia was 
washed off. Notice the retarda- 
tion of pulse. (For a graphic re- 


presentation of Obs. F, see Fig. | 


3 e.) 

Artificial respiration was now had 
recourse to in order to prevent 
deficient oxygenation of the blood 


during the irritation of the nasal | 


mucous membrane. 


The mucous membrane of the nos- 
trils was pricked as before. No 
pulse retardation now. 


Ammonia applied to mucous mem- 


brane of nostrils. Slight pulse ac- | 


celeration. 


Ammonia washed off. 


Ammonia again applied to mucous 
membrane of nostrils. No retard- 
ation of pulse. 


Ammonia washed off. 
(See Fig. 3 f for a graphic represen- 
tation of Obs. I.) 


Yen) 


GN 


NOSTRILS. 


CLOSURE OF 


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PULSI 


OF 


RETARDATION 


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RETARDATION OF PULSE ON CLOSURE OF NOSTRILS. 297 


General Results of Experiments 1, 2 and 3. 1. If the 
nostrils of a rabbit be closed by the fingers, and the breathing 
be thereby arrested, retardation of the pulse speedily ensues. 
The time at which the retardation appears varies; it may be as 
early as the fourth second after the closure (see Obs. B, Exp. 1, 
and Obs. E, Exp. 2), but it frequently begins a few seconds later 
than this’, 

2. The retardation of the pulse does not follow the grasping 
and closure of the nostrils if the respiration be permitted to con- 
tinue through a fistula in the trachea. 


3. When a piece of bibulous paper containing ammonia is 
held before the nose, the animal closes its nostrils und ceases to 
breathe for a time. The closure of the nostrils protects the 
mucous membrane from irritation by the pungent vapour, but 
notwithstanding this the pulse is retarded. 


4. When the trachea is opened and a cannula placed in 
its lower end, retardation of the pulse follows closure of the 
cannula as speedily as it follows arrest of the respiration by 
closure of the nostrils. In the former case the Jocal mechanical 
stimulation of nerves such as those of the nostrils ts out of the 
question. 


5. It follows from 1, 2, 3 and 4 that the retardation of the 
pulse consequent upon closure of the nostrils, and upon the 
~ holding of bibulous paper with ammonia before the nostrils, is 
due to the arrest of the respiration. 


6. The state of the blood resulting from the arrested respi- 
ration is the cause of the retardation of the pulse, and it retards 
the heart by acting upon the cardio-inhibitory nerve apparatus ; 
because if the cardio-inhibitory nerves be paralysed by atropia the 
retardation of the pulse does not follow the arrested respiration 
until death setsin. The cardio-inhibitory centre in the medulla 
is, in all probability, mediately or immediately thrown into opera- 


1 In my former paper, loc. cit., I stated ‘that within three seconds after the 
cessation of respiration the heart comes almost to a standstill, and continues to 
beat very slowly until respiration be re-established.’’ The more accurate and 
graphic method leads me now to put it thus: ‘ Sometimes at the fourth second 
after the cessation of respiration distinct retardation of the pulse may set in.” 


VOL. VII. 20 


298 PROFESSOR RUTHERFORD. 


tion by this state of the blood (deficient oxygen or accumulated 
carbonic acid?), because the retardation of the pulse is not 
observed if the vagi have been divided in the neck previous to 
the arrest of the respiration. (The facts here given (No. 6) were 
ascertained by me six years ago.) 


7. The opinion expressed by me in my previous paper (loc. 
cit.) regarding the cause of the retardation of the pulse receives 
rom these experiments an absolute confirmation. 


8. Irritation of the mucous membrane of the nostrils of 
the rabbit, by pricking or scratching, or by the application of 
ammonia (liquor ammoniae fortissimus), is usually (but not 
always) followed by retardation of the pulse, even when the 
respiration continues through a tracheal fistula. The cause of 
this is probably to be found in the fact that during the nasal 
irritation the respiration usually becomes feeble and slow. And 
this idea is confirmed by the fact that if the blood be kept fully 
oxygenated by artificial respiration the pulse is not retarded 
during the nasal irritation. 


9. Although the application of ammonia to the nasal 
mucous membrane during the maintenance of a thorough arti- 
ficial respiration does not retard the pulse, nevertheless, I twice 
observed (Obs. H and I, Exp. 3) that when artificial respiration 
was stopped, and the ammonia washed off, that retardation of 
the pulse appeared and continued for some seconds. Was this 
owing to retardation of respiration, or could it be due to stimu- 
lation of the cardio-inhibitory mechanism reflexly from the 
nose? Might it not be that the hyperoxygenated condition of 
the blood diminished the excitability of the cardio-inhibitory 
apparatus—thereby preventing the nasal irritant from affecting 
the heart during the artificial respiration ? 


In order to determine this point I divided the vagi of a 
rabbit in the neck-——observed the extent of the retardation of 
the heart’s action that resulted when the lower end of the vagus 
was faradised by a current derived from Du Bois-Reymond’s 
machine (2d. 120mm. distant from Ist coil, one small Grove). 
I then established artificial respiration by means of Richardson’s 
pump and a tube in the trachea (as I had done in the previous 


RETARDATION OF PULSE ON CLOSURE OF NOSTRILS, 299 


experiments). I stimulated the lower end of the same vagus 
with a current of the same strength as before, and the result 
showed that the condition of the blood produced by this mode 
of respiration did not diminish the excitability of the cardio- 
inhibitory mechanism below the level of the middle of the neck, 
and the fact that in experiments 2 and 3 the adoption of arti- 
ficial respiration with the vagi undivided did not accelerate the 
heart’s action, leads me to conclude that the activity of the car- 
dio-inhibitory apparatus in the medulla is not diminished by 
such respiration ; for, were it so, the heart’s action would be 
accelerated as it is when the cardio-inhibitory nerves are para- 
lysed by division or otherwise. I therefore conclude that the 
pulse retardation observed in Obs. H and I, Exp. 3, to which 
allusion has just been made, was probably the result of dam- 
nished respiration resulting from the excitement of the nasal 
mucous membrane. 


20—2 


NOTES OF SOME CASES OF ABNORMAL ARRANGE- 
MENT OF THE ARTERIES OF THE UPPER EX- 
TREMITY. By J. J. Cuarzes, M.A., M.D., &., Demon- 
strator of Anatomy, Queen’s College, Belfast. 


1. Vas aberrans, joining the ulnar artery. A rare variety 
of vas aberrans was discovered last month in the Anatomical 
Rooms of Queen’s College, Belfast, during the dissection of the 
body of a female, aged 30 years. In the right arm a slender 
artery, five inches and a half long, arose from the brachial 
artery, a little below the middle of the humerus, and two 
inches below the lower border of the teres major muscle, and 
accompanied the brachial on its inner side to the neck of the 
radius, where it joined the ulnar a quarter of an inch from 
its origin. It was pervious throughout, being filled with in- 
jection like the brachial, but it gave off no branches. The 
brachial, radial, and ulnar arteries were normal as to their 
position and number of branches, except that the anastomatic 
which should arise from the brachial was absent. 

In the left upper extremity the arteries had a normal 
arrangement. 

Quain, even in his large experience, speaks of having met 
with only nine instances of vasa aberrantia, and these all ended 
in the radial except one which joined the radial recurrent, but 
it was in that case a branch of the ulnar. He states, however, 
that Monro and Meckel had each observed a single instance 
in which the vas aberrans terminated in the ulnar directly’. 
Cruveilhier has, on two occasions, seen a vas aberrans joining 
the ulnar’. 


2. High origin of the radial artery; and vas aberrans 
joining the radial artery. It is a curious coincidence that last 
month there were, in the Belfast Anatomical Rooms, five upper 


1 The Anatomy of the Arteries of the Human Body, pp. 265 and 266; 1844: 
131; 1871. 

2 Traité D’Anatomie Descriptive; 4° Edition; Tome 3°, p. 131, 1871. 
Wenzel Gruber observed in 600 bodies examined vasa aberrantia joining 
the radial artery in five arms, and in one case only did a vas aberrans join the 
ulnar artery. (Eds.) 


DR CHARLES. ABNORMAL ARRANGEMENT OF ARTERIES, 3801 


extremities in which the radial artery had a high origin. In 
the first, the radial arose in the upper third of the arm, close 
to the head of the humerus; in the next three it took origin 
in the middle third of the arm; and in the fifth it arose in 
the lower third, and was joined about an inch from its origin 
by a vas aberrans which was four inches and a half long, and 
had sprung from the inner side of the brachial about half an 
inch below the lower border of the teres major. In all these 
cases the radial arose from the inner side of the axillary or 
brachial, and lay superficial to the pronator radii teres, except 
in one instance where it passed between the two heads of 
origin of that muscle. 

It will be remarked that I have referred above to six cases 
of abnormal arrangement of the arteries of the upper extre- 
mity which occurred in five bodies. One was a case of vas 
aberrans joing the ulnar, in the nght arm; the remaining 
five were instances of high origin of the radial—four in the 
right and one in the left arm. In only one body was the 
disposition of vessels abnormal in both arms; and in that 
body, on the right side, the radial arose from the axillary, and, 
on the left, from the lower third of the brachial, but it was 
afterwards joined by a vas aberrans an inch below its origin, 
This arrangement of arteries in the two arms supports the 
view of Bichat and Quain in opposition to Meckel who held 
that “lateral symmetry, the most powerful of all (kinds of 


1» 


symmetry) is maintained even in malformations’. 


P.S. Most of the specimens referred to above were ex- 
hibited at the Ulster Medical Society, March 15. 


1 Quoted by Quain in his Anatomy of the Arteries of the Human Body, 
p. 268. 


ON THE PLACENTATION OF THE SLOTHS. 
By Professor TURNER. 


(Abstract of a Memoir communicated to the Royal Society of Edin- 
burgh, May 19, 1873.) 


AFTER referring to the paucity of information on the placental 
characters of the Sloths, and to the various inferences which had 
been drawn by anatomists from Carus’s figure of the placenta of 
Bradypus tridactylus, some holding that it was cotyledonary 
and non-deciduate, others that it might have intermingled with 
it maternal deciduous substance, the author proceeded to de- 
scribe his dissection of the perfectly fresh gravid uterus of a 
specimen of a 2-toed Sloth. This specimen only possessed 6 
cervical vertebre, and was referred to the Cholepus Hoffmanna 
of Peters. The author had succeeded in obtaining excellent 
injections both of the fcetal and maternal systems of blood- 
vessels. The placenta consisted of about thirty discoid lobes, 
aggregated together, and occupying about four-fifths of the. 
surface of the ovum. These lobes could be peeled off the 
placental area of the uterus, and carried away with them a 
layer of deciduous serotina, curling arteries, utero-placental 
veins, and a very remarkable system of intra-placental mater- 
nal sinuses, continuous with the uterine vessels, freely anasto- 
mosing with each other within the substance of the lobes, and 
lying between and in contact with the foetal villi. Definite 
walls, distinct from the walls of the foetal villi, could be traced 
around the sinuses. Crowds of red blood-corpuscles were 
situated within the sinuses, and it was observed that many of 
these seemed to be nucleated, an appearance which had been 
recognised a few years ago by Kiihne, Rolleston and Moseley, 
in the blood-corpuscles of the Tardigrada. This sinus system 
possessed a special interest, because it presented a gradation 
between the capillary net-work of the uterme mucous mem- 
brane occurring in the diffused placenta of the mare or the 
cetacean, and the freely anastomosing cavernous maternal 


PROFESSOR TURNER. PLACENTATION OF THE SLOTHS, 3803 


blood-spaces seen in the highly concentrated human placenta. 
The amnion lay in contact with the inner surface of the chorion 
as in the human feetal membranes; its inner surface was smooth 
and serous, but there was no liquor amnii. The foetus possessed 
a very remarkable special envelope like that figured and de- 
scribed by Welcker as investing the feetus of B. tridactylus 
and a few other mammals, which envelope he has named an 
Epitrichium, Numerous additional details respecting the struc- 
ture of the placenta and membranes are contained in the memoir, 

The conclusions drawn from the examination of this pla- 
centa were: that in the Sloths the placenta is not coty- 
ledonary and non-deciduate as in the Ruminants, but in the 
fullest sense of the word deciduate. If the inference drawn 
by Huxley from Sharpey’s observations on the structure of the 
placenta of Manis be correct, then, if the placental system of 
classification is to be of any value, the non-deciduate scaly Ant- 
eaters can no longer be grouped along with the deciduate Sloths 
in the order Edentata, which order will have therefore to be 
subdivided. The author then compared the placentation of the 
Sloths with that of the other deciduate mammals, and- pointed 
out a series of very interesting affinities between its placenta 
and that in the Primates. 


NOTES OF SOME IRREGULARITIES IN MUSCLES AND 
NERVES. By Jonn Curnow, M.D., London; Demonstrator 
of Anatomy in King’s College, London. 


Ty this paper I have recorded only the less frequent irregularities in 
muscles and nerves that have come under my observation, and of 
many of them I have failed to find any mention, although from the 
extensive and scattered literature of the subject, I may have over- 
looked references thereto. I have arranged them in anatomical 
order, taking the museles first. 

1. Lexternal Rectus. In addition to the usual muscles, two extra 
slips were found in the right orbit of a female subject. They were 
apparently differentiations of the external rectus, and arose with its 
lower head from the ligament of Zinn. The inner and shorter slip 
was inserted into the outer half of the cartilage of the lower eyelid, 
where it joined the larger and more external slip, which was inserted 
into the periosteum of the outer wall of the orbit as well as into the 
cartilage. This slip was almost as broad as the external rectus itself, 
and was supplied by a braneh of the sixth nerve equal in size to that 
supplying the external rectus. Both slips passed forwards directly 
beneath the lachrymal gland. Their use is very obscure, for although 
the inner one would slightly retract the lower lid, the outer and larger 
one was chiefly attached to the periosteum. Iam unaware of any 
observations that would throw light on them from a morphological 
standpoint. 

2. Complexus. On both sides a small band of muscular fibres 
arose from the ligamentum nuche at the third cervical spine, in con- 
nection with the upper fibres of the splenius capitis, and ran upwards 
parallel to the ligament nuche to just below the superior curved line 
of the occipital bone. A few of the fibres also arose from the upper 
inch and half of the ligament. It was separated from the complexus 
by a distinct cellular interval, and overlapped the innermost fibres of 
that muscle, 

3. Several anomalies were noticed among the muscles passing 
from the neck to the trunk and shoulder. Besides a typical example 
of the rhombo-atloid of Macalister, and less developed specimens ex- 
tending to the scapula and the fascia over the serratus posticus supe- 
rior from the lower cervical vertebra and the fascia over the splenii, 
two rarer variations occurred. A small ship from the posterior tu- 
bercle of the transverse process of the third cervical vertebra passed 
downwards across the posterior triangle in front of the levator anguli 
scapule to join the first digitation of the serratus magnus. The le- 
vator anguli scapule arose by three digitations from the first, second, 
and fourth vertebra, and missed the third, where its position was ex- 
actly occupied by this abnormal slip. A precisely similar muscle is 
mentioned in the Guy’s Hospital Reports for 1871, and Rosenmiiller 
described a slip from the transverse process of the atlas to the serratus 


DR CURNOW. IRREGULARITIES IN MUSCLES AND NERVES. 305 


magnus. In another subject a small muscle arising from the third 
cervical vertebra, between the tendons of the levator anguli scapule 
and the scalenus medius, passed downwards behind the latter muscle, 
and then turned forwards to be inserted by a bifurcated tendon into 
the first and second ribs just behind the attachments of the serratus 
magnus. Although somewhat resembling some of the additional sca- 
lene muscles described by Albinus, Sémmering, Meckel, and Mac- 
alister, this should be classed among the imperfect varieties of the 
levator clavicule, whose signification has been pointed out so com- 
pletely by Prof. Wood. (Phil. Trans. 1870.) The other slip is one 
of the transitional forms between the levator clavicule and occipito- 
scapularis. 

4. Sterno-scapular or Sterno-chondro-scapular. (Wood.) This 
was a well-developed specimen of the muscle so variously described 
by anatomists as an anomaly of the subclavius, omo-hyoid, and even 
serratus magnus. It arose with the subclavius from the first costal 
cartilage, and passed backwards to the supra-scapular ligament and 
base of the coracoid process, joining the origin of the omo-hyoid. 

5. Ina spare female, on the right side only, the tendons of the 
latissimus dorsi and teres major had this curious arrangement at 
their insertion (see Fig.). The latissimus dorsi (a) did not extend so 
far outwards as the base of the bicipital groove, but was attached to 
a prominent tubercle, three quarters of an inch long, just internal to 
its inner lip. The muscular fibres ceased just as the long tendon of 
the triceps (d) passed behind it, and from this point a tendinous band 
(1) ran upwards in front of the triceps tendon to the axillary margin 
of the scapula. The teres major (6), muscular throughout, did not 
reach the humerus, but was inserted into the band just described, 
and into the long head of the triceps. The insertion of the sub-sca- 


pularis (c) was very large, contrasting markedly with the other ten- 
dons. Besides these anomalies, a small muscle (2) which could act 


306 DR CURNOW. 


as a tensor of the capsular ligament was present. It arose from 
a special groove between the inner lip of the bicipital groove and 
the tubercle for the latissimus, and was lost on the anterior and inner 
part of the capsular ligament, near the insertion of the sub-scapularis. 
I can find no reference to this muscle, or to a similarly aborted in- 
sertion of the teres major. 

6. Biceps and Coraco-brachialis. Tn avery muscular male, the 
right biceps was very complex both at its origin and insertion, In 
addition to long, short, and humeral heads, there was an extra cora- 
coid head, with a few fibres joining it from the capsular and coraco- 
acromial ligaments. It passed downwards and joined the humeral 
head at the junction of the middle and lower thirds of the humerus, 
instead of blending with the normal coracoid as is usually the case. 
The muscle was mainly inserted into the semilunar fascia and the 
tubercle of the radius, but it gave off from its inner side a tendinous 
slip which split into three parts, (a) to condyloid origin of pronator 
teres; (b) a slip giving origin to the fibres of that muscle which 
usually arise from the coronoid process, and separated from the for- 
mer by the median nerve; (c) to upper part of radial origin of flexor 
sublimis digitorum. 

In the same arm was a well-developed chondro-epitrochlearis, and 
an aberrant slip of the coraco-brachialis, as described by Struthers. 
This arose from the capsular ligament, lesser tuberosity, and up- 
per part of the latissimus dorsi tendon, and crossing the brachial 
vessels joined the coraco-brachialis, which was pierced by the mus- 
culo-cutaneous nerve as usual. On the left side, the only peculiarity 
was a biceps, with a third head from its common position on the 
humerus. 

7. Flexor Sublimis Digitorwum. A remarkably thin tendon for 
the little finger was given off from the radial side of the tendon to 
the ring finger, and crossed the latter superficially to its usual 
position. 

8. Flexor Carpi Ulnaris. From its usual insertion a tendinous 
prolongation passed forwards under the abductor minimi digiti, and 
was lost in the ligaments around the metacarpo-phalangeal joint of 
the little finger. I have seen only one specimen of this slip, and I 
can find no mention of it, so that in regard to frequency there is a 
marked contrast between it and the prolongation of the extensor 
carpi ulnaris or ulnaris quinti, which occurs in as many as 12 per 
cent. of subjects. (Wood.) 

9. Ezxtensores Carpi Radiales. Besides the normal radial exten- 
sors of the carpus, a third was present, and equalled either of the 
others in size. It arose by two fleshy bellies from the adjacent sides 
of the radial extensors—that from the brevior being the larger—and 
about 2 inches from the lower end of the radius formed a single ten- 
don, which was inserted into the ulnar side of the index metacarpal 
bone. The longior and brevior were inserted as usual. It differed 
from the extensor carpi radialis intermedius of Wood, which consists 
of a cross slip or slips between the two radial extensors, and also 
from the extensor carpi radialis accessorius, which is inserted into the 


TRREGULARITIES IN MUSCLES AND NERVES. 307 


base of the first metacarpal. An exactly similar muscle is described 
by my late colleague Mr Perrin in the Medical Times and Gazette, 
1872, Vol. 11. 

10. Extensor Quarti Digiti vel Annularis. In the left arm ofa 
male subject, in addition to the common extensor tendon, a special 
extensor of the ring finger was found. It arose from the posterior 
surface of the ulna for three inches to the inner side of the origins 
of the extensor secundi internodii pollicis and extensor indicis. It 
continued fleshy to annular ligament, and having passed through the 
common extensor groove joined the ulnar side of the extensor apo- 
neurosis on the back of the ring-finger. The indicator and extensor 
minimi digiti were normal. The extra tendons to this digit pre- 
viously noticed, have been of two kinds, viz.: differentiations of the 
extensor minimi digiti as in most quadrumana which occur as 
often as thirteen times in 102 subjects (Wood); or more or less abor- 
tive specimens of the extensor brevis digitorum manus, arising from 
the lower end of the radius, the carpus, or metacarpus. Some exam- 
ples of both these varieties were found, but the muscle just described 
differed essentially therefrom, being quite as distinct and special to 
the ring-finger as is the extensor indicis to the fore-finger, which it 
much resembled in size and appearance. 

ll. Latensor Carpi Ulnaris. This was once found double 
throughout, but both tendons were inserted into the fifth metacarpal 
bone. No ulnaris quinti present. Extensor minimi digiti also double. 

12. Psoas. Besides the normal psoas magnus, a muscle arose 
by two origins; an upper from the first lumbar vertebra, and a lower 
from a tendinous arch attached to the fourth intervertebral substance 
and the margins of the adjacent vertebrae. It passed under Poupart’s 
ligament, and was inserted into the capsule of the hip-joint, and into 
the femur just above the lesser trochanter. This seems rather a dif- 
ferentiation of the psoas magnus than a true psoas parvus. On the 
opposite side the psoas magnus was normal, and the psoas parvus absent. 

13. In the left leg of a female subject a thin, penniform muscle 
arose from the posterior surface of the fibula, between the origins of 
the flexor longus hallucis and peroneus brevis, extending from the 
fibular origin of the soleus, to about 1 inch from the ankle-joint. It 
yan with the flexor longus hallucis, grooved the tibia, astragalus, and 
sustentaculum tali, and was inserted into the os calcis slightly anterior 
to that process, being covered by the inner head of the flexor acces- 
sorius. Most of the anomalous muscles in this situation are varieties 
of the peronei (peroneus quartus, Otto, &c.), and are inserted into the 
outer surface of the os calcis, cuboid, or fifth metatarsal bone; but a 
similar muscle is described in the Guy’s Hospital Reports for the 
present year as connected with the flexor longus hallucis. This 
muscle is probably the homotype of the radio-carpus of Fano, occa- 
sionally seen in the upper extremity. In the same leg, beside the 
usual three peronei, there were found a small peroneus quinti, a ten- 
dinous extensor ossis metatarsi hallucis given off from the extensor 
proprius, and a well-developed extensor primi internodii arising 
from the fibula for 2 inches below that muscle. 


308 DR CURNOW. 


14. In the left sole of a male subject a singular arrangement of 
the flexor tendons of the little toe was found. The flexor brevis 
digitorum was inserted into the sides of the proaimal phalanx, after 
having been perforated by a tendon from the flexor accessorius as 
well as by the tendon of the long flexor. The tendon of the acces- 
sorius was inserted into the sides of the middle phalanx, having also 
been perforated by the long flexor tendon, which passed on to the 
distal phalanx. Absence of one or both of the tendons to the little 
toe have been frequently noticed, but I can find no instance of a 
special flexor to each phalanx from the three flexor muscles respec- 


tively. 
Il. Nerve-[RREGULARITIES. 


Inferior Maxillary Nerve. The left foramen ovale was divided 
into two by a plate of bone running obliquely downwards and _for- 
wards. This plate wasa quarter-of-an-inch wide in the middle, and a 
little wider at the ends. Through the lower and posterior division 
passed the sensory portion of the third division of the 5th nerve, 
with separate branches to the external and internal pterygoid muscles ; 
the former in front, and the latter behind. Through the upper and 
anterior division passed the masseteric, deep temporal and buccal 
branches, and a second nerve to the external pterygoid. The small 
meningeal artery was absent. This division of the foramen ovale 
into two, is unnoticed in our English text-books, and Simmering 
describes it as rare. Although present in many skulls, I am not 
aware of its having been noticed in the recent subject, and the rela- 
tive positions of the structures as they passed through described. 
Although this must be quite fortuitous, it is interesting to notice 
that the buccal branch came through the upper foramen with the 
chief motor nerves. 

Tn one case the gustatory or lingual nerve, after its junction with 
the chorda tympani, gave off several small branches to the origins of 
the superior constrictor and buccinator muscles from the pterygo- 
maxillary ligament. I could not, however, definitely make out 
whether they were distributed to those muscles, or passed through 
them to the mucous membrane. 

The inferior dental nerve was divided into two parts by the 
internal maxillary artery as it passed forwards between the pterygoid 
muscles, and the mylo-hyoid nerve arose by a separate branch from 
each division. In one case the fission of the nerve was complete from 
its origin to its entrance into the dental canal, but there the two 
parts joined. In another subject, the left mylo-hyoid arose by two 
branches; one from the motor portion of the inferior maxillary 
trunk, and the other from the deep surface of the inferior dental 
nerve. It gave a branch to the superficial portion of the submaxil- 
lary gland, as well as its usual branches to the mylo-hyoid and 
digastric muscles. 

Inferior Laryngeal Nerve. The right inferior laryngeal has been 
often seen to pass directly inwards to the larynx, instead of turning 
round the subclavian artery. In these cases, as pointed out by Hart, 


IRREGULARITIES IN MUSCLES AND NERVES. 309 


Reid, Demarquay, Turner and Krause, the right subclavian arises from 
the left side of the aortic arch. In one case Turner saw it wind round 
the inferior thyroid artery, and in the case I saw, it did the same, 
although that artery had an abnormal origin from the common 
carotid. 

Spinal Accessory Nerve. In one subject on both sides the external 
branch of this nerve stopped at the sterno-mastoid muscle, while the 
trapezius was supplied by two branches from the third and fourth 
cervical nerves. In another instance, it divided into two branches, 
by far the larger one entered the sterno-mastoid, and the smaller 
twig ran across the muscle superficially to its posterior border, and 
there joined a branch from the second cervical nerve, which with 
another branch from the fourth supplied the trapezius. These two 
cases tend to show that probably the trapezius is wholly supplied by 
the spinal portion of the accessory nerve; for in the first, no fibres 
from the medulla oblongata, and in the second but very few, could 
have entered that muscle. 

Phrenic Nerve. Besides the more frequent varieties in its origin, 
it was once joined by a large branch from the. middle cervical gan- 
glion. In another case it supplied the anterior scalenus muscle, and 
then divided into two branches between which the internal mammary 
artery passed. I have also noticed it divide into two branches imme- 
diately after its origin from the fourth cervical nerve (the outer of 
which was joined by the communicating twig from the nerve to the 
subclavius) and re-unite in the upper part of the thorax. 

Posterior Thoracic Nerve. The serratus magnus was not unfre- 

quently supplied by two nerves, a separate branch from the fifth 
cervical nerve going to its first digitation. Jn one case this differen- 
tiation was carried still further; for besides this branch to the first 
digitation, a second was supplied from the fifth nerve to the second 
and third digitations, while a third branch, wholly derived from the 
sixth nerve, was distributed to the remainder of the muscle. 
- Median Nerve. A small communicating branch between the 
outer and inner heads crossed the axillary artery under the pectoralis 
minor; the heads united in the middle of the upper arm, and the 
nerve then passed behind the brachial artery. In another subject 
the median arose by three heads; the third head was given off from 
an additional origin of the ulnar nerve from the outer cord of the 
brachial plexus crossing the axillary artery. 

Ulnar Nerve. In one case in which the ulnar nerve arose from 
the outer and inner cords as described by Turner in Vol. vi. of this 
Journal, it supplied both sides of the middle, and the radial side of 
the ring finger on their dorsal surfaces, in addition to its usual dis- 
tribution. In another case in which the ulnar arose as usual, it 
supplied the contiguous sides of the ring and middle fingers instead 
of the radial. 

Internal Cutaneous Nerve. From the loop between the two 
anterior thoracic nerves a small twig ran down the front of the upper 
arm, and becoming cutaneous in the lower third, passed over the 
median basilic vein and thus took the place of the superficial branch 


310 DR CURNOW. IRREGULARITIES IN MUSCLES AND NERVES. 


of the internal cutaneous nérve, which altogether passed under the 
vein, and became cutaneous at the bend of the elbow. 

Lumbar Plexus. Besides the more usual varieties in the arrange- 
ment of the branches from this plexus, in one dissection the following 
nerves were all given off separately, presenting a most complex ar- 
rangement. Ilio-hypogastric, ioinguinal, genital branch of genito- 
crural, crural branch of genito-crural, anterior and posterior divisions 
of external cutaneous, branch to iliacus, small branches to psoas, 
anterior crural, middle cutaneous dividing into two branches before 
passing under Poupart’s ligament, obturator, and accessory obturator. 
A more complete differentiation of the branches derived from the 
plexus could scarcely be imagined. I regret that my attention was 
called to the dissection too late to accurately make out from which 
nerve of the plexus each branch was derived. 

I must express my great obligations in the compilation of this 
paper to Prof. Macalister’s catalogue of muscular anomalies, to the 
various memoirs of Prof. Wood on the same subject, to the mono- 
graph of Krause and Telgmann, and the papers of Prof. Turner 
on nerve irregularities. 


CASE OF CONGENITAL ABSENCE OF THE QUADRICEPS 
EXTENSOR CRURIS MUSCLE. Communicated by Prof. 
A. G, DracuMann, of Copenhagen’. 


Miss A. F., aged 28 years, consulted me in November of last year 
(1871), for an affection of the left knee, from which she stated she 
had suffered for a very long time—how long she could not remember, 
but it had increased of late years, and rendered walking more and 
more difficult, while she also usually felt pains in the knee-joint, 
especially when she at all exceeded her ordinary amount of walking. 
The knee-joint, too, became tender and swollen—a condition which, 
however, gradually disappeared if she remained at rest for any length 
of time. On examining the uncovered knee I was not a little sur- 
prised to find the knee-cap wanting; the outlines of both condyles of 
the femur very perceptibly exposed, covered only by the skin and 
some subcutaneous areolar tissue; the anterior inter-condyloid fossa 
similarly covered only by skin and filled with a rather soft, subcu- 
taneous adipose tissue, which however was not present in such amount 
as to completely fill up the whole depression between the condyles, 
but left a visible sulcus behind. No trace of the ligamentum patelle 
existed; but the tuberosity of the tibia, and the outlines of the con- 
dyles of this bone were seen and felt very plainly immediately beneath 


1 Translated from the Nordiskt Medicinskt Arkiv, Vol. 1v., Part 1. 1872, 
by J. W. Moore, M.D., M.Ch, Dub.; Hon. Fell, Swedish Soe. Phys. 


ABSENCE OF QUADRICEPS EXTENSOR CRURIS. abt 


the skin. The whole anterior aspect of the thigh, from its upper 
third down to the knee, had lost its usual roundness and fulness. 
The thigh-bone itself throughout this space was felt immediately 
underneath the skin, without a trace of intervening muscular sub- 
stance. A little above the external condyle of the femur, on the 
outer aspect of the thigh, the atrophied patella was found lying, without 
any attachment to any of the femoral muscles, freely moveable in all 
directions. The skin over the knee-joint was perfectly normal, no 
swelling of the tissues constituting or surrounding the articulation 
was present, no effusion into the capsule of the knee. On rather deep 
pressure being made over and a little above the external condyle, the 
patient felt some tenderness. The kneejoint freely admitted of 
passive motion; as regards active motion, the patient could easily 
bend the knee, but she could by no means extend it, nor was she able, 
while in the recumbent position, to lift the limb. The muscular 
structures on the posterior aspect of the thigh, and in the upper 
third of its inner aspect (the adductors) were strongly developed, 
while the triceps cruris (adductores longus, brevis, et magnus) was 
found more than ordinarily strong and largely developed. Measure- 
ment round the upper third of the thigh gave 18 inches, above 
the knee 104 inches, round the leg 12$ inches. On bidding her 
strip the other, the right knee, of which she did not complain, I 
discovered to my surprise, that even in the most minute particulars 
it corresponded perfectly to the left one. On more closely interrogat- 
ing the patient, I learned that she had not been aware of any 
defect in connection with her knees until her tenth year, although 
her gait had presented some irregularity. This had not however 
prevented her from playing and running about with her playfellows. 
She further stated that at this time she fell and hurt her knees, which 
a medical man had examined and had then declared that she suffered 
from a congenital and incurable defect. Since that time she has 
constantly, as the physician advised, worn a bandage on her knees, 
‘the result of which has been (as she positively affirmed) that the 
patella, which was before situated over the internal condyle of the 
femur, had by degrees glided over to the other, the opposite side, 
where it is now situated, It is only within the past two years, since 
she has been obliged to walk and stand a good deal (she is teacher in 
a large national school), that the inconveniences in walking and the 
troubles above mentioned have become more pronounced, and, as 
stated, have extended as yet only to the left lower extremity. 


REVIEWS AND NOTICES OF BOOKS. 


Principles of Animal Mechanics. By the Rev. 8. Hauauroy, F.R.S., 
Fellow of Trin. Coll., Dublin. London, Longmans, 8vo. 


pp- xix. 495. 


Aut those who are interested in the application of the exact 
methods of mathematical and mechanical science to physiological 
problems, will turn eagerly to a work on animal mechanics by the 
accomplished mathematician who fills the chair of Geology at 
Dublin, And there is a large portion of the book with which they 
will not be disappointed. The important section from p. 164 top. 361 
is occupied with a classification of the muscles according to the 
arrangement of their fibres, and a discussion of their action ; and in 
the next eighty pages the results previously obtained are applied to 
the discussion of the mechanics of various important joints, par- 
ticularly the shoulder and hip-joints of the larger /felidw, and to the 
arrangement of the muscular fibres of the heart. I shall have to remark 
hereafter on one or two special points in these sections; but, as far as 
I am aware, the subject has never before been handled with anything 
like the same completeness and thoroughness; and this portion of 
the book will well repay the labour of careful study, though there 
is little in it suitable for extraction or analysis here. 

The earlier portions of the book, however, and the last section 
pp. 442—485, seem to be of more various, and on the whole very 
inferior merit. 

In problems of purely mathematical interest, the value and 
importance of the results may be but little affected by a certain 
arbitrariness in the assumption of values for the constants introduced, 
or in the introduction or disregard of limiting conditions. Where, 
however, the mathematical problem is intended to represent a 
physical fact, and the actual circumstances are either too complicated 
or too little known to be susceptible of complete mathematical 
treatment, it becomes most important, not only to use the best 
attainable values of the constants, but to form a distinct idea of the 
limits within which these values are to be relied on, and to subject 
each step of the process, and each successive result, to independent 
criticism, in order to make sure that no essential consideration has 
been omitted in the transformation of the problem found in nature 
into one susceptible of being attacked by mathematical methods, 

Prof. Haughton seems at times as if he had no idea of this— 
sometimes he lets himself be carried away by what a moment’s thought 
must have shewn him to be only a plausible analogy’. At other 


1 A curious instance of this is to be found in p. 4. After objecting, 
reasonably enough, to the use of the terms voluntary and involuntary, as corre- 
sponding to striped and unstriped muscular fibre, he makes the following as- 
tounding remarks: ‘‘Judging from the analogy of cleavage I have come to 


REVIEWS AND NOTICES OF BOOKS. 313 


times he forms a theory of the way in which an action takes place— 
takes the best value of the constants involved which he can get 
from materials ready to hand—performs his calculations and gives 
the result as certain, without stopping to enquire how far his theory 
is complete, or what will be the effect of an error in the assumed 
values of the quantities involved. A good instance of this on a 
small scale is found in the curious dissertation on the art of hanging, 
pp. 8—13. 

He says, I believe quite truly, that sudden and painless death 
can be and ought to be produced in hanging by shock of the medulla 
oblongata caused by fracture of the vertebral column. He then 
goes on to say that he has “‘ proved” — apparently by a single obser- 
vation, pp. 11, 12—that 2240 ft. lbs. of work are just sufficient to 
accomplish this. He therefore goes on to calculate a formula 
giving the necessary drop in terms of the weight of the victim, and a 
similar one applicable to the American method of hanging. It never 
seems to occur to him that the force necessary to fracture the 
articulating surfaces of the axis, or to displace the odontoid 
process is likely to vary as much with differences of sex, build, &c., 
as the weight of the body itself—and that the force which would be 
insufficient to break the neck of a brawny ruffian of sixteen stone 
might pull off the head of a woman of half the weight. In the case 

_ quoted, in which the shock was exactly what was necessary, the man’s 
height was increased an inch and a half by the process. 

Another instance may be taken from § 5, on the absolute force of 
muscles, pp. 53—74. Prof. Haughton measured the cross sections 
of the flexors of the arm and leg of a selected subject, a blacksmith 
who had died of cholera. He measured the distances between the 
joints and the insertions of the muscles on another subject, a 


the conclusion that the striped structure in muscles, or tendency to cleave into 
discs, is due to their repeated contraction between two fixed or nearly fixed 
points of origin and insertion.”’ And, after speaking of the observations of 
~ Sorby and Tyndall on cleavage produced by pressure, ‘‘ Whenever, therefore, 
we find in muscular fibres a distinct origin and insertion, then contraction 
between these points will produce the pressure necessary for the develop- 
ment of cleavage at right angles to the length of the fibres,” It is hardly 
necessary to remark (1) That the contraction of a muscular fibre between 
nearly fixed points produces tension and not pressure. (2) That if muscular 
contraction is produced, as some have supposed, by the mutual attraction of 
portions of the fibre and consequent compression of the intervening portions, 
this presupposes the longitudinal differentiation of the fibres which Prof. 
Haughton wishes to make it produce. (3) That there is no real resemblance 
between the structure of striped muscular fibre and true cleavage. (4) That the 
striped structure is perfectly developed in foetal muscles before they have con- 
tracted at all. Indeed I supposed at first that the passage was not intended 
to be taken seriously—but it is solemnly referred to in the index as ‘‘ striping 
of muscle, mechanical cause of,’ and I suppose must be deliberately put 
forward as an explanation. That such an idea should flash through the brain 
of an ingenious man is conceivable enough, but that it should be deliberately 
written down, sent to press, corrected in proof, and published in a book is a 
phenomenon of which I can find no plausible explanation. It is, however, 
hardly more than an exaggerated instance of the way in which the author seems 
to put down any idea which strikes him without the least attempt at self- 
eriticism, 


VOL. VII. ; ral 


S14 REVIEWS AND NOTICES OF BOOKS. 


Frenchman of the same height and length of bones as himself, and 
finally determined by experiments on himself and a friend the force 
that could be exerted by these muscles. It is not quite clear 
whether the results given are the mean of those found by himself 
and his friend, or whether, as I rather infer, the experiments were 
made on his own arm and his friend’s leg. From these he deduces 
the contractile force of muscle per square inch of cross section, 
and, as he finds the results from arm and leg come within about 
15 per cent. of each other, he assumes the mean (taken with 
unnecessary if not misleading accuracy to two places of decimals) 
as the coefficient of muscular force applicable to all muscles of all 
animals, and even to the unstriped fibres of the uterus. In fact he 
assumes that all muscles are capable of contracting with a force of 
102-55 lbs. per square inch of cross section, whereas what he has 
proved is that if the cross sections of his muscles and of his friend’s 
are the same as those of the blacksmith, and the insertions the 
same as those of the Frenchman, the muscles of his arm contract 
with a force of 94°7 lbs. and those of his friend’s leg with a force of 
110-4 lbs. per square inch. He gives results of other authors 
differing by 50 per cent. or more, but always makes use of the value 
he has himself obtained. There seems no reason to believe that 
every cubic millimetre of muscle striped or unstriped throughout the 
animal kingdom is intrinsically equally powerful —and even if we 
confine our attention to the same muscles of individuals of the 
same species it is difficult to believe that there is no difference, except 
in cross section, between the muscles of a trained athlete and those 
of a sedentary valetudinarian. 

Before going on to more complicated questions, I may give two 
more instances of the uncritical way in which the Professor too often 
argues. In pp. 2, 3, he argues, somewhat rashly I think, that the 
muscles which have the smallest fibres must be best provided with 
blood, and therefore best able to endure fatigue. He says accordingly 
that the muscles of women are capable of longer continued work than 
those of men. This is interesting if true—but it is certainly not 
proved by the fact referred to in a note that a mother or nurse 
can carry a child for a time which would be very fatiguing to the 
unaccustomed muscles of the father. Of course this is merely a 
matter of training. In many parts of the Alps men and women are 
trained indiscriminately to carry burdens. I have never heard that 
the women shew more endurance than the men, though I fancy that 
there is not much difference between them. It would be interesting 
to know whether the difference in the diameter of the muscular 
fibres in the two sexes is as great among these people as it is among 
the more sedentary races. 

Again, on pp. 484, 485, after pointing out the extremely large 
“Coefficient of Refreshment” of the heart, he goes on “ This 
interesting result is quite in accordance with the views of those 
anatomists who believe that the heart receives double the supply 
of arterial blood that any other muscle receives, in consequence of 
the semilunar valve, during life, not closing the openings of the 


REVIEWS AND NOTICES OF BOOKS. 315 


coronary arteries during systole.” On which it may be remarked 
(1) That the supply of blood to any muscle depends upon the 
number, dimensions and disposition of the arteries supplying it, 
and of the capillaries in connection with them. (2) That no one 
has yet explained how blood can flow into the coronary arteries 
while the heart is violently contracting, whether the orifices are 
closed or not. The effect of systole can hardly fail to be the expul- 
sion of much of the blood from the substance of the heart, as 
well as from its cavities, leaving it in the most favourable condition 
for being refilled with fresh arterial blood during diastole. 

The short articles on the muscular susurrus, and on the work 
done by the heart are interesting, but do not seem to contain any- 
thing very novel, except the very interesting observations on the 
great diminution of the frequency of the vibrations which give rise to 
the susurrus, in cases of paralysis agitans. In the latter far too great 
weight is given to the essentially unsatisfactory method of inferring 
the pressure in the heart from the distance to which blood is spirted 
from a cut artery. 

The section (pp. 154—164) on the muscular forces employed in 
parturition is also interesting, though several objections may fairly 
be made to the arguments by which the somewhat surprising result 
is obtained that the expulsive force of the voluntary abdominal 
muscles is almost exactly ten times that of the uterus. Indeed, 
when our author comes to discuss more fully the action of the ab- 
dominal muscles (pp. 209—232), he gets results which he has some 
difficulty in reconciling with those he had previously obtained, This 
latter section however is not a favourable example of his method of 
treating the geometrical and mechanical problems of muscular 
action. He applies freely to the abdominal muscles Lagrange’s 
theorem, and the equivalent results which he himself establishes, 
which have reference to the relation between the hydrostatic pressure 
within a closed vessel and the tension of the containing envelope 
~supposed to be homogeneous, and either elastic or inextensible, for- 
getting, it would seem, the essential difference between such an 
envelope and a muscular one in which the tensions depend upon 
the direction of the muscular fibres, instead of being the same in every 
direction, or dependent only on the curvature, as assumed in the in- 
vestigations. If we admit his calculations of the longitudinal and 
transverse strains produced at the navel by the abdominal muscles, 
the hydrostatic pressure which they can produce when acting toge- 
ther, will be the sum of that which they can produce separately, 
and Prof. Haughton would get the mean of the two values he ob- 
tains on p. 228, each of which is about half his former result. He 
goes on to seek for an escape from this difficulty by shewing that the 
pressure producible by the tension immediately above the pubes would 
be greater, forgetting, it would seem, that the pressure produced in 
a closed cavity by the contraction of part of its walls must be that 
due to the muscular forces where these are weakest. After this 
pressure has been reached, the contraction of the more powerful 


21—2 


316 REVIEWS AND NOTICES OF BOOKS. 


muscles would only alter the form of the cavity and forcibly extend 
the weaker muscles without increasing the pressure. 

The “actual experiment” described on p. 163 does not seem to 
prove much. He there determines the greatest weight which could 
be raised by the action of the abdominal muscles, when it was 
put upon a disc placed over the navel of a man lying on* his back. 
If the surface of the abdomen in the neighbourhood of the dise was 
flat the experiment can tell us nothing of the pressure produced by 
the tensions at the navel under normal circumstances. If, as is 
probable, the weighted dise caused a considerable depression, the 
direct action of the muscles would tend to raise it with a foree which 
would be added to that exercised by the pressure of the contents of 
the contracting abdomen and comparable with it in magnitude. 

The substance of a long section (pp. 74—1386) on the compara- 
tive anatomy of the tendons of the Hand and Foot, and their 
mechanical uses, was laid before the Royal Society (Proc. Roy. Soc. 
XvilI. p. 359), and gave rise to some controversy at the time be- 
tween Prof. Haughton and Prof. Humphry of Cambridge (see 
British Medical Journal, 1872, Vol. u. pp. 87, 228, 254, 341). 
Prof. Haughton finds in most animals, and notably in the car- 
nivora, that the sum of the cross sections of the flexor tendons of 
tMe digits of the hindlimb is considerably larger when the ten- 
dons are measured in the toes, than when they are measured above 
their point of division, while in the fore-limb he finds an opposite 
result. 

He attributes this to economy of material rendered possible by 
the existence of an enormous friction amounting sometimes to 
something like forty per cent. of the whole force applied to the 
tendon. He believes the action of a hand and foot to be essen- 
tially different, and that while in a hand the flexor muscles are 
employed in actually flexing the joints, in the foot they are habitu- 
ally employed in resisting forcible extension during the actions of 
walking, scratching, &e. - 

No doubt there are actions in which the muscles of the hind-limb 
are employed in this way, though it seems very questionable whether 
this is their habitual or most important action—but the question is 
one which could hardly be settled without a very careful observation 
and analysis of the gait and action of different animals, some- 
thing like what the Webers did for man in their “ Gehwerkzeuge.” 
It does not, however, seem necessary to answer this question in order 
to attack Prof. Haughton’s theory. It is difficult to conceive how any 
one who has seen and handled a tendon and its sheath can believe 
that there is any considerable friction between them unless he is 
driven to it by the positive evidence of direct experiment. But of this 
there is no trace in the book, and there does not seem to be any 
sufficient reason in the observations quoted for making such a 
supposition even provisionally, Even if we assume, with Prof. Haugh- 
ton, that all tendons contract and lose weight equally in drying, so 
that the weight of a length of “dried” tendon is an accurate measure 
of the cross section of the living one, it does not follow that the 


REVIEWS AND NOTICES OF BOOKS. aig 


strength of tendons is exactly proportional to their cross section. 
Tt may well be, for instance, that the exigences of nutrition (always 
a difficulty in tendons) require a somewhat looser texture in the case 
of large than of small tendons, and it is quite conceivable that the sum 
of the breaking strengths of the tendons of the fingers may be no 
less than that of the breaking strengths of the tendons of the 
muscles which are connected with them, although the sum of their 
cross sections may be decidedly smaller. 

With regard to the extra strength of the tendons distributed 
to the digits of the hind feet there is no difficulty. In many actions, 
and particularly in the tearing and scratching actions of the hind- 
limbs of the carnivora, the resistance to be overcome is by no means 
regularly distributed over the digits—and a tiger which was liable 
to break its tendon every time it applied the full force of its flexors 
to tear open its prey, while only one or two claws had good hold, 
would be soon worsted in the struggle for existence. 

Much might be said, if time and the limits of an article per- 
mitted, on the interesting and suggestive, though not altogether 
satisfactory sections on the work done by muscles, and on general 
laws of muscular action—pp. 24—62, and 442—485. Till we know 
a good deal more of the mode of action of a contracting muscle 
than we do at present, it is probably scarcely possible to discuss 
satisfactorily what Prof. Haughton calls the “statical work done 
by muscles in continued contraction,” i.e. the causes of the fatigue 
experienced when the muscle resists a pressure without doing exter- 
nal work at all. It does not appear that much is gained by Prof. 
Haughton’s method of expressing it as the product of the moment, 
taken about the joint of the forces resisting the muscle multiplied 
by an “angular velocity,” the physical significance of which it is 
difficult to see, particularly as the hypothesis that the extended 
arm begins to fall and is raised again at intervals corresponding to 
the period of the muscular susurrus is discussed and rejected. 

The book concludes with a section on certain “General laws of 
muscular action.” The author says: 

“JT have been led to the establishment of the three following 
laws, the proofs of which I shall give in detail :-— 

Law I. Jn comparing together different muscles, the work done in 
contracting is proportional to the weight of each. 

Law II. Jn comparing the same muscle (or group of muscles) 
with itself, when. contracting under different external conditions, the 
work done is always constant in a single contraction. 

Law III. When the same muscle (or group of muscles) is kept 
in constant action until fatigue sets in, the total work done, multiplied 
by the rate of work, is constant. ; 

For Law I. we are referred to Borelli, who enunciates what is 
equivalent to it, provided the muscles are taken from the same animal. 
This important proviso is tacitly ignored in Prof. Haughton’s 
statement. How far the law, even with the proviso, may be 
treated as an accurate numerical statement seems somewhat doubtful, 
and no evidence on the subject is given. 


318 REVIEWS AND NOTICES OF BOOKS. 


The ‘‘ proof” of Law IT. consists of a discussion of some experi- 
ments of Prof. Stanley Jevons, in which it is shewn that, due 
allowance being made for the work done in setting the arm in 
motion, the work done in throwing weights to the greatest possible 
distance is nearly constant for different weights. 

It is not obvious, nor I think probable, that the law would 
hold good if the time occupied by a contraction varied considera- 
bly in the different cases. 

The third law is deduced from three kinds of experiments. 
(1) Raising a weight by a pulley. (2) Holding the arm extended 
with weights in the hand. (3) Raising weights by holding them in 
the hand and raising the arm from a vertical to a_ horizontal 
position, the weights raised and the time of raising them being 
varied in different ways. Assuming that ‘‘ work” is done by re- 
sisting pressure which may be measured by the moment of the 
forces resisted, multiplied by a constant (the “angular velocity” 
of an earlier section), and that the work done in experiments of the 
elass (3) may be measured by adding the “work” estimated in 
this way to the actual dynamical work performed, the results agree 
very fairly with curves drawn on the supposition of the correctness of 
the law. Considering, however, that the portions of the curves under 
observation do not differ materially in shape from those which would 
result from a different law, and that the results in the cases of the 
slower movements can hardly fail to be complicated by the process 
of refreshment which goes on simultaneously with that of fatigue, 
.it seems very questionable whether much weight can be given to a 
law, of the rationale of which no explanation is offered. 

I have endeavoured in the preceding paperto shew thatasatisfactory 
treatise on animal mechanics cannot be written without an amount 
of cautious and patient criticism, and verification of hypotheses and 
arguments, which I have failed to find in Prof. Haughton’s book ; 
and I have thus been led to dwell almost exclusively upon those 
portions of it which seem to me imperfect or erroneous. I cannot 
close, however, without recommending the book strongly to the 
attention ofall who are interested in the subject. It contains a great 
deal that is both interesting and important on the subject of muscular 
action, particularly in the sections on the Classification of Muscles, 
and on the hip and shoulder-joimts to which attention has been 
called above, and the reader will not fail to find much that is both 
ingenious and suggestive even in those parts of the book from which 
he may be compelled to dissent. 

C. TROTTER. 
Trinity CoLuece, 
CAMBRIDGE, 


Outlines of Physiological Chemistry.—By CHaArLes HENRY RALFE, 
M.A., M.B. Cantab. London, H. K. Lewis. 


Durine the last few years physiological science in England has been 
advancing with remarkable strides. Physiological laboratories are 


REVIEWS AND NOTICES OF BOOKS. 391 


now no longer adjuncts to a few of the large metropolitan medical 
schools, but have been already established in several provincial schools 
of natural science. This is all very gratifying ; but we cannot close 
our eyes to the fact that the chemical side of physiology has been, and 
is, grievously neglected. The old proverb, “ what is everybody’s busi- 
ness Is nobody’s business,” seems to apply with painful force to phy- 
siological chemistry. The science is claimed by two different classes 
of scientific men, chemists and physiologists, each having by their 
education a different method of regarding phenomena brought under 
their notice. The result is easily predicted, and only too apparent in 
the curriculum of our lecturers and the various examination papers 
which are so constantly being set. The fact that animal chemistry is 
in the hands of persons who regard it from two different points of 
view ought to be its great safeguard ; and when chemists and phy- 
siologists come to work more together let us hope that the amount of 
work done may equal that produced by workers in the other branches 
of physiology. ‘The little book before us professes to have been com- 
piled “in the hope that it may furnish students and practitioners 
of medicine with a concise and trustworthy laboratory guide to the 
qualitative and quantitative analysis of the tissues, fluids and excretory 
products of the human body.” The want of such a work in the 
English language as that described above was doubtless very great ; 
but we rather hesitate to say that the author has fulfilled the task he 
has imposed upon himself with entire success. On our first cursory 
glance over the book we were inclined to imagine from the condensa- 
tion of the matter and the profusion with which formule are scat- 
tered over its pages, that it was a production called forth for the 
purposes of “cramming,” rendered necessary by the unfortunate 
examination systems of this country. On more careful examination, 
however, it appeared to be evidently a much more conscientious 
work, intended to give beginners an introduction to the subject, and 
as such all honour is due to the author. We think, however, it would 
~ have been better if he had confined his attention more to making the 
book a laboratory guide, and had sacrificed some of its completeness 
in the matter of formule and some other respects, at the same time 
giving fuller directions as to the practical working of the experiments 
described. To take an example, the directions given for establishing 
a pancreatic fistula are as follows: “The pancreatic fluid may be 
obtained for examination by opening the abdomen of a dog, and 
drawing down the duodenum, and separating the lower and larger 
pancreatic duct and passing a canula into it; the duodenum is then 
returned and the wound closed by a ligature, the canula left 
hanging out.” This is so terse that we venture to predict that an 
operation conducted on these directions would be nearly certain 
to fail; no hint is given as to the position, size, or direction of the 
incision ; in what part of the abdomen the pancreatic duct is to be 
sought for; the canula apparently is not to be tied in; and no pre- 
cautions against accidents of bleeding, &c., which are so liable to 
embarrass the student during an operation, are given. The isolation 
of pancreatin is thus described: ‘‘ Pancreatin is obtained by rubbing 


320 REVIEWS AND NOTICES OF BOOKS. 


down the pancreas of a freshly-killed animal, in full digestion, with 
pounded glass, from which an aqueous solution is made and from 
which the panereatin may be precipitated by alcohol.” We do not 
know who was Mr Ralfe’s authority for the above statement, nor 
can we say whether he had tried the experiment and was satisfied 
with the result, but the product of his operation would certainly be ra- 
ther more than pancreatin. Indeed the purpose of the operations above 
described is not quite obvious, and would seem to do nothing but pre- 
cipitate the soluble proteids resulting from the mashing up of the 
gland. The very interesting action of the pancreatic juice when con- 
tinued for some time is thus dismissed: ‘ According to Dr Kiihne 
the prolonged action of pancreatic juice on newly formed peptones 
leads to the formation of leucin and tyrosin.” This is not the state- 
ment one would expect in a “laboratory guide.” 

An appendix of fifteen pages is added to the book, containing 
a description of the appliances and methods of quantitative analysis; 
but this again is eminently impractical, and if used in the laboratory 
would lead to unreliable results. One method for the estimation of 
nitrogen is given, but no account whatever of the ultimate analysis of 
organic compounds is to be found. On the whole, we think that the 
little book is best suited for examination work, and unless the author 
modifies it very greatly it will not take up a position as an instructor ~ 
to earnest practical students in the difficulties of animal chemistry. 


Descriptive Catalogue of the Teratological Series in the Museum of the 
Royal College of Surgeons of England. By B. THompson Lowne, 
M.R.C.S. London, 1872. 


Ir is unfortunate that we do not possess in English any systematic 
treatise on Teratology. We have indeed many excellent essays on 
certain departments of the subject, such as the article “‘ Hermaphro- 
ditism” by the late Sir J. Y. Simpson, originally published in 7'odd’s 
Cyclopedia, and since reprinted in Simpson’s collected memoirs, and 
Allen Thomson’s well-known essay en Double Monsters, which ap- 
peared in the London and Edinburgh Monthly Journal, 1844. 
Vrolik also furnished a short article to the 4th vol. of the Cyclopedia 
of Anatomy. But there is no treatise in our language to be put on a 
par with Geoffrey Saint Hilaire’s Histoire générale et particuliére des 
anomalies de Vorganisation chez Vhomme et les animaux, or with 
Forster’s Die Missbildungen des Menschen, or with Vrolik’s Vrucht 
van den Mensch en van de Zoogdieren. 

Mr Lowne’s book to some extent supplies the omission to which 
we have referred ; for it is not merely, as its title would lead us to 
infer, a descriptive Catalogue of teratological specimens in a particular 
museum, but many important general principles are expounded. The 
author prefaces his Catalogue with a concise general introduction to 
the subject. He arranges the specimens under the heads of Varia- 
tion, Duplicity, Excess of Growth, Arrest of Growth, Arrest of 
Development, Disease, and then discusses briefly yet clearly the 


REVIEWS AND NOTICES OF BOOKS. 321 


character and extent of each of these classes. In the body of the 
Catalogue also many general observations may be found which will 
interest those engaged in the study of Malformations. 


The Causation of Sleep. A Physiological Essay. By James CAppie, 
M.D. Edinburgh, 1872. 


In this essay the author keeps before him the following questions :— 
In what respect do the condition and action of the brain, or its 
relations, during the continuance of sleep, differ from those which are 
present during wakefulness? what is the physiological sequence of 
change from the one state to the other, and on what special change 
do the more characteristic phenomena of sleep depend? He argues 
that in the causation of sleep there is a combination or succession of 
conditions. The first is a modified nutrition in the nervous texture : 
the last is pressure on the surface of the brain, by an increase in the 
proportion of blood there, due to engorgement of the veins of the pia- 
mater, The connecting link between the two is a weakened capillary 
circulation through the brain itself, owing to diminished activity of 
brain tissue, and a less energetic evolution of nerve force. 


Lessons in Elementary Anatomy. By Sr Georce Mrivart, F.R.S. 
London, 1873. 


Mr Mrvarr has prepared, in the hope that it may serve as a handbook 
of Human Morphology, this volume as one of the series of school 
class-books now in course of publication by Macmillan. He devotes 
upwards of one-half the book to the skeleton, and leaves only the 
smaller proportion for the consideration of the other organic systems. 
His acquaintance with facts and his descriptive style are such as we 
. might naturally expect from so skilful a comparative anatomist and 
so practised a writer as Mr Mivart. We have some doubts however 
how far the book is well adapted to be introduced as a lesson-book in 
schools, unless the teachers are themselves accomplished anatomists, 
and the schools are provided with a well-selected museum of osteo- 
logical and other specimens. Object teaching is the very essence of 
a biological training, and without it instruction in natural science 
partakes too much of the nature of “cram.” 

We can recommend this book however to the notice of those 
teachers of anatomy in our schools of medicine, whose instructions 
are imparted solely from the stand-point of human anatomy. In it 
they will be enabled to recognise that the human body consists not 
only of parts on which the surgeon may be required to operate, or 
the diseases of which the physician may have to diagnose, but that 
the study of its mode of construction teaches important lessons on 
those great principles of organisation which it shares in common with 
other vertebrata. 


322 REVIEWS AND NOTICES OF BOOKS. 


The Comparative Anatomy of the Domesticated Animals. By A. 
CHAVEAU, Professor at the Lyons Veterinary School. Second 
edition, revised and enlarged with the co-operation of S. 
ARLOING, late Principal of Anatomy at the Lyons Veterinary 
School, Professor at the Toulouse Veterinary School. Translated 
and edited by George Friemine, F.R.G.S., M.A.L, Veterinary 
Surgeon, Royal Engineers. 


We welcome the appearance of this book in English as likely to 
raise the standard of veterinary science in this country. The work 
of M. Chaveau is not merely an excellent descriptive treatise on 
Anatomy, but, as stated in the preface, it aims at a philosophical 
character. As an enthusiastic admirer of Cuvier and Geoffrey St 
Hilaire, the author glories in belonging to their school, and observes 
that the prevailing idea in the work has been inspired by their 
labours. Accordingly, the object of the book being the study of 
veterinary anatomy, the Horse is usually taken as the type and 
all the organs of an apparatus are studied in the Horse; afterwards 
the same organs in the other species are studied in the same order, 
and, finally, they are compared with the corresponding parts in man. 
The descriptions are good and clear. A microscopical account of the 
several tissues and organs is added. The translation is well done; 
and it forms a very valuable addition to the library of the veterinary 
student. It is copiously and well illustrated, many of the illustra- 
tions are original; and we regret to be obliged to express the wish 
that the sources had been indicated from which those that are not so 
have been taken. 


Handbook for the Physiological Laboratory. By E. Kier, M.D.; 
J. Burpon-Sanperson, M.D., F.R.S; Micuarn Foster, M.A., 
M.D., F.R.S.; and T. Lauprer-Brunton, M.D., D. Se.; edited 
by J. Burpon-Sanperson. London, J. and A. Churchill, 
1873. 2 Vols. 1 Vol. Text, 1 Vol. Plates. 


Tuts work is one of the most important contributions to scientific lite- 
rature which has appeared during the present year. It isa book which 
will be much appreciated by all investigators in the field of physiology, 
because it contains a description of the methods of research which 
have contributed so much to the advancement of physiological science 
in recent times. The authors are well known as physiologists who 
have devoted special attention to the subjects they profess to teach, 
and they were therefore eminently qualified for the task they have 
on the whole successfully accomplished. Physiology has now taken . 
possession of such a wide area in the field of science, and includes a 
knowledge of so many correlated subjects, such as physics and chemis- 
try, as to render it almost impossible for one man in a short life- 
time to attain to the position of an authority or expositor in all 
departments. One man devotes himself to histology, another to 
experimental physiology, while a third is known chiefly as a physio- 
logical chemist. We therefore think the editor acted wisely in 


REVIEWS AND NOTICES OF BOOKS, 323 


calling to his aid men having special knowledge, and in our opinion 
he could not have made a better selection than in the present 
instance. On the whole, Dr Sanderson has succeeded in producing 
a work which is a credit to physiology, and which has no counterpart 
in any other language. 

Having made these preliminary statements, we shall now notice 
more par ticularly the different sections of the work, We consider it 
unfortunate that the plates were separated from the text, because it is 
always more convenient in reading to be able to refer easily to any 
illustration. The responsibility of this, however, the editor throws 
upon the publishers. It is evident sufficient care has not been taken 
in the text as to references to the plates, and in the same plate 
various figures are mixed up so as to be at the least confusing, more 
especially 4 to a tyro in physiology. These, however, are minor defects, 
which do not seriously detract from the value of the work, 

The section upon histology is by Dr Klein, now Assistant Pro- 
fessor in the Pathological Laboratory of the Brown Institution, and 
formerly Privat Docent in Histology in the University of Vienna. 
Histology, during the last ten years, has made great advances, not so 
much in the way of generalizations of wide scope like that of the cell- 
theory of Schleiden and Schwann, but more in the applications of 
better methods of observation. Attempts have been made success- 
fully to examine tissues in two ways: first, under conditions similar 
to those in which they exist during life; and secondly, after the action 
of re-agents which have the property of “fixing” delicate tissues or 
organs in their normal shape and position, or of staining them in such 
a way as to render minute structure apparent. Full details are given 
by Dr Klein respecting each of these modes of investigation. It 
appears to us that this section is more suitable for the advanced 
student in histology than for the ordinary medical student of our 
universities and colleges. It would be of almost no use to the latter; 
he has not sufficient ‘preliminary knowledge, and when he commences 
- histology he scarcely knows one end of a microscope from the other. 
For him therefore words or phrases such as “objective,” “eye-piece,” 
** Hartnack’s No. 10 immersion,” &c., are unintelligible. He has no 
ideas of the optical principles or mechanical construction of the micro- 
scope. Dr Klein would undoubtedly have added much to the utility 
of this portion of the Handbook by giving a short description of the 
microscope as an instrument of research, and by defining the terms he 
employed. On the whole, however, Dr Klemm has given us an admir- 
able compendium of the methods of histological investigation, which 
will be of great service to those who have not had opportunities of 
becoming acquainted with what has recently been done in this direc- 
tion. This part will often be referred to by advanced students, and 
will be a valuable guide to them. 

The sections on blood, circulation, respiration, and animal heat are 
by the editor, Dr eee aadienson Under the first head, ‘‘the Blood,” 
we have admirable expositions of the chemical and physical proper- 
ties of that fluid. We have also a minute description of the methods 
of obtaining and analysing the gases of the blood, a matter of much 


324 REVIEWS AND NOTICES OF BOOKS. 


importance. These methods cannot be practised by the ordinary 
student of medicine, but for the student of physiology a knowledge of 
them is invaluable. The chapter on the circulation is subdivided 
into two parts, first, the arteries, and, second, the heart. There 
is also a supplement relating to the absorption by the veins 
and Jymphatics. The method of the author with reference to the 
first part is, first, to measure arterial pressure or tension ; secondly, to 
record the variations of arterial tension during each cardiac period 
in two ways: (a) by means of the spring kymograph of Fick, 
and (b) by the sphygmograph of Marey ; thirdly, to watch the cir- 
culation in transparent tissues such as the fish-tail, and the 
mesentery, web, and tongue of the frog, or in the mesentery of 
warm-blooded animals ; and, fourthly, to study the influence which the 
vaso-motor nerves have on the blood-vessels. As the blood-vessels 
during life are not passive elastic tubes conveying fluid, but also 
tubes constantly undergoing changes in calibre from the influence of 
the nerve centres through the nerves distributed to them, Dr San- 
derson’s method is truly philosophical, and is the only one by 
which the physiology of the circulation can be correctly apprehended. 
With regard to the heart, Dr Sanderson describes how the move- 
ments of the heart, the cardiac impulses and the action of the valves 
may be studied. Here, again, he had to deal with a living con- 
tractile organ under the control of various stimuli, and accordingly 
he describes the experiments necessary for demonstrating the influ- 
ence of the intrinsic ganglia, and of the sympathetic and pneumo- 
gastric nerves. We doubt much if all the experiments described 
could be shown to a class of Practical Physiology, but the details 
given will be useful to the investigator who wishes to corroborate 
previous experiments, or to pursue original research, The chapters 
on respiration and animal heat abound in useful practical details. 
This part of the Handbook justifies the high estimation in which 
Dr Sanderson is held as a practical physiologist, and will prove a 
great assistance to all advanced students. 

The functions of muscle-and nerve are dealt with by Dr Michael 
Foster in a very lucid manner, All the experiments detailed in this 
section require, to ensure success, attention to minute arrangements, 
which are here given with great precision. Many cannot often be 
demonstrated to a class of students on the first trial because the 
conditions are complex, but any one carefully following the directions 
given by Dr Foster will no doubt obtain all the results. As an 
illustration take the case of the law of contraction of Pfliiger which 
is confessedly one of the most difficult points to demonstrate in a 
course of practical physiology. Even eminent authorities who have 
published on the subject, differ as to the results they have obtained, 
no doubt while alone and without the distractions of a class. A 
teacher cannot expect to be able to demonstrate these facts off-hand 
to a class, Many of them can only be performed by personal devotion 
of quiet hours of work, and cannot successfully be done for or by 
medical students who have many other classes to attend to and little 
time. But any one who professes to study physiology as such, with 


REVIEWS AND NOTICES OF BOOKS. Sey) 


the view of becoming a teacher or investigator, ought to study care- 
fully this section of the work and train himself by performing all 
the experiments detailed so succinctly by Dr Foster. We miss, 
however, any reference to apparatus for measuring the rapidity of 
nerve current, such as the myographion of Helmholtz and Du-Bois 
Reymond. 

The last section of the work is on the physiology of digestion 
and secretion, by Dr Brunton. Of this part we can only write in 
terms of unqualified praise. Dr Brunton evidently understands the 
difficulties of practical teaching, and he has accordingly specially in- 
dicated those experiments which can be readily done in the presence 
of a class of students, and which are specially adapted for teaching 
purposes. In an appendix we have many useful practical notes on 
manipulation. 

Tn conclusion we have to state our opinion that this Handbook, 
on the whole, is worthy of the reputation of the authors. It has 
defects, but these are not of much importance. There is a want of 
balance between the different parts which was, of course, to be ex- 
pected in a work written by different men. We are surprised at the 
entire omission of the experimental physiology of vision, hearing and 
voice. These subjects are of great importance and can be readily 
illustrated by experiment. They are carefully taught in courses of 
physiology in French and German universities, and it is to be much 
regretted that in this work, which we regard as an expression of the 
state of physiological knowledge in Great Britain, no reference is 
made to them. This ought to be remedied in the next edition. 

This Handbook will now find a place in every laboratory. It is 
not suitable as a text-book for the ordinary student of medicine, but a 
judicious teacher will find in it numerous illustrative experiments 
and procedures to which he can direct the student’s special attention. 
Tt is a handbook for advanced students, for teachers and experi- 
mentalists. Its publication is a stimulus to physiology, and we shall 
- goon expect to see fruits in the shape of substantial work done by 
many of our younger physiologists, 


REPORT ON THE PROGRESS OF ANATOMY. 
By Proressor Turner’. 


OssEous System.—Joseph Hyrtl describes and figures (Denk. der 
Math. Naturwiss. der Akad. Vienna, 1871) specimens of human 
crania which possessed DUPLICITY OF THE CURVED LINE ON THE PA- 
RIETAL BONE. He also figures three feetal crania in each of which a 
suture, beginning at the side of the coronal, traversed the parietal 
bone in the antero-posterior direction, so as in one case almost, and 
in two cases entirely, to subdivide it into an upper and a lower 
segment. Wenzel Gruber records another case of SUPERNUMERARY 
CARPAL BONE (Bull. de (Acad. Imp. de St Pétersb. vit. 705), from 
subdivision of the scaphoid into two secondary bones: and in Virchow’s 
Archiv, uxv. 425, he describes an ELEVATION on the POSTERO-SUPERIOR 
ANGLE of the left scApPuULA, in relation to the insertion of the serratus 
magnus, and possessing In connection with it a synovial bursa. H. 
Wolfermann communicates (eichert u. Du Bois Reymond’s Archiv, 
1872, 312) the results of a comparative enquiry into the ARCHITEC- 
TURE OF THE Bones.—John Struthers gives an account (Lancet, Feb, 
15, 1873), of the occurrence of the Procrssus sUPRACONDYLOIDEUS 
HUMERI in the father of a family and in four out of seven of his 
children. The father and three children had it in the left arm, the 
fourth child in both arms.——Ludwig Stieda publishes a separate 
memoir, Leipzig, 1872, ON THE FORMATION OF Bone. In the first 
part of this essay he describes the observations which he has specially 
made into its development, and in the second part he gives a his- 
torico-critical review of the researches of previous observers. He 
first investigates the bones which form in fibrous membranes, as the 
lower jaw, and concludes that the bony tissue arises out of an osteo- 
genetic substance (osteoplasts), which proceeds from indifferent 
embryonic connective tissue. He then examines into the formation 
of the “cartilage bones,” and his conclusion is that the cartilage only 
possesses a provisional import, that the cartilage tissue atrophies and 
disappears, and that in its place appears the new formed osseous 
tissue, which stands in no genetic relation with the cartilage, but 
which arises as in the “membrane bones” out of the osteoplastic 
cells. Further he maintains that the marrow cells (osteoplasts) do 
not arise from the cartilage cells, but that the medullary tissue is a 
direct prolongation of the osteo- -genetic tissue found lying beneath 
the periosteum. He regards true bone as a tissue swt generis, which 
belongs to the category of the connective substances. Bones grow 
by the apposition of new bony substance to the old: in the long bones 
the increase in length is by apposition at the confines of the carti- 
lage, and in thickness by apposition of new material directly beneath 


1 To assist in making this Report more complete Professor Turner will be 
glad to receive separate copies of original memoirs and other contributions to 
Anatomy, 


REPORT ON THE PROGRESS OF ANATOMY. 327 


the periosteum; whilst the cranial bones grow on their surfaces, 
from the periosteum, and at their margins from the intermediate con- 
nective tissue. A translation of C. Heitzmann’s researches on 
Bone AND CARTILAGE appears in Quart. Journ. Mic. Sc., April, 1873. 


Frero-CartitaGe.—Oscar Hertwig describes (Schultze’s Archiv, 1x. 
80) the structure and development of elastic tissue in the Yellow Car- 
tilages. He examines the cartilage of the human ear and of various 
mammals. The elastic fibres arise immediately after the first appear- 
ance, of an inter-cellular substance, or simultaneously on the surface 
of the protoplasm. The cells which form the earliest elastic fibres 
lie in rows perpendicular to the surface of the cartilage. From the 
commencement the fibres are insoluble in potash-ley. They are not 
derived from a conversion of a homogeneous cartilaginous basis sub- 
stance, which is first formed, but directly from the protoplasm of the 
cells. Subsequent development of the fibres is due to intus-suscep- 
tion in the extra-protoplasmatic substance, so that new fibres either 
enclose the old or grow out of them; or in the immediate vicinity 
of the persisting cells, which continue to exhibit their formative 
activity in various ways. 


Muscurar System.—J. Beswick Perrin records (Med. Times and 
Gaz., Dec. 7, 1872, and Jan. 11, 1873) variations in muscular ar- 
rangements which he has met with in the Dissecting Rooms of King’s 
College, London, during the sessions 1868—69, 1869—70 and 1870 
—71. He classifies them into adventitious, which include muscles 
not common to man; hetero-morphous, where the human arrange- 
ment is departed from; absentaneous, where muscles, which, as a 
rule, are present, are occasionally aberrant. The most important 
varieties are additional fusiform muscles from the scapula to the 
levator anguli scapule: a third head to the biceps brachii from the 
capsular ligament of the shoulder; another from the humerus below 
the insertion tendon of the coraco-brachialis; a levator clavicule; an 
extensor medi digiti; a peroneus quartus; and various modifications 
in the extensor carpi radialis longior. Davies-Colley, F. Taylor and 
B. N. Dalton record variations in muscles observed in Guy’s Hospital 
Dissecting Room from Oct., 1870 to June, 1872 (Guy's Hosp. Rep. 
1873). The most important are: a levator claviculae; variation in 
the rhomboidei and digastricus: a second rectus capitis posticus 
minor; specimens of rectus sternalis and supracostalis: a biceps 
brachvi with four heads: variation in external oblique: additional 
head to adductor longus. John Tweedy relates (Lancet, March 
29, 1873) a case of absence of the thoracic portion of the pectoralis 
major and the whole of the pectoralis minor: and at the meeting 
of the Clinical Society, Feb. 28, a case of absence of the left 
pectoralis major was reported by Burney Yeo.—W. Turner may 
refer here to a subject he dissected in 1865, in which on the right 
side a hiatus existed in the pectoralis major owing to the absence of 
any fibres of origin from the second costal cartilage of the correspond- 
ing part of the sternum. Wenzel Gruber describes (Bull. de? Acad. 
Imp. de St Pétersb. vit.) a case of right m. cleido-hyoideus and left 


328 PROFESSOR TURNER. 


m. supraclavicularis; also a right m. sterno-fascialis which ended in 
the fascia of the neck in the trigonum omo-hyoideum ; also a super- 
numerary m. obliquus eat. abd. from the eleventh right costal cartilage 
to end in the aponeurosis of the external oblique; also a m. protractor 
arcus cruralis from the horizontal ramus of pubis to the crural arch: 
also a tensor of the posterior layer of the sheath of the rectus abdomi- 
nis arising from the tuberculum pubis; also a ¢ensor not only of that 
sheath but of the fascia transversalis; also a m. obliquus internus the 
inguinal portion of which was absent; also, p. 736, a variety of the m. 
tensor fascie suralis, which arising from the semi-tendinosus passed 
down the back of the leg to end in the back of the tendo-Achillis. 
Bruhl has described (Journ. de Zool. 1873, 32) a case of a super- 
numerary long extensor of the great toe inserted partly into the tibial 
side of the base of the 1st phalanx partly along with the tendon of 
the short extensor. Jelenffy, of Pesth, discusses (Pfliiger’s Archiv, 
1873, p. 85) the action of the MuscC. CRICO-THYROIDEUS: he agrees 
with the view that it is a tensor of the vocal cords, but considers 
that it acts in a threefold way; a, through bending over of the 
cricoid cartilage backwards; b, through reciprocal separation of the 
cricoid and thyroid cartilages from each other in the direct sagittal 
direction; ¢c. through a forward movement of the angles of the 
thyroid cartilage. 


Several memoirs on the STRUCTURE OF TRANSVERSELY STRIPED 
Muscutar Fipre have recently appeared. T. W. Engelmann has 
two elaborate articles in Pfliiger’s Archiv, 1873, pp. 33, 155. He 
examines the structure of the fibre during rest, activity and rigidity. 
He recognises in a normal fibre at rest four different kinds of trans- 
verse stripes; I. a clear band, refracting the light very feebly, sub- 
divided into halves by II. an opaque strongly refracting stripe: 
III. a moderately opaque, tolerably strongly refracting band, in the 
middle of which is IV. a clearer stripe refracting the light more 
feebly. In all fibres with very broad transverse striz the opaque 
stripe II. may be subdivided into three transverse striz, a middle 
more opaque and two lateral clearer, and he considers that in those 
fibres where this subdivision has not yet been recognised it must be 
held to exist. I. and II. together form the isotropic band. JII. 
and IV. together the anisotropic band. In the stripe II. of the 
isotropic band, the middle more opaque portion he names interme- 
diate dise (zwischen scheibe), the two lateral, secondary dises (neben 
scheiben), whilst he adopts Krause’s term Grund membran for the 
three collectively. For IV. he adopts Hensen’s term, middle disc 
(mittelscheibe), whilst to the parts of III., in the middle of which 
IV. is situated, he gives the name of cross discs (Quer schetben). 
He then enters minutely into the characters of the individual discs, 
He holds that a fibre at rest is an aggregate of different kinds of 
discs which in the long axis of the fibre adhere so as to form pris- 
matic fibrille, in the transverse direction adhere so as to form in 
general parallel discs. In the second article Engelmann describes 
the fibre in a state of activity, the alterations in form, volume, optical 


REPORT ON THE PROGRESS OF ANATOMY. 329 


appearances and mechanical properties which take place during con- 
traction, and he concludes with some remarks on the cause of con- 
traction. Fr. Merkel completes (Schultze’s Archiv, 1x. 293) his 
memoir on striped fibre by giving a chapter on the process of con- 
traction as it appears with polarized light. C. Sachs commences 
(Reichert u. Du Bois Reymond’s Archiv, 1872, 607) a memoir on the 
structure of striped fibre, the consideration of which must stand over 
until it is completed and the plates are published. E. A. Schiifer 
communicates (Roy. Soc. London, April 3, Abstract in Nature, April 
24), an investigation on the structure of the muscles of the limbs of 
the water beetle. He considers that a muscular fibre consists of a 
homogeneous basis substance, apparently formed of alternate discs of a 
dim and a bright substance, in which are imbedded minute rod-like 
bodies having their axes coincident with that of the fibre itself. He 
calls these muscle-rods, and in the muscle at rest they are uniformly 
cylindrical, but when in action they terminate at each end in a knob 
so as to be dumb-bell shaped. These knobs give the appearance of 
the line of dots existing in the middle of each bright transverse band 
of the fibre (corresponding to stripe II. of Engelmann, Reporter); the 
dim dise again is that in which the shafts of the muscle-rods are im- 
bedded. From several considerations it is argued that the bright 
transverse bands in muscle are produced by the juxtaposition of the 
rod heads. The author states that all the basis substance of a fibre 
is doubly refractive, the rods alone being singly refractive. He re- 
gards the basis substance as the true contractile part, the rods as 
elastic structures to restore the fibre to its original length. P. Ter- 
gast communicates some observations on the nwmerical relations of 
the nerve fibres entering a muscle to the muscular jibres (Schultze’s 
Archiv, 1x. 36). 


Motion and Locomotion.—Marey contributes (Zobin’s Journal, 
Jan. 1873), a memoir on TERRESTRIAL Locomotion both in bipeds 
and quadrupeds. By means of a specially constructed shoe he studies 
the human movements in walking, running, galloping and leaping. 
He also adapts a special apparatus to the feet of the horse, and 
studies by its aid the various paces of this animal, The memoir is 
illustrated by figures of the apparatus, and by diagrams and tracings 
of the oscillations which have occurred during the movements. 
A. W. Volkmann examines (Virchow’s Archiv, 1872, tv1. 467) into 
the conditions under which a man standing erect can, without alter- 
ing the position of his feet, TURN HIS BoDY, so that the face may look 
almost backward. 


SynoviAL Mempranes.—Wenzel Gruber describes (Virchow’s Ar- 
chiv, Lv1. 428) a series of cases of HERNIA-LIKE PROTRUSIONS of these 
membranes: in the scapulo-humeral joint, the radio-carpal, the carpal, 
and the carpo-metacarpal. 


Nervous System.—W. Betz details his experience (Schaltze’s 
Archiv, 1x. 101) on the methods of examining the CenTRAL ORGANS 
oF THE NERVOUS SYSTEM in man. Axel Key and G, Retzius have 


VOL. VIL, SZ 


330 PROFESSOR TURNER. 


studied (Nord. Med. Arkiv, tv. and Schultzes Archiv, 1873, 308) the 
anatomy of the nervous system, more especially in connection with 
the arrangement of the Investine Tissues and the SeRous Spaces. 
The arachnoid membrane with its trabecule, the pia mater, the peri- 
neural and epineural investment of the cerebro-spinal nerves; the 
sub-arachnoid spaces in connection with the spinal cord, the roots of 
the nerves and the spinal ganglia ; and the structure of the Pacinian 
corpuscles, are described and beautifully figured. Anton Spedl 
makes (Reichert u. Du Bois Reymond’s Archiv, 1872, 307) some 
observations on the Purenic Nrerve.——Hagemann undertakes 
(Reichert u. Du Bois Reymond’s Archiv, 1873, 429) a comparative 
anatomical investigation into the StRUCTURE OF THE PINEAL GLAND. 
He distinguishes in it a supporting framework formed of connective 
tissue and a parenchyma. The septa formed by the supporting frame- 
work in the substance of the gland enclose parenchymatous tissue so as 
to form “follicles.” The parenchyma consists of two kinds of cells, 
roundish and spindle-formed. Nerve-fibres enter the gland anteriorly 
from the commissures of the peduncles; he thinks that they ramify 
between the follicles, and that the nerve ganglion cells which the 
pineal gland also possesses le in the same localities. C. Kupffer 
considers (Schultze’s Archiv, 1873, 387) the RELATION OF THE NERVES 
OF GLANDS TO THE GLAND-CELLS. He investigates the salivary glands 
of insects and the larve of Muscide with the view of ascertaining if 
the nerves do, as Pfliiger has maintained, come into direct relation 
with the secreting cells within the acini. His attention was first 
directed to the relations of the trachee to the gland, and he saw that 
not merely did they turn round the organ, but that a not incon- 
siderable number of fine twigs pierced the membrana propria. These 
twigs then ran between the large plate-like cells. From the sheath 
of these intra-cellular trachee fine, pale fibrillee proceeded which 
entered the cells; these fibrilla he considers to be nerves. He holds 
it to be completely established that tracheze enter into the composition 
of the salivary cells of the larvee of muscide. He then examined the 
salivary glands near the cesophagus of Slatta orientalis. These 
glands are provided with a very rich nervous apparatus, which derives 
its roots from the supra-cesophageal ganglion and the abdominal cord. 
In them the entrance of numerous nerves into lobules, the blending 
of the nerve sheath with the membrana propria, and the entrance of 
the nerve fibrillee into the interior of the acini may easily be observed. 
At the first glance it seemed as if the whole /ibrillenstrang entered 
completely into the adjacent or the two adjacent cells immediately 
on entering the acinus, but closer observation showed him that the 
greater part of the fibrille passed further into the interior of the 
acinus. The fibrille did not blend with the substance of the cells 
where they came in contact with them, but passed far into their 
interior so as to be imbedded within the substance of the cells. He 
has not succeeded in tracing a direct connection between the fibrille 
and the nucleus. He describes also certain hollow pear-shaped cap- 
sules, in connection with the secreting cells of these glands. 
J. Schébl describes (Schultze’s Archiv, 1873, 197) the mode of 


REPORT ON THE PROGRESS OF ANATOMY. Sol 


TERMINATION of the Nerves IN THE TacTILE Hairs of the mammalia. 
He recognises two kinds of tactile hairs, a small and a large, with 
intermediate transitional forms; the large possess well-developed 
cavernous bodies which lie between the outer and inner fibrous coats ; 
the small have none. The nervous tactile apparatus consists in the 
tactile hairs of the bat’s wing, of a nervous ring and a partial enve- 
lopment of the cellular body of the root with nerve fibres; in the 
tactile hairs in the mouse’s ear of a nerve ring and glomerulus or coil; 
in others of a nerve ring with which the modified glassy membrane is 
in connection. H. Hoyer gives (Schu’tze’s. Archiv, 1873, 220) a 
long communication on the Nerves oF THE CornEA. He examines 
the arrangement in all the divisions of the vertebrata with chloride of 
gold after Cohnheim’s method, and gives some beautiful figures of the 
extremely delicate nervous plexus found in this structure. In an 
article entitled Cyno-pHRENOLOGY (Boston Med. and Surg. Journal, 
Jan. 23, 1873) B. G. Wilder reports on a collection of brains and 
embryos in the Museum of Comparative Zoology, Cambridge, U.S. 
He considers that real advances into the determination of the co- 
existence of certain mental characteristics with a given pattern of 
brain must be made on the brains of dogs rather than of men, as it is 
more practicable to obtain the brains and ascertain the mental charac- 
teristics of a number of dogs than of a number of men. 


VascuLar SystemM,—Variations in arteries have been described 
(Guy's Hospital Reports, 1873) by Davies-Colley, F. Taylor, and 
B. N. Dalton ; the most important of which aré: left carotid arising 
from innominate, ten cases of high division of the brachial, in one of 
which a vas aberrans arose from one of the arteries just above the 
elbow, and after a course of 24 inches joined the ulnar : once the right 
kidney received an artery from the common iliac, once the external 
iliac divided into profunda and superficial femoral half an inch above 
Poupart’s ligament. M. Duret has described (Archives de Physio- 
_logie, 1873, 97) the ARTERIES OF THE MeEpuLLA OxstoncaTaA. He 
divides them into three sets, lateral to the roots of the nerves, median 
to the floor of the 4th ventricle, and arteries to the olivary bodies, 
pyramids, restiform bodies, &e. 


Kipney.—W. Turner records here two cases of Horsz SHOE 
Kipyey. One was found in a male subject in the dissecting room of 
the University of Edinburgh in November, 1872, the other was 
observed by J. Batty Tuke in the following month in the post-mortem 
examination of a patient who died in the Fife and Kinross Asylum. 
In both cases the isthmus connecting the right and left kidneys was 
situated below, and lay in front of the aorta and inferior cava close to 
the bifurcation. The concavity therefore of the horse shoe was 
directed upwards. The hilum was situated on the anterior surface of 
each lateral half, and in both specimens the ureter passed off from the 
lower end of the hilum on each side. In the Reporter's case a single 
renal artery supplied each half of the double organ, and a special 
branch entered the isthmus where it joined the left lateral half. In 
Dr Tuke’s case each lateral half had two renal branches of the 


22—2 


3 Pe PROFESSOR TURNER. 


abdominal aorta, and a special branch from the right common 
iliac artery entered the isthmus where it joined the right lateral 
half of the kidney: the weight of this specimen was 20 oz. 


Ma.rorMATions.—Three cases of malformation in the human 
foetus are described by Dr Orth (Virchow’s Archiv, trv. 492). They 
belong to the class Acarpiact, and two at least to the subdivision 
named by Forster AMORPHI. M. Roth records (Virchow’s Archiv, 
Lv. 197) several cases of formation of DiverTIcULA in connection 
with the duodenum; and on p. 271 N. Duhay describes a case of 
incarceration owing to ABNORMAL FoRMATION OF THE MESENTERY. In 
the same vol. p. 268, Kuhnt relates a case of DUPLICITY IN BOTH 
Hanps AND Feet. In each foot not only were four toes situated on 
the outer side of the great toe, but on its inner side two well-shaped 
toes were placed. Each hand had 5 digits only, but these were one 
middle, two ring and two little fingers, the thumbs and indices being 
unrepresented, On p. 421, A. Ewald describes a case of CONGENITAL 
HypertTrRopHY OF THE Lert Hann. Jos. Leidy mentions (Proc. 
Acad, Nat. Sci. Philadelphia, May 9, 1871) that in the Museum of 
the University of Pennsylvania is a specimen showing PoLYDACTYL- 
ISM IN THE Horsek, in which an abnormally developed metacarpal 
has a toe with two phalanges, one of which last is inclosed in an 
irregular hoof. He refers also to a case recorded by Mr Mason in 
the Proc. Roy. Asiatic Soc., Bengal, where the usual “splint-like 
rudiments of the metacarpals of the 4th toe in the fore feet, had given 
rise to an additional toe provided with three phalanges, of which the 
last is incased in a hoof.” In an essay on Herrepirary TRANs- 
MISSION OF SrrRucTURAL Pecuuiaritiss (Brit. and For, Med.-Chi. Rev. 
April, 1872), J. W. Ogle collects together a number of cases of defect or 
variation as to digits and other structural peculiarities. H. Gripat 
gives an account (Journal de Zoologie, 1873, 4) of an ACEPHALOUS 
CALF. P. D. Handyside in this Journal, Nov. 1872, communi- 
cates two cases of QuADRUPLE MAmm IN Men, and Max Bartels 
describes and figures (Reichert u. Du Bois Reymond’s Archiv, Nov. 
1872) another case. In Edin. Med. Journal, Jan. and Feb. 1873, 
T. Graham and P. D. Handyside describe a case of Hypospapia 
WITH CLEFT SCROTUM G. B. Ercolani gives a methodical deserip- 
tion (Mem. dell. Acad. di Bologna, 1872) of cases of diverticula of the 
Urinary Buiapper, of double bladder and of dilation of the Urachus. 


Ovum AND Ovary.—C. Weil (Medic. Jahrb. 1. 1873 and Medical 
Record, Jan. 8, 1873) describes the FecunDATION AND DEVELOPMENT 
OF THE OvuM OF THE Rassit. He observed spermatozoa moving in 
a lively manner between the 17th and 46th hours after fecundation, 
within the zona pellucida and in the albuminous envelope. Four 
instances are given of unchanged spermatozoa having been seen within 
the substance of the germ itself, besides numbers between the germ 
and the zona pellucida, When the ovum had reached the uterus, no 
spermatozoa could be seen either within or without the germ. Weil 
concludes that the spermatozoa unite with the germ in an intimate 
way in order that the ovum may be fertilised. Weil’s observations 


REPORT ON THE PROGRESS OF ANATOMY. 333 


confirm those of Bischoff relative to the changes which take place in 
the ovum as it passes along the oviduct. H. Kapff describes in 
Reichert u. du Bois Reymond’s Archiv, 1872, 513, researches into the 
Ovary and its relations to the Perrroneum. He agrees with Wal- 
deyer that a positive difference in colour exists between the surface of 
the ovary and the adjacent part of the peritoneum, He does not 
however confirm the statement that a sharp line of demarcation exists 
between the epithelium investing the ovary and that covering the 
peritoneal membrane, his view being that a gradual transition from 
the smaller peritoneal to the larger ovarial epithelial cells takes place. 
Further, he does not hesitate to say that a sub-epithelial connective 
tissue exists under the epithelium both of the ovary afd the adja- 
cent part of the peritoneum, so that, contrary to some recent state- 
ments, he believes that the entire constituents of the peritoneum are 
prolonged over the ovary. He then proceeds to criticise Waldeyex’s 
observations on the formation of the ovarian follicles and the ova 
from the ovarial epithelium, and his conclusion based on his own 
observations is quite opposed to Waldeyer, for he says that the surface 
of the ovary is in no way concerned in the formation of the fol- 
licles in its interior, therefore also not in the formation of the ova. 
Kapff then communicates some observations on the development of 
the genital gland. 


PuaAcenta.—Several contributions have recently been made _ to 
the anatomy of the human placenta, W. Turner in Proc. Roy. Soc. 
Edinb. May 20, 1872, and in this Journal, vu. 120, J. Matthews 
Duncan in Ed. Med. Journal, Jan. 1873, and F. N. Winkler in Archiv 
Sir Gynekologie, 1872, 238, bring forward various facts in support of 
the view that the placenta contains a cavernous sinus system, through 
which the maternal blood circulates. Braxton Hicks, again, in a 
memoir in 7'rans. Obstet. Soc. London, 1872, argues against such an 
intraplacental circulation of the maternal blood. He re-describes the 
“specimens, which he regards as supporting his views, previously 
recorded by him in this Journal, v1. 405, and adduces a number 
of additional observations. J. B. Pettigrew also in LHdin. Med. 
Journal, Nov. and Dec. 1872, publishes a lecture in which he specu- 
lates on the structure and function of the placenta. He conceives 
that in the human placenta, as in the diffused placenta of a mare or 
a cetacean, the villous surface of the chorion is applied to the mucous 
lining and capillary vessels of the uterus, that the former, in short, 
so far as relates to the foetal and maternal vessels, does not differ 
from other mammals. Further, he believes that the utricular 
glands persist, and that their secretion not only assists in nourish- 
ing the fetus, but acts as an osmotic medium for promoting the 
interchange between the blood in the capillaries of the fcetal villi 
and the capillaries in the uterine mucous membrane. Ingenious 
though this hypothesis undoubtedly is, yet the author does not 
support it by any detailed observations of his own of the presence 
of either utricular glands or maternal capillaries in a fully-formed 
human placenta. And though the structural conditions referred to 


334 PROFESSOR TURNER. 


are undoubtedly met with, as the Reporter has himself shown, in the 
diffused placenta of a cetacean, yet there is no evidence of their 
existence in the mature human placenta. 


MorrHouocy oF THE Liups.—Several essays on this subject have 
recently appeared. Burt G. Wilder draws up a memoir (Proc. Boston 
Soc. Nat. Hist. xtv. 1871) on Intermembral homologies, which he 
intends as an index of what has been done and what remains to be 
done for the elucidation of this ditticult department of morphology. 
In a paper by this author, noticed in our Report, m. 404, it was 
stated that in his opinion the thumb and little toe, minimus and great 
toe, radius and fibula, ulna and tibia are homologous parts. He 
believes, with Folz and Wyman, that the fore and hind limbs are 
antitropically or symmetrically related : but since the publication of 
his former papers he has been led to modify his previous views respect- 
ing the normal position of the limbs, and in so far he concedes 
a point to those who hold that the relation of the limbs is one of 
syntropy or parallelism. Wilder has collected together in a con- 
venient form a large amount of information respecting the opinions 
and statements of the various anatomists who have written on this 
subject. C. Martins, in an article in Dict. Encyc. des Sc. Medic. 
1873, compares the thoracic and pelvic extremities, and repeats his 
well-known theory of their morphology based upon the twisting of 
about 180° which the humerus has undergone.—— Alex. Rosenberg 
investigates the development of the skeleton of the limbs (Stebold uw. 
Kolliker’s Zeit. 1873, 116) in pigs, the len, sheep, horse and various 
birds. The investigation has been conducted with especial reference 
to the Darwinian theory of descent, and the facts which he describes 
are looked upon as of value according to their bearing either for or 
against this theory. ‘The tarsal, carpal, metatarsal and metacarpal 
bones form more particularly the objects of investigation. In G. 
M. Humphry’s Observations on Myology, Cambridge, 1872, the mor- 
phology of the muscles of the limbs is considered. 


CoMPARATIVE ANATOMY. 


QvuapRuMANA.—E. H. Giglioli publishes (Ann. del Museo Civico 
di Storia Nat. di Genova, Dec. 1872) Craniological studies on the 
Chimpanzee. He describes a new species by the name of T’roglodytes 
Schweinfurthir. R. Hartmann continues (Reichert und du Bois 
Reymond’s Archiv, 1872, 474) his observations on the anatomy of 
the anthropoid apes, and considers the cranium of the Chimpanzee. 
——Paul Broca studies (Revue d’ Anthropologie, 1.) the constitution 
of the caudal vertebre in the Primates without tails. He considers 
that the tail may disappear after three different ways ; in one the 
defect in development is due to a proportional atrophy in the true 
and false segments of the caudal apparatus, and is seen in Cynocepha- 
lus, Nycticebus and Loris: in the second it proceeds from the free end 
towards the base, as in the magot: in the third in a modification in 
the first segment much more than in the terminal segment. Here 
the first segment is fused with the sacrum to form a supplementary 


REPORT ON THE PROGRESS OF ANATOMY. 335 


sacrum, whilst the terminal segments remain mobile and constitute 
the coceyx ; this type is found in man and the anthropoids.——Jas. 
Murie gives some observations (Proc. Zool. Soc. June 18, 1872) on 
the Bornean ape. He figures both the pelvis and the cranium. 


Carnivora.—dJas Murie describes some anatomical features (Proc. 
Zool. Soc. June 4, 1872) of the Indian Wild Dog, Cuon primevus. 
He figures the head, soles of the feet, caecum and anal region of this 
animal. From a dissection of two specimens of the Two-spotted 
Paradoxure (Nandinia binotata) W. H. Flower has been able to 
show (Proc. Zool. Soc. June 4, 1872) that this animal is destitute of 
a cecum. It differs from all known Carnivora in the persistence 
throughout life of the cartilaginous condition of the posterior cham- 
ber of the auditory bulla. 


Pinnepepra.—Robert Walker records (Scottish Naturalist, 1873) 
the capture of a specimen, 22nd July, 1872, of the hooded seal, Cysto- 
phora cristata, at St Andrews.——J. E. Gray gives an account (Proce. 
Zool. Soc. May 21, 1872) of the New Zealand Sea Bear (Arctocephalus 
cinerius), and the North-Australian Sea Bear (Gypsophoca tropicalis). 

Crracea.—W. H. Flower has given (Z'’rans. Zool. Soc. 1871) a 
description of the skeleton of Berardius arnouwi, with an introductory 
chapter on the recent ziphioid whales. He defines the common cha- 
racters of the group, and then describes the special characters of 
the genera Hyperoodon, Ziphius, Mesoplodon and Berardius. The 
memoir is illustrated by three large plates representing the skeleton 
of Berardius—J. E. Gray publishes (Ann. Nat. Hist. Jan, 1873) a 
short criticism on Prof. Flower’s memoir on Lerardius and other 
Ziphioid whales.—H. J. Carter notes (Ann. Nat. Hist., March, 1873) 
the presence of the Sperm Whale in the Indian Ocean just within 
the Tropics.—In the Feb. number of the same Annals, J. E. Gray 
notes the geographical distribution, migration, and occasional habitats of 
Whales and Dolphins: and on p. 104 Jas. Hector notes the Whales 

-and Dolphins which frequent the New Zealand Seas, viz.: Neobalena 
marginata, Hubalena australis, Megaptera nove-Zealandie, Physalus 
australis, Catodon macrocephalus, Delphinus nove-Zealandie and 
Forsteri, Electra clancula, Pseudorca meridionalis, Grampus Richard- 
sont, Beluga Kingii, Globio-cephalus macrorhynchus, Epiodon chatham- 
tensis, Mesoplodon Layardii, Berardius Hectori and Arnouait. Dr 
Gray appends some remarks to Dr Hector’s paper. On p. 159 J. E. 
Gray states that Orca stenorhyncha has been found at Bohuslin in 
Sweden. On pp. 157 and 238 W. H. Dall describes parasites which 
infest the cetacea of the N. W. Coast of America. He refers them 
to the following species, Cyamus Scammoni, suffusus and mysticett, 
Coronula balenaris, and diademat, Crypto-lepas rhachianectis, and 
Otion Stimpsoni. W.H. Dall also gives a preliminary description 
(Ann. Nat. Hist. April, 1873) of three species of Cetacea, said to be 
new, from the coast of California: he names them Delphinus Bairdit, 
Tursiops Gillii and Grampus Stearnsii. In the same Annals, Dee. 
1872, is a translation of H. Burmeister’s memoir on Balenoptera 
patachonica and intermedia, and a description by C. M. Scammon of a 


336 PROFESSOR TURNER. 


species of Balenoptera which he names B. Davidsoni. J. E. Gray 
also proposes the name of Hpiodon Heraultii tor one of the specimens 
of Ziphioid whales which has been described. Jas. Murie gives 
(Trans. Zool. Soc. vit.) a long memoir on the organisation of the 
Caaing Whale, Globiocephalus melas, which is illustrated by nine 
beautifully executed plates and several wood-cuts. The various 
organic systems, except the osseous and nervous, are described. 
J. Reinhardt (Vidensk. Meddel. fra den naturh. Forening i Kjébenhavn, 
1872) makes some observations on Pseudorca Grayi. He does not 
think that any appreciable difference exists between it and Ps. 
crassidens, though he distinguishes many points in which it differs 
from Ps. meridionalis (Report, vit. 173). With reference to the 
large dolphins observed by Burmeister in 1850, in the Atlantic, 
Reinhardt considers that the form of the dorsal fin is somewhat in 
favour of the opinion that they were Pseudorce rather than true 
Globio-cephali. E. W. H. Holdsworth notes (Proc. Zool. Soe. 
April 16, 1872) a Cetacean observed on the West Coast of Ceylon, 
which was characterized by the presence of a dorsal fin estimated to 
be not less than five feet high, which stood erect on the highest part 
of the back, and was shaped like the pointed end of an ordinary 
sword, with the anterior edge slightly convex and the posterior 
straight. P. Gervais records (Journ. de Zool. 1. 537) the cap- 
ture of the carcase of a male Cachalot in a state of putrefaction near 
Biarritz in November, 1872; and, on p. 323, after giving copious 
extracts from the Reporter’s memoir on the placentation of the 
Cetacea (Report, vi. 469), Gervais refers to a foetus which he had 
extracted from the uterus of Delphinus delphis, and he figures the 
feetus both inclosed within the membranes and after removal from 
them. On p. 274 of the same vol. P. Fischer describes two species 
of Globio-cephalus, G. macrorhynchus and G. Edwardsi. 


SrrenrA.—P. J. van Beneden communicates (Bull. Acad. R. Bel- 
gique, Xx, 205) observations on the Osteology of the Dugong* and 
Manatee. F. Krauss (Reichert u. du Bois Reymond’s Archiv, 1872, 
257) gives an elaborate description with figures of the pelvic bones of 
the Manatee from Surinam. He had received a number of specimens, 
both male and female, some of which had the muscles still attached. 
Jas. Murie furnishes (Z’rans. Zool. Soc. vit.) an important de- 
eription of the form and structure of the Manatee. The memoir 
occupies 75 quarto pages, and is illustrated by 10 large plates, in 
which the external characters, muscular system, alimentary canal, the 
brain, the pulmonary and generative organs, and the great vascular 
plexuses, are figured. 


ARrtTIO-DACTYLA.—J. Alex, Smith collects together (Proc. Scot. 
Soc. Antiq. 1X.) a number of facts concerning the remains of the Elk, 
Cervus Alces, found in Britain, and notes some instances of the dis- 
covery of the remains of Megaceros hibernicus in Scotland.——John 
W. Clark notes in Proc. Zool. Soc. Feb. 20, 1872, a number of 
observations on the visceral anatomy of Hippopotamus. He figures 
the tongue, larynx, trachea and uterus. 


REPORT ON THE PROGRESS OF ANATOMY. 337 


Dryocerata.—O. C. Marsh communicates additional particulars 
(Americ. Journ. Science and Arts, Feb. 1873) on the very remarkable 
horned fossil mammals, the remains of which have recently been 
discovered in the Eocene beds of Wyoming. Three pairs of horn 
cores are found on the cranium, viz. on the nasals, the maxillaries, 
and the great crest formed by the parietals and supra-occipitals. 
Although the vertebree and limb-bones are in many respects like the 
Proboscidea, yet the cranial characters are so distinctive as to render 
it necessary to constitute a new order for these mammals. 


Marsurpratia.— Alex. Macalister describes and figures (Proc. 
Zool. Soc. March 19, 1872) the cranium of the Broad-headed Wombat 
(Phascolomys latifrons). 


Brrvs.—W. K. Parker describes (Monthly Mic. Journ. Jan. and 
Feb. 1873) the development of the SKULL IN THE Tir AND Sparrow- 
HAWK, and in March the development of the SKULL IN THE GENUS 
TuRDUS. A. H. Garrod and F. Darwin give (Proc. Zool. Soc. 
March 5, 1872) some anatomical particulars of an OstricH which 
lately died in the Zoological Gardens. O. C. Marsh records 
(Amer. Journ. Science and Arts, Feb. 1873) some observations on a 
new sub-class of fossil birds (OponTorNITHES). The type species, Zch- 
thyornis dispar, has well-developed teeth in both jaws, which are 
numerous, compressed, pointed and implanted in distinct sockets. In 
the lower jaw about 20 in each ramus, and apparently about the same 
in the upper. The jaws did not seem to have had a horny sheath. 
Vertebree biconcave; bones of extremities conform to ornithic type. 
He believes that he can distinguish more than one genus, and that 
the discovery of these fossils does much to break down the old dis- 
tinctions between birds and reptiles. Jas. Murie communicates 
(Proc. Zool. Soc. May 21, 1872) observations on the OsTEoLoGy oF 
Topus, and on June 18 he describes the CRANIAL APPENDAGES and 
wattles of the Horned Tragopan (Ceriornis satyra); whilst in Ibis, 
~ Oct. 1872, he discusses the Mormors and their affinities. From 
an examination of the anatomy of the Huia Bird (Heteralocha gould‘) 
A. H. Garrod (Proc. Zool. Soc. May 21, 1872) concludes that it is 
truly Passerine, and not related to Upupa, as had previously been 
supposed. In a paper on the mechanism of the Gizzard in Birds 
(Proc. Zool. Soc. April 16, 1872), A. H. Garrod states that the food 
is thrust between the lateral muscles by the contraction of the su- 
perior and inferior gizzard sacs—upon which these lateral muscles 
contract simultaneously ; and their arrangement is such that all the 
force of their contraction is converted into a compressing force at 
right angles to their direction. 


Reptitia.—J. E. Gray enters (Ann. Nat. Hist. March, 1873) 
into the consideration of the original form, development and cohesion 
of the bones of the STERNUM OF CHELONIANS, with notes on the 
Skeleton of Sphargis. Paul Gervais extracts, in Journ. de Zool. 
1873, 1. from his memoir in Nouv. Archives du Muséum, vil. some 
observations on the OsrroLoGy of the SpHArcis Luru,——A detailed 


338 PROFESSOR TURNER. 


description of the MyoLocy of LioLEpris BELLI is given by Alfred 
Sanders in Proc. Zool. Soc. Feb. 6, 1872. L. A. Segoud com- 
pletes (Robin’s Journ. No. 1, 1873) a memoir on ReEpriLes AND 
BatRACHIANS, classed in five types, based on the configuration of fun- 
damental parts of the skeleton. F. Leydig describes (Schultze’s 
Archiv, 1872, 1x. 1) the stRUCTURE AND DEVELOPMENT OF THE TEETH 
OF THE SNAKES indigenous to Germany. 


Fisn.—In our Report, vi. 447, an account is given of Ercolani’s 
observations 0n PERFECT HERMAPHRODITISM IN THE EEL. He has now 
published some additional considerations on the same subject in 
Ann. dell. Soc. dei Natur. Modena, 1872. An abstract of Ercolani’s 
memoir, and of one on the same subject by G. Balsamo-Crivelli 
and L. Maggi, is in Robin's Journ. No. 1, 1873. Joseph 
Hyrtl describes and figures (Denk. der Kaiser. Akad. der Wiss. 
Vienna, 1872) the cranial arteries and the arrangement of the 
vascular arches in the SHARKS. P. Legouis communicates (Ann. 
des Sc. Nat. 1873, xvi. 17) an elaborate memoir on the PANCREAS 
OF OSSEOUS FISH and the TUBES OF WesER. He writes a historical 
introduction in which he gives an interesting description of the obser- 
vations made at different times to show the co-existence both of 
pyloric cceca, and of a pancreas in the osseous fish; he relates also 
Weber's observations on a tube entering the intestine which was not 
a bile-duct. He then proceeds to point out the general presence of 
Weber's tubes, the general presence of a pancreas, the relation of the 
pancreas to Weber’s tubes and to the viscera, more especially the 
liver. He concludes with some observations on the lymphatic organs. 
Th. Gill states his views (Ann. Nat. Hist. March, 1873) on the 
homologies of the Shoulder Girdle of the Dipnoans and other fishes. 
R. Walker describes (7’rans. Geol. Soc. Edin. 1.) a new species 
of Amblypterus, which he names Am. Anconowchmodus, obtained from 
the shale worked for the distillation of paraffine oil at East Pitcor- 
thie, Fife. R. H. Traquair describes (Roy. Geol. Soc. Ireland, 
Dec. 6, 1871) specimens of Phaneropleuron Andersoni and Uronemus 
lobatus. 


INVERTEBRATA.—E, Ray Lankester communicates (Ann. Nat. 
Hist. Feb, 1873) observations on the development of Loligo, Aplysia, 
of various Vudibranchs, of Terebella nebulosa, together with observa- 
tions on points in the anatomy of Appendicularia, Sipunculus, Ster- 
naspis, Glycera, Terebratula, Phyllirhie, Pyrosoma and Dicyema, 
L. Cienkowski makes some observations (Schultze’s Archiv, 1x. 47) 
on Noctiluca miliaris. In an important memoir on the Anatomy 
of Limulus (Ann. des Sc. Nat. 1872) Alph. Milne Edwards describes 
the vascular, nervous and appendicular system of this animal. The 
memoir is illustrated by twelve beautiful plates, several of which are 
coloured. A. §. Packard gives an account (Mem. Boston Soe. 
Nat. Hist. 11.) of the DEVELOPMENT or LimuLus PoLypHeEmus, and to 
Mem. Peabody Acad. Sc. 1871, the same anatomist communicates 
EmBryo.oaicaL Strupies on Diplax Perithemis, Isotoma, and in same 
Memoirs, 1872, embryological studies on Hexaropous Insects. 


REPORT ON THE PROGRESS OF PHYSIOLOGY. By 
WitrrAM Ruruerrorp, M.D., F.R.S.E., Professor of Physiology, 
Kings College, and Fullerian Professor of Physiology, Royal 
Institution, London". 


Nervous Systen. 


Brain.—The Principles of Psychology. By Herbert Spencer. 
Second Edition. London: Williams and Norgate. See an article 
on this by Douglas A. Spalding (Vatwre, Vol. vu. p. 298).  Heredi- 
tary Genius. By Francis Galton. Macmillan and Co. On Dar- 
win’s Philosophy of Language. By Max Miiller (Vature, Vol. vu. 
p- 145). He criticises the evolution theory in so far as it is by some 
regarded as accounting for the origin of languages, and maintains 
that between the language of man and that of the lower animals 
there is xo natural bridge, and that to account for human language 
such as we possess, would require a faculty of which no trace has ever 
been discovered in lower animals. Darwin admits that articulate 
language is peculiar to man, but contends that animals have, in a 
lower stage of development, the identical faculties necessary to the 
invention of articulate expressions. Miiller replies that no develop- 
ment of mental faculties has ever enabled any animal to connect one 
single definite idea with one single definite word. There is an essen- 
tial difference between the expression of emotions and the expression 
of ideas and abstract conceptions. There is no evidence that mere 
conditional signs and sounds can develope into articulate speech. 
Both man and the lower animals possess emotional, but man alone 
possesses rational language. The latter is to be traced back to 
roots. Every root is the sign of a general conception or abstract idea 
of which the lower animal is incapable. Darwin has stated that 
there are languages which have no abstract terms, but Miiller main- 
~ tains that the names of common objects, e.g. father, mother, &e. are 
abstract terms. Rational language is the true barrier between man 
and beast. 


Tar Causation oF SiEep.—By James Cappie, M.D. pp. 76, 
Edinburgh, James King. For an abstract, see London Medical Record, 
1873, No.9. Dr O'Dea (Quarterly Journal of Psychological Medicine, 
No. 11.) regards.dreams as the “ present mental images of past sensa- 
tions revived by the subjective states of the dreamer, or by the objec- 
tive impressions on his senses. The principal factors of dreams are, 
(a) bodily sensations, whether these be subjective or objective, and 


1 Owing to the short notice in which I have been asked to prepare this 
Report I am obliged to give merely the titles of many papers, and to refer 
the reader to abstracts in other periodicals. 

To assist in rendering this report complete, authors are inyited to send 
copies of their papers to 

King’s College, Strand, ; 
London, W.C. 


340 DR RUTHERFORD. 


(2), our previous waking thoughts, dispositions and prevalent states 
of mind.” H. Quincke (Physiology of the Cerebro-spinal Fluid, 
Reichert’s Archives, 1872, p. 158) injected an emulsion of cinnabar 
into the subarachnoid space of the spinal cord in the region of the first 
lumbar vertebra of the dog. After the lapse of a period, varying in 
different instances from one week to three months, he found the 
cinnabar in the subarachnoid tissue and pia of the brain as well as 
the spinal cord. In ten out of twelve cases it was much accumu- 
lated at the base of the brain. It was found around all the cranial 
and spinal nerves, especially where they pierce the dura mater. Ina 
number of cases it extended for some distance along the nerves— 
especially in the case of the optic, where it penetrated as far as the 
eye-ball. It was also found in the cervical lymph glands. It was 
not found in the proper substance of either the brain or spinal 
cord. In a second series of experiments the cinnabar emulsion was 
injected into the special arachnoid cavity of the cranium. In a few 
days it had largely disappeared from this, and was found in the sub- 
arachnoid spaces and pia of the brain exactly as after direct injec- 
tion into these parts. It was also found in the spinal canal. From 
these experiments Quincke concludes, 1, that there is a connection 
between the subarachnoidal spaces of the brain and spinal cord. 2. 
During life there is a current in the subarachnoid fluid from behind 
forwards as well as in an opposite direction (the pigment passed in 
both directions). He thinks that the respiratory motions of the sub- 
arachnoid fluid (Magendie) are the most probable cause of the diffusion 
of the precipitated particles of the cinnabar. 3. The passage of the 
cinnabar from the arachnoid cavity of the brain into the subarach- 
noid spaces of brain and cord shows that these parts communicate. 
4. The apertures of exit for the cerebro-spinal fluid appear to be 
indicated by these experiments. There seem to be channels in con- 
nection with the nerves through which the fluid escapes. This 
appears to be the explanation of the presence of free cinnabar 
particles and also of lymph corpuscles containing cinnabar in these 
situations. The pacchionian bodies appear also to be places of exit. 
These were strongly pigmented. The cinnabar was never found in 
the central canal of the spinal cord, the “perivascular spaces” of 
brain or cord, the lymph-vessels of the olfactory membrane, in 
Tenon’s space or the perichoroidal space, which, according to the 
injection experiments of Key, Retzius and Schwalbe, communicate 
with the arachnoid space. Quincke concludes that normally these 
parts discharge their fluid into the subarachnoid space, and receive 
nothing from this. 

Fournié.—Recherches Expérimentales sux le Fonctionnement du 
Cerveau. 8vo. Paris. Delahaye. 

“On Instinct,” by Mr Douglas Spalding (Maemillan’s Magazine, 
February, 1873). (Abstract in Lancet, 1873, March Ist.) Also a 
paper by Lewes with reference to this in Nature, 1873, April 10. 

“On the Anatomical and Physiological Localisation of Move- 
ments in the Brain. Excellent lectures by Dr Hughlings Jackson.” 
(Lancet, Vol. 1. 1873, pp. 84, 164, 232.) 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 341 


Sprvat Corp.—The present teaching regarding conduction in the 
spinal cord, may be briefly stated to be this. Sensory impressions 
are conducted by the grey matter, especially near its central part. 
Motor impressions are conducted by the anterior and lateral columns 
and to some extent by the anterior horn of grey matter. Section of 
the posterior column produces hyperesthesia on the same side for 
impressions which give rise to pain, and the posterior columns seem 
to be concerned in the co-ordination of movements. These conclu- 
sions are chiefly based upon Brown-Séquard’s experiments. Schiff 
(Centralblatt, 1872, p. 774) states that he formerly ascertained, and 
Longet confirmed his statements—that the posterior columns of the 
cord conduct only tactile impressions, and not those which give rise to 
pain, or those which result from pressure (“nicht aber fiir die Schmerz 
und Druckempfindungen”)—a fact which agrees with neuropathological 
experience of cases [of tabes dorsalis] in which painful sensation remains 
while tactile sensation is lost. Hence, if the spinal cord be divided trans- 
versely, so as to leave only the posterior columns, the tactile sense re- 
mains while the sense of pain is abolished. Recently Schiff has im- 
proved his operative method so that he can perform the experiments 
with very slight loss of blood, and without producing death. In these 
experiments he divided only the posterior columns of the cord, or one 
of these together with a portion of the lateral column, or the grey 
matter, or the anterior column. At first the symptoms were mixed, 
but after a few days all the other functional disturbances disappeared 
with the exception of the loss of the tactile sense, which was permanent. 
The autopsies showed that the partial recovery could not be ascribed 
to any reunion of the divided parts. Schiff maintains that with the 
exception of the posterior columns lesions of almost all other parts of 
the cord may be compensated for by the portions which remain intact. 
The only exception to this “rule” is the case of the posterior 
columns: ‘ every” lesion of these columns produces a permanent 
loss of tactile sense, which is not compensated for by any other part 
of the cord. In tabes dorsalis, where the loss of tactile sense is the 
most common symptom, the autopsy may show a degeneration 
limited to the posterior columns or extending beyond these, and yet 
the symptoms are almost the same as they are when the posterior 
columns only are affected. It would almost appear as if with the 
exception of the posterior columns the other portions of the cord 
have no definite function. The fact that the tactile sense may be 
much impaired when there is an extensive degeneration of the lateral 
column with but slight affection of the posterior column, has led some 
to believe that the posterior column is not the only part which con- 
veys tactile impressions. Schiff considers the true explanation to be 
this, a lesion of the posterior column is not compensated for, but 
there is compensation in the case of lesions occurring elsewhere. The 
application of this rule must however be limited. It holds good for 
the dorsal and for the lower part of the cervical portion of the cord 
(that opposite the three lower cervical vertebree and extending down 
to the eleventh dorsal vertebra); above and below these points the 
rule requires modification. Sanders-Ezn previously found that in 


342 DR RUTHERFORD. 


the lumbar portion and in the lower dorsal part of the cord the tac- 
tile sensory fibres do not enter the posterior columns transversely, but 
that they pass obliquely upwards and join the posterior columns 
from 6—9 cm. (2—31 inches) above the nerve-root. So that the 
tactile nerves of the posterior extremities traverse the cord with- 
out entering the posterior columns until they reach as high as the 
last but one or the last but two of the dorsal vertebrae (dog, rabbit). 
The posterior columns of the lumbar portion of the cord contain, how- 
ever, the tactile nerves of the organs of generation, pelvis, anus and 
tail. The tactile nerves of the feet lie in yet other regions of the 
cord. Schiff argues with Sanders-Ezn, and further shows that injury 
to the lateral columns of the lumbar portion of the cord has, in rela- 
tion to the posterior extremities, the same result as lesion of the 
posterior columns in the dorsal] and lower cervical portion of the cord. 
The paths of tactile impressions change their position in the lower 
part of the cord, but not their character. Ataxia of the lower extre- 
mities may be associated with a degeneration of the lumbar portion of 
the cord which does not affect the posterior columns. In such a case, 
the tactile sense is still present in the anal region. The converse 
holds true when the posterior columns are degenerated in the lumbar 
regions, Still more important and interesting is the modification of the 
rule regarding the posterior columns, which applies to the part of cord 
above the third cervical vertebra. Here, there is in the lateral column 
a tract of white substance which serves for the conduction of the re- 
spiratory motor influences. A lesion of this tract produces an absolute 
and permanent palsy of respiratory motion. In this case, as in that 
of the tactile fibres of the posterior columns, there is no compen- 
sation such as obtains for the other motor fibres of the cord. The 
lateral columns of the cord between the third lowest cervical and the 
lowest but one dorsal vertebra, are not the conductors of impressions 
which give rise to a sense of pain on pressure in the posterior extre- 
mities. These are conveyed by the central and lateral grey matter. 
While the posterior columns conduct the tactile impressions, the 
lateral grey substance of the right side in dogs and also in man is the 
special, and, it may be, the only conductor of painful impressions from 
the /eft posterior extremity, and vice versa. In cats, however, there 
is an exception. Here, the sensory tracts do not decussate as in man 
and in the dog, but the grey substance conducts the painful impres- 
sions produced on the same side of the body. It is worthy of remark, 
that in all these experiments Schiff found the parts which convey 
painful impressions quite insensible to direct stimulation. 


CuorpA Tympanr.--Vulpian (Gazette Médicale, Feb. 15, 1873) 
has discovered that if the chorda tympani be divided in the dog and 
its peripheral cut end stimulated, there is not only dilatation of 
vessels in the submaxillary and sublingual glands, but also in the 
mucous membrane of the lateral half of the tongue on the same side, 
There is no motion of the tongue, nor any increase of secretion from 
its mucous membrane. As is well known, the salivary glands above- 
mentioned secrete when the nerve is thus stimulated. As is now 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 343 


known, this secretion is coincident with, but not dependent upon, the 
vascular dilatation (see Journal of Anatomy and Physiology, No. xt. 
p- 199). Vulpian has also shown that the chorda contains centri- 
petal excito-secretory as well as centrifugal secretory fibres. The 
former convey their impressions to the secretory centre in the brain. 
(For a fuller abstract of these researches, see London Medical Record, 
1350.0. 17 5) 

By the Wallerian method of investigation, Prevost (Comptes 
Rendus, xxv. 1872, p. 1828) has traced the chorda tympani to the 
tongue. In dogs, cats, rats, and guinea pigs, he divided the chorda, 
and from 6—10 days afterwards examined the terminations of the 
lingual nerve. In all cases, he found degenerated fibres; he also 
observed degenerated fibres in the mucous membrane at the point of 
the tongue. He remarks that the microscopical examination of the 
nerve should not be delayed for more than 10—12 days after the 
section of the chorda, because the granular matter resulting from the 
degenerated white substance of Schwann is, especially in young 
animals, speedily absorbed. [Possibly this is the explanation of 
Vulpian’s inability to find these altered fibres after section of the 
chorda, some years ago. | 


Vacus. Legros and Onimus, “ Experimental Researches on the 
Physiology of the Pneumogastric Nerves,” Robin’s Journal de 0 Ana- 
tomie, 1872, p. 411. This paper contains a number of facts which 
had been previously ascertained. 


INFLUENCE OF THE Vacus on Convuxstons.—Brown-Séquard 
(Archives de Physiologie, 1872, No. 2, p. 204) finds that a strong 
stream of CO, through the throat or larynx can cut short an epileptic 
attack in guinea pigs whose sciatic nerve or lateral half of the spinal 
cord has been divided. He confirms Rosenthal and Leube’s experi- 
ments on the arrest of strychnia convulsions by an apnesic condition 
of the animal [that is, by a hyperoxygenated condition of the blood], 
~ but suggests that the result is not due to apnoea but to mechanical 
irritation of the endings of the vagus in the respiratory mucous 
membrane, by the rapid inflation of the lungs adopted in these 
experiments ; section of the vagi prevents the arrest of the convulsions 
in such a case. A stream of CO, directed against the laryngeal 
mucous membrane, arrests the respiratory motions, as well as strych- 
nia convulsions; moreover, in the case of birds after ligation of the 
large cervical vessels, the convulsions were still arrested when the 
stream of CO, was directed against the mucous membrane of the 
bronchi and that of the lower larynx. He therefore considers that 
the CO, is a powerful irritant of the terminations of the vagus, and 
that it reflexly arrests convulsions due to epilepsy, strychnia, or 
anemia, 


TropHic Nerves.—On this subject see an article in British Medical 
Journal, 1872, Aug. 31, giving a résumé of Fischer, Schiefferdecker, 
Joseph and Vulpian’s researches regarding the influence of nerves on 
nutrition. (Abstracts of these papers will however be found in 


344 DR RUTHERFORD. 


Journal of Anatomy and Physiology, Vols. vt. and vu.) See also able 
articles on this question by Mr Henry Power (Zhe Practitioner, 
Feb. and March, 1873). 


CHANGES IN THE NERVES AFTER SecTion.—See Dr Pye-Smith’s 
abstract (London Medical Record, Vol. 1. p. 198) of a remarkable 
communication to the Académie des Sciences by Ranvier, in which he 
states that the so-called degenerative changes in the distal portion of 
a divided nerve “are, as far as the cellular elements are concerned, 
rather those of hyperplasia: the removal of nervous influence appa- 
rently allowing more unrestrained activity.” [It will be necessary to 
subject this point to careful scrutiny ere a conclusion so novel and so 
opposed to our present views can be accepted. | 


GENERAL PuysioLogy oF Nerve.—Setschenow “on the Beha- 
viour of Nerves during Rapid Irritation” (P#liiger’s Arch. 1872, 
p- 114, also Bernstein (/bzd, p. 318) abstract in Centralblatt), W. 
Filehne, “The Law of Contraction in Dying Nerves” (Centralblatt, 
1872, p. 889). 


The Senses. 


Skin. Sensory Nerves ror Tactin—E AND ParnruL InprEs- 
stions.—The idea that the cutaneous sensory nerve apparatus con- 
cerned in the reception and transmission of tactile impressions differs 
from that which receives the impressions that give rise to pain, 
apparently receives support from observations on ‘cold anesthesia,” 
by Horvath (Centralblatt, 1873, p. 210). He found that after 
immersing the finger for some time in alcohol at the temperature of 
—5°C. he could readily perceive impressions produced by gentle 
contact of extraneous bodies [tactile impressions], while pricks, which 
on other fingers produced pain only, gave rise in the chilled finger to 
a sensation of touch [a tactile sensation]. 


Eye.—Holmgren, “ On Forster’s Perimeter and the Topography of 
the Sense of Colour” (Centralblatt, 1872, p. 823). J. Friinkel, “The 
Apparatus for Accommodation in the Human Eye” (Centralblatt, 1872, 
p- 858). Dr R. J. Lee, “ Further Remarks on the Sense of Sight in 
Birds” (Proc. Roy. Soc. 1873, January 9). Leber, “ Condition of the 
Circulation in the Optic Nerve and in the Retina” (Graefe’s Archiv 
Siir Ophthalmologie, xvit. No. 2). Mandelstamm, “Association of the 
two Retins” (/bid.), Samelssohn, ‘“ Innervation of the Ocular Move- 
ments” (/bid.). F.C. Donders, “On Congenital and acquired Asso- 
ciation” (Lbid.). Dobrowlsky, ‘A number of short Papers on the 
Perception of Colour” (/bid.). Exner, ‘On the Physiological Action 
of Iridectomy” (Abstract in Centralblatt, 1873, p. 17). 


Ear.—Beettcher, ‘“ Critical Annotations and New Contributions 
to the Literature of the Labyrinth of the Ear” (Monograph). He 
treats of the structure of the Lamina Spiralis and details experiments 
which refute the idea entertained by Flourens and Goltz, that the 
semicircular canals are concerned in maintaining the balance of the 


Jt 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 345 


body. (See an abstract by Mr Ernest Hart in the London Medical 
Record, 1873, p. 110.) 


Circulatory System. 


Bioop CoacuLation.— Alexander Schmidt (‘“‘ New Researches on 
Coagulation of Fibrin,” Pjliiger’s Arch. 1872, p. 413. Abstract in 
Centralblatt, 1873, p. 22) modifies his well-known account of the 
coagulation of the blood. Fibrinogen and fibrinoplastic substance 
constitute the material from which fibrin is produced, The quantity 
of fibrin increases with the quantity of either of these constituents— 
it matters not which—within certain limits. But for the production 
of fibrin from these substances, a third body, a ferment, is necessary 
in order to bring about the union of the fibrin-generators. He desig- 
nates this “ Fibrin ferment.” 1. A small quantity of this produces 
in the same fluid as complete a fibrin formation as a large quantity, 
only not so rapidly. 2. The activity of the fibrin ferment, indicated 
by the rapidity of coagulation, increases with the proneness to coagu- 
lation, and attains its maximum at the temperature of the body. It is 
destroyed by the temperature of boiling water, on the other hand it is 
rendered inactive by a freezing temperature. 3. If one filter off the 
serum in which a clot has been formed by the action of the fibrin fer- 
ment, the filtrate can (although not so energetically) anew call forth 
coagulation in a fluid containing fibrinogen and fibrinoplastin. All 
the animal fluids which coagulate, contain fibrinogen and fibrino- 
plastin, but no ferment. This first appears after removal of the 
fluid from the body, rapidly in the case of blood, slowly in that of the 
transudations. [If the coagulation be due to this ferment, then, under 
certain circumstances it must be admitted that it can arise within the 
body.] The ferment may be obtained by precipitating blood serum with 
from 15—20 times its bulk of strong alcohol. Let stand for fourteen 
days, filter, dry the precipitate over sulphuric acid, powder it and ex- 
tract it with cold water. For the activity of the ferment it matters 


-not whether serum or blood be taken. No ferment can be obtained 


from the blood allowed to flow from a vein into alcohol, because the 
ferment is not preformed in the blood. The fermentative activity is 
the greater the longer the time between the removal of the blood 
from the body and its precipitation by alcohol. The accumulation of 
the ferment reaches its maximum however with the completion of the 
coagulation of the blood. After this period there is no further 
formation of ferment. Lowering the temperature to 0°C. retards 
the formation of ferment, but does not entirely prevent it. Schmidt 
is convinced that neither the coloured nor the colourless corpuscles 
take part in the origin of the fibrin ferment. If the solution of the 
fibrin ferment and that of the fibrin generators be treated for some 
time with CO or H, no coagulation follows their admixture. If 
however the fluids be removed from the action of these gases and 
exposed to the air, coagulation sets in. The presence of O therefore 
appears to be necessary for the coagulation. Jf to a fluid containing 
fibrinogen and ferment fibrinoplastic substance be added, it is found 
that a certain quantity is necessary to use up all the fibrinogen in 


WOL, VIL. 23 


346 DR RUTHERFORD. 


the fluid, in order to produce fibrin. When all the fibrinogen is 
removed, the further addition of fibrinoplastic substance to the fluid 
causes no further separation of fibrin, The amount of fibrin formed 
does not increase with the quantity of the fibrin ferment, the rate 
of the formation is alone increased by this. The blood pigment 
or the coloured corpuscles accelerate the appearance of coagulation. 
(Schmidt withdraws his former statement that the blood corpuscles 
are rich in fibrinoplastic substance.) In doing so it does not appear 
to undergo any change, for the same quantity can again and again 
induce coagulation. The blood-pigment shares this peculiar power 
with carbon, platinum, asbestos, animal ferments, and all bodies 
which can destroy hydric peroxide and use its oxygen. The fibrin 
ferment differs from other ferments in the fact that it is unable to 
destroy hydric peroxide. Schmidt is now inclined to regard the 
influence of these bodies on coagulation as an action due to mere 
contact, and not, as he formerly supposed, to the influence of oxygen 
condensed on their surface. 


H#moctiosin.—See Proc. Roy. Soc. 1872, Dec. 12, for a valuable 
paper by Mr E. Ray Lankester on Hemoglobin. In addition to 
numerous original observations, he gives a valuable summary of the 
facts which have been ascertained regarding the distribution of 
Hemoglobin in various animals. Miiller, “Action of Quinine in 
Hemoglobin.” Inaug. Dissert. Bonn, 1872. (Abstract in Central- 
blatt, 1872, No. 40.) 


Carponic OxripE H#mocLopin.—Zuntz (Pfliiger’s Archiv, Vv. p. 
584) finds that carbonic oxide Hemoglobin is not so stable a com- 
pound as has been imagined. The CO is removed by placing the 
HbCO in a vacuum, and the remaining Hb shows the spectrum of 
ordinary reduced Hb. He infers from this fact, that artificial respira- 
tion should be energetically employed in CO poisoning. See also an 
abstract of Podolinski’s researches on this subject (London Jed. 
Record, 1873, p. 70). 


Bioop Corpuscies.—Abstract of a paper by M. Malassez (in 
London Med. Record, 1873, No. 1), “‘On the number of the blood- 
corpuscles in mammals, birds and fishes.” Geltowski, “On the 
action of Quinine on the colowless blood corpuscles” (Practitioner, 
1872, p. 321). 

Tron 1x THE BLoop AND Foop.—Boussingault. (For abstract, see 
Journal of Chem. Soc. Sept. 1872.) 


InorGAnic Constituents oF Broop,—Janisch. (For abstract, 
see [bid.) 


New Test ror Brioop.—Sonnenschein, ‘‘ Action of a New Re- 
agent on Blood and its employment in Forensic Medicine” (Central- 
blatt, 1872, No. 54). 


Bioop Gasrs.—See abstract of a paper by M. Lepine (in London 
Med. Rec. 1873, No. 13). Mathieu and Urbain (Brown-Séquard and 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 347 


Vulpianws Archives, 1872, p. 190) investigate the amount of gas in 
the blood of different arteries. See Mr Power’s abstract in Brit. and 
For, Med.-Chi. Rev. Oct. 1872, p. 524.——Wolffberg, “Tension of 
the Blood gases in the Pulmonary Capillaries,” Pjliiger’s Archives, Iv. 
p. 465 (abstract in Centralblatt, 1872, No. 1).——Wolffberg, “ On 
Pulmonary Respiration” (Pjliiger’s Archives, 1872, p. 23).——Strass- 
burg, “On the Topography of the gaseous tensions in the Animal 
Organism” (Ibid. p. 65). Pfliiger, ‘On the Diffusion of Oxygen, 
the Seat and the Laws of the Oxidation Processes in the Animal 
Organism” (Jbid. p. 43). Abstract of the last three papers (in 
Centralblatt, 1872, No. 40, and in Brit. and For. Medico-Chi. Rev. 
April, 1873). 


EsTIMATION OF THE ABSOLUTE QUANTITY OF BLoop.—Steinberg 
(Pliiger’s Archives, 1873, p. 101), “Minute Moving Particles as 
Constant Constituents of Normal Human Blood” Nedsvetzki (Cen- 
tralblatt, 1873, No. 10). 


INNERVATION OF THE Hrart.—For abstract of recent researches 
by Schiff, see Centralblatt, 1873, Nos. 1, 2, 3, and British Med. 
Journal, 1873, March 8. He denies the existence of accelerating 
nerves for the heart in the cervical sympathetic and cervical portion 
of the spinal cord, and maintains that the only accelerating nerves 
are derived from the spinal accessory. They joi the vagus, but 
afterwards leave this nerve at the ganglion of the trunk passing 
in the pharyngeal or superior laryngeal nerves to the recurrent 
laryngeal through which they pass down the neck to the heart. 
[Schiff no doubt expects to hear something about this from those 
who have furnished him with the ideas which have led him to the 
conclusion regarding which we are for the present silent. | 


“ReFLEX RELATIONS BETWEEN THE STOMACH AND THE NERVE- 
CENTRES FOR THE ORGANS OF CrrcuLATIoN.”—-Centralblatt, 1873, No.13. 


INNERVATION OF THE VESSELS OF THE RaAspit’s-EAR.—Moreau. 
Brown-Séquard and Vulpian’s Archives, 1v. p. 667.—Abstract in 
Centralblatt, 1873, No. 15. 


Respiratory System. 


INFLUENCE OF RESPIRATION ON BLoop-PressurE.—The respiratory 
curves in the blood-pressure have been generally ascribed to the 
mechanical influence of the thoracic movements. Schiff (Central- 
blatt, 1872, No. 48) admits that with exaggerated respiratory 
motions, such as those seen after division of the vagi, a mechanical 
effect upon the blood-pressure, e.g. in the carotid is evident, but 
maintains that in normal respiration the respiratory oscillations of the 
blood-pressure are due to rhythmical excitation of the vaso-motor 
centre in the medulla, causing periodic risés in the blood-pressure by 
inducing contraction of blood-vessels. The excitement of the vaso- 
motor centre is according to him due to the same cause as that which 
exc tes the respiratory centre, that is, a lessening of the amount of 


23—2 


348 DR RUTHERFORD, 


oxygen or an increase in the amount of carbonic acid in the blood. 
The cause of the ordinary respiratory curves is therefore, accord- 
ing to this theory, not mechanical but chemical, If an animal be 
caused to breathe pure oxygen the respiratory oscillations become 
less frequent, so that there may be only one respiratory oscillation in 
the pressure for three or four respiratory movements. If the blood be 
saturated with oxygen the respiratory curves in the pressure entirely 
disappear, although the respiratory movements of the chest be con- 
stantly maintained by artificial means. There are some facts which 
can only be explained with difficulty, or not at all, on the mechanical 
theory, e.g. at times the respiratory pressure curves are extremely 
weak, and may even be entirely wanting, and under these conditions 
a single deep and powerful respiration produces no variation in the 
blood-pressure. Schiff states that the respiratory curves are wanting, 
(1) if the interval between two respirations is not great enough to 
occasion an accumulation of carbonic acid with blood; and (2) if the 
sensibility of the vaso-motor centre be diminished, the respiratory 
curves disappear. The first explanation serves for those cases where 
the respiration is very rapid and the respiratory blood-pressure curves 
are wanting. The second applies to the case of curarised animals 
where the respiratory-pressure curves are much diminished in number, 
and also to the case of animals in which the besoin de respirer is 
diminished by causing them to breathe for some time an atmosphere 
rich in CO, or poor in O. In such a case the respiratory curves are 
wanting. 


INFLUENCE OF ARTIFICIAL RESPIRATION ON THE CIRCULATION.— 
From the fact that during ordinary inspiration the intra-thoracic pres- 
sure is diminished, whereas, during artificial inflation it is increased, it 
has been supposed that the effect on the blood-pressure is such as pos- 
sibly to exert an important influence on the circulation during the per- 
formance of experiments in which artificial respiration is adopted. 
Schiff (ibid.) states that the artificial respiration produces no altera- 
tion in the mean blood-pressure, although it may give rise to oscilla- 
tions of the pressure if the inflation of the chest be excessive. 


INFLUENCE OF ARTIFICIAL RESPIRATION IN CASES OF CONCUSSION 
AnD Compression.—Schiff (ibid.). See abstract in London Medical 
tecord, 1873, No. 1. 


Resprratory Movements.—‘“‘On the Mechanical Conditions of the 
Respiratory Movements in Man,” by Arthur Ransome, M.D. (Proe. 
Roy. Soc. 1872, Nov. 21. Abstract in London Medical Record, 
1873, No. 1.) See Lectures on Human Myology, by Professor 
Humphry (Brit. Med. Journ. No. 619) for opinions regarding the 
action of the intercostal muscles. A Pneumograph invented by 


Prof. Fick. (Centralblatt, 1873, No. 13.) 


Absorption. 


IyFiuEeNce or Nerves on Axpsorption.—PBernstein ‘fon Goltz’s 
Absorption Experiments” (Berliner Klin. Wochenschrift, 1872, No. 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 349 


28). “Onthe Relations of the central parts of the Nervous System to 
Absorption” (Virchow’s Archives, 1872, ivi. p. 248). Bernstein has 
repeated the experiments on absorption performed by Goltz (Journ. of 
Anatomy and Physiology, Vol. vi. p. 480), with the slight variation 
that he removed the heart altogether and tied a cannula in the inferior 
vena cava. He found the same results as Bernstein: to wit, that in 
two curarised frogs suspended by the nose, one having the central 
nervous system intact, and the other having it destroyed, both 
having neutral salt solution poured through a funnel into the dorsal 
lymph sac, and both having the influence of the heart upon the circu- 
lation suspended (in the case of Bernstein’s experiments by removal 
of the heart). Absorption from the lymph sac readily takes place in 
the case of the frog with the uninjured central nervous system but 
not in the other. The evidence of the absorption is furnished by the 
dropping of bloody fluid from the cannula in the vena cava in the 
one case and not in the other. Goltz explained this by supposing 
that owing to vaso-motor palsy in the one and not in the other, the 
blood-vessels are so dilated that nothing flows through them, and also 
that nerves proceeding from the central nervous system to the lymph 
and blood-vessels having the power of causing them to absorb—are 
paralysed in the one and not in the other case. The fact that electri- 
cal stimulation of the frog with the spinal cord and brain accelerated 
the absorption was ascribed by Goltz to stimulation of the nerves con- 
cerned in absorption, just as stimulation of secretory nerves gives rise 
to secretion. Probably very few persons have cared to adopt an 
explanation so startling, and having such important bearings, before 
the advance of less equivocal evidence. Bernstein considers that 
the conditions of absorption are similar in both cases, but that in the 
animal with the cerebro-spinal system intact, the blood-vessels contract 
and so keep up the motion of the blood although slowly. In the frog 
without the brain and spinal cord the vessels are palsied, hence the 
blood stagnates and absorption is not facilitated. This explanation 
is supported by the fact, that if the abdominal blood-vessels be 
opened in both cases, so that the fluid has to pass from the dorsal 
lymph sae through a short vascular path, the fluid is absorbed as 
quickly in the one case as it is in the other. 

Heubel (‘‘On the Relations of the Central parts of the Nervous 
System to Absorption,” Virchow’s Archives, 1872, v1. p. 248) has 
been performing experiments on this subject wnder the direction of 
Goltz, An account of these will be found in the London Medical 
Record, 1873, No. 2. The important feature of his paper is this, that 
he endeavours to explain such facts as the above by the alteration in 
the circulation which follows the destruction of the vaso-motor centres 
in the medulla oblongata and spinal cord, and not by supposing, as 
Goltz did, that there is a special system of nerves for absorption para- 
lysed in the one case but not in the other. [We may fairly infer from 
this therefore, that Goltz has retired from the untenable position—in 
which he asserted that his experiments furnish evidence of the ex- 
istence of such a system of nerves. | 


350 : DR RUTHERFORD. 


Alimentation. 


AumeEntT.—Voit “On the Nutritive value of Gelatine” (Zeitsch. 
fiir Biologie, Vol. vii. Abstract in London Med. Record, 1873, 
No. 3). Article on Food (British Med. Journal, 1872, October 5, 
12, and 26). Carbo-hydrates, and the mode in which they are 
digested and absorbed; Briicke (Wiener Sitz. Berich. Math. Nat. Cl. 
Vol. rv. Part m1. Abstract in London Med. Record, 1873, No. 3). 
——Pettenkofer and Voit “On the Regressive Metamorphosis in 
Animal Bodies during a Flesh Diet” (Zeitsch. fiir Biologie, 1872, 
vu. 3. Abstract in Centralblatt, 1872, No. 46). Schenk, “ Beha- 
viour of Chlorine in the Organism” (Centralblatt, 1872, No. 43). 
F. Hofmann, “ Passage of Fat from the Aliment into the Cells of 
Animal Bodies” (Abstract in Centralblatt, 1872, No. 59)——Falck, 
“On Sodium Chloride” (Abstract in London Med. Record, 1873, 
No. 2). 


INNERVATION OF THE GZSOPHAGUS AND STOMACH OF THE FROG.— 
Goltz (“Movements of the Cisophagus and Stomach in Frogs” 
Phliiger’s Archives, 1872, Vol. vi. p. 616), impressed by the difficulties 
which beset the study of the gastric movements of such an animal as 
a rabbit, in which the stomach is always full, operated on frogs in 
the hope that they might furnish results which might serve as a basis 
for arriving at definite knowledge regarding this matter. He took 
two frogs which had been starved for some days, poisoned them with 
curara, removed the heart (so that in both cases irregularities of the 
circulation might not be encountered), the left lung, the left arm, and 
laid open the abdomen so that the cesophagus and stomach could be 
easily seen. In one case the brain and spinal cord were destroyed, in 
the other these were left intact. Both frogs were suspended by the 
nose. <A solution of NaCl, } per cent. was poured into the mouth in 
both cases. In the frog possessed of the brain and spinal cord it 
rapidly found its way into the stomach ; but in the other frog it was 
quite otherwise. The pharynx was contracted, and so the cesophagus 
and stomach remained quite empty. Nevertheless in this case the 
gullet and stomach both exhibited somewhat irregular peristalsis, 
lasting for hours. A similar phenomenon is observed if the activity 
of the central nervous system be suppressed by the inhalation of 
chloroform, or by a large dose of curara, or if the vagi be divided. 
He infers from these facts that the vagi contain inhibitory nerves for 
the esophagus and stomach, and that the centre for these fibres is in 
the central nervous system. The appearance of the peristalsis after 
division of the vagi, &c. is, according to Goltz, due to palsy of the 
inhibitory nerves. He found that cesophageal and gastric movements 
could be induced reflexly in frogs by faradisation of sciatic nerve, or the 
application of sulphuric acid to the skin. When the application of the 
irritant is continuous, the movements may last for hours, as after 
division of the vagi or removal of brain and spinal cord. Goltz 
therefore thinks that here we have not a reflex action in the ordinary 
sense, but motion of the parts resulting from a cessation of the ac- 
tion of inhibitory nerves, the irritant producing a temporary palsy 


REPORT ON THE PROGRESS OF PHYSIOLOGY. Bol 


of the central ends of the nerves. [But when we remember that 
section of the vagi produces in higher animals at any rate, palsy 
of the gullet and partial palsy of the stomach, and that stimula- 
tion of the dower end causes movement (Goltz even admits that it 
intensifies the irregular peristalsis seen after section of the vagi), it 
must be maintained that Goltz has not made out his case. There 
may be in the vagus of the frog imhibitory as well as motor fibres, 
but why the existence of the latter should be ignored we cannot 
comprehend, To our thinking the point demands further investi- 
gation. | 


Gastric Dicestion.—Von Wittich, “ On the question regarding 
the Peptic Action of the Pyloric Glands” (Pjliiger’s Archives, Vol. vit. 
p- 18. See abstract in London Medical Record, 1873, No. 10). He 
considers the pyloric glands to have no peptic action. This conclusion 
supports the statements of Fick, and Friedinger, &c., and opposes those 
of Ebstein and Griitzner. Wittich’s observations were made on the 
stomach of the pig and rabbit.——Jukes, ‘“ Structure of the Peptic 
Glands” (Centralblatt, 1872, No. 47). Schiff’s statement that the 
energy of gastric digestion is largely dependent on the physiological 
condition of the animal at the time of observation, e.g. whether there 
be hunger, digestion, or exhaustion of the gastric wall owing to 
digestion having been recently going on—although questioned by 
some authorities—has recently (Centralblatt, 1872, No. 50) received 
additional support from experiments performed by him. Through 
fistulze in the stomach of living dogs he introduced known quantities 
of albumen in muslin bags. He found that when digestion had 
recently taken place, a much smaller quantity of albumen was 
digested than was the case when the albumen was introduced into the 
stomach during normal digestion. Seeing that the proportion of acid 
and water in the gastric juice, and the temperature of the stomach 
were the same in both cases, the difference in digestive energy could 
‘only be dependent upon the smaller quantity or the less perfect con- 
dition of the pepsin. Proceeding from the idea that when the 
stomach is removed from the body, its digestive power must largely 
depend upon its condition at the time of death, that is, whether or 
not digestion had been going on, &c., he—as formerly cut the 
gastric mucous membrane into small pieces, and made an infusion of 
1 with 100—500 Cm of acidulated water. Into a measured quantity 
of this infusion a weighed quantity of albumen was placed, and the 
whole kept at the temperature of the body. After a time the amount 
of undigested albumen was estimated so that an indication of the 
digestive power was obtained—suflicient for purposes of comparison. 
Generally, the amount of albumen digested by the entire stomach in 
such a case was estimated at 70—100 grammes, in rare cases 150— 
180 grammes. He finds that the digestive energy depends not only 
on the amount of pepsin present, but also on other conditions. In 
order to obtain the entire digestive power of the pepsin, one must 
carefully ascertain the most suitable quantity of acid and water. The 
quantity of acid varies in different animals and the amount of acid to 


aa DR RUTHERFORD. 


be added to the digestive fluid must increase with the quantity of 
water [that is to say there must be a certain percentage of acid in the 
fluid]. In experiments with the gastric mucous membrane of a cat, 
the digestive power increased with the addition of water to the 
enormous extent of 20—30 litres to the infusion of the single 
stomach. In this much diluted fluid 2000 grammes of albumin were 
digested (a single stomach being employed), while in an infusion 
prepared in the ordinary way not more than 70 grammes are digested. 
When so much water is employed it is necessary to let the gastric 
membrane infuse in it for 10—15 days. The amount of acid employed 
is stated to be the same as that which Schiff in “ his former researches 
found to be the best.” The investigation is not yet completed, but 
he states that for the stomach of a dog 200 litres of water are required 
to make a suitable infusion, and this can digest 75 kilogrammes of 
albumen (a kilo is 2-2 Ibs.). [We presume that coagulated albumen 
was employed. We have not been able to see Mosso’s reports of 
Schiff’s researches published in La Nazione, 1872, Nos. 102, 109, 110, 
116, 8. A. 12mo. p. 59.]|——Manassein, “ Chemical contributions to 
our knowledge of Fever” (Virchow’s Archives, ty. 1872, p. 413. 
Abstract in Centralblatt, 1872, No. 44) investigates digestion in (1) 
healthy dogs and cats, (2) dogs and cats rendered anzemic by deple- 
tion, (3) dogs and cats thrown into a febrile condition by the subcu- 
taneous injection of putrid matter.——Mohlenfeld, ‘On the Peptones 
produced from Fibrin” (Pfliiger’s Archives, 1872, p. 381. Abstract 
in Centralblatt, 1872, No. 43).——Von Wittich, ‘Concerning Pepsin 
and its Action on Blood-fibrin” (Pfliiger’s Archiv, 1872, p. 435. 
Abstract in Centralblatt, 1872, No. 45). 


Liver.—Von Wittich, “On the Physiology of Human Bile” 
(Pliger’s Archives, 1872, pp. 181), finds that fresh human bile ob- 
tained from a fistula contains, like the bile of other animals, a ferment 
capable of converting starch into sugar. The ferment may be isolated 
by precipitating the bile with alcohol and extracting the dried preci- 
pitate with glycerine. He also made the observation tliat if bile be 
filtered through animal charcoal, the bile acids are completely re- 
tained by the charcoal, and could not even be extracted from the 
charcoal by means of alcohol. In a more recent paper (/bid. 1873, 
pp. 28) he shows that the ferment is really formed in the liver, and 
that it may be extracted from it after it has been washed for hours in 
a stream of water (see a fuller abstract of this last in London Medical 
Record, 1873).——“ Oxidation Products of the Bile Pigments pro- 
duced in Gmelins’s Reaction,” Stokvis (Centralblatt, 1872, No. 50), 
[a good paper]. “Determination of the Sulphur of the Bile,” Kiiiz 
(Archiv fiir Anat. und Phys. 1872, pp. 98. Abstract in Centralblatt, 
1872, No. 56).——“Cholie Acid,” Baumstark (Berlin. Klinik. Wo- 
chensch. 1873, No. 4: Abstract in Centralblatt, 1873, No. 16).—— 
“Biliary Caleuli.” Ritter (Journ. d. P Anatomie, 1872, No. 1. See 
Abstract in Brit. and For. Med.-Chi. Review, April, 1873).—— 
“ Action of Mercury on the Liver.” Leading Article (Lrit. Med. 
Journ, 1873, January 4th). 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 350 


PancrEas.—|It is well known that, in the rabbit the pancreatic 
duct opens into the intestine twelve inches, or more, below the bile 
duct, and that Bernard observed the striking fact, that after the 
ingestion of food containing fatty substances, the absorption of the 
fat takes place after the food has passed the orifice of the pancreatic 
duct and not before this, although it has been subjected to the action 
of the bile. In other animals where the pancreatic duct opens with 
or near the bile duct, a short way below the pylorus, the absorption 
of fat begins much higher up than in the rabbit. Unabsorbed fat 
has been, in several cases of pancreatic disease observed in the feeces 
in the human subject. These and many other well-known facts have 
led to the prevailing belief that the pancreatic juice has to do with 
the digestion of fat, and indeed that if the flow of the juice into the 
intestine be prevented, the digestion of fat is incomplete. Bernard 
failed to preserve dogs alive after the extirpation of the pancreas ; so 
to prevent the secretion from entering the intestine, he injected 
melted tallow into the pancreatic ducts. In two cases the animals 
lived, and although they had voracious appetites, they became 
emaciated. The feces contained undigested fat.|——Schiff (Cen- 
tralblatt, 1872, No. 50. Abstract by Boll of Schiff’s researches 
reported in Za Nazione, by Dr A. Mosso) considers the non-absorption 
of fat in these dogs to have been probably due much more to the 
interference with the entrance of bile than of the pancreatic juice into 
the intestine. He thinks that the inflammation set up in the pan- 
creatic ducts by the decomposing fatty matter may have led to partial 
or complete occlusion of the bile duct.. His reasons for thus think- 
ing are derived from the following. He injected 16—27 Cm. of 
melted paraffin into the chief excretory duct (Hauptausfiihrungsgang) 
of the pancreas of full-grown dogs. (He selected paraffin because 
it does not like tallow decompose and so tend to produce inflamma- 
tion.) ‘In many cases the pancreas became swollen and tense 
throughout its whole extent. The autopsies performed after some 
-time, showed the epithelium of the acini to be in a state of fatty 
degeneration or quite atrophied and gone, while the remainder of the 
gland preserved its form.” Jn the most favourable cases the animals 
had good appetites, their weight increased during high feeding, and 
the feces had a normal appearance. The digestion of fat was 
scarcely affected in any way. A normal dog can digest daily 12 
grammes of fat for every kilogramme of body weight. Schiff found 
that on three successive days a dog on which he had operated took a 
larger quantity of fat than this, and in the excrements were found 
only ‘quite faint traces of fat.” Then he has extirpated the pan- 
creas in birds, and he finds that digestion appears to be scarcely at all 
disturbed. Colin and Bérard have done the same with like results. 
Although they always found that adult mammals die from the effects 
of such an operation, yet in newly-born mammals the gland may be 
extirpated, and notwithstanding this, the animals may live and grow. 
Schiff concludes from all this, that even in adult animals after elimi- 
nation of the action of the pancreatic juice upon the food the other 
digestive fluids can carry on digestion perfectly. [It occurs to us to 


354 DR RUTHERFORD. 


say, that although Schiff’s statements regarding this matter obviously 
demand serious attention, nevertheless it is a pity (we judge from 
Boll’s abstract) that in the dog experiments he should have blocked 
up only one of the pancreatic ducts with the paraffin. Why should 
we suppose that the lesser pancreatic duct did not continue to pour 
pancreatic fluid into the intestine, and why should we believe (in 
the absence of direct statement) that the portion of the gland in 
connection with this smaller duct was degenerated. On the 
contrary, its action may have become greater, and so may have 
secreted juice enough to digest the fat. If Schiff infers from these 
experiments that in Bernard’s dogs the failure in the digestion of the 
fat was owing to the interference with the flow of bile into the intes- 
tine, it seems to us a very gratuitous assumption. Why may it 
not have been owing to inflammatory occlusion of the smaller pan- 
ereatic duct? We have not been able to procure La Nazione.|—— 
Brunner’s Glands. Krolow (Inaug. Dissertation. Berlin, 1872. 
Abstract in London Medical Record, 1873, No. 10). 


Lymph. 


H. Nasse (‘Researches on the Influences which govern the 
Formation of Lymph,” Marburg, 1871. Abstract in Centralblatt, 
1873, No. 10), collected and measured the lymph obtained from the 
cervical lymphatics under various conditions. He found that ligature of 
the carotids diminished the lymph-stream to the extent of 24 to 40 
per cent. After removal of the ligatures from the arteries the original 
quantities were seldom reached. With the diminution of the lymph 
there was a relative increase in the amount of its watery constitu- 
ent. Compression of the external jugular veins increased the 
lymph-stream to the extent of 10 to 31 per cent. Experiments on 
the effect of venous depletion yielded the unexpected result that after 
depletion, either to a large or to a small extent, the lymph stream 
increased to from 10 to 31 per cent. [one would have anticipated the 
contrary]. A diminution first set in, in such cases when the coagula- 
bility of the lymph increased. The amount of water in the lymph 
after venous depletion, remained, contrary to expectation, in most 
cases unchanged; on the other hand, the amount of fibrin was dimin- 
ished, and the amount of Na Cl was but slightly lessened. Researches 
on the influence of section and stimulation of the cervical sympathetic 
were not carried out to the entire satisfaction of the investigator, 
because of the difficulty of separating it from the vagus in the dog. 
In most of the experiments stimulation of the cervical sympathetic 
produced a great diminution of the lymph stream, in others the lymph 
stream was accelerated during the period of stimulation. Still less 
decided were the results of section of the sympathetic. Stimulation 
of the undivided vagus always increased the lymph stream, Stimu- 
lation of the central end of the divided nerve was followed by a 
considerable increase, while stimulation of the peripheral end of the 
divided nerve caused slight diminution. Stimulation of sensory 
nerves (which?) considerably increased the lymph stream. This he 


or 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 35 


attributes to reflex contraction of vessels, The injection of 1 per 
cent. solution of Na Cl into the blood-vessels [how much ?] enormously 
increased the lymph stream and diminished the coagulability of 
the lymph. Injection of water had—on the other hand—compa- 
ratively little effect. Carbonates of the alkalies increased the lymph 
stream, but little alkali, however, passed from the blood into the 
lymph. He also made researches with defibrinated blood, and found 
that this had no influence on the amount of fibrin in the lymph, but 
increased the solids of the lymph generally. He concludes from his 
researches that the lymph is a simple transudation. [On this explana- 
tion how is the effect of depletion to be accounted for ?] 


Milk. 


Soxlet, “Contributions to the Physiological Chemistry of Milk” 
(Centralblatt, 1872, No. 53. Abstract in Journ. of Chemical Soe. 
Feb. 1873). Boussingault. (See Jowrn. ef Chem, Soc. Dec. 1872.) 

Schwalbe (Schultze’s Archives, vul. p. 269) says in opposition to 
Kehrer that milk globules have a membrane. If the oil be removed 
and the vesicles then be subjected to the action of osmic acid, the 
membrane exhibits a double contour. [It is surprising how anyone 
can doubt the existence of a membrane. The manner in which the 
globules run together and form masses of irregular shape after they 
have soaked for some time in acetic acid—under the microscope— 
appears to be only explicable on the view that there is a membrane 
which acetic acid is capable of dissolving. | 


Urine. 


Fr. Hofmann, ‘On the Passage of Free Acids through the Alka- 
line Blood into the Urine” (Zeitsch. fiir Biologie, vu. 3. Abstract in 
Centralblatt, 1872, No. 38). See also on the same subject Gaethgens 
(Centralblati, 1872, No. 53). Soborow, “On the Excretion of Lime in 
the Urine” (Centralblatt, No. 39). Falck, on the effect upon the 
urine of the injection of water into the blood (‘Contribution to the 
Physiology of Water;” abstract in Centralblatt, 1873, No. 12). 
Rabuteau, “On the Physiological effects and the Elimination of Urea 
introduced into the organism” (L’ Union Médicale, 1872, No. 142, 
abstract in Centralblatt, 1873, No. 8). Salkowski, ‘On the Estima- 
tion of Urea and Alkaline Chlorides in Urine containing Potassium 
Iodide” (Pjliiger’s Arch. 1872, v. Abstract in Centralblatt, 1873, 
No. 4). Yvon, “Ona New Method of determining Urea” (1? Union 
Médicale, 1873, January 11. Abstract in Lond. Med. Record, 1873, 
No. 13). Schultzen and Nencki, ‘“‘The Antecedents of Urea in the 
Animal Organism” (Zeitsch. fiir Biologie, 1872, vit. p. 124. Abstract 
in Centralblatt, 1872, No. 50). Salkowski, “Estimation of Potas- 
sium in the Urine” (Pyliiger’s Arch. 1872, v1. Abstract in Central- 
blatt, 1873, No. 2). Maley, “Estimation of Uric Acid” (Pfliiger’s 
Archives, 1872, vi. p. 201. Abstract in Centralblatt, 1872, No. 48). 
Pawlinoff, “On the Sources of the Uric Acid in the Organism” 
(Centralblatt, 1873, No. 16). 


356 DR RUTHERFORD. 


Decomposition oF Uric Acip, by Bacteria.—R. Lex (Cen- 
tralblatt, 1872, No. 33) finds that if a solution of Sodium Phosphate 
and Uric Acid be kept at a temperature of 20 to 30° c., with a 
moderate supply of air, Bacteria develope within afew days. When 
they make their appearance the previously acid reaction of the fluid 
begins to decline, and in the course of from eight to fourteen days 
from the commencement of the experiment the acid reaction com- 
pletely disappears, and alkalinity takes its place. Then, the muroxide 
test fails to detect the slightest trace of Uric Acid, but on the other 
hand the presence of Urea and Ammonium Carbonate may be readily 
ascertained by the usual methods. This decomposition of Uric Acid 
into Urea and Ammonium Carbonate is ascribed by Lex to the fer- 
mentative action of the Bacteria. He supposes that Oxygen and 
Water are taken up during the decomposition. 


Source or Urine Piement.—R. Maly (Annal. der Chem. 
und Pharm. Vol. 163, p. 77) communicates further information 
regarding his statement (Journ. of Anat. and Phys. Vol. v1. p. 468), 
that when Bilirubin suspended in water is treated with a strong 
reducing agent, such as Sodium Amalgam for 2 to 4 days, a substance 
is produced which is identical with Jaffe’s urine pigment Urobilin. 
This substance Maly names Hydrobilirubin, because it contains more 
water than Bilirubin. Its composition is C,, H,, N, O, It is a 
feeble acid, readily soluble in alkaline solutions, from which it may 
be precipitated by the addition of acids. It is sparingly soluble in 
water, but dissolves readily if the water contain Sodium Phosphate, 
Its colour changes under the action of acids and alkalies: its spectral 
characters, etc. all show its identity with Jaffe’s Urobilin. Maly 
found Hydrobilirubin abundantly in the pigment separated from the 
urine after Scherer’s method. He asserts that in the intestine— 
especially the large intestine—Hydrobilirubin is formed from the 
colouring matter of the bile, that it is absorbed from the intestine 
and is excreted by the kidneys. This absorption is not complete 
however, for Hydrobilirubin occurs in the feces. From the spec- 
trum, Maly ascertained the existence of traces of Hydrobilirubin in 
the serum of the blood of the ox, and he ascertained that if this pig- 
ment be injected subcutaneously it is speedily absorbed and excreted 
in the urine. Masius and Vanlair supposed (Journ. of Anat. and 
Phys. Vol. vi. p. 468) that the substance named by them Stercobilin 
differs but little from Hydrobilirubin. According to Maly this sub- 
stance of theirs has no existence. Lastly he cannot agree with 
Heynsius and Campbell in thinking that Urobilin is identical with 
the final product of the oxidation of Bilirubin—to wit, Choletilin ; 
for this, unlike Urobilin, shows no fluorescence, and moreover it 
differs in other essential particulars. Stokvis, “Identity of Choletilin 
and Urobilin” (Centralblatt, 1873, No 14). 


Lachrymation. 


Demtschenko (Pfliiger’s Archives, 1872), from experiments on 
dogs, cats and rabbits, narcotised by morphia, finds that electrical 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 357 


stimulation of the cervical sympathetic causes an increase of the 
lachrymal secretion ; it moreover increases the secretion from the 
conjunctiva, even after removal of the lachrymal gland. Electrical 
excitement of the lachrymal nerve increased the secretion. The 
sympathetic tears were cloudy—the lachrymal nerve tears were clear 
and watery [analogous to the sympathetic and chorda saliva]. The 
increased lachrymal secretion which follows stimulation of the frontal 
infraorbital, nasal, lingual, glossopharyngeal and vagus nerves stops 
if the lachrymal nerve be divided, but section of the sympathetic has 
no effect. Nevertheless in cases of palsy of the fifth nerve, the con- 
junctiva remains moist although the person cannot shed tears. 


Skin. 


Aubert, “On the amount of CO, excreted by the human skin” 
(Pjliiger’s Archives, 1872, vt. p. 549. Abstract in London Med. Kec. 
1873, No. 4). 


Bone. 


Ollier, ‘‘On the Growth of Bone” (Archives de Physiologie, 1873. 
Abstract in London Med. Rec. 1873, No. 5). Philipeaux, ‘On the 
Formation of Bone from Periosteum” (London Med. Rec, 1873, No. 
6). Weiske-Proskau, ‘“ Effect on the Composition of Bone of 
different Earthy Phosphates in the Food. (Abstract in Centralblatt, 
1872, No. 57). Aeby, ‘“‘ Composition of Bone Phosphates (Central- 
blatt, 1873, No. 7). 


Muscle. 


Schenk, “On the Amount of Nitrogen in Flesh” (Abstract in 
Centralblatt, 1872, No. 45).—Salkowski, ‘On the Composition of Mus- 
cular Substance of the Heart” (Pjliiger’s Arch. 1872, p. 213. Abstract 
in Centralblatt, 1872, No. 56). Marcet, “On the Nutrition of 
Muscle and Lung in Health and in Phthisis” (Philosoph. Mag. Nov. 
1872. Abstract im London Med. Rec. 1873, No. 7). Fick, “On 
the Measurement of Muscular Power.” See abstract (in London 
Med. Rec. 1873, No. 6). Preyer, ‘Myophysical Researches” 
(Pjliiger’s Arch. v1. p. 237). A Criticism of these, by J. Bernstein 
(Ibid. vir. p. 90). Volkmann, “ Relations of Elasticity to Muscular 
Activity” (Pjliiger’s Arch. Jan. 1873). 


Animal Heat. 


Siemens, “On measuring Temperatures by Electricity” (Nature, 
1872, May 16). Casey, ‘ Diurnal Variations of the Temperature” 
(Lancet, 1873, Feb. 8). Senator, ‘‘ Researches on the Production of 
Heat and Tissue Metamorphosis” (Reichert und Reymond’s Archiv 
1872, p. 1. Abstract in Centralblatt, 1872, No. 42).——J. Rosenthal, 
“ Regulation of Temperature in Warm-blooded Animals” (Zrlangen, 
1872. Abstract in Centralblatt, 1872, No. 53), among other things 
states the important fact that he—like Riegel (see Academy for Sept. 1, 
1872) has been unable to confirm Nauyn and Quincke’s statement, 


358 DR RUTHERFORD. 


that section of the spinal cord is followed by an increased production 
of heat in the paralysed parts. [On this has been founded the rash 
conclusion that there are nerves which inhibit the production of heat, 
and owing to section of these nerves in the spinal cord, an increase in 
the amount of heat is the result.] Section of the spinal cord is, as one 
would expect, followed by increased loss of heat owing to dilatation 
of blood-vessels. When the temperature of the surrounded medium 
is raised, the temperature of animals thus injured does not rise faster 
than in normal cases. Some considerable time after the operation 
there is indeed an increase in the temperature of the animal, but this 
is apparently owing to a febrile condition resulting from the wound. 
Winternitz, “Regulation of Temperature” (Virchows Arch. 
1872, vi. p. 181. Cenétralblatt, 1873, No. 12). Horvath, “On 
the Physiology of Animal Heat” (Centralblatt, 1872, Nos. 45, 46, 47. 
Abstract of No. 45 in Journal of Anat. and Phys. Nov. 1872, p. 185). 
“On the Behaviour of Frogs under the Influence of Cold” 
(Centralblatt, 1873, No. 3). Stricker and Albert, “On the Tem- 
perature of the Heart” (see abstract in London Med. Record, 1873, 
No. 8), find the blood in the right warmer than that in the left 
ventricle, and that this difference is not due, as Heidenhain and 
Korner supposed, to the proximity of the right ventricle to the liver. 
They found that when the abdominal cavity was opened and the liver 
separated from the under surface of the diaphragm, that the blood in 
the right ventricle still remained warmer than that of the left ven- 
tricle. See also Bernard, “ Lectures on Animal Heat” (Revue 
Scientifique, 1872). 


Generation. 


“Theory of evolution in Germany” (Nature, 1873, March 6 and 
April 3). 


Becinnincs oF Lire.— Dr Burdon Sanderson (Nature, Jan. 9, 
1873) gives an account of experiments on the origin of Bacteria 
performed by Dr Bastian and himself. The experiments show that 
Dr Bastian was right in stating that Bacteria appear in turnip infu- 
sion containing cheese, although it be kept in sealed glass flasks after 
the infusion has been boiled in the flask for as much as ten minutes. 
See leading article (in Brit. Med. Journal, Feb. 1, 1873, and London 
Med. Record, 1873, No. 10). Bastian, “On the Temperature at 
which Bacteria, vibriones, and their supposed germs are killed, when 
immersed in fluids or exposed to Heat in the moist state” (Nature, 
March 27 and April 3, 1873). Roberts, ‘Criticism of Bastian’s 
Experiments” (Nature, Feb. 20, 1873). Huizinga, “‘ New Experi- 
ments on Abiogenesis” (Nature, March 20, 1873). 


Miscellanea. 


Bert, “Influence of Barometric Pressure on the Phenomena of 
Life” (Abstracts in London Med. Rec. 1873, Nos. 10 and 13. 
Liborius, “ On the Quantitative Estimation of Albumen” (Abstract in 


REPORT ON THE PROGRESS OF PHYSIOLOGY. 359 


Centralblatt, 1872, No. 55). Salkowski, “‘ Reaction of Cholesterine 
with Sulphuric Acid” (Pfliiger’s Arch. 1872, v1. Abstract in Central- 
blatt, 1872, No. 55). Hiifner, “‘ Amorphous Ferments” (Journ. 
Sir practische Chemie, v. p. 372. Abstract in Centralblatt, 1872, No. 
37). Bohm, “ Influence of Arsenic on the Action of Amorphous 
Ferments” (Abstract in Centralblatt, 1873, No. 6). Vulpian, “ On 
the Septic Virus” (see London Med. Rec. 1873, No. 9). Davaine, 
“On the same subject” (bid. No. 8). Rabuteau and Papillon, 
* Physiological and Antifermentative action of Sodium Silicate” 
(Comptes Rendus, 1872, uxxv. p. 755. Centralblatt, 1872, No. 54). 
——“ Article on Flowers and Fevers” (Brit. Med. Journal, 1873, 
March 29). 


THE VARIETIES IN THE MUSCLES OF MAN. The ab- 
stract of three Lectures delivered by Professor HUMPHRY 
at the Royal College of Surgeons of England on June 2nd, 
4th and 6th, 1873°. 


LEcTURE I. 


In the three lectures on Myology which Professor Humphry 
delivered last year at the College of Surgeons’, he showed that 
the several muscles in man are modifications of the simple 
pattern which is found in certain lower animals. In the present 
course of Lectures he proposed to show that the varieties in the 
muscles are to be viewed in the same light and that they are, 
generally, the result of an imperfection in those processes of 
segmentation from the simple type and of concentration, by 
which the more complete specialization of the several parts of 
the muscular system in man is attained. These varieties, as 
well as the modifications of which they are varieties, have a 
relation to utility in this way: taking the normal standard _ 
of muscular disposition, as the most perfect and therefore the 
most constant, those variations from it are the most frequent 
which least interfere with the movements of the body. Those 
muscles, that is to say, are the most frequently wanting and, on 
the whole, the most liable to variety which can be best spared 
and the variations in which are least detrimental, such as the 
pyramidalis abdominis, the palmaris longus and the psoas parvus. 

The principle of subdivision or segmentation of muscles for 
the purpose of varied action, and that of concentration upon 
particular points for the purpose of definiteness of action, is 
carried to its greatest extent in man, particularly in the limbs, 
and especially in the upper limb; and it is in these parts that 
varieties most frequently occur. The want of proper segmenta- 
tion is also most frequent in the case of muscles which lie 
parallel and have similar action, as the radial extensors of the 
wrist, which are often united ; whereas the peroneus longus and 


1 These Lectures will be published at length in the British Medical Journal 
during this and the ensuing month. 
? Published in the British Medical Journal of June and July, 1872. 


THE VARIETIES IN THE MUSCLES OF MAN. 361 


peroneus brevis which pass to different parts, and which, though 
parallel in great part of their extent, have different actions, are 
rarely found to be preternaturally blended with one another. 

The imperfection in concentration was illustrated by the 
extension of the coraco-brachialis upon the humerus, as in the 
case of many lower animals, also by the presence of supernu- 
merary muscles, as in the case of a cleido-occipitalis or an addi- 
tional coraco-brachialis ; such muscles being usually adjuncts 
to, or reduplications of, ordinary muscles. 

With regard to the correlation of varieties, the lecturer ob- 
served that several are often present in the same subject, and 
they are often associated with varieties in other structures; and 
the same kind of variety is often repeated in the same person. 
They are also often symmetrical; but the corresponding varie- 
ties are not commonly found in the upper and the lower limbs, 
that is, the serially homologous muscles are not usually affected 
in the same person. 

Males and females, and also the two sides of the body, are 
equally liable to be affected; and the other races of mankind 
and some of the lower animals are, as far as the evidence goes, 
as liable to muscular varieties as ourselves. 

The varieties are themselves very varied, that is, an addi- 
tional muscle or head of a muscle is scarcely alike in any two 
instances of its occurrence. There are however certain lines or 
directions, as they have been called; and these are often in 
the direction of the normal form of some lower animal and 
chiefly of those nearest to man, but also, not unfrequently, of 
those remote from him in the animal scale. This the lecturer 
regarded as an illustration of the similarity of the forces which 
are in operation in the evolution of the several animals, and 
not necessarily as an evidence that the one animal has been 
evolved from the others, and that the varieties are therefore to 
be regarded in the light of reversions to the form of some other 
animal or animals. 

The muscles of the abdomen were first considered. The 
varieties in most of these were described as few and infrequent, 
which accords with the simple disposition of these muscles and 
the little variation they present in the different classes of 
animals. In the obliqui and transversalis they are almost 


VOL. VII. 24 


362 PROFESSOR HUMPHRY. 


limited to the occasional persistence of inscriptions over the ribs 
or the costal cartilages. The most frequent varieties are in 
the two muscles which are most easily spared, viz. the pyramid- 
alis and the psoas parvus. These are often absent; yet one of 
them, the psoas parvus, is very generally present in Mammals, 
while the other is more frequently absent. In the rectus 
also the varieties are few. In the lower animals this muscle 
is often prolonged upon the thorax as far as the first rib. This 
is not the case, even as a variety, in man. Still muscles not 
continuous with the rectus, but in other respects resembling this 
continuation of the rectus, and forming what the lecturer called 
the ‘rectus thoracicus profundus, are sometimes found beneath 
the pectoral muscles. They are to be distinguished from the 
‘rectus thoracicus superficialis,’ also called ‘ sternalis, which hes 
upon the surface of the pectoralis, is often continuous with the 
sterno-mastoid, and which the lecturer believed to be, not as 
supposed by some, a part of the platysma series, but a partial 
representative of that extension of those superficial strata of the 
mesial portion of the abdominal wall into the sterno-mastoid 
which are found in some of the lower vertebrates. It was 
pointed out that this last-named anomalous muscle—the rectus 
thoracicus superficialis—has no representative in other Mam- 
mals, or in Birds or higher Reptiles; and it was suggested that 
its frequent occurrence in Man may have some relation to the 
flatness of the front of his thorax, affording space for its 
development. 

The muscles of the neck were next discussed. The pla- 
tysma and thesterno-cleido-mastoid do not often present varieties ; 
though the former is sometimes developed upon the trapezius or 
over the parotid gland, or furnishes slips to the deeper muscles, 
and the latter is sometimes imperfectly separated from the tra- 
pezius. The varieties in the digastric and stylo-hyoid, which are 
numerous and frequent, were given at some length and were 
shown to be confirmatory of the views elsewhere’ expressed by 
the Professor, that the anterior belly of the digastric is derived 
from a superficial cervical stratum, and the posterior belly and 
the stylo-hyoid from a deeper stratum, and that the intermediate 


1 This Journal, v1. 324, and Observations in Myology, by Professor Humphry, 
p. 136, : 


THE VARIETIES IN THE MUSCLES OF MAN. 363 


tendon is the remnant of the hyoidean intermuscular septum. 
The occasional presence of an inscription in the sterno-hyoid and 
sterno-thyroid was noted as a remnant of one of the primitive 
transverse intermuscular septa; some other irregularities in these 
muscles were noted. 


Lecture II. Delivered June 4, 1873. 


The muscles connecting the upper limb with the trunk—the 
superior ventro-appendicular muscles—were described as in two 
sheets; a superficial and a deep sheet. 

The superficial sheet which, in some of the lower animals, 
as Lepidosiren, forms an almost continuous funnel-like expan- 
sion from the trunk upon the limb, without defined divisions or 
boundaries, is, in Man, divided into three segments or sectors— 
the pectoralis, the latissimus dorsi and the trapezio-deltoid. 
These have tolerably definite boundaries at either end; though 
the segmentation from surrounding muscles is still not quite 
complete. Thus, the pectoral muscle usually presents evidence 
of imperfect separation from the external oblique in the form of 
connecting slips; and the latissimus dorsi is commonly connected 
with the triceps. The varieties in the components of this sheet 
are, for the most part, due to a still less complete separation of 
them from one another, or from the contiguous muscles, at their 
proximal or their distal ends. The bands so often found more 
or less bridging over the axilla and passing from the latissi- 
mus dorsi or the pectoralis, or the ribs contiguous to them, to 
one or other of those muscles or to the humerus or the coracoid, 
and constituting the various ‘axillary muscles’ or ‘achselbogen’ 
are of this nature; also the blendings of the pectoral with the del- 
toid, and of the latter with the trapezius. The costo-epitroch- 
lien was described as a segmented extension of the pectoralis. 
Various muscles described as cervico-humeral, cervico-clavicular, 
masto-scapular, occipito-scapular, &c., were shown to stand in a 
similar relation to the trapezius, and to form, with the rhomboids, 
fragments of a deeper layer of the trapezius. The varieties in the 
deeper layer of the pectoralis, which forms the pectoralis minor, 
are frequent, and consist chiefly of extensions of the muscle, 
over the coracoid, to the humerus, the capsule of the shoulder, 

24—2 


364 PROFESSOR HUMPHRY. 


the supra-spinatus tendon, &c., simulating the disposition of 
the pectoralis minor in many of the lower animals and of the 
levator humeri in birds. The imperfection of concentration or 
limitation, distally, is shown by the extension of the latissimus 
dorsi and of the pectoralis down the arm, where they blend with 
the several muscles and more particularly with the triceps; and 
the deltoid is sometimes prolonged to the supinator longus, as in 
Manis and Birds. 

The deeper layer of the ventro-appendicular sheet consists 
of the subclavius, the omo-hyoid, the levator-scapule, and the 
serratus magnus. Or; these the subclavius is sometimes pro- 
longed in both directions, forming the sterno- or costo-scapularis. 
The omo-hyoid presents many varieties of imperfect concentra- 
tion, being often extended upon the clavicle and sometimes being 
unsegmented from the sterno-thyroid; or it may spread upon 
the upper edge of the scapula as far as the levator scapule; 
and one or both bellies may be double or variously disposed. 
The levator scapulz and the serratus may present the want of 
separation so common in lower animals; or they may be over 
segmented, either being in two or more parts; and they may 
extend at their origin or insertion beyond the usual area. 

The varieties in the deep muscles of the shoulder are few 
and unimportant. 

The adducto-flexor muscles—coraco-brachialis, biceps and 
brachialis anticus—offer many examples of imperfect separation 
and concentration. The two former retain a union at and near 
the coracoid in the normal state; and all three are lable to be 
variously blended and extended. The coraco-brachial is some- 
times spread upon the inner side of the humerus, more or less 
approaching to its disposition in Monotremes and Reptiles. The 
biceps has often two or more origins from the humerus, and 
below is often connected with some of the muscles of the fore- 
arm. It rarely acquires a partial insertion into the ulna, though 
this is often the case in lower animals, its action as a supinator of 
the forearm in Man requiring its limitation to the radius. The 
brachialis anticus is sometimes imperfectly segmented, above, 
from the coraco-brachialis and the biceps, and, below, from the 
muscles of the forearm. The triceps presents few irregularities. 

In the forearm the palmaris longus varies often and in 


THE VARIETIES IN THE MUSCLES OF MAN. 365 


every conceivable way. The other pronato-flexor muscles pre- 
sent many examples of deficiency of separation. In the nor- 
mal state most of them retain some of the union which in the 
primitive form is more extensive; and the irregularities present 
many conditions of union intermediate between the normal and 
the primitive form. ‘The internal condyle of the humerus and 
the coronoid process of the ulna were pointed out as the two 
chief meeting points or starting points of these muscles, espe- 
cially the coronoid process; and many of the irregularities con- 
sist of unusual, or unusually extended, connections with this 
process. The flexor longus pollicis, which is more segmented 
from other muscles in Man than in the lower animals, still in 
him retains its hold upon this process, and often fails to be 
separate from the flexor digitorum. The numerous irregulari- 
ties in the flexors of the fingers were alluded to in connection 
with the varied disposition of these muscles in lower animals. 
The flexor sublimis and the flexor profundus are, together with 
the flexor longus pollicis, divisions of one muscle, and the re- 
tention of the union is often presented by slips passing between 
them and by union of their tendons, and especially of those 
going to the little finger. The lumbricales, though usually re- 
maining with the flexor profundus alone, sometimes are con- 
nected with the flexor sublimis, or with the flexor longus pollicis; 
or they may extend up to the coronoid process. The Hexor carpi 
radialis sometimes extends, above, upon the coronoid process, 
and is blended with the other muscles; and, below, 1t may 
range upon the carpal bones, and upon the third and fourth 
metacarpals. A flexor carpi radialis brevis, passing from the 
radius to the carpal bones or the metacarpals, has in several 
instances been met with, and is a reminder of the extensive 
attachment of the pronato-flexor mass to the bones of the fore- 
arm and to the carpus and the metacarpus in the Cryptobranch 
and other Urodelans. 

On the dorsal aspect of the forearm, the supinators and the 
radial extensors of the wrist are often incompletely separated 
from one another and from the adjacent muscles. ‘The exten- 
sor digitorum is often not separated from the extensor carpi 
ulnaris; and the extensor minimi digiti is an incomplete seg- 
ment from between them, which often ranges upon other fingers 


366 . PROFESSOR HUMPHRY. 

besides the fifth. In the deeper layer, the extensors of the 
thumb, which in Man only are devoted each to its particular 
bone of the thumb, often fail to attain this individuality in him. 
They are frequently blended. Especially is this so with the ex- 
tensor primi internodii, which has less expressed function than 
the others and is often absent. In like manner the extensor 
indicis is frequently not separate; or it ranges upon other fin- 
gers; or it is absent. 


Lecture III. Delivered June 6. 


In the lower limb the muscular irregularities are fewer than 
in the upper limb, owing, partly, to the less complex disposition 
of muscles in the limb and to the absence, in consequence of the 
fixity of the pelvic girdle, of muscles, the homologues of which, 
in the upper limb, are the seat of many varieties. The sarto- 
rius is the most erratic of those passing from the pelvis, and it 
is the most variable in lower animals. The biceps, by its occa- 
sional connections with the gluteus and extension to the sacrum 
and the crista of the ilium, harks back, as it were, to its primi- 
tive relations with these parts’, The adductors may be va- 
riously segmented. The semitendinosus, semimembranosus and 
popliteus do not vary much. 

The irregularities in the calf-muscle are on the side of ex- 
cess rather than of deficiency; although that muscle is in Man 
more complicated than in any other animal. They consist 
chiefly in the presence of additional heads to the gastrocnemius, 
or in extensions of the origin of that muscle and of the soleus. 
The plantaris is sometimes absent or abnormally connected with 
adjacent parts above or below. 

In the deep flexor muscles the most mterestmg varieties are 
in connection with the accessorius, which is a remnant of the 
primitive wide attachment of the simple unsegmented flexor 
mass in the Urodelans to the bones of the leg, the tarsus and 
the metatarsus. Accordingly extensions of it, or derivatives 
from or adjuncts to it, are often found spreading up the leg 


~ 1 See British Medical Journal, July 20, 1872, Journal of Anatomy, v1. 355, 
Observations in Myology, p. 167. : 


THE VARIETIES IN THE MUSCLES OF MAN. 367 


along the fibula or widely connected with the tarsus. Below, 
it is sometimes more largely united with the flexor digitorum 
than usual, or it joins the flexor hallucis or the lumbricales, 
or it gives off one of the flexor tendons of the toes. The 
varieties in the flexors of the toes consist chiefly in the 
closer union of the flexor digitorum and the flexor hallucis, as 
is the case in lower animals, and in the partial absence of the 
flexor brevis, or the blending of it with the flexor longus, which 
more particularly happens with regard to the division to the 
little toe. 

On the dorsal aspect of the limb the quadriceps is but little 
liable to varieties. The portion of the tibialis anticus attached 
to the metatarsal bone is sometimes separate; and, further, is 
sometimes subdivided, one portion passing to the first pha- 
lanx, thus resembling the disposition of the extensor tendons 
to the pollex. Sometimes the tibialis anticus extends to the 
plantar fascia; or it ranges upon the dorsum of the foot, as 
in the Hippopotamus. The peroneus longus seldom varies ; 
though sometimes attached to other metatarsals, in addition 
to the first, it rarely or never fails to reach the first. The 
other peronei often show their imperfect segmentation from 
the extensor digitorum by spreading upon the toes. Par- 
ticularly is this the case with the peroneus tertius, the com- 
plete separation of which and its devotion exclusively to the 
tarsus is a human feature, and is related to the mode in which 
the fore part of the sole is planted fully upon the ground in 
Man.—The extensor digitorum sometimes shows a persistence 
of connection with the metatarsal bones with the extensor hal- 
lucis and with the extensor brevis, which are reminders of the 
primitive oneness of the extensor mass, and which are evidences 
of imperfect segmentation of it. No instance had presented 
itself however of that extension to the fore part of the femur 
which is so frequent in other Mammals.—The varieties in the 
small muscles of the foot and hand were also discussed. 

In indicating the relation between utility and variability, 
and showing that that which is most useful has on the whole 
the greatest stability, the Professor did not express any view as 
to the connection between the two, or wish to prejudge the 
great questions associated with this subject and with the allied 


368 PROF. HUMPHRY. VARIETIES IN THE MUSCLES OF MAN. 


problem how that which is most fitted for its purpose in each 
animal and each part comes to be present. Other questions 
were alluded to. Whether, for instance, the variety in muscles 
which are of least importance, on the one hand, and in those 
which are peculiar to Man, on the other, is an indication that 
those muscles are in process of being fixed in, or expunged 
from, the economy. Also whether varieties are more or less 
frequent in the more advanced and more civilized members of 
the human family than in others. At present there is not suffi- 
cient evidence to furnish an affirmative answer to any of these 
questions, or to establish the hereditary transmission of muscu- 
lar varieties, which must be regarded as probable.—The nerve- 
supply to the supernumerary muscles, in the few instances in 
which it has been described, seems to corroborate the view “that 
nerve-course is somewhat too arbitrary or too much regulated by 
convenience in each instance for us to be able to rely upon the 
disposition of the nerves as sure guides to the discernment, in 
difficult cases, of the homological relations of muscles and 
other structures *.” 


1 Journal of Anatomy, v1. 56, Observations in Myology, p. 56. 


INDEX TO VOLUME VII. 


A 


Abiogenesis, new experiments on, 358 

Abnormalities, see ‘‘ Variations ” 

Absorption, cutaneous, 188 ; of heat in 
incubation, 185; phys. reports on, 
348, 349 : 

Acid, carbonic, an antidote for strych- 
nia, 196, amount of from human skin, 
357; uric, sources of, 355, decomp. 
of by Bacteria, 356; sulphuric, 
cutaneous nerves irritated by, 177, 
reaction of cholesterine with, 359 ; 
cholic, 352; phosphoric, in serum, 
190; passage of free acid, into urine, 
through alkaline blood, 355 

Aeby, comp. of bone phosphates, 357 

Africa, arrow poison of, 139 

Albert, on cardiac temp., 358 

Albini, on Thiry’s Fistula, 187 

Albumen, quantitative estimation, 358 

Atimentation, 350—354 

Allbutt, T. Clifford, exercise on bodily 
temp., 106 

Amblypterus Anconozchmodus, 338 

Anderson, Dr E. A., Potass. Bromid. 
in snake bites, rgt 

Anomalies, see ‘‘ Variations” 

Antagonism of poisons, 197, 198 

Apes, anthropoid, 171, 334 

Aplysia, develop. of, 338 

Apomorphia, an emetic, 194 

Appendicularia, anat. of, 338 

Aptenodytes Humboldtii, eye of, 170 

Arctocephalus cinerius, 335 

Arloin, action of r. and |. vagus, 180 

Arterial system, see “‘ Circulatory ” 

Articulation of cerv. vert. of Balzeno- 
ptera musculus and B. rostrata, 1 

Artio-dactyla, comp. anat. 336 

Arsenic, influence of on amorphous fer- 
ments, 359 

Arytenoid eartilages, fixation of during 
phonation, 190 

Ashmead, physostigma, an antidote to 
strychnia, 198 

Atropia, an antidote to physostigma, 
198 ; morphia, an antidote to, 198 

Aubert, amount of CO, excreted by 
human skin, 357 


B 


Bacteria, decomp. of uric acid by, 356; 
temp. destructive to, 358 

Balena, primigenius, 173 

Balznoptera, musculus and rostrata, 
cery. vert. of, 1,172; B. patachonica 
and intermedia, 335; Davidsoni, 336 

Balzxnotus, insignis, 173 

Balenula, balzenopsis, 173 

Bartels, Max, quadruple mammez in 
Man, 332 ~ 

Bastian, temp. destructive to bacteria, 
vibriones, &c., 358 

Batrachia, see ‘‘ Reptilia ” 

Baumstark, cholic acid, 352 

Baxt, cutaneous nerves irritated by sul- 
phuric acid, 177 

Bears, New Zealand, and North Austra- 
lian sea-, 335 

Beaunis, injections into brain, 177 

Belladonna, checks and prevents sweat, 


Beneden, P. J. van, on various fossil 
whales from Antwerp, 173; osteology 
of the Dugong and Manatee, 336 

Berardius arnouxii, skeleton of, 335 

Berger, Paul, internal structure of um- 
bilical vessels, 171 

Bernard, lect. on animal heat, 358 

Berstein, nerves, phys. of, 344; ab- 
sorption, influence of nerves on, 348, 
relation of nerve centres to, 349; on 
Preyer’s myophysical researches, 357 

Bert, barometric pressure and pheno- 
mena of life, 358 

Betz, W., examination of central organs 
of human nervous system, 329 

Bile, pigment of in urine, 165; oxida- 
tion products of, 352; determination 
of sulphur of, 352; biliary calculi, 
352; phys. of human, 352 

Binz, Prof., Quinia, 196 

Birds, ciliary muscle in, 170; sight in, 
344; blood-corpuscles in, 346; anat. 
reports on, 173, 174, 337 

Bladder, see “ Genito-urinary organs” 

Blake, Dr James, action of inorganic 
substances introduced directly into 
blood, 201 


370 


Blood, ferment-action of, 187; inor- 
ganic constituents of, 346; gases of, 
346; absolute quantity of, 347 ; iron 
in, 346; new test for, 346; physical 
nature of coagulation, 210; direct 
introduction into of inorganic sub- 
stances, 201; minute moving parti- 
cles in, 347; urine after injection of 
water into, 355; free acids, through 
alkaline b., into urine, 3553; coagula- 
tion of, a4: 346, corpuscles of, 346 

Bock, on inducing diabetes, 187 

Bohm, effect of arsenic on amorphous 
ferments, 359 

Feettcher, labyrinth of ear, 344 

Boll, vision with compound eyes, 179 

Bone, phys. reports on, 357 

Boulaud, P., curvatures of the spine, 168 

Boussingault, iron in blood and food, 
346; chem. of milk, 355 

Bowditch, H. P., irritability of muscu- 
lar fibres of heart, 182 

Brain, galvanisation of, 175; forms of, 
in Carnivora, 172, in Marsupialia, 
172; injection into substance of, 177; 
localisation of movements in, 340; 
grey matter of, instrument for inea- 
suring, 169; minute structure of in 
brain-wasting, 170; hypertrophy of 
hemisphere of, with atrophy of oppo- 
site side of body, 257; relation of 
to mental characteristics in dog, 331; 
fonctionnement du cerveau, 340; 
phys. reports on, 339 

Bremond, M., absorption of medicinal 
substances by skin, 198 

Broca, Paul, “deformation Toulousaine” 
of skull, 167 ; caudal vert. of tailless 
Primates, 334 

Brown-Séquard, strychnia, 196; influ- 
ence of vagus on convulsions, 343 

Briicke, carbo-hydrates, their digestion 
and absorption, 350 

Bruhl, supernumerary muscle to great 
toe, 328 

Brunton, T. Lauder, effect of digitalis, 
on the blood-vessels, 134; report on 
phys. 175 ; Handbook for the Physio- 
logical Laboratory, 322 

Burdon-Sanderson, Dr J., Handbook 
for the Physiological Laboratory, 322 

Burmeister, H., Globiocephalus Grayi, 
and Pseudorea Grayi, 173 
3urtinopsis similis, 173 

Buzzard, ciliary muscle i in, 170 

Byasson, Dr H., absorption of soluble 
salts of mercury, 191; chloral-sul- 
phydrate, 192; formic ether, 193 

C 

Cesium, salts of, introduced directly 

into blood ‘205 


INDEX. 


Calculi, biliary, 352 
Campbor, little energy on Man, 197 
Cappie, Dr Jas., Causation of sleep, 321, 


339 

Carbo-hydrates, digestion and absorp- 
tion of, 250 

Carbonic acid, see ‘‘ Acid” 

Carlet, G., locomotion in Man, 169 

Carnivora, forms of brain in, 172; comp. 
anat., 335 

Carter, H. J., Sperm-whale in tropical 
Indian Ocean, 335 

Cartilage, fibro-, structure and develop. 
of elastic tissue in, 327 

Casey, diurnal var. of temp. 357 

Cells, passage of fat from aliment into, 
350 

Ceratodus, myology of, 169 

Ceriornis satyra, cranial appendages of, 
337 

Cervus Alces, remains of in Britain, 
336 

Cetacea, comp. anat. reports on, 172, 
173, 335, 336 

Cetotherium hupsebii, brevifrons, dubi- 
um, burtinii, 173 

Charles, Dr. J. J., art. var. in upper 
extremity, 300; 

Chatin J., myology of Hycemoschus, 
172 

Chaveau, Prof. A., Comparative Anatomy 
of the Domesticated Animals, 322 

Cheiroptera, myology of the, 171 

Chelonians, sternum of, 337 

Chimpanzee, 171, 334 

Chloral, 192; sulphydrate, 192; strych- 
nia an antidote to, 197 

Chlorides, urea and alkaline, in urine 
with potassium iodide, 355 

Chlorine, behaviour of in the organism, 
350 

Chloroform, combined effects of opium 
alkaloids ‘and, 194 

Cholesterine, reaction of with sulphuric 
acid, 359 

Choletilin, identity of, and Urobilin, 356 

Cholic acid, see ‘‘ Acid” 

Chorda Tympani, 342, 343 

Cienkowski, L., Noctiluca miliaris, 238 

Circulatory system; heart, innervation 
of, 180, 347; arteries, variations of 
in upper extremity, 300; cranial, in 
sharks, 338; veins, valves in renal, 
163, changes in liver from obliteration 
of portal, 187; placental cavernous 
sinus system, 333; blood-vessels, ef- 
fects of digitalis on, 134, innervation 
of, of rabbit’s ear, 347; blood, a fer- 
ment-action of, 187, action of inor- 
ganic substances introduced directly 
into, 201, physical nature of coagu- 
lation of, 210, pressure of in respira- 


INDEX.. 


tion, 347; circulation, in optic nerve 
and retina, 344, relation of stomach 
to nerve centres of, 347; influence of 
respiration on, 348; pulse, Jaw regu- 
lating frequency of, 219, cause of *re- 
tardation of on closure of nostrils 
of rabbit, 283; of Globio-cephalus 
melas, 336; reports, anat., £71, 331, 
phys. 180—185, 345—347 

Clark, John W., visceral anat. of Hip- 
popotamus, 336 

Cochlea, rods of, 170; preparation of, 
for microscopy, 170, of organ of Corti 
for ditto, 170 

Cold, behaviour of frogs under influence 
of, 358 

Coleoptera, injection of, 185 

Colius, the genus, 174 

Colour, topography of, 344; perception 
of, 344 

Compensator, use of the round, 179 

Compression of brain, effects of artificial 
resp. on, 348 

Concussion, effects of artificial resp. on, 
348 

Convulsions, influence of vagus on, 343 

Cornea, permeable to fluids, 189; nerves 
of, 331 

Coronula, balznaris, and diadema, 335 

Corpora quadrigemina, phys. of, 175 

Corpuscles, blood, in Mammals, Birds 
and Fishes, 346, colourless, action of 
quinine on, 346 

Corti, organ of, preparation of, for mi- 
croscopy, 170 

Crypto-branchus, myology and nerves 
of, 169 

Crypto-lepas rhachianectis, 335 

Cunningham, D. J., nerves of the head 
and neck, 94 

Cuon primeevus, 335 

Curnow, Dr John, var. in muscles and 
nerves, 304 

Cyamus Scammoni, suffusus and mysti- 
ceti, 335 

Cyanates, not poisonous, 193 

Cynocephalus, Hamadryas, anat. of, 171, 
334 

Cyno-phrenology, 331 

Cystophora cristata, 335 


D 


Dall, W. H., Cetacea, on the parasites of, 
of the N. W. coast of America, 335, 
on three new species of, 335 

Dalton, B. N., muscular var., 327; art. 
var. 331 

Darwin, F., ostrich, 337 

Davaine, septic virus, 359 


371 


Davies-Colley, muscular var., 327; art. 
var. 331 

Death, on a sure sign of, 190; changes 
in electrical excitability in muscles 
after death, 191 

Delphinus Bairdii, 335 

Demtschenko, lachrymation, 356 

Dentition of Narwhal, 75 

Development, see ‘‘Embryology” 

Dewar, James, phys. action of light, 
275, 278 

Diabetes, new plan of inducing, 187 

Dicyema, anat of, 338 

Diet, regressive metamorphosis during 
fiesh diet, 350 

Digestive system, stomach and nerve- 
cent:es of cireulation, 347; duodenal 
diverticula, 332; cecum of Indian 
Wild Dog, 335, absence of in Para- 
doxure, 335; gizzard, mechanism of, 
337; pancreas, of osseous Fishes, and 
tubes of Weber, 338; visceral anat. 
of Greenland Shark, 233, Globio- 
cephalus melas, 336, Hippopotamus, 
336; phys. reports on, 186, 187, on 
alimentation, 350—454 

Digitalis, effeets of on blood-vessels, 

ok 

Dinocerata, 267, 337 

Diplax Perithemis, develop. of, 338 

Dipnoans, homologies of shoulder-girdle 
of, 338 

Dobrowlsky, perception of colour, 344 

Dock, formation of glycogen in liver, 
187 

Dog, gases of lymph of, 189; rela- 
tion of brain to mental characteristics 
in, 331; the Indian wild, 335; extir- 
pation of pancreas in, 353 

Dolphin, Risso’s, 173 

Donders, F. C., congenital and acquired 
association, 344 

Donitz, W., structure of kidney - of 
African Elephant, 171 

Drachmann, Prof. A. G., congenital 
absence of quadriceps ext. cruris, 310 

Dreams, 339 

Duhay, N., abnormal formation of me- 
sentery, 332 

Dugong, osteology of, 336 

Duncan, Dr J. Matthews, placental 
cavernous sinus system, 333 

Duret, M., art. of med. oblong., 331 

Dwight, Thos., Baleenoptera musculus, 
172 


E 


Eberth, C. J., smooth muscular fibres im 
human kidney, 171 

Ebstein, formation of pepsine in stomach, 
186 


372 


Edwards, Alph. Milne, Placenta in Ta- 
mandud, 172; anat. of Limulus, 338 

Edwards, H. Milne, La Phys. et ? Anat. 
Comp. de VHomme et des Animaux, 
166 

Eel, hermaphroditism in, 338 

Elasticity, relations of to muscular ac- 
tivity, 357 

Electricity, measuring temp. by, 357 

Elephant, Indian, anat. of 60; African, 
structure of kidney of, 171 

Embryology, develop. of ova of river 
Trout, 174, and fecundation of ovum 
in Rabbit, 332, of Loligo, Aplysia, 
Terebella nebulosa, Nudibranchs, 338, 
of teeth of Snakes, 338, of skull of 
Sparrow, Tit and genus Turdus, 337, 
osseous, of limbs, 334; malformed 
human feetus, 332 

Emys lutaria, action of right and left 
vagus in, 180 

Enge!mann, movements in nerve-fibres, 
178, striped muscular fibre during 
rest, activity and rigidity, 328 

Epiodon Heraultii, 336 

Ercolani,G. B., var in bladder, 332; fur- 
ther remarks on perfect hermaphro- 
ditism in eel, 338 

Eschrichtius robustus, 173 

Eulenburg, electrical stimulation of 
nerve aud muscle, 179 

Evolution, theory of, in Germany, 358 

Ewald, A., congenital hypertrophy of 
left band, 332 

Exner, iridectomy, 344 

Eye, nerves of, 331; vision with com- 
pound, 179; cornea, permeable to 
fluids, 189; reports on, auat., 170, 
171, phys. 344 


F 


Falck, sodium chloride, 350; urine after 
injection of water into blood, 355 

Fat, passage of, from aliment into cells, 
35° 

Felide, claw at end of tail of, 271 

Feltz, V., absorptive power of bony me- 
dullary tissue, 167 

Ferment, amorphous, 359; effect of 
arsenic on, 359 

Ferrier, Prof. David, report on Phys., 
17 

Beee chemical contribution on, 352; 
flowers and, 359 

Fibrin, coagulation of, 345; peptones 
from, 3523; action of pepsin on blood, 


352 
Fick, Prof., pneumograph, 348; mea- 
surement of muscular power, 357 
Filehne, W., law of contraction in dying 
nerves, 344 


INDEX. 


Finny, J. Magee, morphia an antidote 
to atropia, 198 

Fischer, P., sperm-whale on the coasts 
of France, 173; Cetacean bones trom 
Léognan, 173; Globio cephalus ma- 
crorhynechus and Edwardsi, 336 

Fischer, trophic nerves, 343 

Fishes, blood-corpuscles in, 346; comp. 
anat. reports on, 174, 33 

Fistula, Thiry’s, 187 

Fleming, George, translation of Cha- 
veau’s Comparative Anatomy of the 
Domesticated Animals, 322 

Flower, W. H., Diagram of the Nerves 
of the Human Body, 166; Risso’s 
Dolphin, 173; sub-fossil whale in 
Cornwall, 173; carpus of Sloths, 255 ; 
Nandinia binotata, 335; skeleton of 
Berardius arnouxil, 335 

Flowers and fevers, 359 

Follet, Formic ether, 193 

Food, iron in, 346; article on, 350 

Formic ether, 193 

Forster, Perimeter, &c., 344 

Foster, M., inhibition of beart’s action 
in Mollusca, 184; Handhovok for the 
Physiological Laboratory, 322 

Fournié, le Fonctionnement du Cerveau, 
340 

Frankel, accommodation in human eye, 
344 

Fraser, Thomas R., the Kombé arrow- 
poison, 139; reports on phys. action 
of medicinal and poisonous substances, 
191 ; Atropia, antidote to physostig- 
ma, 198 

Frog, termination of nerves in bladder 
of, 170; on the current in gastrocne- 
mius of, 179 ; esophagus and stomach 
of, innervation of, 350; behaviour of 
under influence of cold, 358 

Fuchs, laws of nerve stimulation, 179 


G 


Gaethgens, free acids into urine through 
alkaline blood, 355 

Galton, Francis, hereditary genius, 339 

Galton, John C., translation of Roser’s 
Manual of Surg. Anat., 166. 

Garrod, A. H., Sphygmography, 98 ; law 
regulating frequency of pulse, 219; 
source of nerve-force, 251; Dinoce- 
rata, 267; Ostrich, 337; Huia Bird, 
337; Gizzard, 337 

Gegenbaur, Carl, zur vergleichenden 
Anat. der Wirbelthiere, 166 

Gelatine, nutritive value of, 350 

Geltousky, Dr, on Quinia, 195; qui- 
nine, on colourless blood-corpuscles, 
346 


INDEX. 


Generation, phys. reports on, 358 

Genito-vrinary organs of Elephas Indi- 
cus, 60, of Greenland Shark, 233; 
placenta, structure of human, 120, of 
Tamandua, 172; bladder, structure 
and termination of nerves in, of frog, 
170, sphincter of, 191, var. in, 332; 
hypospadia with cleft scrotum, 332 ; 
reports, anat. on, 333, 334, on kiduey, 
171, 331, on ovum and ovary, 332, 
333, phys., on urinary system, 187, 
188, on urine, 355, 356. 

Genius, hereditary, 339 

Gervais, Paul, forms of brain, in Car- 
nivora, 172, in Marsupialia, 172; on 
the Baleen whales, 173: carcase of a 
male Cachalot and a foetal Delphinus 
delphis, 336; osteology of Sphargis 
Luth, 337 

Giglioli, E. H., Chimpanzee, 334 

Gill, Th., homolo-ies of shou!der-girdle 
of Dipnoans, &c., 338 

Gillette, Dr, sesamoid bones in Man, 
167 

Glands, Brunner’s, 354; mammary, 
quadruple, 56, 332; parotid, stimuli 
on secretion of, 161; submax., secre- 
tion of, 186; pineal, comp. anat. of 
structure of, 330; peptic, structure 
of, 351; pyloric, peptic action of, 
351; relation of nerves to gland- 
cells, 330 

Globio-cephalus, Grayi, distinguished 
from Pseudorca Grayi, 173; melas, 
336 

Glycera, anat. of, 338 

Glycogen, formation of, in liver, 187 

Goltz, innervation of cesophagus and 
stomach of Frog, 350 

Graham, T., hypospadia with cleft 
scrotum, 332 

Grampus griseus, 173, Stearnsii, 335 

Gray, J. E., New Zealand Sea-Bear, 
335; Prof. Flower’s memoir on Berar- 
dius, 335; distribution, migration, &c. 
of Whales and Dolphins, 335; new 
species of Ziphioid Whale, 336; ster- 
num of Chelonians and skeleton of 
Sphargis, 337 

Gripat, H., acephalous Calf, 332 

Gruber, Wenzel, supernumerary carpal 
bones, 167, 325; muscular var., 108, 
327; art. thyroidea ima, int. mam- 
mary, &c., 171; var. in postero-sup. 
angle of scapula, 326; hernia-like 
protrusion of synovial membranes, 
329 

Griitzner, formation of pepsine in sto- 
mach 

Guyon, on combined effects of opium, 
alkaloids and chloroform, 194 

Gypsophoca tropicalis, 335 


373 


H 


Hagemann, comp. anat. of structure of 
pineal gland, 330 

Hair, see ‘‘ Integumentary system” 

Hammersten, gases of Dog’s lymph, 189 

Hand, hypertrophy of, 332 

Handyside, P. D., Quadruple Mamma, 
56, 332; hypospadia with cleft scro- 
tum, 332 

Hare, hybrid of Rabbit and, 172 

Hailey, Dr John, camphor, 197 

Hart, Ernest, abstract on Beettcher’s 
labyrinth of ear, 345 

Hartmann, Rt., anthropoid Apes, 171, 

34 

He C., Anatomische Studien, 166 

Haughton, S8., animal mechanics, 168; 
Prirciples of Animal Mechanics, 312 

Heart, temp. of, 358; innervation of, 
180, 347; imbibition of action of, in 
Mollusea, 184; muscular fibres of, 
irritability of, 182, comp. of, 357 

Heat, see ‘‘ Temperature” 

Hector, James, New Zealand Bottle- 
nose, 173; Whales and Dolphins of 
New Zealand Seas, 335 

Heidenhain, R., secretion of submax. 
gland, 186 

Heitzmann, C., bone and cartilage, 327 

Hermann, effect of galvanic currents on 
muscle and nerve, 178 ; phys. of vomit- 
ing, 186 

Herpetocetus scaldiensis, 173 

Hertwig, Oscar, structure and develop. 
of elastic tissue in yellow cartilage, 


327 

Heteralocha gouldi, 337 

Heubel, relations of nerve-centres to 
absorption, 349 

Hicks, Braxton, objection to the theory 
of placental cavernous sinus system, 
333 

Hippopotamus, visceral anat. of, 336 

Hitzig, on galvanisation of brain, 175 

Hemoglobin, 346 

Hofmann, F., passage of fat from ali- 
ment into cells, 350; free acids, into 
urine, through alkaline blood, 355 

Hoffmann, on inducing diabetes, 187 

Holdsworth, E. W. H., peculiar Ceta- 
cean, 336 

Hollis, W. Ainslie, tissue metabolism, 
80 


Horvath, animal heat, 185, 358; be- 
haviour of Frogs under influence of 
cold, 358 

Hoyer, H., nerves of the Cornea, 331 

Hiifner, amorphous ferments, 359 

Huia Bird, 337 

Huizinga, new experiments on Abio- 
genesis, 358 


374 


Humphry, Prof., 00s. in Myology, 166, 
169, 334, 348; varieties in muscles, 


360 
Hybrid of Hare and Rabbit, 172 
Hycemoschus, myology of, 172 
Hypertrophy of left hand, 332 
Hyrt!, J., pelvis of kidney in Man and 
Mammals, 171; var. in human crania, 
326; cranial arteries in Sharks, 338 


I 


Ichthyornis dispar, 337 

Incubation, absorption of heat during, 
185 

Insects, preparation of eyes of for mi- 
croscopy, 170; flight of birds and, 
173; development of hexapodous, 338 

Instinct, 340 

Integumentary system ; skin, perception 
in, of lower limb, 180, amount of CO, 
excreted by, 357, absorption of me- 
dicinal substances by, 198; sweat, 
checked and prevented by belladonna, 
197; tactile hairs, termination of 
nerves in, 331; teeth of snakes, struc- 
ture and development of, 338; pow- 
der downs of Rhinochetus jubatus, 
174; phys. reports on, 188, 189, 344 

Tridectomy, phys. action of, 344 

Tron, in blood and food, 346 

Trregularities, see ‘* Variations’ 

Isotoma, develop. of, 338 


’ 


J 


Jackson, Dr H., localisation of move- 
ments in Brain, 340 

Janisch, inorganic constituents of blood, 
346 

Jelenffy, fixation of arytenoid cartilages 
during phonation, 190; action of m. 
erico-thyrod., 328 

Joseph, trophic nerves, 343 

Jukes, structure of peptic glands, 351 


K 


Kapff, H., the ovary, 333 

Kerner, Quinia, 195 

Keuchel, Paul, action of various sub- 
stances on nerves of submax. glands, 


9 
Key, Axel, on anat. of nervous system, 
329 
Kidney, see “Genito-urinary organs,” 
Klein, Dr E., Hand-book for the Physio- 
logical Laboratory, 322 
Knoll, phys. of corpora quadrigemina, 


175 
Kolliker, Albert, typical absorption 
surfaces in bones, 167 


INDEX. 


Krauss, F., pelvic bones of Manatee, 
336 

Krolow, Brunner’s Glands, 354 

Kubnt, duplicity in hands and feet, 332 

Kiiltz, determination of sulphur of bile, 
352 

Kiintzel, on inducing diabetes, 187 

Kupffer, C., relation of nerves of glands 
to gland-cells, 330 

Kupressow, sphincter vesicz, 191 


L 


Labbe, combined effects of opium alka- 
loids and chloroform, 194 

Labyrinth of ear, 344 

Lachrymation, 356 

Lagenorhynchus clanculus, 173 

Lang, suppression of perspiration, 189 

Langer, Karl, Growth of skeleton, 168 

Language, philosophy of, 339 

Lankester, E. Ray, development and 
anatomy of several invertebrata, 338; 
hemoglobin, 346 

Laqueur, cornea permeable to fluids, 
189 

Lavdowsky, M., termination of nerves 
in bladder of frog, 170 

Leber, circulation in optic nerve and 
retina, 344 

Lee, R. J., ciliary muscle in birds, 170; 
eye in Rhea, Phenicopterus and Ap- 
tenodytes, 170; sight in birds, 344 


Legg, I. Wickham, bile-pigment in 
urine, 165 


Legouis, P., pancreas of osseous fishes, 
and tubes of Weber, 338 

Legros, vagus, 343 

Leidy, Jos., polydactylism in horse, 
332 

Lens, anat., phys. and path. of crystal- 
line, 171 

Lepidosiren, myology and nerves of, 169 

Lepine, blood-gases, 346 

Lesser, to obtain lymph in large quan- 
tities, 184 

Lewes, instinct, 340 

Lex, R., decomp. of uric acid by Bac- 
teria, 356 

Leydig, F., structure and development 
of teeth of snakes, 338 

Liborius, quantitative estimation of al- 
bumen, 358 

Life, beginning of, 358; barometric 
pressure and the phenomena of, 358 

Light, physiological action of, 275, 278 

Lime, on estimating, in serum, 190; in 
urine, 355 

Limulus, anat. of, 338; develop. of L. 
Polyphemus, 338 

Liolepis Belli, myology of, 338 

Lion, claw at end of tail of, 271 


INDEX. 375 


Lithia, salts of introduced directly into 
blood, 202 

Liver, see ‘‘ Digestive system”’ 

Locomotion, in Man, 169; terrestrial, 


Bao 

Loligo, develop. of, 338 

Loris, 334 

Lowne, B. Thompson, Descriptive Cata- 
logue of Teratological Series in Museum 
of Royal College of Surgeons, England, 
320 

Lung, see ‘‘ Respiratory system” 

Lymph, to obtain in large quantities, 
184; gases of dog’s, 189; phys. re- 
ports on, 354, 355 


M 


Macalister, Alex., catalogue of human 
muscular anomalies, 168; anat. of 
Cynocephalus Hamadryas, 171; my- 
ology of Cheiroptera, 171, of Sarco- 
philus ursinus and Phascolarctos ci- 
nereus, 172; cranium of broad-headed 
Wombat, 337 

Magnus, sure sign of death, 190 

Major, C. J. F., Fossil Monkeys in 
Ttaly, 171 

Major, H. C., Tephrylometer, 169; 
minute structure of cortical substance 
of brain, 170; 


Malassez, blood-corpuscles in mammals, 


birds and fishes, 346 
Maley, estimation of uric acid, 355 
Malformation, of leg and foot, 156; 
anat. reports on, 332 
Maly, R., source of urine pigment, 356 
Mamme, quadruple in Man, 56, 332 
Mammalia, blood-corpuscles in, 346; 
comp. anat. reports on, 17I—173 
_Manatee, osteology of, 336; form and 
structure of, 336 
Manassein, chemical contributions on 
Fever, 352 
Mandelstamm, association of the two 
retine, 344 
Marcet, nutrition of muscle and lung in 
health and in Phthisis, 357 
Marey, M., flight of birds and insects, 
173; terrestrial locomotion, 329 
Marsh, O. C., Dinocerata, 337; new 
sub-class of fossil birds, 337 
Marsupialia, form of brains in, 172; 
cranium of broad-headed Wombat, 337 
Martius, C., thoracic and pelvic extre- 
mities, 334 
Mason, polydactylism in horse, 332 
Masoin, action of right and left vagus, 
180 
Matthieu, blood-gases, 346 
McKendrick, Dr J. G., Physiological 
action of light, 275, 278- 


McRae, Dr A. Edward, Chloral, 192 

Mechanics, animal, 168 

Mechanism of muscles, 169 

Megaceros hibernicus, remains of in 
Scotland, 336 

Megapteropsis robusta, 173 

Mercury, absorption of soluble salts of, 
1gt; action of on liver, 352 

Merkel, Fr., contraction of striped mus- 
cular fibre under polarised light, 329 

Metabolism, 80 

Metamorphosis, regressive during flesh 
diet, 350; heat and tissue metamor- 
phosis, 357 

Meyer, Adolph Bernhard, digitalis on 
blood-vessels, 134; action of right and 
left vagus, 180 

ae _ phys. ‘chem. of, 355; globules of, 


Moret, St George, Lessons in Element- 
ary Anatomy, 321 

Mohlenfeld, peptones from fibrin, 352 

Moitissier, absorption of heat in incuba- 
tion, 185 

Mollusca, inhibition of heart’s action in, 
184 

Monkeys, fossil, in Italy, 171 

Monodon monoceros, dentition of, 75 

Moore, Dr J. W., translation of Prof. 
Drachmann’s report of congenital ab- 
sence of quadriceps ext. cruris, 310 

Moreau, innervation of vessels of rabbit’s 
ear, 347 

Morphia, antidote to atropia, 198 

Morphology of limbs, 334 

Moseley, H. N., preparation, of organ 
of Corti for microscopy, 170, of eyes 
of Insects, for ditto, 170; injection of 
Coleoptera, 185 

Mosso, reports on Scehiff’s gastric di- 
gestion, 352; researches on function 
of pancreas, 353 

Motion, and lecomotion, 329 

Motmots, and their affinities, 337 

Miiller, Max, Darwin’s Philosophy of 
Language, 339 

Miiller, Quinine in hemoglobin, 346 

Murie, Jas., genus Colius, 174; powder 
downs of Rhinechetus jubatus, 174; 
vegetable organisms in thorax of living 
birds, 174; Bornean Ape, 335; In- 
dian wild Dog, 335; Caaing Whale, 
336; form and structure of Manatee, 
336; Todus, Ceriornis satyra and 
Motmots, 337 

Muscle, see ‘* Muscular system” 

Muscular system, of Cheiroptera, 171, 
of Hycemoschus, 172, of Sarcophilus 
ursinus and Phascolarctos cinereus, 
172, of Globiocephalus melas, 336, 
of Liolepis Belli, 338; muscle, ciliary, 
in Birds, 170, trophic action of nerve- 


376 


centres on, alterations in after nerve- 
lesions, 178, effect of galvanic cur- 
rents on nerve and, 178, electrical 
stimulation of, 179; irritability of 
fibres of heart, 182; var. in, 360, 
304; smooth fibres of in human 
kidney, 171; congenital absence of 
quadriceps extensor of thigh, 310; 
sphincter, of bladder, 191; malforma- 
tion of leg and foot, 1565; observa- 
tions in myology, 166, 169, 334, 3483 
reports, anat., on, 168, 169, 327—- 
329, phys., on, 357 

Myrmecophaga jubata, vert. column of, 
172: 


N 


Nandinia binotata, 335 

Nasmyth’s membrane, 17! 

Nasse, H., formation of lymph, 354 

Nawrocki, sensory impressions in spinal 
cord, 177 

Nedsvetzki, minute moving particles in 
human blood, 347 

Nencki, antecedents of urea, 355 

Nerves, of touch and painful im- 
pressions, 344, phrenic, 330, tro- 
phic, 343, 344, of head and neck, 
94, of cornea, 331, histology and 
phys. of, 170, 3443; structure and 
termination of in bladder of frog, 
170; cutaneous, irritated by sulphuric 
acid, 177; movements in, 178; effect 
of galvanic currents in muscle and, 
178; electrical stimulation of muscle 
and, 179; action of various substances 
on; of submax. gland, 199; source 
of force, 251; numerical relation of 
fibres of, to fibres of muscle, 329, 
ditto, of glands, to gland-cells, 330; 
var. in muscles and, 304; changes in 
after section, 344; influence of, on 
absorption, 348; circulation in optic, 
344; chorda tympani, 342, 343; ac- 
tion of right and left vagus, 180; 
phys. of cerebro-spinal fluid, 340; in- 
nervation of, heart, 180, 347, of ocular 
movements, 344, of cesophagusand sto- 
mach of Frog, 350, of vessels of Rab- 
bit’s ears, 347; Diagrams of nerves of 
the human body, 166 ; reports on, anat. 
169, 170, 329—331, phys. 175—180, 

Nervous system, of Cryptobranchus and 
Lepidosiren, 169; brain, forms of, in 
Carnivora, 172, in Marsupialia, 172; 
cerebral hemisphere, hypertrophy of 
with atrophy of opposite side of body, 
257; relation of nerve centres of cir- 
culation to stomach, 347, to absorp 
tion, 349 


INDEX. 


Nitrogen, amount of in flesh, 357 
Noctiluca miliaris, 338 
Nudibranchs, develop. of, 338 
Nycticebus, 334 


oO 


Obersteiner, cause of sleep, 176 

Oellacher, J., on the ova of river Trout, 
174 

(Esophagus, see “‘ digestive system” 

O’Dea, Dr, on dreams, 339 

Odontornithes, 337 

Ozle, J. W., Hereditary transmission of 
structural peculiarities, 332 

Ollier, growth of bone, 357 

Onimus, on the vagus, 343 

Opium, 194; combined effects of opium 
alkaloids and chloroform, 194 

Oré, M., is strychnia antidote to chloral? 
197 

Orth, Dr, malformation in human feetus, 
332 

Osseous system, of, young Balznoptera 
musculus, 172, Risso’s Dolphin, 173, 
Berardius, 335, Dugong and Mana- 
tee, 336, Todus, 337, Sphargis, 337; 
Cranium of anthropoid Apes, 171, 
334, 335, Baleen Whales, 173, Sal- 
mon, 174, Grey Seal (edentulous) 
273, Sparrow, Tit and Turdus (de- 

~ velop.) 337; cartilaginous auditory 
bulla of Paradoxure, 335; vertebral 
column of Myrmecophaga jubata, 
172; vertebrae, cerv. (and articula- 
tions) in Fin-Whales, 1, caudal, in 
Primates, 334; sternum of Chelonians, 
337; carpus of Sloths, 255; shoulder- 
girdle, homologies of, in Dipnoans, 
&e., 338; anat. reports on morpho- 
logy of limbs, 334; duplicity in 
hands and feet, 332; malformation of 
leg and foct, 156; remains of Elk 
and Megaceros hibernicus, 336; bones 
of sub-fossil Whale in Cornwall, 173 ; 
anat. reports on, 167, 168, 326, 327 

Otion Stimpsoni, 335 

Ova, development of, of river Trout, 
174; fecundation and development of, 
in Rabbit, 332; anat. reports on, 
332, 333 

Ovary, anat. reports on, 332, 333 

Owen, Richard, on the genus Phasco- 
lomys, 172 

Owl, ciliary muscle in Eagle, 170 

Oxygen, diffusion of, seat and laws of 
oxidation, 347 


P 


Paalzow, von, on cutaneous respiration, 
188 


INDEX. 


Packard, A. §., develop. of Limulus 
Polyphemus, Diplax Perithemis, &c., 
338 

Papillon, phys. and anti-fermentative 
action of sodium silicate, 359; flowers 
and fevers, 359 

Paradoxure, two-spotted, 335 

Parasites, on cetacea of the N.W. coast 
of America, 335 

Parker, W. K., skull of Salmon, 174; 
skull of Tit and Sparrow and of genus 
Turdus, 337 

Parotid gland, effects of stimuli on se- 
eretion of, 161 

Pancreas, see “ Digestive system” 

Paulus, perception in skin of lower 
limb, 180 

Pawlinoff, sources of uric acid, 355 

Pepsin, formation of in stomach, 186; 
peptic action of pyloric glands, 35r1, 
on blood-fibrin, 352 

Periosteum, formation of bone frem, 357 

Perrin, J. Beswick, var. in muscles, 

27 

Be earadon, suppression of, 188, 189 

Petienkofer, regressive metamorphosis 
during flesh diet, 350 

Pettigrew, J. B., structure and function 
of placenta, 333 

Pfliiger, diffusion of oxygen and laws 
&e. of oxidation, 347 

Phenicopterus antiquorum, eye of, 170 

Phaneropleuron Andersoni, 335 

Phascolarctos cinereus, myolozy of, 172 

Phascolomys, the genus, 172; p. lati- 
fions, 337 

Philipeaux, formation of bone from peri- 
osteum, 357 

Phonation, fixation of arytenoid earti- 
lages during, Igo 

Phosphates, difference of various earthy 
in formation of bone, 357 

Phosphoric acid, see ‘“‘ Acid” 

Phrenic nerve, 330 

Phyliirhée, anat. of, 338 

Physostigma, atropia an antidote to, 
198; p-, an antidote to strychnia, 
198 

Eaeacat bile, in urine, 165 ; oxidation 
products of bile, 352; source of in 
urine, 356 

Pinnepedia, comp. anat. 335 

Placenta, structure of human, 120, of 
Tamandua, 172, of Sloths, 302 

Plesiocetus garopli, 173 

Pneumograph, 348 

Podolinski, carbonic oxide hemoglobin, 

46 

Poisons ; the Kombé arrow, 139; re- 
ports on phys. action of medicinal 
and poisonous substances, 191, 200 ; 
anat. reports on, 333, 334 


VOL. VII. 


377 


‘Potassium, bromid., in snake-bites, 191; 


urea and alkaline chlorides in urine 
with p. iodide, 355 ; estimation of, in 
urine, 355 

Pouchet, G., vertebral column of Myr- 
mecophaga jubata, 172 

Power, Henry, on tropic nerves, 344; 
abstract of Mathieu’s and Urbain’s 
blood-gases, 347 ; 

Pozzi, S., on m. peroneus brevis, 168 

Prevost, chorda tympani traced to 
tongue, 343 

Preyer, myophysical researches, 357 

Pribram, estimating lime and_phos- 
phoric acid in serum, Igo 

Pritchard, Urban, rods of the cochlea, 
170; preparation of cochlea for mi- 
croscopy, 170 

Probalena du Buisii, 173 

Pseudopus pallisii, myology of, 169 

Pseudorca Grayi, distinguished 
Globio-cephalus Grayi, 173, 336 

Psychology, principles of, 339 

Pye-Smith, Dr., changes in nerves after 
section, 344 

Pyrosoma, anat. of, 338 


Q 


Quadrumana, comp. anat. reports on, 
334) 335 

Quincke, H., phys. of the cerebro-spinal 
fluid, 340 

Quinia, arrests movements of white blood- 
corpuscles, 195, 196 


from 


R 


Rabbit, hybrid of Hare and, 172; cause 
of retardation of pulse on closure of 
nostrils of, 283; innervation of ves- 
sels of ears of, 347 

Rabuteau, Dr, cyanates not poisonous, 
193; Opium, 194; combined effects 
of opium alkaloids and chloroform, 
194; phys. effects of urea introduced 
into organism, 355; phys. and anti- 
fermentative action of sodium silicate, 
359; flowers and fevers, 359 

Ralfe, Chas. Hy. Outlines of Physiolo- 
gical Chemistry, 318 

Ransome, Dr A., respiratory move- 
ments in Man, 348 

Ranvier, Louis, histology and physi- 
ology of the nerves, 170 © 

Reinhardt, J., on Pseudorea Grayi, 336 

Reptilia, comp. auat. reports en, 337, 
338 

Respiratory system ; respiration, cuta- 
neous, 188, pulmonary, 347; pulse, 
regulating frequency of, 219; cause 
of retardation of, on closure of nes- 


25 


trils of rabbit, 283; phonation, fixa- 
tion of arytenoid cartilages during, 
190; lung, nutrition of im health and 
phthisis, 357; phys. reports on, 347, 
348 i 
Retina, circulation in and optic nerve, 
344; association of the two, 344 
Retzius, G., anat. of nervous system, 


Reymond, Du Bois, current in the gas- 
trocnemius of frog, 1793; use of the 
round compensator, 179 

Mthea Americana, eye of, 170 

Rhinochetus jubatus, powder downs of, 
174 

Riegel, regulation of temp. in warm- 
blooded animals, 357 

Ringer, Dr Sydney, Belledonna, 197 

Ritter, changes in secretions under in- 
finence of certain agents 199 ; biliary 
calculi, 352 

Rivington, Walter, valves in renal veins, 
163 


Roberts, on Bastian’s experiments, 358 


Robinski, anat., phys. and path. of 
crystalline lens, 171 
Rohrig, cutaneous respiration, 188; 


cutaneous absorption, 188 

Rosenberg, Alex., osseous development 
of limbs, 334 

Rosenthal, M., changes in electrical ex- 
citability in muscles after death, 191 

Rosenthal, J., regulation of temp. in 
warm blooded animals, 357 

Roth, M., duodenal diverticula, 332 

Lubidium, salts of, introduced directly 
into blood, 204 

Rutherford, Prof. W., cause of retarda- 
tion of pulse on closure of nostrils of 
rabbit, 283; report on physiology, 
3397359 


S 


Sachs, C., structure of striped muscular 
fibre, 329 

Salkowsky, potassium, estimation of in 
urine, 355; urea and alkaline chlo- 
rides, in urine with iodide of, 355; 
comp. of cardiac muscle, 357; reac- 
tion of cholesterine with sulphuric 
acid, 359 

Salmon, develop. and structure of skull 
of, 174 

Samelssohn, innervation of ocular move- 
ments, 344 

Sanders, Alfred, myology of Liolepis 
Belli, 338 

Sanderson, Dr Burdon, Beginnings of 
Life, 358 

Sanson, A., hybrid of Hare and Rabbit, 
172 


INDEX. 


Sarcophilus ursinus, myology of, 172 

Scammon, C. M., Balzenoptera David- 
soni, 336 

Schafer, E. A., muscular fibre of water- 
beetle, 329 

Schiefferdecker, trophic nerves, 343 

Schiff, on conduction in spinal cord, 
341; innervation of heart, 347; blood 
pressure in resp., 347; artificial resp., 
effects, on circulation, 348, on con- 
cussion and compression, 348; gastric 
digestion 351 ; extirpation of pancreas 
in dog, 353 

Schenk, chlorine in the organism, 350; 
nitrogen in flesh, 357 

Schmidt, Alex., coagulation of fibrin, 


345 

Schlagdenhauffen, M., 
muscles, 16g 

Schmicdeberg, innervation of heart, 180 

Schobl, J., “termination of nerves in 
tactile hairs, 330 

Schultzen, antecedents of urea, 355 

Schwalbe, membrane of milk-ylobules, 
355 

Seal, grey, edentulous skull of, 273; on 
the hooded, 335 

Secretion, of parotid gland, 161, of 
submax. gland, 186; formation, of 
pepsin in stomach, 186, of glycogen 
in liver, 187; effects upon, of certain 
agents that modify blood-globules, 
19); peptic action of pyloric glands, 
351; action of pepsin on blood-fibrin, 
352; phys. of human bile, 352; phys. 
chem. of milk, 355; phys. report on 
urine, 355, 356; on lachrymation, 356 

Segoud, L. A., reptiles and batrachians, 
338 

Senator, production of heat and tissue 
metamorphosis, 357 

Setschenow, nerves during rapid irrita- 
tion, 344 

Shark, Greenland, visceral anat. of, 233 

Siebert, Dr, on apomorphia, 194 

Siemens, measuring temp. by elect., 357 

Sight, in birds, 344 

Silver, salts of, introduced directly into 
blood, 206 

pps Th., abnormal parietal foramina, 
107 

Sipunculus, anat. of, 338 

Sirenia, comp. anat. reports on, 336 

Skin, see ‘‘Integumentary system” 

Sleep, cause of, 176; causation of, 321, 


mechanism of 


339 

Sloths, carpus of, 255; placentation of, 
393 

Smee, A. H., physical nature of coagu- 
lation of blood, 210 

Smith,J. Alex., remains of Elk and Me- 
gaceros hibernieus, 336 


INDEX. 


Soborow, lime in urine, 355 

Socoloff, suppression of perspiration, 183 

Sodium, chloride, 350; silicate, phys. 
and anti-fermentative action of, 359 

Solowieff, changes in liver from gradual 
closure of v. porta, 187 

Sonnenschein, new test for blood, 346 

Soxlet, phys. chem. of milk, 355 

Spalding, D. A., Spencer’s Psychology, 
339; Instinct, 340 

Sparrow, development of skull of, 337 

Spedl, Anton, phrenic nerve, 330 

Spencer, Herbert, principles of psycho- 
logy, 339 

Sphargis, skeleton of, 337 

Sphygmography, 98 

Spinal-cord, sensory impressions in, 177 ; 
insensibility of, 177; phys. of the ce- 
rebro-spinal fluid, 340; phys. reports 
on, 341, 342 

Steinberg, absolute quantity of blood, 
347 

Sternaspis, anat. of, 338 

Stieda, Ludwig, formation of bone, 326 

Stokvis, oxidation products of bile-pig- 
ments, 352; identity of Choletilin and 
Urobilin, 356 

Stomach, see ‘‘ Digestive system” 

Stoney, P. Butler, stimuli on secretion 
of parotid gland, 161 

Strassburg, topography of gaseous ten- 
sions, 347 

Stricker, cardiac temp., 358 

Strophanthus hispidus, the poison of, 


139 

Struthers, S., Prof., cervical vertebrze 
and their articulations in Fin- Whales, 
I; processus supracondyloideus hu- 
meri, 326 

Strychnia, carbonic acid an antidote to, 
196; physostigma, an antidote to, 
198; s. an antidote to chloral, 197 

Sulphur, determination of, of bile, 252 

Sulphuric acid, see ‘‘ Acid” 

Synovial membranes, 329 


4t 


Tamandua, placenta in, 172 

Taylor, F., muscular var., 327 ; arterial 
var., 331 

Teeth, anat. reports on, 171 

Temperature, exercise on bodily, 106; 
phys. reports on, 185, 186, 357, 358 

Tension, topography of gaseous, 347 

Tephrylometer, 169 

Terebella nebulosa, develop. of, 338 

Terebratula, anat. of, 338 


Tergast, P., numerical relations of nerve- © 


fibres to fibres of muscle supplied, 329 
Thallium, salts of, introduced directly 
into blood, 203 


379 


Thiry, fistula of, 187 
Tiegel, ferment-action of blood, 187 
Tissue, metabolism, 80; metamorphosis 


of, 357 

Tit, develop. of skull of, 337 

Todus, osteology of, 337 

Tomes, C. T., on Nasmyth’s membrane, 
171 

Topography, of sense of colour, 3443 
of gaseous tensions, 347 

Traquair, R. H., Phaneropleuron An- 
dersoni and Uronemus lobatus, 338 

Tripier, action of right and left vagus, 
180 

Troglodytes Schweinfurthii, 334 

Trotter, C., review of Haughton’s Ani- 
mal Mechanics, 312 

Trout, ova of river, 174 

Tubes of Weber, 338 

Tuke, Dr J. Batty, hypertrophy of 
cerebral hemisphere with atrophy of 
opposite side of body, 257 

Turdus, develop. of skull in genus, 337 

Turner, Prof., dentition of Narwhal, 
75; placenta, human, 120, of Sloths, 
302, Cavernous sinus system of, 333; 
report on anat., 167, 326; Ziphius 
Cavirostris in British seas, 173; vis- 
ceral anat. of Greenland Shark, 233; 
claw at end of Feline’s tails, 271; 
edentulous skull of Grey Seal, 273; 
hiatus in m. pect. major, 327; horse- 
shoe kidney, 331 

Tursiops Gillii, 335 

Tweedy, John, absence of portion of m. 
pect. major and the whole of m. pect. 
minor, 327 


U 


Urbain, blood-gases, 346 

Urea, phys. effects of, introduced into 
organism, 355; estimation of, and 
alkaline chlorides, in urine with po- 
tassium iodide, 355; new method of 
determining, 355; antecedents of, 
355 

Unic acid, see ‘‘ Acid” 

Urine, bile pigment in, 165; phys. re- 
ports on, 355, 35 

Urobilin, identity of choletilin and, 
356 

Uromastix spinipes, myology of, 199 

Uronemus lobatus, 338 

Uterus, see ‘‘ Genito-urinary organs” 


Vv 


Vagus, action of, 180, 343; influence 
of on convulsions, 343 
Valves, in renal veins, 163 


Variations, in, human crania, 326, 


350 


postero-superior angle of scapula, 326, 
human muscles, 168, 304, 327, 360, 
arteries, 171, 300, 331, nerves, 304, 
bladder, 332; supernumerary carpal 
bones, 167 ; abnormal width of parie- 
tal foramina, 167; processus supra 
condyloideus humeri, 236; ‘‘defor- 
mation Toulousaine” of skull, 167; 
horse-shoe kidney, 331; duodenal 
diverticula, 332; hypospadia with 
cleft scrotum, 332; congenital hyper- 
trophy of left hand, 332; duplicity in 
hands and feet, 332; polydactylism in 
horse, 332; malformation, of leg and 
foot, 156, in human feetus, 332, in 
mesentery, 332 ; hereditary transmis- 
sion of structural peculiarities, 332 

Vascular system, see ‘‘ Circulatory” 

Venous system, see ‘‘ Circulatory” 

Vertebrate animals, disposition of mus- 
cles in, 169 

Vibriones, temp. destructive to, 358 

Virus, septic, 359 

Viscera, see ‘‘ Digestive system,” also 
“* Genito-urinary” 

Voit, nutritive value of Gelatine, 350; 
regressive metamorphosis during flesh 
diet, 350; article on food, 350 

Volkmann, A. W., turning motion of 
human body, 329; relations of elas- 
ticity to muscular activity, 357 

Vomiting, phys. of, 186 

Vulpian, trophic action of nerve centres 
on muscular tissue; alterations in 
muscles after nerve lesions, 178; 
chorda tympani, 342; trophic nerves, 
343; septic virus, 359 

Vulture, ciliary muscle in Egyptian, 
170 


W 


Wagstaffe, W. W., malformation of 
leg and foot, 156 

Walker, Robt., hooded seal, 335; Am- 
blypterus Anconozchmodos, 338 


CAMBRIDGE : PRINTED BY C. J. CLAY, M.A., AL THE UNIVERSITY PRESS. 


INDEX. 


Water, effect upon urine after injection 
of, into blood, 355 
Watson, M., Anat. of Indian Elephant, 


60 

Weil, C., fecundation and develop. of 
ovum in rabbit, 332 

Weiske-Proskau, differences of various 
earthy phosphates in formation of 
bone, 357 

Whales, Fin- (see also “‘ Baleenoptera”), 
cervical vertebree of, and their articu- 
lations 1, 173; Baleen, 173; sub- 
fossil, in Cornwall, 173; fossil, from 
Antwerp, 173; New Zealand Bottle- 
nose, 173; Sperm, in tropical Indian 
ocean, 335, stranding of, on coasts of 
France, 173 

Wilder, B. G., brain relative to mental 
characteristics in dog, 331 ; intermem- 
bral homologies, 334 

Willy, electrical stimulation of nerve 
and muscle, 179 

Winkler, F. N., placental sinus system, 


333 

Winternitz, regulation of temp., 358 

Wittich, von, gastric digestion, 331 ; 
pepsin on blood-fibrin, 332; phys. of 
human bile, 352 

Wolfermann, H., architecture of bone, 


32 
Wolffberg, tension of blood-gases, 347; 


pulmonary respiration, 347 
Wolski, insensibility of spinal cord, 177 


BG 
Yeo, Burney, absence of m. pect. major, 
27 
Yvon, new method of determining urea, 
355 
Z 


Ziphius cavirostris, in British seas, 173 
Zuntz, carbonic oxide hemoglobin, 346 


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