^ p ^

i\^

THE

JOURNAL

OF THE

COLLEGE OF SCIENCE,

IMPERIAL UNIVERSITY,

jj^:pa.n.

VOL. VI.

-•—

# ® :;'c ^ fP If

PUBLISHED BY THE UNIVEESITY.

TOKYO, JAPAN.

1894.

MEIJI XXVII.

2J2^

CONTENTS.

Determination of the Temperature of Steam arising from Boiling
Salt Solutions. By J. Sakukai, F.C.S., Professor of Chemistry,
Science College, Imperial University. flVith Plate I.J 1

Notes on an observation by Gerlach of the Boiling Point of a Solu-
tion of Glauber's Salt. By J- Sakurai, F.C.S. , Professor of Chem-
istry, Science College, Imperial University 21

Modification of Beckmann's Boiling Method of determining Mole-
cular Weights of Substances in Solution. By J. Sakurat, F.C.S.,

Professor of Chemistry, Science College, Imperial University 23

A Simple Experiment in Chemical Kinetics. By K. Ikeda, Biqakushi... 43

ImidoSUlphonates. By E. Divers, M.D., F.E.S., Prof., and T. Haga, F.C.S.,

Asst. Prof., Imperial University 49

On the Anatomy of Magnoliaceae. By S. Matsuda, Science College, Im-
perial University. (With Plates II.-V. J 115

Researches on the Multiplication of Elliptic Functions. By Pt. Fuji-

SAWA, Professor of Mathematics, Imperial University 151

On the Process of Gastrulation in Chelonia. (Contributions to the
Embryology of Eeptilia, IV.) By K. Mitsukuri, Ph. D., Fdqakuhakitshi,
Professor of Zoology, College or ^ie^ice^ Imperial University. (Wit It
Plates VI.—VHI.J 227

Note on the Eyes of Cardium Muticum Beeve. By K. Kishinouye,
Ei</ah-iishi, Zoologist to the Department of Agriculture and Commerce.
(With Plate IX. J 279

Note on the Ccelomic Cavity of the Spider. By K. Kishinouye,
Eigaliiishi, Zoologist to the Department of Agriculture and Commerce.
(With Plate X.J 287

Studies of Reproductive Elements. II. Noctiluca miliaris, Sur.; its

Division and Spore-formation. By C. Ishikawa, Pli. D., Pùijah-ushi,
Bifiakiihakiishi, Professor of Zoology, Agricultural College, Imperial
University. fU'iih Plates XL-XIV.J 29T

CONTEXTS.

On the S aro- Amniotic Connection and the Foetal Membranes in the

Chick. By S. HiROTA. lU(jaliiishi, Zoological Institute, Imperial
University. (With Plates XV.-XVII.J

On a New Human Tape-Worm (Bothriocephalus sp.) By I. Ijiha

J'li. IK, liiii(i/:iis/ti, lluiakuhakitsld, Prof, of Zoology, Imperial University
and T. KuKiMOTO, L/akushi, Prof, in the Fifth Higher Middle School
Nagasaki. (With Plate XVIII. J

837

371

TRIXTED AT I'nit TôKIÔ-SEISHIPUSSnA, TCKlS.

Uy

Determination of the Temperature of Steam
arising from Boiling Salt Solutions.

Joji Sakurai, F. C. S.

Professor of Cliomlstry, Imperial University.

Introduction.

Although it is well kQO\Yn that the boiling point of a salt solution
is .'ilwnys higher than that of water under the same pressure, tlie tem-
perature of steam arising from sncli a solution has l)een the siihject of
much dispute, and, even at present, different (opinions seem to prevail
among cliemists and pliysicists.

According to one set of observers, the temperature of steam arising
from a boiling salt solution is the same or nearly the same as that of
the solution : Gay-Lussac and Faraday, and, subseqnentl}^, AVuhner,
Mnofniis, and lYaimdler, rank amono- these. According' to the other,
it is the same or nearly the same as that coming from pure water
boiling imder the same pressure, whatever may be the temperature of
the boiling sohition : tliis is the view held by Rudberg and, to a
certain extent, by ]\[iiller. Experimental ]n'oofs are adduced on both
sides, but these, as I shall show, are none of them conclusive, and
establish, T think, neither the one view nor the other.

The determination of the real temperature of steam arising from
a boiling salt solution presents many experimental difficulties ; for,
besides othei's to l)e mentioned later on, steam condenses upon the
bulb of the thermometer and also upon the sides of the vessel al)ove

2 J. SAKÜT^M: DETETîMTXATTOX OF THE TEMPEILVTÜRE

the vapour, unless sper'i;il precautions l^e taken, so that the tempera-
ture indicated will not be liigher tlian 100°. Tf, on the other hand,
the walls of the containing- vessel be maintained at a higher tempera-
tiu-e, the thermometer will be exposed to the heat radiated from them,
and will indicate a higher temperature.

For some time past, I have been engaged in an experimental
investigation of the determination of the temperature of steam escaping
from boiling salt solutions, and have devised a method which is, as
far as I can see, free from such objections. V>y tliis means, it can be
readily shown that the temperature of the steam escaping from the
boiling salt solution is exactly the same as that of the solution itself.

I'efore proceeding, however, to a description of my own experi-
ments, I shall briefly refer to the history of tliis question, discussing
the metliods employed and the results obtained by previous workers.

Historical Summary and Short Critical Review of the Work

already done.

Faraday, as long ago as 1822, published the results of his experi-
ments upon this question in the Annales de Chimie et de FJujsiquc for
that year. Fie found that Avhen the l)ulb of a thermometer was
sprinkled over with a salt and then introduced into steam coming out
of boiling water, the th(;rmometer showed a temperature higher than
100°, especially Avhen it was held horizontally, so as to prevent the
salt from ])eing washed awav too quickh'. Still higher temperatures
were observed l)y him when, the experiment being otherwise the same,
the thermometer l)ulb was wrapped up in a linen or woollen cloth.
From these experiments, Faraday concluded that since a salt solution
was heated up to its boiling point by the action of steam at 100°
upon the salt, therefore the steam generated from a boiling salt solu-
tion had only the temperature of 100°.

01<' STEAM AIMSING FROM BOILING SALT SOLUTIONS. 3

riay-Lussac, as the editor of the Frcncli journal, appended a note
to tliis paper, in Avhich he first pointed out that facts similar to those
observed by Ftiraday had long been known in France, namely, that
when steam from boiling pn re water was passed into a cold concen-
trated solution (jf a salt, the latter could be heated up nearly to its
boiling point. Then, witli regard to Faraday's view that steam
generated from a boiling salt solution has only the temperature of
100^5 Gay-Lussac remarked : '* Sans irivoquer ici le secours de la
théorie, nous pouvons afiirmcr, d'après le iémoignage irrécusable de
l'expérience, que la temperature de la vapeur fournie par un li(|uide
quelconque, sons une pression quelconque, est exactement celle de la
couclie liquide immédiatement en contact avec la vapeur."

Faraday then undertook more researches on this <|uestion, and in
the Quarlerly Journal of Science for 1823, he published the results of
his experiments, and stated that he had proved Gay-Lussac's asser-
tions to be correct, but that he had been astonished at the difficulty
of obtaining definite results. Only when he used a double-walled
vessel, which contained the experimental solution botli between the
Avails and also abfjAC them, only when he heated the thermometer
previously to a temperature higher than that of the boiling solution,
and only after repeated observations, had he been able to convince
himself that no anomaly existed in this j'henomenon. These experi-
ments of Faraday will be criticised presently along witli those of
MaQ:nus.

Ivudberg (Ann. Chcm. Vhijs. [Vogi].\ 34, 257), in 1835, pul^Iishcd
the results of a long series of observations on the temperature of
steam evohed from difierent solutions, boiling under different pres-
sures, and pointed out that it is always the same as that of steam
arising from pure water l)oiling under the same pressure. Some of
his numbers are quoted below.

J. SAKUllAI: DETERMIN' ATIOX OF THE TEMPERATUllE
Temperature of steaui.

'ressure in umi.

Water.

Salt solutior.

Salt.

701-54

100-06°

100-07°

C<XO,),

763-37

1()0-12

100-13

5?

7G9-Gi

l()0-35

100-34

5?

7G9-17

] 00-33

100-3G

KXO,

763-20

1U0--19

100-50

5 J

Tlie experiments of Kudberg are always regarded as coiiclnsixe
evidence tliat tlie temperature of steam issuing from a boiling salt
solution is only 100°; but as tliey were carried out in the ordinary
way without proper precautions against the steam cooling before
reaching the thermometer, we must regard his results as unsatis-
factory, in presence of the fact that with such precautions tlie tem-
pera tm-e proves to he higher.

AViillner (Ann. Chan. Plii/s. [Pogg.] 110, 387), from determina-
tions of the tension of steam arising from boiling salt solutions, pointed
out that such steam, having less tension than that evolved from pure
water at the same temperature, must be iion-saturated, and that
theoretically, therefore, it could not have the temperatine of 100"".

Magnus (Ann. Clem. VJnjs. [I'ogj-] 112, 408) is generally regarded
as having experimentally proved that steam from a boiling salt solu-
tion has a temperature equal, or nearly equal, to that of the solution
itself. It will, therefore, be interesting to know exactly the nature of
his evidence, and I propose to refer to his pa])cr somewhat in detail.
In tlie first part, ^lagnus menti«ms that IJudorff, who was working
in his lal)orat(<ry, had devised two methods of ascertaining the real
temperature of steam coming out of a salt solution, the results show-
ing that the temperature of the steam is above 100°.

One of these methods consists in dipping the thermometer into a
boiling salt solution and, when it has attained the lemperature of the

Ui' STEAM AKlÖlNO) FROM BOILIXU SALT SOLUTIONS. 'i

o

latter, in lioklino* it in tlie sterim over the solution. This method of
ascertaining the temperature of tsteam is, as ^lagnus liimself allows,
by no means convincing ; fur, as already observed by Faraday and
others, when such a. thermometer is held in the steam arisini»- from
boihng pure water, temperatures higher than 100° are always in-
dicated.

The other method consists in previously heating the bulb of a
thermometer to a temperature higher than that of the boiling solu-
tion, and then introducting it into the steam. It is said that, in this
wav, temperatures a])proaching the boiling point of the solution are
indicated. I have, however, repeatedly tried this method, and liave
fully convinced myself that if sufficient care is taken to protect the
bulb of the thermometer against the splashing up of the hot solution,
the temperature rapidly falls to 100°.

After referring to these two methods proposed by liudortf, and
after giving a good liistorical summary of the subject, to which I am
greatl}'^ indebted for this part of my paper, Magnus goes on to describe
his own method and the results obtained by its means. His apparatus
consists of a double-walled metallic vessel, filled to a convenient lieight,
both within and without the inner walls, with the experimental
solution. Steam, generated by heating the solution from bel<jw, rises
in the inner as well as in the outer chamber, and escapes by means of
a lateral tube, also metallic and provided with two holes, one of
whicli opens into the inner clinmbcr and the otlier into the outer
chamber. The object <jf li:n iug iloublc chambers is to kec]) the inrier
one surrounded by hot steam, so as to prevent loss oi heat by radia-
tion. The temperature of the steam is indicated by means of a
thermometer held horizontally and inserted in the lateral tulje, its bullj
Ijcing placed in the centre of the inner chamber. The following are
the results oljtained Ijv jMau'nus with a solution of calcium chloride.

J. SAKUliAi: DETEilMlNATION OF THE 'J'EMPERATUllE

Temperature

Of the solution.

Of the steam.

Difference.

107-0°

105-25°

1-75°

107-5

105-5

2-00

108-0

105-8

2-2

109-2

lOG-5

2-7

llO'O

107-0

. 30

111-0

107-G

3-^

112-0

108-1

3-9

113-0

108-8

4-2

lM-0

110-0

4-0

115-0

110-9

4-1

116-0

111-2

4-8

Mao-nus confesses that these numbers have no absokite value: but,
at the same time, he regards it as proved that the temperature of the
steam is nearly the same as that of the solution, the difference being-
due to the unavoidable loss of heat by radiation. Such is the con-
clusion Magnus has draAvn from his experimental results, and this
seems to have lieen accepted by chemists and ])hysicists as being
convincing. Even those who had held the opposite view seem never
to have criticised the method Magnus employed, or the results lie
obtained.

It seems to me, however, that the conclusion he draws from the
results of the experiments is open to criticism. As the double-walled
vessel, which he employed, was made entirely of metal, its walls must
have been heated up by conduction nearly to the temperature of the
boilino- solution it contained, and the steam enclosed by such walls
must have had nearly the temperature of the latter ; that is, a tem-
perature more or less approaching that of the solution, and that
quite independently of its temperature wlien generated. His num-

OF STEAM ARTSTN«; rnOI\r TIOTLTNT! SALT SOLUTIONS. 7

]")ors in the seoonrl column of the talile may have indicaterl, therefore,
not so miK'li the temjierature of the natural steam as that of the sur-
rounding walls and of tlie steam superlieated l)y them. Strange to
say, Magnus seem:^ to have heen aware of this element of uncertainty
in his results, and yet to have disregarded it. He writes : — " Soviel
ist indess durch diese Versuche bewiesen, dass die Dämpfe, welche
auskochenden Salzlösungen aufsteigen, eine höhere Temperatur als
100° lial^en, und eine um so höhere, je höher die Temperatur der
kochenden Lösung ist. Dass sie aber dieselbe Temperatur wie diese
Di)Suno' haben, ist mir nicht 2:elun2:en nachzuweisen, und ich zweifle
dass diess möglich sein wird. Denn wenn die Wände des Dampf-
raumes auf der Temperatur der kochenden Lösung erhalten werden,
so ist man, wie schon oben bemerkt, nicht sicher, dass nicht die
Erwärmung von diesen Wänden hervorgebracht ist, haben dagegen
die Wände eine niedrigere Temperatur, so wird aucli die Temperatur
des Thermometers niedriofer ausfallen."

What then, it may pertinently l)e asked, is the value of his ex-
})erimental results? It seems to me that Magnus erred in having the
Avails of his vessel of a highly conducting material, so that the thermo-
meter would indicate the temperature of these walls, derived as it was
from the solution and even from the heat of the lamp, and then to
regard the temparature observed as ])eiHg that of the steam. In fact,
it can be readilv sliown that l)y keeping the walls of a vessel at a
temperature Idgher tlian 100°, say at 110°, the steam issuing from
water itself indicates a temperature almost equal to that of the walls.

The results obtained by IVIagnus, which have been regarded as
tlie most weighty and cc^nclusivc experimental proof of the view that
the steam arisino- from a boilin"" salt solution has the same or nearlv
the same temperature as the latter, thus appear to me t«^ be valueless,
and the same remark applies to Faraday's later experiments already

g. J. SATvURAT : DETERMIXATIOX OP THE TEMPERATURE

referred to, in which he employed a double-walled vessel containing
the experimental salt solution in both chambers.

Müller {Berichte d. Deutsch. Chem. Ges. 9, 1629), apparently not
acquainted witli I'nn'.day'.s earlier experiments, made observations
similar to his, and came to the same conclusion as Faraday first did,
nnmelv, that the steam g-enerated from a boiling salt solution hns only
llie temperature of 100°. He regards this view as being confinned by
the following considerations.

1st. The solution begins to boil (that is, steam bubbles freely ri>e
to the surface and there burst) below its proper boiling point, and the
temperature then slowly rises. The steam formed at first at 100°
acts on the solution in the same manner as that passed from without,
and heats it up to the boiling point.

2nd. Higher temperatures are indicated Ijy salt solutions when
boiling gentl}^ than when boiling violently ; in the latter case, m<ire
stenm bubbles comino: in contact with tlie bulb of the thermometer.
Similarly, a rise of temperature is observed on removing the flame
after violent ebullition.

Mliller's account of the first effect of heating a salt solution is so
remarkable that I feel it incumbent to adhere closely to his words.
He says : " A solution of calcium chloride whose boiling point is
126°, for example, is already completely boiling at 110°, and the
temperature then rises to 126° in about half a minute" (he. cif.,
1631). I have repeatedly tried similar experiments, but have never
been able to confirm the statement that tiie solution, whose proper
boiling point is 12o°, is completelij boiling at 110°, though it is per-
fectly true that the solution begins to boil at about this temperature.
1 confess, therefore, tliat it is a relief to me to be able to dispense
with any discussion of ^liiller's interpretation of these phenomena,
for whatever their sio-nification mav be, the fact that the observed

OP STEAM AllIÖlXG PllOM BOILIXG SALT 80LUTIUX«. 9

phenomena of incipient ebullition of a «ult solution differ in no
respect from those of ])urc water, except lliat of higher temperatures,
deprives them of all vnlue for throwing light upon the question at
i.ssuc. A similnr remnrk may Ije made with regard to Midler's other
observations, namely, th()se which I'elate to the difference of tempera-
ture in the boihno- solution, accordinii' as it is freelv evolvini"' tl e
steam or not. It is a welhknown fact that a thermometer dipped
into a boiling liquid, say water, gives very irregular indications,
higher temperatures being always indicated wlien steani is less
freely escaping. In short, the facts observed by Midler are by ikj
means limited to salt solutions, and his aro-ument that the steam j.t
the moment of its formation has only the temperature of 100° cannot,
therefore, be regarded as in any degree satisfactory.

Wiilhier (Berichte d. Deutsch. Chcni. Ges. 10 256) wrote a note
on Midler's paper, pointing out thtit the fact that a solution of calcium
chloride may be heated up to its boiling point by the action of ordinary
steam had already been known to Gay-Lussac and observed by
Faraday, who on more careful experimental enquiry changed his first
opinion, and came to the conclusion that the steam escîiping from a
boiling salt solution must have the temperature of the latter. He also
points out again that such steam is not saturated, and that, therefore,
it could not have the temperature of 100°.

Pfaundler (Berichte d. Deutsch. Chem. Ges. 10, -163) objects to
Midler's conclusion on much the s-ame grounds as WiUlner, and fidly
supports the views of the latter. Further on, Pfaundler attempts to
explain- the fact observed by Magnus that the temperature of steam
arising from a boiling salt solution is not quite the same as that
of the latter ; this explanation, which is not easy to accept, is also not
needed, as the facts observed by Magnus, and upon which this ex-
planation is Ijasedj are, as already pointed out, by no means convin-

J_U J. ÖAKUKAI: DETEllMINATlOX Ui'^ THE TEMPÈRATUUE

cing. My own experiments prove, moreover, tliat there exi.sts no
difference between the temperature of the boiling solution and that of
the steam issuing from it.

The above smnmary should o-ive some idea of the vao'uenes., and
unsatisfactory character of tlie experimental evidence now existing
as re^'ards the temperature of tlie steam arising from a Ijoiling salt
solution. The following is a description of ]ny own experimented
methods, and the results oljtained by their means.

Experhnenlal lYlelhods.

For success in establishing the true temperature of the steam
escaping from boiling salt solutions, certain conditions must be
observed.

1. The thermometer must be kept clear of all contact with the
s(jlution, even the smallest dro])s thrown up Ijy ebullition, as other-
wise it is evident that the experiment hjses all chiiui to accuracy.

2. The elfect of the radiant cooling of the thermometer must either
be prevented, or rendered inappreciable in proportion to the heating
np by the stcimi. Ik^fore considering these alternative conditions, it
may be well to call attention to the familiar and striking evidence we
have that loss of heat by radiation from the bulb of the thermometer
does occur and cannot be neglected. Whenever distillation of water,
or any other liquid with no fixed matters dissolved in it, is going on,
the thermometer immersed in the vapour to record the " boiling
point " is seen to be always condensing some of the vapour, drops
falling from it into the boiling liquid.

The former alternative seems hardly practicable, or, at least, has
proved to be impracticable under various conditions in my hands.
Tlie latter alternative, that of overpowering tiie 1(JSs (jf heat by

or STEAM ARTSIXa FROTNT TIOTLTXG SALT SOLUTTOXS. H

radiation by rapid removal of the vapour in contact with the thermo-
meter, can be easily effected by the expedient of com])ining the in-
troduction of steam from without with the boiling of the solution by
the lamp, the combination lieing regulated by maintaining steady the
temperature marked by the tliennometer in tlie solution. Ebullition
alone should suffire for thi.-; purp(i;e, hut the practicnl difficulties in
the way render it insuffi(,'ieiit. Tlie boiling would liave to be tumul-
tuous to generate mucli vapour, and in a short time the solution
would become too concentrated for the experiment to be continued
with convenience. On the other hand, with due regulation of the
steam entering from without and the height of the flame, an abun-
dant supply of steam can be got without impediment. Moreover,
irregular boiling and bumping of the solution are hoth entirely pre-
vented when tlie operation is worked in this manner.

3. The walls of that part of the vessel which serves as the steam
chamber for the thermometer must lie sufficiently protected from
cooling externally, and yet, :>.t the same time, not be heated to the
teinperature of the steam. For if, as in Rudberg's experiments, the
former condition is not observed, so much of the steam is condensed
in keeping the walls at 100° that it is liardly possible to keep enough
passing over the thermomeler bulb ; while if the latter condition is
ignored, as in Magnus's experiments, the indications of the thermo-
meter may be illusory.

The arrangement which I adopted after several trials and modifica-
tions in order to meet these conditions was as follows. It consisted of
an elongated, round-l^ottomed flask, F, with a long neck ; this flask
contained the experimental snlt solution. The moutli of the flask
was provided with a cork, through which passed two thermometers,
a and h, and a tube .s for tlie escape of steam ; the thermometer a was
used to indicate the temperature of the steam, and the thermometer

12

T. S A KÜR AT: DETEliMTXATTOX OP THE TEMPEILATURE

/Mhnt of tlie. solution. The nock nnd n jKirf of tlic fbisk F wns cn-

f'io.sod in :i o-l:iss ovlinder, ;//, fixed :i1)ove Iw menns of a eork nnd
below 1)y nie:'. RH of an iii(]i;!-rn1)])er l)a7id. Through the eork of t lie
cylinder, and hy the side of the neek of the flask ¥, passed a third
thermometer, c, whose hnlb was kept at the same height as that of
the steam thermometer a. I'he body of the flask, finally, was pro-
vided Avith a hole in its side, through Avhich a glass tube,*/, somewhat
drawn ont at one of its ends and bent as shown in the figure, Avas
passed a,nd jointed to the flask by means of a short pieee of india-
rubber tubing slij^ping over it. This tube, whose drawn-out end
nearly reaehed the bottom of ihe flask, Avas used for supplying steam
from the boiler Tl. The gl;!ss eylinder j}" Avas connected on one hand
with another flask, II, and on the other with a condenser, K ; these
connections were made as before by means of shoi-t pieces of india-
ruliber tubing sli])])ed on over the glass tubes u and v res]iectively.
Tiie flask M conlained acclii^ acid somewhat dihiled witli water.

OF STEAi\r APJSTxri rnoi\r iiotltnt; salt solutions. 13

Tlie flrrsks F, (r, niid IT wore smpportorl on stnnrls, and F rested on n
piece of nsl)estos r:i]'dl)o:ird, liavinii; u hole in tlie centre, so that only
tlie l)ott<^in miulit ])v heated by tlie direct flame. The stoppered
funnel' m a.iid 7/ : erved to aipply the flask.'. (J and }I with water and
acetic acid respectively, wheri necessary, without dismounting the
apparatus. The funnel iii further served, at the end of an experiment,
to restore the equilibrium of pressure between the outer air and the
inside of the flask G, thus preventing the salt solution from being-
sucked into the latter. The whole arrarjgeinent is exceedingly simple,
and can be fitted u]) by any one with materials found in every chemi-
cal laboratory. The three thermometers a, />, arid c had, finally, been
carefully verified, including tlie exposure correction for a and e.

The salt solution, whose boiling point had been approximately
determined and known to be higher than that of the dilute acetic acid,
was introduced into the flask F by means of a long funnel, and the
inside of the neck of the flask then Aviped with a cloth, so that no
particle of the salt solutiori should be left adhering to it. Some
cotton-wool was loosely tied round the stem of the tliermometer h,
and below the bulb of the thermometer a ; this served as a very effec-
tive screen against splashing up of the solution on to the bulb of the
second thermometer a. A little cotton wool was also tied round the
stem of the thermometer a, and above its Indb ; this prevented any
water which might condense in the steam-issue tube s, from flowing
down the stem of this thermometer. The cork with its two thermo-
meters thus prepared was then fitted into the mouth of the flask F ;
this cork projected a little below that of the jacket, in order that there
miofht be no diance of steam condensino" within the neck of the flask.
The tubes u and t were covered with (^itton-wool in order to lessen
llie cooling action of the air on the acetic acid vapour and the steam
respectively.

]^4 T- SAKURAT: DETERMTNATTOX OF THE TEMPERATURE

The acetic acid was now made to boil in the flask H. Its vapour,
passing into the jacket jj, heated up the walls of the neck of the flask,
that is, of the steam chamber over the solution, then passed into the
condenser K, and collected in the receiver R. The flame under the
flask II wns so i-eo-ulated that the acetic ncid only sl(->w]y distilled ; in
this mnnnor, a stondy supply of v^ponr of almost constant temperature
could be readily mnintained in the jacket. The thermometer c rnpidly
rose to about 11 1° under the influence of the acetic acid vapour, whilst
the thermometer a more slowly rose, until ultimately it also showed
about the same temperature.

The salt solution was next hen ted to boiling. The water in the
flask Ct, which had been kept nearly boiling, was then also made to
boil regularly, and a rapid current of steam was passed into the salt
solution boilinof in the flask F. The thermometer a now beo-an to
rise above that in the jacket, until it indicated the same temperature
ns that of the solution. The flame under the boiling solution Avns so
refï'ulated that the temperature of the latter sliould either remain
constant or rise xery slowly. The results obtained in one exjieriment
with a solution of calcium chloride are shown below.

Température

of the

Diff.

(ir.)-(i)

Solution.
(II.)

Acetic acid vapour.
(III.)

'vonoo.

St.(^a.m.

(!.)-( ni.)

111-2°

112-5°

110-8°

1-3

0-4

111-7

112-5

110-9

0-8

0-8

112-2

112-6

111-1

0-4

1-1

112-5

1127

111-3

0-2

1-2

112-7

112-Î)

111-5

0-2

1-2

113-0

113-0

111-fi

00

1-4

113-1

113-2

111-8

0-1

1-3

113-3

113-3

111-9

0-0

1-4

OF ÖTEAM AKIÖINU rilÜM BOILIXG SALT SOLUTIONS. J^

The time occupied was about 20 minutes. The bulb of the steam
thermometer a remained perfectly clean ,• indeed, by w;ishin<>' it with
a little water and adding- a drop of silver nitrate solution to the wash-
ings, not a (race of cloudiness was produced, shoMing- that the
splashing of the solution on to it had been completely prevented.
In fact, the u])per portion of the cotton-woul remained perfectly clean
and dry.

For the following two experiments, still more dilute acetic acid
was employed. The results obtained with a solution of'sodiimi nitrate
are as follows : —

Temperature of tlie

Difference.

Ste:ini.
(I.)

106-0°

lOG-U

107-2

107-0

107-7

107-7

107-8

Solutiou.
(II.)

107-5°

Acetic acid vapovir.
(III.)

105-9°

(ii.)-{i.)
0-9

(I.)-(III-)
0-7

107-G

105-9

0-7

10

107-G

106-0

0-4

1-2

107-7

106-1

.0*2

1-4

107-8

106-2

0-1

1-5

107-8

106-2

0-1

1-5

107-8

106-3

0-0

1-5

The time occupied was again about 20 minutes. The following
are the observations made with a solution of potassium nitrate : —

j[g J. ttAKÜllAI: DETEllMINATlOX OF 'J'HË TËMPEllATUllE

Temperature of the

,-

Sokitiuu.

or.)
109-4°

Acetic acid vapour.

(in.)
107-8"

Difference.

IS team.

107-8°

(II. )-(!.)

1-G

(I.)-(IIl.)
0-0

los-;}

109-5

107-8

1-2

0-5

108-7

109-7

10<S-0

1-0

0-7

109-1

109-8

108-1

0-7 .

1-0

109-3

109-N

108-2

0-5

1-1

109-:)

109-9

108-2

0-4

1-3

1U9-8

110-0

108.2

0-2

1-«

110-0

110-2

108-3

0-2

1-7

110-2

110-:;

108-5

0-1

1-8

110-2

110-3

108-5

0-1

1-7

The time occujned was iibuut half an h<jur.

In the following experiment a Htronger solution of calcium
chloride was taken, and aniyl alcohol (containing- a little uf the lower
alcohols) was employed instead of acetic acid.

Teuii:)erature of tlie

Diflereuce.

Steam. Solution. Amyl alcohol vapour.

(I.) (II.)

127-5^ 128-3°

128-1 128-5

128-6 128-9

129-1 129-2

129-4 129-5

129-7 129-8

129-8 129-8

The time occupied was about 20 minutes.

The ex]: erimenîs above described pro\ e, beyond any possible doubt,
that ^/i<3 temperülare of tJie steam escap'uuj from a hoiling sail solution is

(III.)

126-5'

126-8

127-2

127-8

128-1

128-3

128-5

(ii.)-(i.)

(I.)-(III.)

0-8

1-0

0-4

1-3

0-3

1-4

0-1

1-3

0-1

1-3

0-1

1-4

0-0

1-3

OF STEAM ARISING FllOM BOILING SALT SOLUTIONS. IT"

exactly the same as that of the solution. This, I believe, is the first
occasion on which the above important fact has been experimentally
established. Philosophers had, indeed, asserted that the temperature
of such steam should be, or would Ije, the same as that of the solu-
tion, but without any experimental proof. Gay-Lussac's assertion,.
" the temperature of the vapour furnished by any liquid is exactly
the same as that of the liquid layer in immediate contact with the
vapour," is based upon the knowledge of the facts derived from the
study of simple liquids, such as water, alcohol, &c. ; for no direct
experiments had been made by him, or any other investigator before
him, upon the temperature of the steam arising from a boiling salt
solution which established the truth of that assertion. So far as
direct experimental evidence is concerned, he had, I think, no good
reason for extending his remarks to the case of a salt solution..
Faraday thought he had proved Gay-Lussac's assertion to be correct
in the case of salt solutions also, but upon evidence not at all con-
vincing. Magnus could not show that the temperature of the steam
is exactly the same as that of the solution. " That, however, it (the
steam) possesses the same temperature as the solution I cannot prove,,
and I doubt whether it can possibly be proved," he says. The grounds
for such doubt have now been remo\ed by the experiments above
described, the secret of the success lying in the fact that the walls of
the steam- chamhcr must he ahovi 100^, and yet below the temperature of
the solution, and that, at the same time, a sv^cient quantity of steam must
escape from the solution to ensure that these walls shall hare no material
cojling ef'ect vpon the steam. To meet this condition the quantity of
steam evolved from the boilin«- solution itself is not sufficient, and
hence the necessity of passing steam from without into the boiling
solution.

That the quantity of steam merely arising from a boiling salt

J 3 J- SAKURAT: DETERMINATION OF THE TEMPERATURE

solution is not sufficient to overcome the coolino* effect of the walls
will best be seen from the following numbers. These refer to a
solution of calcium chloride experimented upon in exactly the same
manner as before, but witliout introduction of steam from an external
source. The acetic acid was also more rapidly distilled.

Temperature

of the

Acetic acid vapour.
(III.)

Differenc

StCcam.
(I)

Solution.
(II.)

',e.

(II.)-(I.)

(III.)-(I-)

103-0"

111-9°

103-2°

8-9

0-2

103--1

114-4

103-5

11-0

0-1

103-7

115-2

103-7

11-5

0-0

104-1

117-0

104-2

13-0

0-1

104-5

118-4

104-5

13-9

0-0

104-8

120-5

105-1

15-7

0-3

104-9

122-6

105-3

17-7

0-4

105-1

125-2

105-5

20-1

0-4

In spite of the fact that the solution was boiling briskly with a
free and uninterrupted evolution of steam, and in spite of the additional
fact that the temperature of the boiling solution rose from 111*9° to
125-2°, the temperature of the steam only rose from 103° to 105*1°, and
never above that of the acetic acid A'apour in the jacket. The rise
that did occur is evidently due to the heating effect of the acetic acid
vapour ; for, in this case, the outer surface of the steam-chamber was
constantly wet, owing to the condensation of the acetic acid va])our
upon it. Acetic acid vapour, Jind not steain, was then acting as the
heater.

This result was so remarkable and apparently so conclusive, that
I was for some time induced to believe that the temperature of the
steam escaping from a solution of calcium cldoride boiling at the
temperature of even 125° is lower than 105^ and probably only 100°.

OF STEAM ARISING FROM BOILING SALT SOLUTIONS. X9

l»y slightly modifying the experiment, namely, by keeping the distil-
lation of the acetic acid at a very slow rate, while the solution was
kept boiling as briskly as possible, the steam-thermometer could be
made to indicate somewhat higher temperatures than the acetic acid
vapour, but far below the temperature of the solution. It was this
■<3bservation, however, that led me to try the introduction of stenm
into the solution from without.

y>y the introduction of steam into the boiling solution from
without, evaporation and condensation of steam in the solution can be
so readily and exactly counterbalanced, that its boiling temperature
may be maintained constant for any length of time and within a few
thousandths of a degree (_ entigrade. I am developing this part of my
observation f<3r the exact determination of the boiling points of vari(3us
solutions, and also for simplifying the determination of molecular
Aveights l)y tlie boiling method, which has, of late, been made tlie
subject of an extensive study by Beckmann.

In conclusion I wish to express my warmest thanks to my
colleague Dr. E. Divers, F. 11. S., for many valuable criticisms ;ind
suggestions from time to time while this investigation was in progress,
and also to Dr. C G. Knott and Professor K. Yamakawa fnv ihe
interest which they have takei: in my work.

Note on an observation by Gerlach of the Boiling
Point of a solution of Glauber's salt. *

by

Joji Sakurai, F. CS.

Professor of Clieiuistry, Imperial University.

A few years ago Dr. G. T. Gerlach (Zeit. anal. Cliem., 26? 113)
published a paper in which he mentions that steam, escaping' from a
boilino' solution of Glauber's Salt containini'' a crystalline maii'ma of
the anhydrous salt, sliows the temperature of 100°, Avhilst the liquid
is boiling at 82° or even at 72°. This observation appeared to me so
curious and so anomalous that I was induced to repeat his experi-
ments : the results, on the whole, confirmed Gerlach's observations as
to temperatures but, at tlie same time, deprived them of all exceptional
character. Gerlach describes his experiments in the following words :

" 700 gr. of crystallized Glauber's salt were melted in an iron
vessel and kept boiling for some time. The vessel was then removed
from the lamp, and the li(jnid portion of its contents was poured otf
as completely as possible, whilst the separated anhydrous salt was left
in the vessel. The whole was allowed to cool to about 50°, before
the vessel was again heated. This vessel was now provided with a
tin-plate cover having two holes, through one of which a thermometer
passed nearly reaching the bott<:)m of the vessel, and through the
other a second thermometer which hang in the steam chamber over
the heated crystalline magma."

" The liquid completely boiled at 82° (Die Flüssigkeit kam schon
bei 82°c. vollständig in's ivochen), whilst the escaping steam showed

22 J- SAKURAI.

the temperature of 100°. Only very slowly did the temperature of
the heated magma rise and attained the temperature of about 100° as
the mass got almost dry."

" To this almost dry mass I added 100 gr. of crystallized
Glauber's salt, which melted on shaking."

" The crystalline magma now boiled at even 72°c., whilst the
steam indicated 100°c." Qoc. cit., 422).

It is not correct to say that " the liquid completely boils at 82°,
whilst the escaping steam shows the temperature of 100°", for it is
only a wet mass of anhydrous sodium sulphate that is heated. I found
that steam, in such a case, does not arise from the heated mass
uniformly, but escapes through a number of channels produced in
those portions of it which are in contîict with the sides of the vessel
and which are, therefore, most heated. The central portion of the
magma, where the thermometer bulb finds itself, is more slowly
heiited, and hence the fact that while the steam indicates the tempera-
ture of 100°, the wet magma shows a lower temperature.

In the second experiment, where some crystallized Glauber's salt
is added to the heated magma, the temperature of the latter is, for a time,
very much lowered, not only because a quantity of a cold body is
introduced, but also because the fusion of the crystallized salt absorbs
much lieat. The central portions of the mass, therefore, show as low
a temperature as 72°, whilst steam which rapidly forms and escapes
along the sides of the vessel shows the temperature of 100°. That
the latter does not indicate a higher temperature is easy to understand
irom the form of the experiment. It is also needless to mention
that the temperature of the whole mass soon rises.

Gerlach's observations are erroneous, then, in so far as they
imply that a substance can evolve a vapour hotter than itself.

Modification of Beckmann's Boiling Method

of determining Molecular Weights of

Substances in Solution.

by

Jöji Sakurai, F. C. S.

Professor of Cliemistry, Imperial University.

JVith Plate I.

Recent investigations in Physical Chemistry have extended our
means of determining molecular weights of substances. Besides the
ordinary method, which depends upon the determination of the
specific gravity of a substance in the gaseous state — the so-called
" vapour density " method — we have now, thanks to the labours of
Kaoult and others, various means of ascertaining molecular weights of
substances in solution. Diminution of vapour tension, lowering of
the freezing point and rise of the boiling point, of a solvent, are the
changes which accompany the dissolution of a solid body in it, and
each of these changes supplies us with a means of determining the
molecular weight of the dissolved body. These are mutually
connected together by theoretical considerations, but for purposes of
determining molecular weights the method, which depends upon the
change in the boiling point, fnv surjxasses the others on account of the
simplicity of the manipulation and of the greater exactness in the
results obtained. To lîeckmann (Zeitsch. pkysik. Chan., 3 603
[1889] J 4, b'62 [1880J; 5, 76 [1890]; 6, 4H7 [L890]; 8, 223
[1891]) we owe the application of this method for laboratory pur-
poses; but, in spite of his indefatigaljle labours, there are some points

24 J- SAKURAI: MODIFICATION OF BECKMAXN'S BOILING

in his method which are capahle of improvement and simpKfication.
The object of the present paper is to describe a modification of the
method which renders it practicable in every laboratory and, at the
same time, able to yield results which are even more concordant than
those obtained by Beckmann.

It may, however, not be out of place to first briefly refer to the
principle which underlies this method of determining molecular
weights. By the study of the lowering of vapour pressure of various
solvents by bodies dissolved in them, Raoult was ultimately led to
the following law :

" The relative lowering of vapour pressure is proportional to the
ratio of the number of molecules of the dissolved substance to the total
number of molecules in the solution."

Expressed in a formula, we have

p N+n

where p and p' are the vapour pressures of the solvent and of the

solution, and N and n, the number of molecules of the solvent and of

the dissolved body respectively. The constant c may be taken as

equal to unity. If, then, G and g be the weights taken, and M and m

the molecular weights, of the solvent and of the dissolved body, all

expressed in grams, we have

9
p—p' _ m

P ^ _; ^ '

or vi=M.-4t~' — ^•

(r p—p

Hence, if M, G and g are known, the molecular weiglit of the dis-
solved substance, m, is given by determining p and 2_>', the vapour
pressures of the solvent and of the solution.

Now, the diuiiuution of vapour j>ressure is proportional, on the

METHOD OF DETERMINING MOLECULAR WEIGHTS. gg

one liand, to tlic ](nveriiig of the freezing point and, on the otlier, to
the rise of the ])oilini>' point. The former relation, which was
established by lîaoult in a purely empiric:d way (Compt. rend., 87
[1878]; 103 [1886]; 104 [1887]; 105 [1887]; 107 [1888];
Ann. Clihn. Plnjs. [6], 15 [1888]), liad already been deduced by
Guldberg- {Compt. rend., 70 [1870]) from tlie mechanical theory of
heat. Moreover, van't Hoff (Zeitsch. plujsik. Chcm., 1[1887]) theore-
tically deduced the laws of the lowering of the freezing points of
solutions from the relation which exists ljetv;een vapour pressure and
osmotic pressure.

The determination of, the other relation, the rise of the boiling
point also affords us a means of ascertaining the molecular weight of
the dissolved substance, inasmuch as it is possible to determine the
temperatures at which the solvent and tlie solution exhibit equal
vapour pressures, instead of ascertaining their vapour pressures at an
equal temperature. This method was lirst practically carried out by
Beckmann.

The calculation of the molecular weight is made by taking into
consideration the projiortionality which exists between the difference
of pressure and the difference of tempeniture. î^ow, since the solutions
subjected to examination are all very dilute, the equation

p—p' _ n
p ~ N+n

may assume the following form witliout causing any material dif-
ference in the results :

p—p' _ n
^} ~ A' ■

The difference of tem])orature prodnred in tlie l)oiling point by the
dissolution of one gram-molecule of the substance in one gram mole-
cule of the solvent and which corresponds to the above pressure-

2 g J. SAKURAI: MODIFICATIOX OF BECKMAXX'S BOILING

difference is called the mokcular elevation of boiling point. This con-
stant is ohtained expérimental 1 y by determinino; the rise of boiling
point of ÎI solvent, fo]l(3wing* the dissolution of a known weight of a,
substance of known molecular weight in a known ({uantity of the
solvent. Thus, for example, Beckmann (Zeitscli. phijsil'. Cheiu., 3,
603 [1889]) found that 4.740 grams of ethyl benzoate dissolved in
100 grams of ether raised the boiling point of the latter by 0.605°.
The weight of ethyl benzoate dissolved in one gram-molecule of ether
(74 grams) is, therefore, — — tää =3.5076 grams. The rise of boil-
ing point corresponding to the dissolution of one gram-molecule of
eth}^ benzoate (150 grams) in one gram-molecule of ether would

then he

0.66.5x1.50 ,,o i
8.5076

Experiments with other sul)stances give nearly the same value for the

molecular elevation of the boiling point of ether. 'J'he molecular

weiglit of any dissolved substance is, then, obtained from tlie equation,

A
where, in case of ether, 7>=28.4, f/= weight of tlie substance dis-
solved in 74 grams of ether, and A= the observed rise of boiling
point.

In practice, it is more convenient to refer the constant B to 100
grams, instead of to the molecular weight, of the solvent ; the calcula-
tion is thus very much simplified. The formula then becomes

A

74
where, in case of ether, jB' = 28.4x^r7— -='21.0, and ^'=weight of the

substance dissolved in 100 grams of the solvent. As the mean of

several determinations, lîeckmann (he. rit.) o])tained the number, 20.9

for the value of 2^.

METHOD OF DETERMINING MOLECULAR WEIGHTS. 2J

The constant B or B' may, moreover, be accurately calculated in
the following manner from the latent heat of vaporization and the
boiling point of the solvent, as has been pointed out by Arrhenius
(Zeitsch. physik Chem., 4, 550 [1889]).

Suppose we have a solvent of molecular weight, il/, and of
boiling point, T (in absolute temperature), at pressure, ];. By dis-
solution of n gram-molecules in 100 grams of the solvent, the boiling
point rises to T+dT. The relation between this rise, dT, and the
dissolved mass, n, may be calculated thus :

The solution boils under pressure p at the temperature T+dT.
The vapour pressure of the solvent at T is equal to p, it is equal to
p + dp at the temperature T+dT. Between dT and dp the following
relation exists according to the second law of thermodynamics, as
already shown by van't Hoff ^Zeitsch. phjsih. Chem., 1^ 494 [1887]):

^P =clT. ^'^ ■

p '27'-

where m= latent heat of vaporization of a gram-moleciile of the
solvent at temperature, T.

Now, according to Kaoult's law, the relation between the diminu-
tion of vapour pressure and the number of molecules of tlie solvent
and of the dissolved substance is expressed by

p—p' n

p

~ N+n'

ip + dp)-

■p n

p + dp

N+n

dp

11

or

or

p + dp N+n

But, as the solution is very dilute, dp is very small compared to p,
and n very small compared to A^ The last equation may, therefore,
be written as

gg J. sAKURAi: Modification of ëeokmann's boiling

dp _ n

or, as 100 grams of the solvent is employed, and M is its molecular
weight, ^ve have - .

ch) M

— n.

P 100

Equating this value of — ^ with that already obtained, we have

1

n. M CO. clT .

100 ~ 2r2 '

In this equation, o) being the latent heat of vaporization of a gram-
molecule of the solvent {M grams) in grani-calori, it follows that
-^= latent heat of vaporization of one gram of the solvent. Calling
this IK, we liave

n. M M. W. dT

100 '2r-

02 r^

or dT^^ny. — ^^^ —

If s= weiirht of the substance dissolved in 100 orams of the solvent
and m= molecular weig-ht of the dissolved substance, then n=^—'
Substituting this value of n in the above equation and solving for ///,
we have

.0-2I- .«?

7)1-

W clT

In this equation, s nnd dT have the same meaning as (j and A

respectively in one of the preceediug equations, and the constant,

02 r-
^ > for ether is found to be equal to

.02 X (273 + 84.97)- _,-,,
yo.l ~^ '

the value which we already obtained for B' in the same equation.

Now, turning to the practical side of the question, the great and

METHOD OF DETEIJMIXIXJ MOLECULAR WEIGHTS. 99

the only difficiiJt\' which lies on tlic way is the exact deteiiinriation
of the boiling points, as has ah-eady been pointed out by Beckmann.
Irregular boiling and bumping, attended with sudden alterations in
the temperature of a boiling livpiid are phenomena which are too well
known to be mentioned. Even when the liquid seems to be regularly
boiling, there are still constant changes in the temperature of the
li<]uid, as may be readily observed by the use of a delicate thermo-
meter. A'arious means have been tried to overcome these difficulties.
]5its of metallic wire or foU, of broken glass, etc. are almost useless.
JJaoult tried coating the thermometer bulb with a layer of palladium
recently charged witli liydrogen gas, but it did not lead to the
desired result, as the evolution of hydrogen gas soon slackened and,
in about 20 minutes, stopped altogether.

Beckmann has succeeded in gi-eatly overcoming the difficulties.
He uses a piece of stout platinum wire which is fixed in the bottom of
tlie boiling vessel l)y means of fusible glass. On account of the
superior thermal conductivity of tlie metal, the boiling is said to take
place exclusively from the piece of platinum, and bumping is avoided
entirely. I regret to say that I have not yet had an experience witli
Beckmann'« apparatus. In fact, we ordered one from Germany, but
on my first attempt to make trial of it, the boiling vessel was found
t(j be cracked at the point of insertion «jf the platinum. Tliis was
merely an accident, but it is an accident wliich nray often occur. A
small p(jrlion of the bottom of the boiling vessel is thickened by the
insible glass, and there is no wonder if the bottom is cracked by the
slightest carelessness in heating.

Tiie means wliich I adopt in producing a constant temperature
in tlie boiling li(juid has already been described in my paper, on " The
delerminati(jn of the temperature of steam arising from boiling salt
solutions," and consists in ])assing a current of the vapour of the

30 J- SAKURAI: MODIFICATION OF LECKM.VXX'S BOILIXG

.solvent into the IjoiJing Jiqiiid or doJiition, the ainoiint of tlie v;i[);*ni'
tliu.s liaised in li-oiii Avitliont Ijcinij' roii'iilated liv tli(î lieiu'ht of tlio
lamp. The degree of constancy of the température attained in tlii.s
manner l)y tlie l:)oilinL;- ]i(|uid i:s really a.jtoni.shing, a moot delicate
thermometer, capahle of showing xo-Vo^th of a degree, I'emaining a.]mor>t
stationary.

Dc-'icn'ption of Apparalan. Iiiotead of the tlvree-iieH<:ed ii:!.~-k wiili
a piece of platinum wire fixed into tlie bottom and with a c-)mplex
condensing arrimgement, used by J Beckmann, I eiP.|>loy, for the hoiliiig
vessel, an ordinär}^ U-tube, A, about '2 centimeters in interna!
diameter and 21 centimeters in height. Xear the top (jf one hmb of
this U-tube, a hole is l)l(jwn out l)y means of the 1)lowpi[>e. wliicli
serves for jointing on, with a piece of india-rul)ber tubing, a small
side-tube having a glass «topper-cock s. The side-tube is, in its turn,
connected with an ordinary Liebig's condenser C. " The boiler for
generating the vajjour of the solvent is an ordinary round-bottomed
flask T>, provided with a cork, through which |>as.^ the stem of a
tapped funnel / and a delivery tube d. The latter is connected, l)y
means of a piece of thick india-rubber tubing /, witli w tube e, drawn
out and slightly Ijent, at its lower end, so tliat tl)i.< end of tlie tube
may just reach the bott<jm of the boiling vessel, wlien tl>e cork,
through Avhich it ])asses above, is ßxed into the moutli of the U-tube,
neiirest to the b.->iler. This tulje e thus establislies comnuuiication
between the boiler and tlie boiling vessel. A tin-pla.te vessel pp with
a hole in its bottom and resting upon a tripod serves to enclose the
boiler and to protect it from draughts. The boiling >essel is, on the
other hand, enclosed by a box made of thick asbestos card-board qq,
witli a UKjvable bottom. When the solvent is of low boiling poiiii,
such as carloii l)isulphide or ether, a j)iece of plain a..:bcsto:^ card-l)<>ard
serves as the botO)m of I he box. and the lioilini;' ^essel is lieaied bv

METUüD OF L)Ei'EKMIXrN(i MOLECULAR WEIGHTS. 3|

indirect flmne, us sliown in the figure. In tlie case, liowever, of less
volatile R<ilvent:^, hucIi iva Avnter or even alcoliol, a piece of asbestos
car(l-])oar(l havins; a hole in the centre is n.ed instead, and the hoilino-
is produced hv direct flame.

The height of tlie flame l)nth under tlie boiler and the b(.)iling
vessel has to be carefully regulated for each particula.r solvent a.ccord-
ing to its volatility ; and, in order to avoid tlie effect of any alteration
in the ga..^ pressure owing to constant openings and shuttings of gas
ta.ps in other parts of the lal^oratorv, the fjllowino- arrano-ement is
adopted and is found to answer admiral)ly. The gas tube is connected
witli tln-ee Ihuisen Inn-ners and, fully opening the gas tap, the burner
nearest tlie tap is liglited. This acts a,s a ga.s regulator. The supply
of gas to the other two Ijurners is adju, -ted by means of screw-clips
k and / ; a.iid, since the amount of gas supplied to the regulator-
burner is incompara.bly grea.ter than that supplied to the two others,
no effect of alterations of ga.s pressure in the mains can be perceived
in the latter. Thi.:! circum: taiice is of considerai>le importance,
because I am thereljy enaljled to use the budlj of the thermometer
always naked, there being no necessity to envelope it with asbestos
fibres, as when working vrith 15eckmann's apparatus.

The therinometer which I use includes onlv a ranofe of 6°c. and
is graduated into jijyth. of a degree. Each of these divisions (about
0.1 mm.) can be readily esthnated into rVth by means of a reading
telescope, which I ahvays use, so tha.t the thermometer can indicate
T^V^th of ti degree. There is an arrangement in tlie instrument by
which the quantity of mercury in the bulb may be increased or
diminished, so that it can be used either for hnv or high temperatures.
'1 hi.^ instrument has alread}^ been described by Beckmann.

The whole arrangement is exceedingly sim|)le, and can ])e set up
by any one with materials commonly found in :dl l:d)oratories. This,

32 J. SAKÜKAI: MODIFICATION OF BECKMANX'S BOILING

I veiitnre to say, is a great advnntnge. The oij]y costly part of the
apparatus is tlic thermometer, hut for good and accurate woi-k in
most of tlie physico-chemical in(]uiries, ii delicate tliermometer is
indispensahle and, after all, its cost is not disproportionate to the
important services it can rerider.

]}[o'Jc of Worhiurj. For working with tlie apparatus descrihcd
above, I beoin bv inserting the tube e into one limb of the br,iling
vessel, lixiiif it ])y meisns of the cork c. Small glass beads are then
introduced into the boiling vessel through the other limb, till they
quite fill tlie bent ])ortion of the latter. (IV^-kmann recommends tlie
nse of small garnets or glass beads for preventing local differences of
temperature, and I find them also very effective for thoroughly
mixing the vapour passed in with the boiling liquid and thus
establishing a perfect uniformity of temperature.) The experimental
liquid is now poured in, so that it occupies a space of 3 or 4 centimeters
above the level of the glass beads in the oj'cn limb, and the latter is
then closed by the cork carrying ilie thermometer. The lid of the
asbestos box is next slipped down the two lim1)s <^f the U-tube, which
pass through two holes previously bored in it in proper positions, and
the boiling vessel, after having been properly enclosed in the asbestos
box and supported on the ring of a retort-stand, is connected, on the
one side, with the boiler and, on the other, Avith the side tube and
condenser, as shown in the figure.

First the boiler, half filled with the solvent, and then the boiling-
vessel are heated by carefully reguLited lamps, the stop-cocks r and s
both open ; and, when the li(juid in both vessels begins to boil [pro-
perly, the stop-cock v is closed, and the two lam])s Aery carefully
regulated, so that the height of the column of the boiling lixpu'd
remains, as nearly as can be judged, level with the lid of the asbesto.s
box for, at least, a quarter of an hour. This circumstance is used as

METHOD OF DETERMTXIXG MOLECULAR WEIGHTS.

33

the criterion f<^r ja<lging that the boiler and tlie boiling vessel are
each being heated by a properly adjusted lamp, so that evaporation and
condensation balance each other in the boiling vessel. The thermo-
meter rises raj)idly at first but more slowly afterward and attains the
maximum temperature in 10 or 12 minutes, which then remains per-
fectly constant. The top of the tliermométer is now lightly tapped for
some seconds, and tlie observation of the exact temperature made by
tlie aid of a reading telescope. Finally, the volume of the distillai e
collected in the sm:dl flask F is noted and, after Avithdrawing the
lamps, the stoji-cock r is opened and the stop-cock s closed.

I make it a rule to repeat this observation of the exact boiling
])f)int of the solvent once or, in some cases, twice more in order to
make quite sure of the temperature. For this purpose, the distill-ite
is returrjed to the boiler arid the whole of the al)Ove process repeated
under exactly the same conditions as before, taking the final observa-
tion of the temperature "when the distillnte reaches the n^ark previously
made (^n the receiver flask.

The next ste]) is to introduce the suljstance into the solvent.
This may l)e d(^ne in two ways. The first and simplest method,
which should be used when the solvent is of low boiling point, con-
sists in taking out the cork, which carries the thermometer, adding
the solid substance to the solvent, and then immediately replacing
the cork to it original position. The other method, which is employed
in the case of solvents of high boiling point and where, therefore, if
the first method were adopted, the highly heated thermometer would
have to he suddenly exposed to the cold air, is as follows : After
the exact determination of the boiling point of the solvent has been
mnde, the boiler is ;i little cooled witiiout opening the stop-cock r,
when a small (|u:intity (n"" the solvent runs up the tube e and fiows
down into the Ijoilei-. The stop-cock r is now opened and then, dis-

34 J' SAIiURAI: MODIFIOATIO-X OF BECKMANX'S BOILTXG

connecting' the delivery tul^e d from the tul^e e, a small quantity of a
concentrated solution of the substance, made beforehand, is introduced
into the boilino- vessel bv means of a small funnel attached to the
india-rubl'Cr tubing /. After removing the funnel, communication is
rc-estabbshcd between the boiler and the boiling vessel.

Tjie lam];s, which have been put aside but not blown out, are
next brought under the two vessels as before, and the boiling point of
the solulion determined in exactly the same manner and under
exactly the same conditions as those in which the boiling point of the
solvent was determined, collecting, of course, the same quantity of the
distillate. When the exact temperature has been ascertained, the
lamps are put out, the stop-cock r opened, the stop- cock s closed, and
the whole apparatus allowed to cool.

After the solution has completely co()led, the percentage composi-
tion of the solution is determined ])v transferrin 2: some of the solution
into a weighed stoppered ])ottle ]>y means of a pipette and, after
ascertaining the weight of the solaii(3n taken, evaporating the latter
at a o'entle heat on a water l)alh and weijihin*:'' the fixed residue. In
case of iodine, it is determined vojumeti'lcally by a standard solution
of sodium thiosulphate.

From the account above detailed of the process I adopt, it will ])e
seen that it differs from the method used In' Beckmann essentially in
this that, whilst Beckmann takes a weighed quaiitity 1)oth of the
solvent and of the solid substance, arid determines tlie boiling p(;ints
in an apparatus provided with a reflux coîidenser and, therefvre,
requires a complicated arrangement, the meth(-d liere de; criljed re<[uires
no special apparatus, as it determines ihe boihug poiiits in the
ordinary manner, l)ut o1)viatirjg the u,^n:d (liflicubies. of such det(>r-
minations and, at the same time, kee])iiig llie (jiiaiiiity of the boiliiig
liquid constant, b}^ ])a,-siiig a current of tlie \a])v>ur of tlie solvent

METHOD OF DETERMIXIXG MOLECULAR WEIGHTS. q X

from without. Another dittercncc which, I vcntiii-e to s'ly, i.s also an
iiuprovcineiit i.s that the Ijoiliii!^' point of the .<ohition hi {i.scertained just
before its compo.sition is (leteriuined and it, therefore, render« the
roniieetion between these properties more certain tli-an in the case of
Jîecknianr/s method. The following tables contain the results of mv
determiijatioiis.

Ü6

J. SAKUllAI: MODIFICATIUX OF BECKMANN ö BOILING

r;

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METHOD OF DETEllMINIXG MOLECULAR WEIGHTS.

37

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38

J. SAKURAI : MODIFICATION OF BECKMAXX'S BOILIXG

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r^

"^

(6

o

CO

Xr-

»o

CO

o

r-H

X

X

CO

o

CM

X

Cî

o

X

CO

CO

CO

CO

CM

1—1

iC

CO

1—1

M

Ö

ö

d

d

d

d

d

d

^

ri

o

ce

-H

i-o

CM

.^H

1-H

o

n^ /

ö

o

^H

X'

o

o

-^

Ci

L-

^ ^

QC>

i:^

^1

'X

L-^

X

'3

O}

CO

CO

-*

CO'

CO

't

c?

CO

m

,-,

>o

c^

o

c^

1 — 1

o

C>T

-M

œ

X)

X

X

o

»it«

Tl

X

-t;

CO

X

'-0

t^

lO

-*

-o

\cß

CO

CO

CO

CO

CO

CO

'CO

CO

00

c^

1 — 1

II

6
o

CO

o

d

^

Ü

<*

CM

œ

CD

-

CM

'L

r^

œ

o

h-H*

^

^

^

^

1ö

:-i

r^

ïlj

g

^

o.

^

ce

o

h- 1

!3
CG

40 J- SAKUEAI : MODIFICATION OF BECKMAXX'S BOILIXG

Fii ooiiclunoii, a word may be waid with regard to the molecidar
weights of iodine and of nidpliur in solution. Paterno and Xasini
(Ber. d. Deutsch. Chem. Ges., 21, 2153 [1888]), l^y the method of
tlie freezing point depression, found that the molecular weight of
iodine in very dilute benzene solutions is represented hy To, whilst it
possesses higher vrdiies in more concentrated solutions. They found
also that iodine is more or less dissociated into the atomic state in its
solution in glacial acetic acid. Loeb (Zeit, phjslk. Chem., 2, 60(i
[1888]), however, could not verif}' these conclutions by tlie same
method; and, both on account of the small solubility of iodine in
either benzene or Macial acetic acid and of the o'reat variations in the
results according to concentration, he idtimately gave up these ex-
]ieriments. By determining the vapoiu^ ])ressures of solutions of
iodirje in ether and in carlDon l^isulphide, Loel") showed, on the other
li;uid, that the molecular magnitude of iodine in lirown etheria.l
solutions is represented by I^, whilst in carbon l^isulphide it is less
complex and corresponds to the formula lo.j. Lastly, Ijeckmann
(^Zett. fi]iysik. Ghein., 5 76 [ISOfl]) found, by the aid of his boiling-
method, that no appreciable ditterence exists in the molecular magni-
tude of iodirie, whether dissolved in ether or in carbon bisulphide,
and that it corresponds to L, in both of the solutiorjs. The numbers
he obtained varied between 235 and 25() in etherial solutions, and
between 250 and 272 in carl)on Insulphide solutions (1-2 = 254).
More recently. Hertz (Zeit, phiisik. Chem., 6, 358 [1890]), by the
freezing method, obtained numbers varying between 255.5 and 276,0
f<3r the molecular weight of iodine in its dark red solution in
naphthalene, and pointed out that the red colour of the iodine solution
is not due to the existence of complex molecules, as already shown by
Beckmann. My own determinations also fully conûrm the results
obtained by the latter.

METHOD OF DETERMINATION MOLECUI;AR WEIGH7'S. 4 [

With regard to the molecular weight of sulphur in carbon
bisulphide solution Reckmann, by the boiling method, oljtained
numbers varying between 245 and 281 and, therefore, pointing to the
existence of complex sulphur molecules represented by 8^=256. The
formula Sg had generally been regarded as the mnximum molecular
formula of sulphur, and the results obtained by Ijeckmann, therefore,
needed confirmation. The experiments by Hertz (loc. cit.) who obtained
numbers varying between 262.3 and 279.4 and, still more conclusively,
my own determinations, put Beckmann's views beyond any doubt.

A Simple Experiment in Chemical Kinetics.

by
K. Ikeda, Rièakushi.

The usual lecture experiment of absorbing oxygen from air by
a piece of phosphorus presents a very simple case of ciiemical kinetics.
For, if the lump of phosphorus employed be not too small, the nctive
surface remains practically constant, and the chemical action must h^
proportional to the density of the oxygen left. The course of the
chemical change can be followed by observing the diminution of
pressure or of volume. The chief disturbing effect is the elevation
of temperature consequent on the oxidation, but this can be prevented
to a great extent by keeping the phosphorus cool.

To this end a sufficient quantity of pure clear phosphorus was
melted between two test-tubes ; the inner tube was
next filled with cold water when phosphorus solidified
and could be easily drawn out of the external tube.
In this manner a uniform cylinder of phosphorus was
formed around the lower part of the smaller test-tul)e.
Any irregularities in surface of the phosphorus were
removed with a sharp knife, and the whole carefully
smoothed. Tlie phosphorus cylinder thus prepared
was about 30 m.m. long, a little less than 1 m.m.
thick, and 12 m.m. in external diameter, so that the
total surface was very nearly 11 square cm. To
keep this cylinder cool, the test-tube was fitted with Fig. 1.

44

K. IKEDA.

a cork furnished with two thin tuhes, through which a current of
water at ahout 19°C. was maintained. The annexed figure shows the
phosphorus cylinder thus equipped. The surface remained perfectly
smooth to tlie end of the experiment, even some thin striae left by the
knife remaining perfectly distinct. The diminution of phosphorus
during the experiment was, of course, very slight, in the second case
not amountino- to more than one-hundredtli of a m.m. in thickness.

The first arrangement.

A large two mouthed bottle (1.5 litre capacity) containing air and
a layer of water, was surrounded
with water, as shown in fig. 2.
The side mouth was closed with a
caoutchouc stopper holding a
manometer and a short glass tube
with india-rubber tube and pinch-
cock. The phosphorus tube was
now introduced into the bottle
through the central opening
which it closed air-tight by
means of a stopper surrounding
its upper part. To make the air-
tio-htness certain, the mouths
were both covered with water.
Before beg-innino; the observation,
the pinch-cock was opened for a Fig. 2.

moment to equalize the internal and external pressures. As oxida-
tion' .proceeded the internal pressure gradually diminished, and the
reading 'of the manometer repeated every five minutes gave the
following numbers : —

A SIMPLE EXPERIMENT IN CHEMICAL KINETICS. 45

^■=-:^log

.

Diminution of

Pressure of

L ime.

pressure.

Oxygen left,

â

X

A—x

0'

0.0

155.3

5'

5.0

150.3

10'

10.9

144.4

15'

16.6

138.7

20'

21.5

133.8

25'

2(^.5

128.8

30'

31.1

124.2

85'

36.3

119.0

40'

40.7

114.6

45'

44.9

110.4

50'

49.1

106.2

55'

53.0

101.8

60'

57.3

98.0

65'

61.3

94.0

70'

65.5

89.8

75'

69.7

85.6

80'

73.8

81.5

00

155.3

0.0

â '^^ A-x

0.00284
0.00316
0.00327
0.00324
0.00325
0.00324
0.00330
0.00330
0.00330
0.00330
0.00333
0.00333
0.00335
0.00340
0.00345
0.00350

A, the initial pressure due to oxygen, was not directly measured

but cnlculated out in the following' manner :- —

A = (760 m.ni.— 17. in. in.) 0.209 = 155.3 m.ni.

Where 760 m.m. : I^>nronietric height,

17 ,, ,, : The vapor-tension of water, the teTn))erntnre of

the surrounding water being 20.*'0 — 20.°1 during the

experiment.

20.9 7o : the percentage of oxygen in air.

As the volume of air remained constant, the active mass of

oxygen at time & is proportional to A—x, so that the equation
dx

=^Ti{A — x) ought to be satisfied in this case. But ■ n did not

-L 1 ^

diminish regularly as the time extended, and therefore —j- log —^

46

K. IKEDA.

is not constant as it should be, but grows gradually larger. This
may be owing to the heating effect of oxidation, or to some other
cause. For one thing the manometer being hastily extemporized did
not allow of any exact reading. On the whole, the experiment was a
hasty and rough one, and the result must not be regarded as final.
Still the general agreement with the law of mass action is apparent.

With a strong solution of mercuric iodide in potassium iodide
solution, or some other heaving liquid properly coloured, as the
manometric liquid, this arrangement may be employed in a lecture
experiment to illustrate the simple principle of chemical kinetics.

The second arrangement.

A burette B of 100 c.c. capacity and 18 m.m.
internal diameter was connected with a water reservoir
C and surrounded with water as shown in Fig. 3. The
phosphorus cylinder was introduced into the upper part
of the burette ; and after adjusting the level of water in
B and C, the volume of air in the former was accurately
observed which gave, with proper correction, tlie initial
volume ^0=93.5 c.c. The observation was repeated
every five minutes, the pressure in 1> being always kept
equal to that of the external atomos]:>here, so that the
diminution of the volume x here denoted the amount
of oxygen absorbed. The result is shown in the follow-
inof table.

Fig. 3.

A SIMPLE EXPERIMENT IN CHEMICAL KINETICS. 47

Time.

&

0'

5'
10;
15'
20'
25'
30'
35'
40'
45'
50'
55'
60'
65'
70'
75'
80'
85'
90'
95'
00

During- the experiment the temperature of the surrounding water
did not vary more than three- tenths of a degree (20.°5— 20.°8) and as
this affects only the second place of the decimals in the reading of
volumes, no correction has been introduced for it. The temperature
of the water current passed through the test-tube remained constant
at 18.°5. The final volume of air or rather nitrogen was observed
after the lapse of 48 hours and gave on proper correction 76.0 c.c.
This subtracted from the initial volume gave ^ = 93.5 — 76.0 = 17.5 c.c.
for the initial volume of oxygen.

olume.

V

Diminution
of volume.

X

Volume of

oxygen left.

A—x

li=-rr\x+,l\-A)

^""^ A-x\ .

93.5
91.9

0.0
1.6

17.5
15.9

1.78

90.5

3.0

14.5

1.73

89.5

4.0

13.5

1.58

88.3

5.2

12.3

1.60

87.3

6.2

11.3

1.58

86.3

7.2

10.3

1.58

85.4

8.1

9.4

1.58

84.6

8.9

8.6

1.58

83.8

9.7

7.8

1.58

83.1

10.4

7.1

1.58

82.5

11.0

6.5

1.57

81.9

11.6

5.9

1.57

81.4

12.1

5.4

1.56

80.9

12.6

4.9

1.56

80.4

13.1

4.4

1.5S

80.0

13.5

4.0

1.57

79.6

13.9

3.6

1.58

79.3

14.2

3.3

1.57

79.0

14.5

3.0

1.56

76.0

17.5

0.0

48

K. IKEDA.

A — x

In this case the volume of air diminished gradually, and the
chemical action at time â was proportional not to A —x, but to
so that

dx , A-~x , A—x
K = k-

dä V v^)—x

or M& = ^^^f^dx = (l + ^^f^)dx

A—x \ A—x /

which on integration between o and x gives

Here it is, of course, necessary to employ the natural logarithms.

As the last column in the foregoing table shows, -Y\x + iVo—A)\og
—^ [ is practically constant with the exception of the first two

A — X J

numbers. This deviation allows of several explanations and suggests
various further experiments ; but speculations had better be omitted,
especially since there is but the single experiment recorded above to
go upon. Still it would be highly interesting to see what influence
various diluents have on the velocity of the oxidation, especially as
some gases are said to retard the action greatly.

The Chemical Laboratory of the

Higher Normal School, Tokyo.
August, 1892.

Imidosulphonates.

by

Edward Divers, M.D., F.R.S., Prof.,

and

Tamemasa Haga, F.C.S., Asst. Prof.

Imperial University.

This paper is based upon an investigation of the s(^diiim imidosul-
phonates and their derivatives, all hitherto unknown, but includes,
besides, some new things about the ammonium and potassium salts.
Several mixed tribasic salts are described in it, particularly those of
lead, of silver, and of mercury, which show peculiarities in their
constitution. It also contains the determination by experiment of the
nature of sulphatammon (Rose), a compound of one molecule of
sulphur trioxide with two of ammonia.

Histonj.—noHe (1834-1840 ; !>/.(/., 32, 81; 47, 471 ; 49, 1S3),
Jacquelain (1843: Ann. Cliim. Phys., [3] 8, 293), Fremy (1845; Ann.
Chim. Phjis. [3] 15, 408), Woronin (1859; J. Buss. Ch. Soc. 3, 273),
Claus and Koch (1869; Ann. Chem., 152, 336), Berglund (1875; /..
miiv. Arsk, 12 and 13; null. Soc. Ch., 25, 455; 29, 422; Ber. Ch. Ges.,
9, 252 and 1896), Rascln'g (1887 ; Ami. Chem., 241, 161), and Mente
(1888; Ann. Chem., 248, 232) are the chemists who have worked
upon the inorganic salts now known as imidosulphonates.* The
constitution of these s.alts luis been established mainly through in-
vestigations by Rose, Jacquelain, and Berglund.

* Woronin's paper we only know from abstracts in the Jahreshericht and Watts's Dictionary.
Berglund's memoir in the TAind:^ iiniirrsitäts Ar>i]irift we have also not seen.

50 DIVERS AXD HA^A.

Rose disccwered in 1884 that the compounds of sulphur trioxide
with ammonia, his sulj)]iafammon and jxircLsidphatamiuoit, give, when
freshly dissolved in water, the reactions of a sulphate very incom-
pletely or not at all, and do not yield up all their nitrogen as ammonia
to chloroplatinic acid and other reagents.

Jacquelain discovered the then unlikely fact that the stable com-
pound of sulphur trioxide and ammonia, liis sulphamide, is derived
from two molecules of the oxide and three of ammonia.

Berglund determined the constitution of this stable compound to
be imidosulphonic, by ascertaining that it behaves ns an ordinary
ammonium salt only to the extent of two-thirds of its nitrogen.

To Fremy is owing the discovery that hydrolysis of the main
product of the action of a nitrite upon a sulphite yields a salt, his
s'ldphaDiidate, now known to be an imidosulphonate ; and that this salt
exhibits the reactions of neither nitrite nor sulphite and can be so far
hydrolysed as to yield ammonia.

He and, after him, Claus and Koch having attributed to this salt
a composition which allowed of its being written down as an addition-
compound of ammonia, lierglund proved them to have erred by
himself establishing the identity of P^remy's ])otassium sulphamidate,
Claus's disulplunnmonati', witli potassium imidosulphonate prepared
from ammonium imidosulphonate coming from chlorosulphonic acid
and ammonia. Mendeléelî" (al)s. in Ber. Cli. Ges., 3, «S72) had just
before suggested such a constitution as ])ossibly belonging to Fremy's
sulphami dates. In his Vrinc'qdcs of CJiciiiislnj, (18i)l), however,
apparently unaware of Berglund's results, he treats parasulphatam-
mon as being ammonium amidosulphonate. More recently, Raschig
showed that the properties of Fremy's salts are inconsistent with
Chius's views as to their being quinquevalent nitrogen compounds.

Claus recognised the sulphonic constitution of these salts, while

IMIDOSULPHOXATES. 5 J

Berglund's discovery that they issue from the interaction both of
sulphur trioxide and ammonia, and of sulphiu- dioxide and a nitrite,
supplied a beautiful, if not needed, confirmation of Claus's theory.

Although, as we shall demonstrate. Rose, Jacquelain, Fremy, and
Woronin had all prepared imidosulphonates with three equivalents of
base, as well as witli two, to one of nitrogen, yet no clear conception
of these compounds as salts of a nioi'e than bibasic acid had been
formed l)efore they were examined by Ikrglund and shown by him
to be derivatives of a tribasic acid, one in ^^hich the aminic, as well as
the sulphonic, hydroo-en is basic. He prepared for the first time
tripotassium and some other trimetal salts.

It will be seen that the existence of imidosulphonates with three
equivalents of base, first made evident by Berglund, constitutes a
misnomer the term imidosulplionate, which he liimself introduced,
since' in these salts the third hydrogen of ammonia is gone. But no
simple way out of the difficulty presents itself, and we adopt it, there-
fore, as Raschig and Mente have done before us.

Imiilosulphonic Acid.

Iraidosulphonic acid, IiN(S03H)2, has been obtained only in
solution. It was first prepared by Jacquelain. We have followed his
process, which consists in treating the lead salt in water with hydrogen
sulphide, and made it our special care to have the lead salt as free as
possible from sodium. The lead salt was decomposed by the hydro-
gen sulphide with a very sensible elevation of temperature, and this
strengthened our expectation that the acid must hydrolyse almost as
fast as it forms. But such is not tlie case, and i-apid filtration over
the pump gave a concentrated solution containing very little sulphuric
acid, and therefore very little amidosulphonic acid.

The imidosulphonic acid was fully identified by its giving an

52

DIVERS AND HAGA.

abundant precipitate with excess of baryta water, freely soluble in
nitric acid, exclusive of a little sulphate ; and particularly by its yield-
ing a crystalline precipitate, also aljundant, with potassium acetate.
'Tacquelain converted his ])reparation into the very soluble dibarium
salt l)v just neutralising- the acid with the Ijaryta, knowing, however,
that excess would give him the insoluble salt. Un leaving the solution
of the acid in a vacuum-desiccator it was found by us to have almost
wholly hydrolysed into amidosulphonic acid in eighteen hours or less.

Fremy attempted to prepare the acid by adding hydrofiuosilicic
acid to j)otassium imidosulphonate and found it to decompose almost
immediately. He made out the products of its decomposition to be
sulphuric acid, sulphurous acid, and ammonia, but in this he must
have erred. J^erglund prepared the acid as Jacquelain had done, and
found it very unstable.

We find that dry dijjotassium imid(jsulphonate may be mixed
with dry potassium hydrogen sulphate, and fused with it without
being affected. At temperatures high enough, 420°-450°, the
imidosulphonate suffers the decompijsition proper to it when alone.
To obser^'e the inactivity of the salts ordinary moist air has to be
excluded, and we therefore \\()rked in a Sprengel-pump vacuum.
From the fused mixtui-e of the two salts, cooled and powdered, alcc^hol
dissohes out only a very little sulphuric acid ; ether does the same.

Dipotassium iniidosul])honate, in dry, tine powder, mixed with
two mc^lecular proportions of sulphuric acid, sp. gr. 1.84, forms a
thin jiaste, which kept in a dry atmosphere S(jon scjiidities to a hard,
somewhat translucent mass. Powdered and exposed to air, this mass
does not deliquesce. Left foi* days in dry air it experiences no evident
change. On dissolving such a preparation in water, to which a little
potassium hydroxide has been added to guard against much hydro-
lysis ol" any component, it yields much imidosulphonate unchanged,

IMIDOSULPHONATES. 53

together with aniidosulphonate, hut no aiiiiiinni;!. The amidosul-
phonate would do doubt be ub.seiit but lor water taken up from the
air during mixing, from tlie sulphuric acid itself, and lastly, to some
degree, during dissolution in the Avater in s])ite of the alkali present.
When heated, tlie mass very readily fuses and gives off sulphuric acid
and sulphur dioxide. Jac(|uelain observed similar behaviour with
sulphuric acid, hot and cold, in diammonium imidosulphonate.

The little action of concentrated sulphuric and nitric acids was
noticed by Fremy, and by Claus and Koch. Evidence, therefore, has
not been obtained of any action of sulphuric acid upon a, dry imido-
sulphonate such as would be expressed by the equation. — H]S[(803K)2
+ 2S0 A= HN(803H),+ 280,HK.

Aiiiinoniiuii iviidosiilpJio'tKites.

jyiaiinuoNiuin mridosulplionate. — Diammonium imidosulphonate,
HN(603Am)., can be ]>i-epared from ammonia and sulphur trioxide
(Rose, Jacquelain, Woronin), or chlorosulphonic acid (Berglund,
Mente), or sulphuryl chloride or pyrosulphuryl chloride (Mente). It
is obtained when nitrous gas is passed into an ice-cold solution of
ammonium sulphite, and the product hydrolysed (Fremy). Along
with ammonia, it is the product of heating ammonium amidosul-
phonate to 160° (Berghuid).

The decom])osition of l)arium imid«Jsulphonate by a soluti(jn of
anmionium sulphate can only be very imperfectly accomplished, but
by treating it .tirst with dilute sulphuric acid barely sufficient to
convert it into the soluble barium hydrogen imidosulphonate and
then adding annnonium sul])hate until no barium remains in solution,
diammonium imidosulj)honate can be ])repared satisfactorily, except
f(jr the incumbi-ance of the bidky barium imidosulphonate and sulphate.
Another procedure which can be followed is to decompose either of the

54

DIVERS AXD HAGA.

hydroxy-lead imidosulphonates with a solution of ammonium car-
bonate. Triargentum imidosulphonate (p. 93) stirred with a solution
of its equivalent of ammonium bromide has-been our source of the
diammonium salt. Triammonium salt is thus produced, and the
solution decanted from the silver bromide needs to be evaporated on
a water bath until amuKjnia almost ceases to escape, and then to be
filtered from a little silver bromide. To crystallise the salt the
solution generally requires further evapcjration and, in (^rder to guard
against hydrolysis into amidosulphonate during the operation, is kept
alkahne by occasionally adding a droj) of strong ammonia-water.

Diammonium imidosulphonate- forms monosymmetric prisms,
isojuorplious with the potassium salt (Mlinzing and Raschig). In
its general reactions it is like the sodium and p(jtassium salts. It
reddens blue litmus, even while still quite free from sulphuric acid
(Jacquelain).

The effect of heating it was tried by Rose, and is described in the
first of his papers upon the sulphatammons. iVccc^rding to him the
products differ little from those coming from heating ammonium
sulphate, except in the absence among them of water. Anmionia
and sulphur dioxide are evolved, first the former, then ])rincipa]ly
the latter. Acid ammonium sulphate makes the residue, at not too
high a temperature. In the retort-neck ammonium sulphite and
sulphate are formed; in the receiver the former salt <3nly. He also got
a little of the yellow compound of the same gases. When the salt is
heated in ammonia gas, he found that but little sulphate is obtained.

This account of Rose's <3f the effect of heating diammonium
imidosulphonate is essentially incomplete, but is true to the extent
thîit anmionia first, and then sulphur dioxide are evolved, that these
two partly condense as a sublimate, and that acid-sulphate is formed.
Heating in a, roomy retort, and thus permitting the action of moist

IMTDOSULPHOXATES.

56

air, fîivoni's the chang-e here described, and brings it still nearer to that
caused l^y heat in ammonium sulphate.

Jacquelain found it fusible without decomposition and capable of
being kept in fusion without change in a current of dry ammonia,
and that, a little above its melting point, it decomposes into ammonia,
ammonium sulphite which sublimes, and ammonium acid-sulphate as
a residue.

We have found matters otherwise. Diammonium imidosul-
phonate, heated in a vacuum, suffers no change (except that it yields
a very minute liquid sublimate at 190°) until very near to 357°, when
it melts and also effervesces or boils. Most of tlie vapours condense
as unchanged imidosulphonate just above the heated part of the tube,
but ammonia of very low tension is given off. At last, inconsiderable
sublimates, white and yellow, appear some distance up the tube, con-
sisting of compounds of ammonia with sul})hur dioxide, and then
with the formation of these, minute quantities of nitrogen escape.
Stopping the operation in an hour or two, when the evolution of gas
is hardly noticeable, the residue is found to be unchanged imidosul-
phonate, mixed with a little acid-sulphate. When the salt is subjected
to a stronger heat, so as to be kept in violent ebullition, most of the
unchanged salt condenses and flows back, and but little of it is got as
a sublimate. On the other hand, the sulphite-Jimmonia sublimates
are more considerable, though still not abundant, and nitrogen is now
given off more freely, l)ut also, as well as free ammonia, still in small
quantities, even though the heating be maintained for a long time. In
one experiment, after about an hour's violent ebullition, the residue was
found, by dissolving it in sodium-liydroxide volumetric solution in a
closed vessel and titrating with acid, to contain about one-fifth of its
weight of ammonium pyrosulphate. From what precedes it is evident
that diammonium imidosulphonate is a comparatively stable body.

56

DIVERS AND HAGA.

boiling :ind condensing unchanged for the most part. A^ery slowly,
however, even when more strongly cheated, it decomposes into nitro-
gen, ammonia, water, sulphur dioxide, and sulphur trioxide, thus : —

6HN(S()3Am)â='2N2+14NH3+60H„-|-6S02+6S03,
these ])roducts then ]mrtly condensing and yielding finally —
'2N2+ 2NH,+ SSaOsAmoH- HOCSOsAm),.

The little ammonia first given off without niti-ogen is no doul)t
attriljutable to the action of moisture in the somewhat deliquescent salt.
This water would cause the hydrolysis of its molecular equivalent of
the salt into amidosulphonate and acid-sulphate, and then, by further
heating, int(j pyrosulphate and ainmoni;i : —

HNlSOBÂm),-!- H2O = HsNSOsAm-t- HOSOsAm = H3N + OiSO^Am).,

Experiments, as yet unpublished, by one of us h.ave fully estab-
lished the existence of the undescribed salt, 0(lS():5Am)o. aimnonium
pyrosulphate.

Trianimoniwii imidosulithouaU' Ciifdrated). — This salt, hitherto un-
described, is formed by the union of ammonia with diammonium
imidosulphonate in the presence of water. The ammonia must l)e in
excess as else the salt is |)artly decomposed by \Nater. Tlie most con-
centrated ammonia-water ] )reci pitates it from a saturated solution of
the diammonium salt ; also^from a concentrated solution of the diso-
dium salt, but then it is mixed with sodium ammonium salt (p. 75)
Ammonia-gas acts similarly and much more effectively. Thus ob-
tained, it forms a crystallinej)owder, l)ut it «'an be got in large crystals
by dissolving it, or the diammonium salt, to saturation in moderately
stronn- ammonia-water, in a closed vessel bv the aid of heat, and
settinj2r the solution aside to cool. It is also üot, and in the laro-est
crystals, by evaporating its anunoniacal solution in a desiccator,

IMIDÜSULPHONATES. 57

over solid potassium hydroxide and in a stroiig-ly ammoniacal
atmosphere.

The crystals resemble those of the tripotassium salt and are,
therefore, asymmetric. They are clear and lustrous, but gradually
eliloresce ("mostly through loss of ammonia) and become opaque and
damp. The saJt when dry smells mildly of ammonia, and its crystals
in a dry ammoniacal atmosphere over solid potasssium hydroxide very
slowly become anhydrous and opacpie throughout, while retaining
much of their lustre and hardness.

Heated,. the salt becomes, mainly, the diammonium salt, but to
some extent it melts, loses both ammonia and water, and hydrolyses
into sulphate.

Dissolved in water it decomposes, but very incompletely, into the
diammonium salt and ;inim(niia, and the solution evaporated in the
open air yields crystals of only the diammonium s'ait. Hence, when
eithei- the trisodium or the tripotassium salt is mixed with ammonium
chloride, much ammonia is liberated.

It was analysed sufficiently by heating it to 160° very gradually,
in a current of dried air, in order to determine the water and third
atom of ammoni;i. The sulphur was also estimated, and the salt Avas
thus proved to liave a composition analogous to that of the potassium
salt, namely, AmN(S03Am).2, OH.,. AVhen analysed the crystals
were damp but transj^arent, after rapid pressure in filter paper and
weighing.

Calc. Found.

Sulphur 26-0 1^5*4

Loss at 160° 14-2 16-9

A nhi/droiis triam.iiioniuni, imidosulphonate. Sulphatanimon. — An-
hydrous triammonium imidosulphonate is, as we shall show, Rose's
sulphatammon. Rose found that dry ammonia conducted into a flask

58 DIVERS AND HAGA.

Jiiied with .sulphur trioxide coiiibines with the trioxide yieldino- two
products, one a hard vitreou« iiiatter replacing the lining of sulphur
trioxide, the other a loose flocculent deposit. Jacquelain got the saine
flocculent body when he mixed vapour ol" sulphur trioxide with dry
ammonia in excess. Rose found that both the flocculent and vitreous
matters when dissolved in watei" and evap(jrated vielded crNsta.ls of the
same substance, but that the vitreous matter was often acid through,
as he considered, imperfect action of the ammonia. The crystals from
solution he called jiarasulphataimiiuH : they are ikjw known to be diam-
monium imidosulphonate, and from what we ourselves have seen of
sublimed ammonium imidosulphonate, we have very little doubt of the
vitreous matter being the same salt, and of its occasional acid reaction
being due to its hydrolysis. The crystals are therefore formed from
three volumes of amnKjnia and twf) of sulphiu" trioxide. but Rose made
them out to have the same composition as the undissolved flocculent
mattei', and to be formed therefore from two volumes oi ammonia and
one of sulphur trioxide. He recognised, however, such differences in
their properties that he held them to be distinct bodies, and called the
flocculent matter siilphakumiiuH. Jacquehiin {»repared parasulphatam-
mon from the flocculent matter by two metliods. 1)oth without the use
(jf watei'. He found this matter to be variable in ccjmposition, and (jnc
of his methods of getting crystalline j)ure parasulphatammon from it.
was to fuse it in a current of ammonia, let it solidify, and then sto[> the
further entrance of ammonia before it cooled. His other method was
to expose the flocculent matter, tirst, over sulphuric acid in a vacuum
till it ceased to lose weight, and then to a moderate heat, which caused
a further loss.

Having in a most careful manner proved the strange erroneous-
ness of Rose's analysis of parasulphatammon, Jacquelain himself made
a, n(j less strange misconstruction of the facts he had correctlv observed.

IWIDOSULPHONATES. 59

He \vc\à found that paras ulphatammon, as it cooled after ])eing' prepar-
ed by heat, took uj) ammonia if in contact with it, and tliat the
flocculent matter, named sulphatammon by Rose, would lose from six
to nine percent, of its weight, in a vacuum over sulphuric acid, leaving
a residue giving up more ammonia Avhen heated to 100,° and then
consisting of parasiilphatammon, as already mentioned. But he treat-
ed these phenomena as a case of the physical abs()r])tion of gases by
porous solids, and likened the ammonia lost or taken up to hygroscopic
moisture. Yet a compound of two molecules of sulphur trioxide with
four of ammonia has to lose as ammonia only .seven and a half per
cent, of its weight in order to become a compound of two molecules
of sulphur trioxide with three of ammonia. As for the high loss of
nine per cent, recorded, that if correctly observed may have been due
to loss of moisture, for, as Rose found, sulphatammon is markedly
hygroscopic and loses the moisture again at 100.°

Sixteen years later, Woronin first established the accuracy of
Eose's analysis of tlie flocculent matter, sulphatammon, and then
showed that this and parasulphatamnK^i were different salts of the
same acid. Another sixteen years later, Berglund claimed for sul-
phatammon that it nuist be triammonium imidosulphonate, since
pnrasulphatanimon had pnn'ed to be diammonium imidosulphonate.
But then, opposed to this conception of its nature tliere were the facts,
first, that Hose had made no admission of its evolving ammonia when
dissolved in water and evaporated so as to yield parasulphatammon;
secondly, that llaschig much more recently had dissolved an old s])eci-
men of parasulphatammon in anuiionia -water and on evapoi-ation oN-er
sulphuric acid got only ])arasulpliatanunon again ; and, lastly, known
\n ourselves at least, that trisodiinu, or tripotassium, or triargentum
imidosulphonate when decom])osed by ammonium-chloride solution
sets free much ammonia. It seemed to us. therefore, at first to be as

60

DIVERS AND HAGA.

evident as it was probable that imidosiil phonic acid c<:)uld not fix a
third ntoni of ammonia along with its imidic hydrogen, and that
Berglund's view might not be right.

Experiment has, however, established th;it sulphatammon is,
as Berglund considered it, the triammonium salt. Crystals of the
diammonium salt were coarsely ground (they are somewhat hy-
groscopic, as just stated, and slip under the pestle, and so are not
readily ground fine). Of this powder a quantity, equal to 2*29
grammes if dried at 100°, wns exposed to a current of carefully dried
ammonia, of which it absorbed in two hours 5f per cent., and in two
hours more an .additional quantity amounting altogether to 6^ ])er
cent., the temperature being 20.° The calculated quantity to ])e
absorbed is 8 per cent., but considering the coarseness of the particles
and the temperature, the result obtained is sufiicient. The absorption
of the ammonia was attended with a very considerable change in tlie
volume of the solid, and gave it the form of n loose, non-coherent,
amorphous powder. Tts odour was only mildly ammoniacal, and it
was easily preserved without care in a liottle. Dissolved in water it
became strongly ammoniacal, :ind the solution gave a copious ])re-
cipitate with barium chloride and otherwise behaved as a solution of
the crystalline, hydrated salt, which salt indeed it yielded wlien
treated with ammonia gas. Evaporation <^f the solution gave crystals
of the dinmmonium snlt, which is just what Woronin observed
in the behaviour of sulphatammon which thus became parasul-
])hatammon.

Exposing it in a desiccator over sidphuric acid for three days at
a temperature of or about 20,° we found it to lose only about a half
per cent, of its weight so that prob.ably the ])ure dry salt loses am-
monia only in a damp atmosphere, ahhough indeed Jacquelain has
found sulphatammon (probably damp) sometimes give up much am-

IMTDOSULPHONATES. 6 J.

moiiia in vacuo over sulphuric acid. Having observed the loss of
ammonifi to l)e so slight at common temperatures, we heated the same
portion in a current of well-dried air for If hours at a temperature
of 100-120,° and then found the issuing air t(^ be still carrying
ammonia The salt cooled and reweighed had now lost only 5*4
per cent, in weight by this heating and gave with water a solution
which by tests showed very fully the presence of some triammonium
salt still.

From what ])recedes it is evident that triammonium imidosul-
phonate is comparatively stable in the anhydrous state, and that Rose's
sulphatammon is this salt. Coming, as hei'e described, from another
imidosulphonate without hydrolysis, it cannot be ammonium amido-
sulphonate, as it is commonly assumed to be : (as in Mendeléeff' s
Principles of Cliemistri/, and Ramsay's System of Itumianic Chemistrif).
Further, its conversion by heat into parasulphatammon and ammonia
affords no proof that it is ammonium amidosulphonate, for this salt
would only change tlius at temperatures above 160.° A salt which is
certainly ammonium amidosulphonate can be ])repared by processes
which include the hydrolysis of an imidosulphonate, and this is quite
a, different body. According to Berglund, it crystallises in large plates,
and only at 160° is converted into imidosulphonate and ammonia. It
is not readily decomposed by water, is neutr;il in reaction, and does
not precipitate barium salts, tacts verified for us by our ]jupil, Mr. Y.
Osaka.

Imidosulphonainide. — Mente has found that i midosulphonamide,
H jSr(S 00X112)2, ii^ ])roduced by the action oi' ammonia upon pyrosul-
phuryl chloride. Diammonium imidosulphonate, when heated to its
subliming ])oint, may possibly yield it, but as at common temperatures
it reacts at once with watei" to form diammonium imidosulphonate,
its presence can not be tested for.

Q2 DIVERS AND HAGA.

Potassium imidosulp horiates.

THpotasskim imidosîilpho7iate. — This yjilt was obtained by Fremy
by tlie hydrolysis of potassium nitri]osul})honate, and this by the
action of sulphite upon nitrite. Claus and Raschig have both ])ub-
lished accounts of the preparation of the salt in this way. it has
also been obtained by double decomposition between diammonium
imidosulphonate and a potassium salt (Fremy, Bergiund): also from
the dia,mmomum s;ilt by Woronin, by converting this first into
Jacquelain's barium salt and then decomposing that with potassium
sulphate.

Imidosulphonates are derived fr<^m nitrilosul])honates by hy-
drolysis readily enough, but as the two potassimn salts are but
sparingly soluble in water, and the imide salt almost as i-en.dily passes
into the amide salt l)y further hydrolysis, the pre))aration of dipotas-
siiun imidosulphonate as hitherto carried out has ] »roved of uncertain
productiveness. On the other hand, the conversion of sodium nitrilo-
sulphonate into disodium imidosulphonate can he very easily ;ind
exactly effected, and, since the disodium imidosulphonate is very
soluble, the insoluble ])otassium salt can be prepared from it by double
decomposition and ])recipitation withc^ut any trouble. Tlius ])re]>ared
it is also verv pure ;ind, consequently, sttible. It can also be i-ecovered
from mercuric dipotassium imidosulphonate by the action of nitric
acid (p. 96)

Dipotassium imidosulphonate results also from the action of heat
upon ])otassium amidosulph<mate. The first knc^wledge of this in-
teresting chîinge— :^H.,NS()Jv=H3N + HN(S0,K),— is due to IJerg-
lund. We ourselves find that at oi- just beloAv 350° this change
proceeds readily and productively, the gas evolved being nmmonin
unmixed with any nitrogen. At this temperature the salt, in our

lMli)U8ULPHÜXATES. g 3

exjDerimeiits. fused, und then gi-adiuilJy solidified ;is the decomposition
proceeded. The salt used was not, howeNer, pure, being mixed with
potassium sulphate.

The effect of heating di potassium imidosulphonate has heen
studied by us. Xot below the softening point of soft lime-glass does
the dry diijotassiinn salt, in a dry atmosphere, sutfer any notable
change. In a ISprengel-])umj) vacuiuii it evolves gas only very
slowly and of Nery small tension, and yields besides only some very
slight sublimates between o 60° and 440." At the temperature of its
actual decomposition it melts and boils. The decom])osition which
then occui's is expressed by the equation —

The ammonia and some of the sulphur dioxide condense to form a
rather vol;itile yellow-brown pseudo-sublimate and a less volatile white
pseudo-sublimate, both of which readily yield ammonia and sul])hur
dioxide aiiain when tested in water. The brown sublimate aives a
clear and colourless solution. Ijoth sublimates ap))ear to be bodies
ol)tained by Rose from su]j)hur dioxide and ammonia, i'robably,
from wliat Meute has done, the yellow" body is nitrogen sulphide and
the wdiite is sulphamide. The permanent gases consist of one measure
of nitrogen to somewhat more than two of sulphui- dioxide. We had
expected to be able to get evidence of the intermediate formation of
nitrilosulph(jnate — 3HN(S03K)2 = H,N-f-2X(803K)3— by arresting the
decomposition betöre it liad gone very tar, and by careful regulation of
the temperature, but we could not do so, any more than in the case of
the diammonium salt.

Fremy (jbserved the formation by heat of potassium sulphate,
sulphur dioxide, and ammonia, and of the coloured sublimate. Claus
and Koch recorded the decomposition of the salt above 200" sharply
into the same bodies. Hut neitlier chemist mentions the still more

g4 DIVERS AND HAGA.

abundant white sublimate and, what is of in<jre «ignificance, the nitro-
o'en. Had either done so, it would then have been seen that the salt
is not an addition compijund of ammonia, as they supposed.

One of the remarkable properties we find imidosulphonates, as
also oxiniidosuiphonates, to possess, is that of hydrolysing at a suffi-
riently high temjjerature by fixing the moisture of the air. Claus had
observed slow increase in weight of ])otassium nitrilosulphonate and in
oximidosul]>hon;ite (his trisulphammonate and disulphydroxyazate)
when they \Nere heated in the air, but wrongly ascribed tliis increase
to oxidation. Oxidation it could not be, for this would (jnly give rise
to volatile products, such as water, or oxide of nitrogen, or nitrogen
itself, and not increase the weight of tlie fixed matter. We have, be-
sides, fuUv j)roved by experiment that at the temperature, or at a highei-
one than that even, at which any imidosulphonate increases in weight
in ordinarv air, that salt remains for hours unchanged in w^eight and
appearance in a current of dried air, and yet is ready, when exposed at
the same temjjerature to undried air, to begin increasing in weight.
By the hydrolysis in ordinary air an imidosulphonate loses crystalline
lustre, cakes together, and becomes acid. Salts, such as the disodium
imidosulphonate, which contain water of crystallisation, i-equire slow
drying at a gentlv rising temperature t(3 prevent hydrolysis, and even
then are difficult to dehydrate, Di])otassium imidosulphonate, the
crystals of wliich are anhydrous, is particularly stable and may be
heated for hours to 140° or higher in undried air, without the
least sensible change. But at 170-180° it slowly fixes water and
hydrolyses.

As the tripotassium imidosulphonate is mucli more soluble than
the dipotassium salt the latter dissolves readily in potassium hydrox-
ide scjliition, from which it can be precipitated by carbonic acid
(llaschig). Other and new reactions of the dipotassium salt, observed

IMIDOSULPHONATES. 65

by us, are so similar to those we record of the sodium salt, as to make
it profitless to give ])articulars of them.

Tripotassiuiii imidosidpliofitik'. — Tripotassium imidosulphonate was
prepared by Berglund, and again by Haschig, by dissolving the
dipotassium salt in solution of potassium hydroxide, evaporating, and
crystallising.

Sodium imidosidp honates.

Disodiiim iiiiidosidplwnate. — The preparation of this salt depends
upon the same changes as those which give the potassium salt by
Fremy's method, but the details of working differ materially. Sodium
nitrite, as nearly pure as possible, and sodium carbonate in crystals, in
the proportion of two molecules of nitrite to three molecules (six
equivalents) of carbonate are, with twice their weight or more of water,
treated freely with sulphur dioxide in a capacious flask, better fitted
with cork and gas tubes. After the carbonate crystals have dissolved,
and acid-carbonate has been precipitated and redissolved with con-
tinuous effervescence, the liquor, still quite alkaline to litmus, begins
to grow warm when more sulphur dioxide enters it. To prevent
any considerable rise of temperature, which might cause premature
hydrolysis and spoil the operation, the flask is, from this point,
kept in motion in cold water, and the stream of sulphur dioxide
somewhat lessened and held under watchful control. Neutralisa-
tion of the alkaline liquor by the sulphur dioxide entering it is
markedly retarded by the action of the nitrite, in the presence of
which neutral sulphite cannot continue to exist. When, at length,
neutrality to lacmoïd paper is almost reached, the flask is actively
agitated without intermission, and the flow of gas first made quite

Ö6

DIVERS AXD HAGA.

slow and then at once stopped when the pajjer i« just permanently
reddened. The hquor at this stage contains, besides a very httle
sulphate, nothing but the two salts, nitrilosulphonate and metasulphite,
which, omitting notice of intermediate stages, have been derived as
follows : —

2NaNOo+3Na2C03 + 8S02=2N(S03Na)3 + Na2SA+3C02

The apparently redundant metasulphite is necessary in order to get
the nitrite converted wholly into nitrile salt. Its presence is almost
equally essential for the preservation from destruction of the imidosul-
phonate about to be formed.

When full neutrality is reached, or perhaps just passed — it is
impossible to say — the liquor after standing a few minutes suddenly
becomes strongly acid and warm, and evolves much sulphur dioxide.
Hydrolysis has occurred, but thanks to the metasulphite present, only
through one stage, and the liquor now holds only imidosulphonate,
sulphate, and sulphur dioxide —

2N(S03Na)3+ Na^S A + OH2 = 2HN(S03Na)2+ 2Na2S04+ 2SO2.

Sulphurous acid appears to have no immediate hydrolysing action upon
the imidosulphonate, but as it oxidises to sulphuric acid it must be
removed. A ra])id current of air is, therefore, sent through the liquor
for half an hour or so, after which the addition of a very little sodium
carbonate suffices to render it slightly alkaline and quite stable. It is
evaporated, so far as may be necessary, on the water-bath and then
cooled down to 0° or a little below, whereby in the course of some
hours most of the sulphate is made to crystallise out. A second
evaporation and cooling is generally needed to separate still more
sulphate, after which renewed evaporation and ordinary cooling brings
about the separation in well-formed crystals of much of the disodium

IMIÜOSULPHONATES. 67

imidosiilphonate. The mother- liquor can be furtlier worked f(3r sul-
phate and imidosiilphonate, if worth the trouble.

The reaction proves to be almost quantitative, for the salt obtained
in crystals is 80 per cent., while there is considerable loss of salt in the
mother-liquors adhering to the very voluminous masses of sodium
sulphate. The crude salt can be purified by recrystallisino- from
warm water rendered slightly alkaline as a precaution against
hydrolysis.

Instead of sodium carbonate the equivalent quantity of normal
sulphite, or of metasulphite. pure and therefore freshly prepared, may
be used along with the additional sulphur dioxide indicated by the
equation already given. The mixture of nitrite and metasulphite be-
comes at once alkaline and rapidly heats up, so as to need cooling, in
accordance with what has been stated. Thh mode of procedure is
certainly less simple and less practicable than that described. It may
also be mentioned that a sufficient excess of metasulphite will of itself
convert all nitrite to nitrile salt, but as then addition of another acid
is required to eifect liydrolysis into imidosulphonate the liquor obtain-
ed is unsuited to give a good yield of imidosulphonate crystals.

It is not unfrequeritly convenient to ])re])are disodium imidosul-
phonate from the trisodium salt. The latter salt can be preserved
without need of taking any precautions, which is not the case with
the disodium salt. Again, where the direct process for preparing
sodium imidosulphonate has been carried out with impure salts, or
with imperfect observance of the proportions to be used, and of other
details, it may not be so easy to crystallise out the disodium salt,
and here the very ready separation and purification of the trisodium
salt becomes advantageous. From the trisodium salt the disodium
imidosulphonate is prepared by treating the crystals with sulphuric
acid slightly diluted until neutrality to litmus is reached. The solu-

68

DIVERS AND HA<-iA.

tion with any crystalline precipitate (sulphate) is kept for some hours
at or about 0° and the mother-liquor decanted, while still at that
temperature, from the sodium sulphate. Evaporation, as already
described, then yields the disodium salt in good crystals, pure or
almost so.

Disodium imidosulphonate crystallises readily in large rhombic
prisms. It is very soluble in water, slightly acid in reaction, and
devoid of sulphurous taste. Its crystals, which contain water, do not
effloresce, even in dry seasons, so that crystals of sodium sulphate
among them may be easily recognised. In air kept dry by sulphuric
acid, they slowly become opaque, but retain their shape, along with
much of their lustre and hardness ; while the rate of loss, even when
they have been crushed, is so slow as to make a water estimation in
this way almost impracticable. They can generally be kept for many
weeks in bottles, and apparently for any time in a sufficiently dry
atmosphere, but are liable to undergo hydrolysis into an acid mixture
of sulphate and amidosulphonate. The solution of the salt is also
unstable, but it may be heated for a short time to a moderate degree
and yet escape change. The crystals can hardly be heated to 100"
without some decomposition.

The decomposition of disodium imidosulphonate that almost
unavoidably attends the process of removing the water of crystallisa-
tion, also complicates observation of the effects of a high temperature
upon it. For, having been to some extent converted by this water
into sulphate and amidosulphonate, it gives at first when heated only
ammonia, which comes from the reconversion of the amidosulphonnte
into imidosulphonate. But at a higher temperature, this is followed by
products the same as in the case of the dipotassium salt, namely,
nitrogen with more than double its volume of sulplnir dioxide, small
brown and white sublimates, and sodium sulphate. The sodium salt

IMIDOSULPHONATES. 69

n.ppears to be much more fusible thun the potassium salt, bat then in
the state examiued the latter was practically pare while the former
was mixed with the products of its hydrolysis, which must have
affected its fusibility.

Disodium imidosulphonate in aqueous solution gives, like the
much less soluble dipotassium salt (Berglund), no precipitates with
many of the usual metal salts. But it gives when in moderately con-
centrated solution a precipitate of dipotasssium imidosulphonate with
solutions of potassium salts ; of trisodium imidosulphonate with
sodium hydroxide; and of a barium sodium imidosulphonate with
barium hydroxide. Concentrated ammonia-solution also gives abun-
dant precipitation. The precipitate, when the disodium imidosulphonate
is unmixed with other salts, is a sodium ammonium salt (p. 75).
But when a httle sodium nitrate, chloride, or other sodium salt is
added before the ammonia, the latter, in concentrated solution, pre-
cipitates pure trisodium imidosulphonate. The two forms of pre-
cipitation are expressed by the respective equations : —

5HN(S03Na)2 + 5NH, - 2NagAmN2(S03)4 + AmNCSOgAm)^ :
HN(S03Na)2 4- NHg -f-NaNO^ - NaNCSOsNa), + AmNOg.

The precipitation by ammonia is in both cases greatly increased by
excess of the reagent, and with the disodium imidosulphonate added
in powder to strong ammonia-water, much of the triammonium salt
also precipitates.

Hydroxy-lead acetates, it was known, give copious precipitates.
Normal lead acetate also gives scanty precipitation of a lead imidosul-
phonate, even when the acetate is truly normal as described, and the di-
sodium salt in solution faintly acid to litmus (pp. 86, 88). Lead nitrate
occasions no true precipitation, but, unlike the acetate, it rapidly
effects hydrolysis and consequent precipitation of a little lead sulphate.

70

DIVERS AND HAU A.

Both iiiercury nitrates ure also precipitating agents, acting l\ere
as they do when added in excess to a solution of trisodinm imidosul-
phonate (p. 101). Mercuric oxide, especially the precipitated form,
reacts with it to give mercuric disodium imidosulphonate (p. 102).
Warmed with a moderately concentrated solution of the salt, the
mercuric oxide dissolves and, on cooling and standing, the solution
deposits crystals of the double salt. Silver hydroxide is rapidly
converted by solution of disodium imidosulphonate into the sparingly
soluble argentum disodium salt (p. 95) Cupric hydroxide is without
action.

Evaporated on the water-bath with S(idium carbonate, or even
with acetate, disodiutn imidosulphonate is partly converted into triso-
dium imidosulphonate, while carbon dioxide or acetic acid escapes.
In the case of using acetate, therefore, there is presented the striking
phenomenon of a strongly acid vapour rising from ;i well-marked
alkaline liquor.

In analysing the salt, we hydrolysed it by heating it with hydro-
chloric acid to 150° in sealed tabes for some hours, here following
Raschig in his annlysis of the potassium salts. The composition of
the salt is expressed by the fornuila — HN(S03N"a)o, (OHs)., the results
of analysis being —

Sodium

Sulphur

Nitrogen

Trisodium imidünalphoriate.— Thin salt is prepared from the di-
sodium salt by adding sodium hydroxide to its strong solution. It is
unnecessary to have the disodium salt pure and in crystals. After
following the process for getting this salt so far as to separate one
good crop of crystals of sodium sulphate, the liquor is diluted with

Calc.

Found.

17-90

17-77

17-7(5

24-90

— 24-88

24-78

5-45

5-40

—

IMID0SULPH0XATE8. 71

twice its volume, or more, of water, treated with sodium hydroxide
solution till it tastes slightly caustic, and then cooled in an ice-l)ox.
Tlie trisodium salt begins to separate almost at once after the addition
oi the sodium hydroxide, but the water previously added retards the
separation and enables the crystals to grow in the cooled liquor suffi-
ciently Jarge to be afterwards l)etter freed from their mother-liquor,
Strained out, and pressed in calico, the salt becomes almost pure, ;ind
can be rendered fully so by recrystallisation.

Trisodium imidosulphonate is a very stable and easily prepared
salt. It forms thin, overlapping, hexagonal plates, which may grow
to considerable size,:. but are .seldom if ever to be seen single and per-
fect, or with much thickness. The crystals readily effloresce in dry
air, iuid have a mildly alkaline taste. The salt is sparingly soluble in
very cold water and very soluble in hot water, taking for dissolution
at 27^-° about 5'4 parts of water. Its solution readily shows super-
saturation. It is alkaline to litmus, and even to phenol-phthaleln, but
exercises no action whatever on iodine-solution, and is |)recipitated
from its aqueous solution by alcohol without decomposition. It can
be repeatedly recrystallised without any loss of alkali. Aqueous solu-
tions are quite stable even when continuously boiled. The crystals
melt when gently heated, and their water may be ra])idly boiled off
without causing any decomposition of the salt. In a vjicuum over
sulphuric acid they lose only eleven molecules of the twelve they
contain. The salt loses more water when heated, but even at 160° it
retains some, ])Ossibly through hydrolysis, so that heated until decom-
position begins, it yields always a very little water and ammonia (gas
and sublimate). Heated in a vacuum, liowever, it yields even still
less, so that atmospheric moisture seems to be active in hydrolysing it
when heated in air.

riie pi'esence of hydrogen in the dried salt might have particular

^^ DIVEES AND HAUA.

significance, since Fremy und, after him. Clans have represented the
dipotassium salt to contain three atoms of hydrogen instead of the one
given by the imide constitution and displaced by metal in the normal
salts. But careful combustion of some grammes of the dried salt with
copper oxide gave us in one case only 0*22 per cent., and in another
only 0*24 per cent, of water, calculated from the hydrated crystals, and
such a percentage corresponds to only about one-ninth of an atom,
instead of the extra two, of hydrogen required l)y the sulphammonate
constitution given by Claus.

When dried trisodium imidosulphonate is heated somewhat
strongly in an open tube it yields, with fusion and effervescence,
nitrogen and some sulphur dioxide, and sulphur whicli sublimes.
The saline mass becomes also very dark-coloured from the presence in
it of sulphur, and gradually solidifies, until at a commencing red -heat
it forms a semi-fused hepar sulphiiris. By heating in a vacuum, proof
is obtained that the sulphur dioxide comes only from the action of
the air upon the sulphur. From 350° to 440,° gas of only very
low tension is given off, along with sulphur vapour forming a sub-
limate of drops. Just below the softening point of good soft lime-
glass, the salt fuses and efi'ervesces. The gas consists entirely of
nitrogen. Some of tlie sulphur remains in the fused mass, partly free,
partly as thiosulphate. A trace of the ammonia compound with
sulphur dioxide forms a sublimate. Apart from this the reaction is
expressed by the equation- —

2NaN(S03Na)2=N2-|- Ö -1- 8 SOiNag

Acids readily dissolve trisodium imidosulphonate, yielding neutral
solutions of the disodium salt when the quantity of acid is equivalent
to one-third of the sodium. Remarkable is the action of concentrated
sulphuric acid which, being not in excess, dissolves the crystals with
marked fall of temperature, although when in quantity more than

IMIDOSULPHONATES. 73

enonsrh to form the disodiniii salt, its iidditioii causes heating'. From
tlie neutralised solution sulphate and disodium imidosiilphonate can
be crystallised out. Carbon dioxide also decomposes the trisodium
salt in water, sodium acid-carbonate being precipitated, if the water
present is not too great. Concentrated ammonia-water precipitates
the trisodium salt free from ammonia, and with its usual water of
crystallisation. Ammonium salts suffer double decomposition with the
trisodium salt, ammonia then becoming free by the decomposing action
of water upon some of the triammonium salt. When concentrated solu-
tions of the trisodium salt and of a potassium salt, such as the nitrate,
are mixed, nothing is observable, but on neutralising with an acid,
there is precipitated the dipotassium salt in crystals.

Trisodium imidosulphonate, unlike the disodium salt, precipitates
many metallic snlts, giving in some cases, however, onlv hydroxides.
Precijntates of imidosulphonates are ol)taiiied with silver nitrate, the
mercury nitrates, lead salts, and barium salts, while with calcium
chloride crystals may slowly form which are of characteristic appear-
ance. S])ecially noteworthy is tlie fact that silver nitrate added in
different j)roportions yields three precipitates distinct both in appear-
ance and in composition. The compounds formed by these reagents
are described later in this paper. Mercuric chloride has apparently no
action upon trisodium imidosulphonate, for nothing separates, and the
reaction with lituius remains alkaline. Mercuric oxide dissolves to a
limited extent in solutions <^f tlie trisodium imidosulphonate, the more
in cjuantity (in proportion to the trisodium salt) the weaker these
solutions are, and makes them somewhat caustic-alkaline (see, in this
connection, the reaction of mercuric sodium imidosulphonate with
alkali hydroxide, p. 104).

It was not practicable to get the crystals of trisodium imidosul-
phonate exactly dry for analysis, because the thin plates adhering

74

DIVERS AND HAGA.

together retained mother-liquor, and, on the other hand, during
much crushing and pressing with [Kiper tliey effloresced. Avoiding
the latter source of error, our results were 12;^ ÜH2, the i representing
one per cent, of moisture. The numerical details of the ^vater deter-
minations, which are a little complex, are the following : —

C^alculated. Found

Moisture
1 1 mois.

1-00
42-70

43-6()

43-56

Loss in vacuo

1 mol.

3- S S

( 3-71
(0-23

47-60

3-64
0-27

Loss at ILP
Retd. at 110°)

Total

47-59

47-47

We have already given (p. 72) the'^percentage of retained water,
as actually found in the salt dried with special cure, as 0-22 and
0-24 per cent. I'he above numbers, 0-23 and 0-27, are difterence
numbers.

The analysis of the dried salt ga\e —

Calc. Found.

Sodium 28-40 28-24

Sulphur 26-34 26-46

Nitrogen 5-76 5*92

The formula of the salt is therefore NaN(S03Na)„ (OH.)(OH,),„

Soiliuni (immomnm imido^nlphonates.

Possibly a salt two-tliirds sodium and one-third ammonium-
imidosulphonate might be got in presence of an excess of the triammo-
nium salt, but this is doubtful because of the sparing solubility of the
followino- salt in presence of ammonia, and because ammonia and an-
other scxiium salt convert disodium to trisodinm imidosulphonate

rp. 69).

IMIDOSULPHONATES.

75

Pentasodiwn ammonkim hmdosulphonate. — From dilute solution this
«alt f.Tystallises with seven molecules of water, NagAmNg (803)4, (OH..)';,
but from concentrated saline liquors it crystallises with less water. To
prepare the fully hydrated s^alt, strong ammonia- water is added in large
excess to a solution of disodium imidosulphonate, which must be free
from sulphate or other salt. Separation of the salt begins at once or
very soon and continues ibr some time if the liquor is kept nearly ice-
cold. The salt forms minute prisms which bear moderate washing,
with concentrated ammonia-water, and can be drained on a tile un-
changed if under close cover. W]ien dry, the salt does not smell
noticeably of ammonia. The numbers obtained l)y calculation and
experiment are as follows : —

Sodium

Ammonium

Sulphur

A salt with ^^^H./) only, falls as a crystalJine precipitate on
adding strong ammonia-water to a solution of trisodium imidosul-
phonate and its equivalent at least of ammonium nitrate (3 mois.).
l^)eing but very little soluble in ammonia-water, it can be properlv
washed with this, and niay then be drained dry on a tile under cover
without change. Heated it loses water and ammonia without suffer-
ing aqueous fusion. It contains no nitrate. Analysis gave —

Calc. Found.

Sodium 21-8G 99.01

Ammonium 3-42 3-40

Sulphur 24-34 24-40

For notice of a monohydrogen sodium ammonium imidosul-
phonjite, one in which the ammonium greatly predominates over the
sodium, see p. 77.

Calc.

Fouuii

18-95

19-19

2-97

2-85

21-09

20-95

76

DIVERS AND HAGÀ.

Hydrogen sodium ammonium imidosidplionate nitrate.

On dissolving trisodium imidosiilphonate and then its equivalent
of annncjuium nitrate (three molecules) in half their combined weig'ht
of hot water in ;i- closed nearly full vessel, a cold solution is obtained,
which sometimes remains unchanged, super-saturated, sometimes
s!(3wly deposits the trisodium salt again of the ordinary form, but in
thicker, crystals than Usual. The attempt to redissolve the crystals
by dipping the vessel in liof water readily succeeds l)ut is generally
attended with the copious separation of some white «jpaque salt
(sodium ammonium imidosulphonate?). On the vessel being left for
some days in a cold place, with occasional agitation, the trisodium salt
re-forms in crystals, and the opaque salt redissolves. When proceeding
differently, the fresh mixed solution of the trisodium salt and am-
monium nitrate in half their weight of water is placed on the steam
bath in an open glass capsule, anunonia freely escapes, and soon groups
of small prisms form, evidently in consequence of the loss of ammonia
by the solution and not its mere evaporation, for the addition of some
water has little effect up<jn tlie quantity of these crystals. The crystals
could not be separated i'rom their highly concentrated mother-liquor in
a: state lit for trustworthy analysis, Ijut we are pretty confident of their
being tlie salt now to be described.

A well-crystallised d<nil)le salt of definite composition is ol)tainable
by evaporating a soluti(jn of trisodium imidosulplionate Avith a.t least
its equivalent of ammonium nitrate, until the liquor is nearly neutral,
then adding water to redissolve any crystals which have formed, and
leaving the product to cool sl(jwly. Through(3ut, care must be taken
that the liquor does not 1)ecome acid during evaporation, by adding if
necessary a drop or two of ammonia. Small flat, thick prisms are

IMÎDOSULPHOXA'Jl^S. 77

ohtaiiied whirh arc nnhydrous and stable in the air. They cannot be
washed, and must be freed from their mother-liquor bv pressure in
paper. The new ;>alt i« of a composition which may l)e represented as
being that of one molecule of diammonium imido.>ul[>lionate with one
of sodium nitrate. Prepared from a good excess of ammonium nitrate
the crystals pr()\'ed to be almost pure, but holding a little water (a);
with the use of less ammonium nitrate, the salt showed on analysis the
presence of a very slight excess of clisodium imidosulphonate. The
yield of salt was three-f(xu-ths of the possible quantity. The mother-
liquor on eva[)oration gave crystals of sodium nitrate and some more
of the salt. Tlic double salt cannot be recrystallised, its solution
yielding, first, good crystals of a salt which inay he described as (anhv-
dioiis) diammonium imidosulphonate contaiiiing o per cent, of sodium,
;ind. af'er this salt, crystals of the double salt. Since the mother-li<pior
of the double salt yields sodium nitrate, it would seem that double
decomposition occurs between sodium imidosulphonate and ammonium
nitrate.

The presence of nitrate in the d(juble salt cannot be directly
indicated in the ordinary way, Ijut may be ar once detected by a new
reaction of delicacy and of some general interest. The salt freely
effervesces when immersed in strong sulphuric acid, the gas being-
nitrous oxide with a little nitrogen. iS'o nitrosyl sulphate is jiroduced.
Quite a minute quantity of nitrate mixed or combined with an
imidosulphonate is shown by dropping the dry solid into a little of
tlie acid, and not mixing tliem up t<jo much. The reaction needs
careful study. V\e ventured to determine the nitric acid in the salt
by means of this reaction, taking the gas as nitrous oxide only, and
got a good result, l)ut do not ieel justified in entering it in the tabular
statement below

In one preparation we determined the nitric acid; in the other it

78

DIVERS AND HAGA.

v>'as undetermined, except in the experiment above referred to. The
nitric acid was estimated by evaporating the salt with barium hydrox-
ide to expel ammonia, filtering ott' the barium imidosulphonate,
removing barium as carbonate, and then patting the concentrated
solution of sodium nitrate into the nitrometer. The result was low,
but that is hnrdlv to be wondered at. Water, in that preparation in
which it was present in any quantity, was estimated by heating the
salt in dried air, and the result can be only approximate, for slight
hydrolysis, fixing water, is hardly to be avoided.

Calc. a. b.

Sodium I'll 8-00 8-42

Ammonium 12" 16 11*71 11 •37

Sulphm- 21-62 21-62 22-08

Nitric acid, XO3H. 21-28 19-08 —

Water O-QO

The preparation a was formed in presence of good excess of
ammonium nitrate, and h in presence of little more than enough of
it. The calculation is for HN(S03Am)o, NaNOs.

Hydrogen sodiwn potassium imidosulphonate nitrate.

T^he very sparingly soluble dipotassium imidosulphonate is hardly
affected by digestion with a cold dilute solution of sodium nitrate, and
w^hen dissolved in it by heating crystallises out again almost un-
chano-ed, l)ut when left in a concentrated solution of sodium nitrate it
removes this salt from it. Taking about equivalent quantities of the
salts, HN(S03K)2 : SISTaNOs, and leaving the solution of the nitrate
standing over the dipotassium imidosulphonate in a loosely covered
beîiker for some days, tVie latter gives place to a caked mass of crystal-
line irrnnules, appearing under the microscope homogeneous but \\\\h

Cale.

Found.

3-32

3-61

28-23

27-34

18-48

18-89

18-20

18-01

IMIDOSÜLPHONATES. 79

no well-detined Ibrins. The decanted mother-liquor evaporuted a
little gives more of the granules, but no crystals of sodium or potas-
sium nitrate. The granular mass drained and pressed on a tile has a
composition closely approaching that of one molecule of dipotassium
imidosLiIphonate to one of nitrate, the nitrate being of potassium and
sodium in single atomic proportions, NaN'03,KN03, 2HN(S03K)o. It
is decomposed by water, and is anhydrous.

Sodium
Potassium
Sulphur
Hydrogen nitrate

The two potassium salts or the two sodium salts do not form
double salts together. Disodium imido.sulj^honate and potassium
nitrate suffer double decomposition. The formula of the salt just
described shows also that, to some extent, this double decomposition
is reversible.

In oiu- next connnunication we shall have to describe still more
remarkable combinations of nitrates with oximidosulphonates.

Bariwii imidosulplionates, simple and double.

Barium imidosidphonate. — This salt has been prepared by Berglund,
but no particulars of it have been published, except possil)ly in
Swedish. It is obtained as a, voluminous, coherent precipitate when
trisodium imidosulphonate is added gradually to good excess of
barium chloride. Drained on the porous tile, after thorough washing,
it is still very bulky, and hangs togetlier in soft flocks, more like
some organic salt than an inorganic barium salt. It retains moisture
in the interstices (jf its matted texture with remarkable obstinacy, so

30 DIVERS AND HAÖA.

that it can not ])e drained dry on the tile, or rendered dry in the
desiccator exce[)t at tlie surface of its masses. Yet it is not in the
least delit^^nescent, and becomes dry in the air when spread out in very
thin layers ; indeed in very dry air it is even sliglitly efflorescent.
Analysis shows it to contain water, hut it loses this only very slowly
even at 115.° Under the microscope it is seen to consist of intei-lacin<i-,
vei'v long, slender needles. It is only very slightly soluble in water,
only sufficiently so to cause the water to give a very slight opalescence
with sulphuric acid. It is nlknline to litmus. It is decomposed a
little by solutions of alkali carbonates. It may be dissolved in dilute
nitric acid, and, with very rapid treatment, re precipitated unchanged
by barium hydroxide. This is a property which, witli precautions,
may be taken advantage of to remove sodium sah, whicli is nearly
always present in the original precipitate, but always absent after
second precipitation. Only once did we get the original preci])itate
free from sodium. Analysis gave —

Calc. Found.

Barium 48'8,S 47-«)l

Sulphur lo'O.S 15-04

Nitrogen o • 2 9 o • H 1

The calculation is for Ba.N/SC),),, (Oil.,),,. The alkalinity of the salt,
measured by decinormal acid and methyl orange, proved equal to one-
third of the bariiun.

Ammonia added to a solution of the salt next described converts
it into tribarium imidosulphonate insoluble, and ammonium imidosul-
phonate.

Barium htjdroijen iiiiiiloxulj)]ioiinli'. — This salt has been briefly
noticed by Jacquelain and by P)erglund. We have prepared it from
the tribarium salt, thoroughly free from scxliiim. by cautious treatment
with dilute sulj)huric acid equivalent to slightly less than one-third

IMIDOSULPHONATES. gl

of the barium, filtering off barium sulphate, and evaporating in a
desiccator. Minute, brilliant, orthorhombic crystals are obtained which
are moderately soluble in water. The salt is stable in the desiccator,
or in the open air in dry weather, but soon hydrolyses in a bottle.
In dried air it is stable even when heated considerably, but it hydro-
lyses in ordinary air at and above ]40°. Eapidly heated it decom-
poses suddenly into a cloud of barium sulphate. Its solution is acid
to litmus but neutral to methyl orange.

Proceeding in almost the same way as that followed by us,
Woronin ol)tained apparently the same salt, but he gave to its com-
position a molecule of ammonia more, calling it hariiini snlpliamate.
Since our preparation contains a molecule of water, the determination
of l^arium and of sulphur .agrees with both fornuili\î — r)a(H2NSÜ3)2
' sulphamate,' and l)aHN(;S03)o, OH^ imidosulphonate.

Quantitative analysis and the properties of the salt agree with the
l^jrmula, BaHNCSO.,),, OH,

Calc Found.

Barium 41-51 41-67 41-65

Sulphur 19-39 19-51 —

In warm solution barium hydrogen imidosulphonate reacts with
mercuric oxide to form the double salt.

Barium (luunouium iiiiidosulplioiiate. — By adding one molecule of
l:)arium hydroxide to one of triammonium imidosulphonate (' neutral
îimmonium sulphamate') Woronin observed the formation of a
'soluble' barium salt, with the evolution of ammonia. He made no
statement as to its composition. On adding Ijarium hydroxide
gradually to triammonium imidosulphonate until it nearly ceased to
o-ive a precipitate we obtained a very slightly soluble salt as a pre-
cipitate which was a Iwrium ammonium imidosulphonate. It was
not fully analysed, but- gave a percentage of barium of 41-87 while

v^2 DIVERS A^'D HAGA.

calculatioii for BaAmN(S03)2 requires 41'64 7o- ^^^^t it may, not
improbably from what follow«, have l^een a hydrou« salt with less
ammonia.

The experiment repeated with diammoniiim imidosulphonate
gave ns a precipitate at first gelatinous but becoming granular on
standinf^". Under the microscoj'C it was seen to consist of granular
crystals intermixed with a very few slender needles of the tribarium
imidosulphonate insignificant in amount. Its alkalinity determined
bv decinormal acid and methyl orange showed it to be a normal salt,
the alkalinity being eqiT^l to one ntmn of univnlent base to two atoms
of sul])hur present. After hydrolysis the totnJ îimmonia, as well as
the sulphuric acid, was estimated. The results agree well with the
calculation for Ba5Am2N4(S03)8(OH2)a, a five-sixths barium salt, cor-
respoiidiiig, therefore, to tlie sodium amuKniium salt : —

Calc. Found.

Barium 43-88 43-84

Sulphur 16-40 16-53

Nitrogen 5-38 5-27

It is here assumed that the observed presence of a very little
tribarium salt could be disregarded. The assumptiori that the pre-
paration was a mixture Bn3N2(SO.,)4, (OHo), + 2[I^»aAm¥(S03)2, (0E,'),_,]
could not possibly be admitted as in accordnuce with the microscopic
a]>penrnnre of the preparation.

Jncquelain ol)l;nned, by adding diammonium imidosulphonnte to
a slight excess of baryta water, a precij)itate which he very fully
analysed, and found to be composed in accordance with the empirical
formula, (NH3)2(BaO)2(S03)3. Instead of this impossible formula we
venture to write Ba4AmN3(S03)6, (0112)3, the formula of an eight-ninths
bnrium nmmoniuin imidosulphonate, since the calculated numbers
agree still better witli his auiilysis than tliose for his formuhi

IMIDOSULPHONATES. ^3

Cale. Found.

(Jacquelain.)

Barium 47-99 48*26

Sulphur 16-81 16-81

* Nitrogen 4-90 4-97

Hydrogen 0-88 ()-88

Using junnioiiiated solution of barium chloride in.^tead of baryta water,
he got nearly the same results.

Baritdii potassiimi inudo.sulplionatt'. — This is a, nearly insoluljle,
crystalline salt, prepai-able by heating dipotassium hydrogen imidosul-
phonate with baryta water. It has been described by Berglund, and
was known to Freni}'. We have not analysed it, and Ijelieve it has
not been analysed by others.

Inirliim .soditiin ivtidosidjilwnate. — Barium chloride, in dikite
solution, very slowly added with constant stirring to an excess of
trisodium imidosulphonate gives a crystalline precipitate, wliicli under
the microscope is not seen to contain any tribarium imidosulphonate
in admixture. It is alkaline to litmus, very sparingly soluble in
water, readily soluble in nitric or hydrochloric acid, and hirgely ])ut
never completely decomposed by ammonium carbonate. It contains
water, but does not noticeably lose v.eight even jit 120.° Heated
quickly to a higher temperature, it is dissipated as a cloud of barium
sulphate and gases. Five preparations made at different times show
close agreement in composition, with the exception of that first pre-
pared (which is entered in our note-book as not pure). The numbers
agree with those calculated lor Ba^Na^N, „(803)20, (Ulln),., wliich may
be written as 8BaXaX(S03)., OH, + Ba,i\,(SO,),. (OH,), :

Calc. a h (■ il I'

Barium 41-12 42-85 40-05 4117 40-45-41-07 40-85-40-91
Sodium 5-02 5-08 5-2(i 4-93 — 5-25

Sulphur 17-46 18-80 — — . 17-54 17-56

g^ DIVEKS A^"D H AG A.

The barium and .sodium were determined by caiitioiis ignition of the
silt alone, then with «ul[)huric acid, and l)oiling out with water. The
residue was weighed as barium sulphate, and the soluble inatter after
re-io"nition weighed as sodium sulphate. The sodium of d was lost.

O c5 J-

The sulphur was estimated l3y hydrolysis followed by precipitation with
barium chloride, but the barium sulphate was here weighed in two
quantities, that formed by hydrolysis and that l)y the barium chloride.

Calcium iiuidustdpli ouates, simple and doidile.

Normal ealciiiiii imldosidphoiiate, C-dJs.^Q^O-^i, is not forjned by
reaction between sodium or ammonium imidosul})honate and calcium
chloride. When a sc^lution of diammonium imidosulphoijate is treated
with pure, soft, calcium hydroxide in the calculated quantity, am-
monia is at once liberated and is all expelled by two or three eva}'ora-
tioiis of the solution to dryness. The siüt left dissolves and forms
concentrated solutions crystallising in transparent prisms. J]ut
although the salt dissolves at first freely, the last ])ortions are more
difficult to bring into soluti(3n, and it may be that water partly de-
composes the salt into the calcium hydrogen salt and calcium
hydroxide. The salt was not analysed, except that a calciLun deter-
mination was made, according to wdiich the crystals would have ten
molecules of w^ater to two atoms (jf nitrogen in the salt.

Calcium liijdroijen Imidosidplioiiate is obtained in radiating groups
of fine needles when diammonium imidosulphonate and the calculated
(piantity of calcium hydroxide are mixed, and the solution repeatedly
evaporated to expel all ammonia. The calcium hydroxide dissolves
quickly after it is added. The salt is soluble and has not been
analysed.

Calcium ammonii(iii imidosulphonate is a sparingly -soluble salt
obtained wlien the calculated quantity of calcium hydroxide is dissolved

IMIDOSULPHO .NATES. 85

in diammonium iiiiidosiilphonate aoliition, this quantity being in the
propoj-tion, Cii(OH)o : Am2HN(S03),,. The salt rapidly crystallises
after the dissolution of the lime. It has not been analysed.

Galcmm sudiuni iniidosidplioiiak'. — To a hot (-(jncentrated solution
of trisodium iniid(jsiilph()nate about the ciilculated quantity of calcium
chloride solution to form the tricalcium salt is added, and the mixture
left to cool. N^ot the tricalcium salt Ixit the calcium sodium salt
crystallises out in groujis of hard brilliant prisms. The salt can be
recrystallised, and thus purified if necessary. It is sparingly soluble
in c<jld water, and reacts alkaline. It only very slowly takes carbonic
acid from the aimosphere. Analysis corresponds to the composition
CaNaN(S03)2,(OH,)3—

Calo. Found.

Calcium 13-75 18-95 1407 —

Sodium 7-90 7-72 — —

Sulphur 21-99 — •22-21 —

Nitrogen 4-81 — — 4-88

The relative positions of the calcium and sodiimi are not very

certain. For sake of symmetry the sodium might be put in the

imide relation, but such a procedure has nothing to support it.

Taking the sodium to act as the more basylous element, the formula

becomes —

^N

'SOsNa

oa

SOsNa

Lead iiiiidosidphonaies.

A lead imidosulphonate was used l)y Jacquelain, and again by
lîergluad, to prepare imidosulphonic acid ; but we know of n(3
publication of particulars concerning the sail employed.

86

DIVERS AND H AU A.

Di sodium or other bibasic imidoHulphonate does not precipitate
with lead nitrate and only feebly with normal lead acetate. It is
therefore necessary to take at least either the trisodium imidosul-
phonate, or else a basic lead acetate, to get an insoluble salt in suf-
ficient quantity. This salt is always a basic or hydroxy salt, so that
its separation renders its mother-licjuor much less basic than the
mixed solutions employed. Hence the common retention of lead salt
and imidosulphonate together in the mother-liquors, and also the fact
that either one or the other must Ije present in them. In fact, only
by the use of a basic lead salt is it possible to throw all imidosul-
phonate out of solution, and only by using the trisodium salt is it
possible to precipitate all lead. Oxy-lead acetate and lead nitrate or
normal acetate precipitate different salts with trisodium imidosul-
phonate, but all the metal of either salt precipitated is lead. One
other ])oint to l3e here noticed as affecting precipitation, is the solu-
bility in solution of normal lead acetate of the lead imidosulphonates
insoluble in water. The solution is alkaline and freely precipitable by
carbonic acid, and m;iy be regarded as containing basic lead acetate
and the soluble lead hydrogen imidosulphonate. Altliough a super-
ficial examination of the reactions of imidosulphonates with lead salts
may suggest the view of their being ill-defined, fuller investigation
proves them to be quite definite.

The normal salt, Vh^l^.^C^O-^^^, appears not to exist. The first
addition of lead acetate or nitrate precipitates a basic salt with tri-
sodium imidosulphonate ; iind basic lead imidosulphonate treated
with small quantities of nitric acid dissolves as a whole to yield a
neutral or slightly acid solution, the undissolved part remaining
unchanged (p. 89).

The lead Injcli-oycii salt, ]:\1^(Ü0-^.2V]), appears to exist in solution,
but the attempt to separate it leads to decomposition. Its solution can

lMIDO«ULPHONATEÖ. g7

be prepared by treating' either basic salt with somewhat less than
enough dilute sulphuric acid, and decanting the clear solution from
the sulphate and undecomposed basic salt. The solution can be
preserved unchanged in absence of acetates for a very short time only.
It has a slight acid reaction with litmus. Left in a desiccator it
suffers decomposition into amidosulphonic acid and lead sulphate,
slowly at first, more rapidly as the .solution grows concentrated.
Heating the solution effects the same change. Alcohol gives a volu-
minous precipitate gradually giving place to a crystalline de{josit
]>artly adhering to the sides of the vessel, nearly insolul)le in \v:!ter,
and p.pparently sulphate.

Hemihiidroxji-lcad iiiiidosulplionate, HOPb]S[(80;.Pb()H)o — Lead
nitrate and trisodium imidosidphonafe in solution brought together
in widely varying proportions yield this salt as a precipitate, volumi-
nous at first, but soon becoming dense and granular. Disodium
imidosulphoiiate and s(jdium nitrate are the otlier pr(^ducts of the
reaction : —

3Pb(NO,)2 + 3OÎI2+ 4Na3N(S03)2 = (HOPb)sN(S03)o + 3HNaoN(S03)2

+ 6NaN0;,
The precipitation is closely quantitative. With the trisodium salt not
used in excess the mother-li([uors ai-e neutral in reaction with metliyl
orange.

ISTormal lead acetate may be used in place of the nitrate, hut only
with less perfect results. The best way to proceed is to mix the
solutions rapidly together in soinething approaching the right propor-
tions ; or the two salts may be ru))bed together in the solid state and
only then treated Avith water. In either of these ways, when the
proportion of either salt is not very many times greater than it should
be, a product of constant composition is obtained, which differs
however from the pure lead salt in having about one-seven ty-third of

88

DIVERS AND HAÜA.

the lead replaced by hydrogen. The weight of the precipitate
obtained from a given quantity of trisodium salt, is greater with lead
acetate than with tlie nitrate, and greater as the excess of acetate used
is greater. Tlie acetate mother-liquors, though alkaline to methyl
orange, are acid to phenol-] )hthaleïn, and after replaciijg the lead in
them by sodium, Iw ])reci[)i fating with normal sodium oxalate, can be
titrated h\ sodium liydroxide and phenol-] )hthaleïn as indicator. 'I'hus
tested these liquors j)roved to contain free acetic acid, and in increasing-
quantity with that of the acetate used, and, therefore, of the jDrecipitate
]3roduced. In these experiments |3articularly excellent normal lead
acetate was used. It ap]3eared from tlie ex]3eriments made that about
one-eighth of the excess of lead acetate used must react with disodium
imidosul])honate, a ])roduct of the main reaction, as already ]winted
out, the change being as follows : —

3Pb(OÄ^)2+ 3OH2 + HNa.N(S03\, = (HOPb)3N(S03)2-|- 2NaOÄ^ -1-4H(3.4^

The acetic acid tlnis generated, acting so as to reverse the reaction,
is ]irobably the cause of the very slight replacement of lead by hydro-
gen in the salt ]^recipi tated by acetate.

When the trisodium imidosul])liona.te solution is added by degrees
to the lead acetate, a gelatinous jjredpitate like lesid hydroxide forms
and then redissolves in the lead acetate. When enough of the
sodium salt has been added to cause a permanent jjrecipitate this is
somewhat slimy and only slowly becomes pulverulent, while the
walls of the vessel get coated with crystalline preci])itate. The mixed
])rt)duct is unfit for analysis. When the lead acetate is added
gradually to tlie sodium imidosulphonate, the ])recipitate also remains
^■olu!llillous and somewhat gelatinous, and is jirobablv not a ]3ure
|>V()(luct. When after adding not too much sodium salt, tlie liquor is
quickly filtered, it slowly yields a few brilliant crystals of what is

IMIDOSULPHOXATES. g 9

evidently, hoth from its nppenraiife nnd nn imperfect analysis, the
hvdroxy-lead salt.

The hydroxy-le:!d salt in perfect brilliant micr(3scopic prisms
can he obtained very pure by treating the more basic lead salt next
described, with little more nitric acid than that calculated to remove
tlie excess of lead. The nitric acid can be clearly seen to diss(:>lve a
[>ortion of more basic salt, and almost at once to deposit n crystalline
precipitate raid incrustation of the hemi-basic salt. If instead oi"
stopping the addition of nitric acid when enough has been used, more
is added, the crystalline salt proportionately dissolves without repre-
cipitating. The nitric solution is neutral to methyl orange, so that
the reaction is, in the dissolution of the more basic salt, —

(HOPb)3NiÖÜ3)2, Pb(OH)2 + 6HN03-HPbN(S03)., + 50H2 + 3Pb(N03),
and then, this solution reacting with a further quantity of the more
basic sîdt : —

2(HOPb)3N(S03)2, Pb(OH)2 + HPbN(S03)2-3(HOPb)3N(S03), + OH2
This, in a manner, can be shown by pouring, the nitric solution into
a large excess of hydr(3xy-lead acetate solution, when the hemi-
hydroxy-lead imidosulplionate also precipitates in the pure state.
There is also the f ict oljserved that the nitric liquor when soon poured
olf from undissolved more-basic salt contains much lead, whereas
when it is left over the undissolved more-basic salt it goes on deposit-
ing a crystaJline precipitate of the hemi-basic salt.

The modification of this process expressed by the equation —
8(HüPb)3N(Ö03)2, Pb(OH)2-fHN(S03Na),-H'2HN03 = 4(HOPb),N(S03)2

-—has also been quite successful, ]jy adding the nitiic acid very slowly
and shaking well.

The hemi-liydroxy-lead iuiidosulplionate is an anhydrous salt,
not counting its hyclroxyl, very })ermanent, losing nothing at 100°,

90 DIVERS AND HAGA.

insoluble in water, and scarcely, if at all, alkaline in reaction when in
contact with wet litmus paper.

A number of preparations by different methods have l^een
analysed with closely concordant results : —

Calc.

Found.

Lead

78-40

78-21-73-35

SuJphur

7-56

7-50

The lead acetate precipitates gave from 72-81 to 72'78 7o lead,
and from 7'52 to 7*65 7o «idphur. The sodium was determined in
some of these preparations, and was found to be no more than
0-075 7,.

The pure salt was also titrated with volumetric nitric acid and
its basicity found to agree with calculation. One gramme of salt took
48-71 CCS. of acid, instead of 47*28 ces. calculated to just dissolve it
and furnish a solution neutral to methyl orange.

Five-eight] I s-ox y -lead imidosulphonate, (R0Vl))^l^(K>i).).2, PbO or
Pb(0H)2. — This salt is precipitated on adding trisodium imidosul-
phonate to excess of basic lead acetate. The quantity of basic lead
acetate which is needed is very large. Trial taught us to use not
less than six molecules of hemi-hydroxy-lead acetate to one of tri-
sodium imidosulphonate. The calculated quantity is five molecules,
but an excess helps to keep sodium out of the salt. The reaction is as
follows : —

NaeNlSOg)^ + öHOPbÜAc = (HOPb)3N(S03)2,Pb(OH)2 + SNaOAc + Pb(0Ä^)2
and therefore it would be better probably to use a more basic lead
acetate. Using as we did, about six molecules of hemi-hydroxy-lead
acetate, the mother-liquor of the precipitate proved to be markedly
basic still, l^oth precipitate and mother-liquor were quite free from
sulphate. A very little sodium, but no acetic acid, was found in the
precipitate, after it had been well -washed. It is readily soluble in

IMIDOSULPHONATES. 9I

dilute nitric acid, insoluble in water. Its partial dissolution in nitric
acid leads to the formation of the hemi-hydroxy-lead imidosulphonate.
It has scarcely any action on moist red litmus with which it is left in
contact. It is insoluble in solutions of its mother-salts, and unaffected
in composition by them.

Heated even at 130° it loses no water. When sufficiently hot it
gives off, first water, then ammonia, then again water, and then it
blackens through formation of sulphide, and then it evolves sulphur
dioxide. The loss of water and ammonia is explained by the
equation : —

8(HO).5Pb,N(S03)2 = 6PbO + aPbSOg + -SPbSOi + NH3 + N2 + 60H
Then the les'.d sulphite becomes, as usual by the rising temperature,
partly lead sulphide and sulphate, partly lead oxide and sulphur
dioxide. At a fusing heat the sulphide and oxide would of course
react.

Two preparations were analysed, a and h : —

Calculation. Pound.

( H0Pb)3N (803)2, PbCiHsO <i h h

Lead 7(v81 76-71) 76-05 76-82

Sulphur 5-94 5-82 6-00 —

Sodium — ()-15 0-15 —

Behaviour of imidosulphonates as compound amines to other bodies.

We collect together here some facts already recorded in the paper
which appear to show that imidosulphonates form compounds in
which the nitrogen becomes quinquevalent as in ammonium salts.

First, in order of observation, there is the retention of one
molecule of water at 110° by the trisodium salt. The potassium salt
which crystallises with one molecule of water (Raschig) lias not been
examined in this regard. Next, there is the union of diammonium
salt with sodium nitrate to form a well-crystallised compound which

92 DIVERS AND HAÖA.

H. /SOsAm),
may lie formulated as J ; /^\ , aiid fhe similar compound

■(NO3K ^Na
of dipotassium imidosulphonate. Lastly, there is the moi-e-basic lead

salt described in the Inst part of the pi'evious section of this paper.

Tills may be written | )^^ or (H())N^ .if we,

(V \PbOH ^(PbOH)2

allowably by the an:dysis, reject or admit tlie presence of one molecule
of water.

Silver imidosid'plionates^ singk and double.

Silver nitrate and tri sodium imidosulphonate will yield three
different compounds, in which one. two, and three atoms of sodium
are replaced by silver. Such an nnnsna.l behaviour in the salt of a
polvbasic acid gives to imidosulphonic acid a special interest.

• The addition of siher nitrate in limited quantity onuses a white
precij)itate which redissolves to be ra])idly followed, unless the
solutions are very dilute, by the separation of another comp(3und.
This consists of interlacing fibrous crystals filling the liquid as a
felted mass, even when pi-esent in only small quantity. Tluis, the
wliole m:iv be well stirred and yet afterwaixls j^rove a])le to support
the stirrei" awuv from the sides of the Ijeaker, th(jugh the actual volume
of the salt, separated frojn its liquor and pressed, may be a small
fraction of the wlxjle.

Further additions of silver nitrate to the liquor and crystals
cause geuerally more of -the first white precipitate, and after that,
when in sufiicient quantity, the disap[)e;jrance of both the preci])itates
together. The new solution becomes almost immediately full of n
third precipitate, this time of very dense, h-u'd, minute, separate
crystals, which rapidly subside into a deposit that in appearanci^, in
grating under the stirrer, and in movement when disturbed, most

IMIDOSULPHONATKS.

93

closely resembles wlute sea-sand (silver-saud). A salt of the same
composition as this often occurs ns a htird, crystalline incrustation on
the sides and bottom of tlie beaker, when to the mother- H(juor of the
felted crystals a little silver nitrate is added.

(Some experiments with potassium imidosulphonate have o-iven
us salts apparently analogous to those got from the sodium salt, but
we have not further examined them).

Tn'argentum imidosulphonate.* — By adding the S(^dium salt in
limited quantity to silver nitrate, the white precipitate is also formed,
and is now permanent in its mother- liquor, This precipitate, which
is triai'gentum imidosulph<mate, is very httle soluble in water, and after
long v>ashing on the filter yields a milky filtrate. It is •> relatively
voluminous preci]>itate forming chalk-like masses when drv. It is
without water of crystallisation, as was to be anticipated, but even
wlien kept long at 110° retains 0*5 5 per cent, of water (possiblv fixed
by hydi-olysis from the atmosphere), ;is determined by combustion
with copper oxide. It bears heat well, but at a comparatively high
tem])eratui'e rapidly decomj^oses with or without fusion, giving in
the open tul^e first ;i very little ammonia g;is and sublimate, then
sul])hur dioxide, nitrogen, silver, silver sulphide, and silver sul])hate.
When heated in a vacuum, scarcely any sublimate is fornied. and no
change takes ])lace till the temperature reaches 440°, when gas comes
oif, and blackening without fusion occurs. The residue consists of
silver sul])hate, siher sulphide, and silver, and the gas of nitrogen
and sulphur dioxide. The proportions of these ])roducts vary a, little,
and a])parently accoi'ding to the mode of heating, a iiigher temperature
giving more silvei- and sulphur dioxide. The first moietv of gas Avas
thus found to consist of four volumes of nitrogen to fi\'e of sul{»hur
dioxide, and the second moietv of two volumes of nitroo-en to thi-ee oï

* May possibly have been already described l>y Berf:?lund in the Lunds univ. Arsk. 12.

94

DIVERS AND HAGA.

sulphur dioxide, so that the decomposition lies between those expressed
by the equations : —

4AgN(S03Ag\,=2N2+2SO., + 5>^(),Ag,+ 8Ag2
4AgN(S03Ag),= '2N2 + 4S0, + 4SO,Ag2 + t^Ag^,

but nearer the iirst one.

The salt was pre])ared for analysis Iw addino- the sodium salt
gradually, with stirring, to excess of silver nitrate. In spite of the
strongly alkaline reaction of the former, the mother- liquor of the
precipitate proved to be neutral, as it should be, while the precipitate
was not in the slightest l)ro\vii, but brilliantly white. Being unde-
composed by \Yater it was washed. Its analysis gave, besides the
0*55 per cent, water mentioned above, the following results : —

Ai^^NiSOgAg).

Foiin<l.

Silver

65-06

64 92

64-53

64-58

Sulphur

12-85

—

—

12-94

Nitrogen

2-81

3-16

Diargentuin sodimn imidosiilphonate. — The sandy precipitate, de-
scribed above, is a salt with two atoms of silver to one of sodium. It
is sparingly soluble in water and very slightly decomposed by it,
silver hydroxide being one of the products. It has a faint buff colour.
Its crystals are seen under the microscope to be hexagonal plates,
always single, which rotate polarised light. This salt, though other-
wise anhydrous, retains a little water with the same firmness as that
shown by the wholly silver salt. Its behaviour when heated closely
reseml)les that of the latter, except that, naturally, sodunn sulphate
forms }\art of the residue, as well as silver sul|)hate.

To prepare the salt for analysis, the sodium salt was added
gradually to a little less than two-thirds of its equivalent of silver
nitrate. The sandy precipitate obtained was only slightly washed,

'IMIDOSULPHONATES. 95

•Hid then drained and dried on a tile. An incrustation (washed in
situ) was also analysed. The results were : —

Sodium

AgaXaNCSOg)^

5-57

Incrust.

5-()l

Precip.

5-67

Silver

52-30

5 2-02

51-S7

Sulphur

15-50

15-()0

Nitrog-ea

3-31)

3-45

Argentuiii disodiuni imidosnlphonate. — This salt, which occurs as
crystalline fibres, is the most soluble of the three salts, but is still
only s{)aringiy so. It is gradually and visibly decomposed by water,
the change consisting apparently, partly in the formation of silver
hydroxide and hydrogen disodium imidosulphonate, partly in resolu-
tion into trisodium salt and the diargentum sodium salt. Although
readily produced it is, therefore, difficult to get pure. It retains a
very little water, as usual, and behaves when lieated almost like the
preceding salt. For analysis it was pressed froni its mother-liquor on
a calico filter, and then between ])orous tiles. In preparino- it the
sodium salt was added to a little less than one-third of its equivalent
of silver nitrate, and the ndxture agitated in order to redissolve the
white precipitate of silver salt first thrown down before the fibrous
crystals began to form. Analysis gave —

Ag-XasNlSOg).^ a h c

Sodium 14-02 14-75 — — 13-81

Silver 32-93 33-07 33-49 33-50 33-49

Sulphur 19-51 19-18 19-52 —

Nitrogen 4-27 — — 4-29 —

Double mcrcurtj iinidosulphonates.

Single mercury imidosulphonates seem to be inca])able of ex-
istence.

96

mVEES AND HAGA.

Mercurij jtoiassiimi iiiiidosidphonafes. — Berglund jR-epared mercury
dipotassiiun imidosulplKjnate, Hg]Sr2(S03K)4, from tripotassium imido-
8iil])lionate. We liave prepared it from dipotassmm imidosulplioiiate
by adding mercuric oxide to it and heating the mixture with water,
filtering hot if necessary, and Jen vi ng the solution to cool. The salt,
which is nearly insoluble in cold water, separates out in very minute
needles, sometimes arranged loosely, sometimes in s[)herical tufts, and
is when dry of silvery lustre.

Au oxyinercKfic potassium salt, analogous t(j the sodium sîdt
fp. 105), seems to be umong the products of the reaction between mer-
curic nitrate and potassium imidosulphonates, but we have not made
any study of it.

The behaviour of mercur}' dipotassium imidosulphonate Avith
acids, as observed by us, is of interest in connection with the theory
of "the constitution of these salts, and we shall refer to it later on when
discussing that point ([>. 110). When the salt, in a state of paste
with water, is treated with nitric acid, it is converted into the very
slightly soluble dijMjtassium imidcjsidphonate, all its mercury dissolv-
ino- as mercuric nitrate. Washed and drained on a tile, the salt left
undissolved has been found to be free of mercury and of sulphate.
The same experiment can l)e carried out by the use of dilute «idphuric
acid, but less satisfactorily. The nitric acid used may be strong and
in excess, but i]ni sul|)huric acid nuist be dilute and not in excess, and
is apt to give a small precipitate of mercuric salt. Hydrochloric acid
exchana'es hydroiten foi" both metals, and effects extremely rai)id
hydrolysis.

Mercunj Iniriiini, HgN.(k5(J:i).J)a., and other double niercunj iiuidosul-
plionaies. — Mercury dipotassium imidosulphonate gives, according to
Berglund, a series of doubles salts in wiiich the potassium has been
replaced by another 1)ase. It foll(jws that tliese salts can also be

IMIDOSULPHONATES. 97

prepared from the mercury sodium salts, but we have only made
ourselves familiar with the mercury barium salt, particularly noticed
by Berglund. It is a lustrous, crystalline, dense salt, almost insoluble
in cold water.

Mercuni (Hhiiayotjcn imidosulphonate. — When mercury barium
imidosulphonate is treated with dilute sul})huric acid, not in excess of
the barium, it appears to be sbirply converted into liarium sulphate
and mercury hydrogen imidosulphonate. This observation made by
Bero'lund we have confirmed and extended. We worked quantita-
tively and were careful to use the sul})huric acid in slight deficit, and
to filter quickly, \'erv easy to do in consequence of the dense state of
the barium sulphîîte. But already hydrolysis of the imidosul])honate
into sulphate and amidosul]3honic acid had begun when we tested the
filtrate. It progressed steadily, so that in ten minutes the sulphuric
acid had grown to a considerable quantity, showing that mercury
hydrogen imidosulphonate is stable to a less degree even than sim])le
hydrogen imidosulphonate. In about ten minutes also, the solution
began to grow turbid, and let fall an oxymercuric unit, probably
amidosulphonate. The fresh solution when almost neutralised with
potassium hydroxide gave a slight white turbidity, doubtless of oxy-
mercuric potassium imidosulphonate, which dissolved when a little
more alkali was added. The slightly alkaline solution, thus prepared,
soon began to deposit minute crystals which we fully identified as
mercury dipotassium imidosulphonate. Thus, we had proved, so far
as might be done, that the fresh filtrate from the barium sulphate is
really a solution of mercury hydrogen imidosulphonate.

Oxijmercuric hiidrcxjeu imidosulphonate. — A second salt of mercury
and hydrogen can be readily obtained, which is at once ])asic and acid
salt, as its name indicates. Highly concentrated mercuric nitrate
solutions, even when free as possible from nitric acid, dissolve, to clear

98

DIVERS AND H AG A.

and comparatively stable liquors, solid potassium or sodium imidosul-
phonates. When sufficiently he;ited these liquors allow the imidosul-
phonate to hydrolyse into amidosulphonate, but ^^'hen, without heating,
they are diluted, they deposit oxy mercuric hydrogen imidosulphonate.
The same salt is almost immediately precipitated when tri.sodium
imidosulphonate in solution is added to mercuric nitrate.

To prepare this salt take about five parts l)y weight of mercuric
nitrate solution, undiluted and equal to about half its weight of mere
curie oxide, and pour into it with stirring a cold solution (necessarily
dilute) of one part of trisodium imidosulphonate. Precipitation of th-
snlt begins almost at once and is finished in a few minutes. The
mother-liquor contains still nuich mercuric nitrate and some imidosul-
phonate, besides sodium nitrate and much nitric acid. Addition of a
little more trisodium salt causes scarcely any more precipitation, while
that of a large quantity throws down the oxymerciu-ic sodium salt.
The brilliant w^hite and voluminous oxymercuric hydrogen salt hydro-
lyses only very slowly in its acid mother-liquor, because of the in-
hibitory action of the mercuric nitrate. It is to be washed repeatedly
by subsidence and décantation with abundance of cold water and
drained till dry on a tile or filter. Where less mercuric nitrate has
been taken, and the precipitate contains sodium (as oxymercuric
sodium salt), either digestion for a, day with concentrated mercuric
nitrate solution and washing, or else, without use of mercuric nitrate,
continued and thorough washing with water will convert the
precipitate into the ])ure oxymercuric hydrogen salt. After digesting
with the mercury nitrate, the first washing waters used must con-
tain a, little nitric acid to guard against the formation of any
oxynitrate.

Oxymercuric sodium imidosulphonate when long washed with
much water leaves a much smaller weight of oxymercuric hydrogen

IMIDOSULPHOXATES. 99

salt, and imparts coiirmiKJUsJy to tlie \Nasb-waters «mall quantities of
a mercury sodium imidosnlphonate — aj)[)ai'ently the same as that got
by digesting mercuric oxide in a sohition of mercuric disodium imido-
snlphonate and perhaps 0[HgN(S03]Sr;»),,]^,. These waters are neutral
or faintly alkahne, and when eva.porated a- little on the water-bath
yield small qusmtities of micaceous crystals. This reaction appears to
be expressed by the equation : —

2Hg( OHgS03)2NHgN(803Na),,+ UHo- OLHgNlSOsNa)^,

-f2HN(ö03HgO),,Hg.

Oxymercuric hydrogen imidosuljjhonate is an anhydrous salt.
Already dried in an ordinary desiccator, it loses, when further dried at
100° or above, 0'7 to 0*8 per cent, of water. It may be heated in
dried air to 180° or higher Avithout changing. Only a little below a.
dull red heat does it decompose, and then slowly gives water, nitrogen,
and sulphur dioxide. At this temperature it is yellow, but Avhen
cooled it resinnes the white colour of the undecomposed salt. At the
softening point of hard glass it melts to a, dark red liquid and eifer-
vesces, yielding sublimates of mercury metal, mercurous and mercuric
sulphates, and some other mercuric salt not sulphate and apparently
nitrogenous. It seems impossible, even in a vacuum, to decompose it
completely before the mercury sulphates themselves partly decompose.
The first o-avses sfiven off consist of nitroîxen with half its volume or
more of sulphur dioxide — those coming after still contain nitrogen along
with sulphur dioxide and oxygen. When rapidly raised to a red heat,
the s:dt, at the moment of melting, effervesces almost explosively.

Its reactions with sodium hydroxide and chloride, and with
trisodium imidosnlphonate are similar to those of oxymercuric sodium
salt ([). 106). Its basic or oxysalt character is thus clearly demon-
strated, independently of the evidence from its quantitative analysis.

lOÔ

DIVERS AND HAGA.

The presence of hydro^-en, not merely as water of hydration, is shown
by first heating the salt for hours at 1<S0° in a current of dried air, and
then afterwards getting, at a nuich higher ftemperatu re, water vapixu-
condensed (containing sulphuric acid). As the salt is calculated to
yield only a little more than one-thousandth of its weight of hydrogen,
and the isolation of the water from all mercury would be difficult,
hydrogen was not quantitatively estimated.
The analytical results obtained were : —

HNtSOglsHggOa

(a)

(b)

Merciu'v

74-35

74.03

74-01)

Sulphur

7-93 .

8-06

8-00

Nitrogen

1-73

1-76

l'6o

Hydrogen

0-12

(Sodium)

0-06

0-06

(Moisture)

0-20

0-13

The preparation, (a), was made by the principal and direct
method. The other, (b)^ was digested with mercuric nitrate to remove
sodium which it at first contained, and then washed with water
slightly acidulated witli nitric acid.

On writinof down the reaction bv which the salt under considéra-
tion is formed from mercuric nitrate and trisodium imidosul{>honate: —

8Hg(N03),-l-Na3N(S(),)2-F20H2=HN(S03)2HgA+-^NaN03-i-8HNÜ3

— and comparing it with the equation wliere oxymercuric sodium
imidosulphonate is the product, it will be seen that in both cases the ni-
trate comes out half as sodium nitrate and half as nitric acid, and that
the oxymercuric hydrogen salt may be represented as resulting from
the reaction of the sodium salt with mercuric nitrate, thus : —

Hg(N03)2+NaN(S03)2Hg2() + OH2 = HN(SÜ3)2Hg302-hNaN03-|-HN03.

IMIDOSULPHOXATES. IQl

Now this reaction can actually be realised, as already stated, but only
with difficulty :!nd in the presence of great excess of mercuric nitrate
solution, active probably by virtue of its free acid. In the direct
method of preparing the hydrogen salt, it is formed simultaneously
with half as much again of nitric acid as is produced when the sodium
salt is fanned. From this it would appear that there is a point —
difficult to determine by direct experimentation with nitric acid,
becjiuse of its liability to cause hydrolysis after a time, and of the
uncertainty in knowing when the sought-for change has occurred —
where, the nitric acid being in sufficient quantity along with the
mercuric nitrate, nitj-ogen takes or keei)s hydrogen in place of mercury,
and the second half of the sulphonic radicals, as well as the first, takes
mercuric oxide in place of sodium, just as it does in other cases even in
presence of free acid.

Mercury sodinni inridosulplionates. — There are two mercury sodium
imidosulphonates. The monosodium salt is oxymercuric or basic, but
tlie disodium salt is normal, and corresponds in composition with the
potassium salt obtained Ijy Berglund.

unless gradually added, mercuric nitrate causes, almost immedi-
ately, a white crystal !o-flocculent precipitate in solutions of the
trisodium imidosulphonate, which disap|)ears on agitation so long as
enough of the sodiinn salt remains to keep the mixture alkaline or
neutral. When this point is passed, preci])itati()n is pei'manent, and
ox ij mercuric sodiuiu salt is tlie product. When the nitrate is only added
till neutrality is reached or nearly reached, the liquor soon begins to
form small brilha.nt crysttils, or else will do so after some evaporation.
These crystals are normal mercuric disodium imidimdjdioiuite. Adding
solution of trisodium imidosnlphonate to that of mercuric nitrate, free
from any unnecessary excels of nitric acid, causes again, as already
stated, a white crystal lo-tl<3Cculent ])recipitate which, with sufficient

102

DIVERS AND HAGA.

excess of iiitrnte rem-nuing, is oxijmercunc Injdrugeu salt. With too
Jittle nitrate remaiiiiiig, some oxyiiierciiric sodium salt will be
deposited. Mercuroiis nitrat.^ behaves in the main like mercuric nitrate
but gives besides a j)recipitate of the metal.

Mercuric disodiuiii hiiidosiiljHioNaie. — Little more needs to l)e stated
concerning the preparation of this salt from the trisodium salt. Here,
as in other cases, the mercuric nitrate solution is preferablv to be
highly concentrated, because then the excess of nitric acid necessary
becomes very small. To form such a solution, moderately concentrated
nitric acid should be so far saturated with mercuric oxide that oxyni-
trate beg-ins to form, and then decanted from excess of oxide and left
to clarify by subsidence. When too much mercuric nitrate has been
added to the s<^dium salt, the proportions can be rectified quite suc-
cessfully by adding more sodium salt. The mercuric disodium salt
can be purified if necessary, liy recrystallisation from hot water.

Mercuric disodium iinidosulphonate can also be readily prepared
from disodium imidosulphonate and mercuric oxide. The two sub-
stances may be triturated together in al)Out the right proportions
mixed with water, and then warmed witli it. The solution is to be
filtered if necessary, and then set aside to crystallise.

The crvstals of mercuric disodium imidosulphonate are small bril-
liant prisms, always separate, quite permanent in the air, and spar-
ingly soUible in cold water. The solution has a, neutral reaction.
The crystals contain six molecules of water of which only four are lost
in a vacuum at common temperatures.

Heated to 100°, after exposure' in a vacuum desiccator, it loses most
of the remaining water, but not all, tor then (and at higher tempera-
tures, such as that of 130°, more quickly) it also increases slowly in
weight by fixing atmospheric moisture, becoming hydrolysed and
thereby strongly a(nd (see effects of heating dipotassium imidosulpho-

TMIDOSULPHONATES. IO3

nate, p. 64). Hen ted more strong-ly in an o])en tube, it yields a
small sublimate of an ammonia-sulphite salt, Jiiercurous sulphate, and
mercury, along with sulphur dioxide and nitrogen as gases, and
mercurous, mercuric, and sodium sulphates as residue.

Heated slowly in the vacuum of the Sprenii'el-pump, it suffers
change in a way that can be more closely studied. Even at 444:° no
material alteration t tikes place in the salt, but just below a red heat, it
decomposes steadily, temporarily blackening through formation of
mercuric sulphide, and giving much mercury as a sublimate, nitrogen
and sulphur dioxide in the proportions of two volumes of the former
to three of the latter, a little, very volatile, white ammoniacal sublimate
along with a very little water, and another white sublimate volatilising
sigain at o5*)-4()().° The residue is sodium sulphate. The margin,
remote from the heai, of the sublimate last mentioned consists of mercu-
rous sulphate, but the rest of this sublimate, which adheres to the glass
firmly, is of a peculiar nature but imperfectly made out. It is, how-
ever, a mercury compound scarcely affected l)y potassium hydroxide,
boiling water, or dilute nitric acid, and is api)arently a nitrogen-hold-
ing derivative of mercuric sulphate.

The main changes by heat appear to l)e —

'2HgN2(S03Na),=4SO,Na2-l-HgSO,-l- HgS -H 2SO2+ 2N„
and then by further heat —

4S04Na2+2Hg + 4S02-t-2N2.
The ammoniacal sublimate and moisture are evidently due to water
retained by the salt, prol:)al)ly tin'ough hydrolysis ; while mercurous
sulphate will have come from mercury and mercuric sulphate.

With soluti(jns of ordinary mettiUic salts, the mercuric disodium
salt gives the various double mercury imidosulphonates noticed by
Berghmd. For exam[)le, with barium ch'oride it gives a precipitate of
mercury Ijarium imidosulphonate.

JQ4 DIVERS AND HAGl.

Sodium hydroxide precipitates mercuric oxide from the pure salt,
hut not in the presence of sodium imidosuJphonate, and under any
circumstances the precipitation of the mercury is far from complete.
According to Raschig, Berglund foimd that mercuric di])otassium
imidosiilphonate gives no precipitate with potassium hydroxide, hut
we find that, in this res|)ect, the ])otassium salt behaves like the sodium
salt, except that the precipitation is perhaps less. Dilution lessens
precipitation (see behaviour of mercuric oxide with trisodiiim imido-
sulphonate, p. 73).

Anmionia gives a white precipitate. So, too, does amnioniiun
chloride, whir]i in this case is probably amidomercuric cliloride.
Mercuric oxide dissolves slightly in solution of the mercury sodium
salt, rendering it alkaline. Nitric acid and, still more so, hydr(3-
chloric acid dissolve the salt freely. Nitric acid does ncit immediately
decom]3ose it ap])arently, but the action of hydrochlorie acid effects
the complete decomposition of the salt. If the (juantity of this
acid is insufficient for the whole of the salt, its action is confined
to its equivalent quantity, the rest of the salt being left undissolved,
and no preferential replacement of the sodium or the mercury Ijy
hydrogen taking ])lace. By extraction with ether, by evaj »oration,
and by other tests, the change effected has been ascertained to be the
formation of mercuric sodium chloride, sodium acid sulpliate, and
amidosulphonic acid.

Analysis has given the following results : —

HgN2(S08Na)„(OH2)e Found.

Mercury 2&74 27-16

Sodium 12-3() 12-35

Sulphur 17-11 17.19

Nitrogen 3*74 3-9â

Water 14-44 13- Ifjj^'^^ ^""'^ '" '""■

IMTDOSULPHONATES. JQ5

The exposure in the vacuum wa.s for 40-45 hours. The furtlier
loss at 100° was the o-reatest obtainable in undried air, lons"er heating-
being followed very slowly by increase of weight. As analysed, the
salt shows nearly a tenth less than the water calculated. Efflorescence
of the sample was not noticed, but may have occurred to a slight
extent. The main cause of the deficient finding is without doubt,
however, fixation of some of the crystallisation- water or of atmospheric
moisture by hydrolysis.

Oxjimercuric sodinm imidosulphonate. — In the preparation of this
salt, when the sodium imidosulphonate is taken in double molecular
proi)ortion to the mercuric nitrate, that being the calculated proportion,
the process works well. The highly acid motlier-liquor retains much
of the salt in solution, but will let it fill if nearly neutralised with
sodium hydroxide, nnd then contains scarcely any other mercuric or
imidosulphonic salt. There is, however, no necessity to adhere closely
to the calculated proportions. Provided that the sodium salt is neither
in quantity great enough to redissolve the precipitate as mercuric
disodium salt nor so small as to leave the nitrate in much excess, the
process will succeed ; but it is better to use too little than too much of
the nitrate, above .all should its solution contain any quantity of free
acid. Any precipitation which water alone would cause of oxynitrate
in the solution of mercuric nitrate, is prevented by the presence of the
sodium imidosulphonate, since this salt forms its basic mercury
derivative by liberating half the nitric acid of the nitrate that it de-
composes, and this acid is much more than enough to keep any
remaining nitrate from passing into oxynitrate, and, consequently, the
washed precipitate proves to be always free from nitrate. It may be
washed sufficiently with water, which only very slightly acts upon it,
and may then l)e dried, either on paper or better on a tile.

On attemj)ting to form a third and intermediate mercury sodium

IQQ DIVERS AND HAGA.

salt, which was to have the formula, 0[FIgN(S03]sra)2]2, analogous to
the oxynitrate, 0(HgN03)2, we got, instead, only the other two salts,
one in solution, the other as a precipitate. The oxymercuric sodium
salt was thus obtained from a liquor which from first to last was
never acid, even the mother-liquor being still faintly alkaline. The
sulphonate and nitrate were used in the proportions — 2Na]Sr(S03]S[a)2
to Hg(N 0.3)2 ; the nitrate was added gradually, with stirring, to
the dilute solution of the sulphonate, and simultaneously a solution of
sodium hydroxide run in in the proportion of nearly five-sixths of a
molecular quantity. By this procedure, much yellow mercuric oxide
was formed along with the white precipitate, but by prolonged stirring,
the precipitate lost all tinge of yellow. The mother-liquor now con-
tained much disodium mercury imidosulphonate, and the precipitate,
washed and drained on a tile, ])roved to be pure oxymercuric sodium
imidosulphonate (analysis, d). Allowing for sodium hydroxide used
in neutralising the free nitric acid present in the mercuric-nitrate
solution, the reaction appears to be that expressed by the equation : —

6NaN(S03Na)2 + 6Hg(N03)2 + 4NaOH=
= 2HgN2(S03Na),-fHg(OHgS03)2NHgN(S03Na)2-t-12NaN03+2H20.

Oxymercuric sodium imidosulphonate contains water of crystal-
lisation and appears to be a little efflorescent ; otherwise, it is for a
long time permanent, wet or dry, though ultimately becoming yellow-
brown })y decomposition. Protracted washing with water decomposes
it, as does als(3 digestion with concentrated mercuric-nitrate solutions,
a residue being left in both cases, which is the hydrogen oxymercuric
salt (p. 97 ). It is very stable when but moderately heated, only
losing its water, with the exception of a very little. At about 1 35"
it permanently changes colour shghtly, and at a much higher tempera-
ture melts to a dark red-brown liquid and effervesces, decomposing in

IMIDOSULPHOXATES. IQ7

a way essentially the same as that followed by the normal mercury
sodium salt. Most gradual heating in a current of dried air to a
temperature of 170° fails to expel more than four-hfths of the water,
in consequence no doubt of hydrolysis.

It is much more readily dissolved by hydrochloric acid tlian by
nitric or sulphuric acid. From its hydrochloric-acid solution it can not
be recovered by neutralisation, being almost instantly decom])osed, like
the normal mercuric sodium salt. It is converted by sodium hydrox-
ide into mercuric oxide insoluble, and mercuric disodium imidosid-
phonate dissolved. Its basic composition is at once demonstrated by
the action on it of sodium chloride, which leaves insoluble mercuric
oxide, and dissolves the rest, probably, as the two salts, mercuric
sodium imidosulphonate and mercuric sodium chloride. Trisodium
imidosulphonate dissolves it, but not to a large extent. When tlie
solution of this salt is concentrated, a little mercuric oxide may separate,
but enough water added causes this gradually to dissolve. When the
trisodium salt is added to the oxymercuric sodium salt still in its
mother-liquor, free dissolution at once occurs; but without the mother-
liquor the reaction is as above stated.

The composition of the salt is expressed, as the following analytical
results show, by the formula — OHg2N(S03)o]Sra,(OH2).2— which has,
however, to be doubled to display its constitution. The water comes
out low, partly because of loss by efflorescence, but mainly by getting
•fixed through hydrolysis.

Calc.

UO

(fl)

('')

(<■)

('?)

Mercury

61-68

62-18

61-22

61-56

—

61-73

Sodium

8-54

3-87

4-14

8-97

8-57

8-56

Sulphur

l)-8(i

10-28

10-06

10-08

—

9-79

Nitrogen

2-16

—

2-27

2-24

—

■ —

Water.

5-55

—

4-44

—

4-88

4-58

1Ö8

Divers and haga.

The- preparation, (a), was precipitated from three molecules of mercuric
nitrate by two of imidosulphonate, while (h) was got by adding sodium
hydroxide to the decanted mofher-liquor of (a). The preparation,
(c), was precipitated from two molecules (3f nitrate by one molecule of
imidosulphonate, and (d) was formed in a non-acid mother-liquor.
All preparations, as well as their mother-liquors, were free from
sidphate.

The formation of oxymercuric sodium imidosulphonate is ex-
pressed by the equation : —

4Hg(N03)2 + 2NaN(S03Na)2-|-20H2=Hg(OHgS03)2NHgN(S03Na)2 +

4NaNÜ3+4HN03.

This reaction is noticeable for being one in which sodium is withheld
in the precipitate from nitric acid ; but, if imidosulphonic acid be re-
garded as a weaker acid than nitric acid, the precipitation of potassium
nitrate by tartaric acid or perchloric acid is quite as remarkable, while
if, as is most probable, it is like sulphuric acid, then the retention of
sodium from the nitric acid is only natural. It is hardly noticeable
for the precipitation of a b;isic salt \v\t\\ production of nitric acid, since
similar reactions are connnon with oxygenous mercuric salts.

In its constitution, as regards the oxylic mercury, oxymercuric
sodium imido8ul})honate resembles oxymercuric sulphate and oxymer-
curic sulphite, as the following • formula display (J. Coll. Sei., t,
101):—

Imidosulphonate (NaO'SOolg iN-Hg-NKSOs-O-Hg-O), :Hg

Sulphate HgilO-Hg-ü-Süg-ü-Hg-O)^ :Hg

Sulphite Hg:tSÜ2-0-Hg-ü)2:Hg

Constitution, of the Mercinif Jmidosulplwnates.
From the widely established character of the relation between

IMIDOSULPHONATES } Q f)

mercury and the nitrogen of ammonia and cyanogen, and from that of
the mercury in its oxygenous salts, particularly those of sulphuryl, such
as sulphites and sulphates, the relations of mercury in its imidosulpho-
nates become of much interest, as likely to vary from those of other
basylous elements.

Firstly, as to the mercury disodium and mercury dipotassium
imidosulphonates, no hesitation will be felt in accepting for these salts —
say that of sodium — the formula, HgN2(S03Na), (l^erglund), yet this
point is not so simple as it seems. Calcium forms the salt, CaNa]S[(S03)^;
silver forms the salt, Ag2NaN'(S03),, ; and mercury itself the salts,
OHg2NaN'(.S03)2 and 0.,Hg3HN(S03)2. All these salts liave only one-
third of the bases either sodium or hydrogen, and it will be well, there-
fore, to briefly review the reasons for writing Hg]Sr2(S03Na)4.

Disodium hydrogen imidosulphonate is a salt neutral or only
slightly acid to litmus, although active as an acid, and tlierefore, it
cannot hold tlie group, SO3H, since this aPways gives to its com-
pounds sourness and strong action on blue litnnis. It must therefore
be written HN(S03Na),. Mercury readily takes the place of the
hydrogen of this salt, or of one atom of the sodium of the alkaline
trisodium imidosulphonate, and the resulting mercury disodium salt is
neutral in reaction. Transposition of the metals cannot be admitted
to take place in its formation for two reasons. One is that the alkaline
reliction of the trisodium salt disappears when it becomes the mercury
sodium salt, and there is no accounting for this if the mercury dis-
places the sulphonic sodium. The other reason is that if the mercury
takes the sulphonic relation in the salt, there is to be seen in this salt
an exception to the observation that all oxylic mercuric salts existent
in presence of water are insoluble basic or oxysalts.

This theory of the constitution of the normal mercury double
sulphonates accords with the interesting behaviour of these salts

no

DIVERS AND HAGA.

towîirds acids and alkalis. The latter only partially precipitate
mercuric oxide from them, because the mercury is in immediate
relation with (unoxidised) nitrogen. Nitric acid, which can replace
the mercury by hydrogen, cannot do the same with the potassium or
sodium, because this is in oxylic relation with sulphuryl, as it is in
sulphates, and therefore irremovable by this acid. Sulphuric acid also
first replaces the mercury by hydrogen, when acting on the mercury
potassium salt, although from the mercury barium salt it takes first
the barium away.

Berglund would have chemists regard mercury imidosulphonates
as salts of an independent acid, in which mercury is combined with
special force. A fuller knowledge of the imidosulphonates does not
tend to support him in this. There are other series of double imido-
sulphonates besides that of mercury, ai)parentl3' the only one observed
by him ; normal mercury hydrogen imidosulphonate has less stability
than imidosulphonic acid itself ; oxymercuric hydrogen imidosulpho-
nate is a far more stable body than it ; dipotassium and disodium
hydrogen imidosulphonates have equal or greater claims to be treated
as particular acids ; lastly, it is highly probable that, powerfully as
mercury takes the place of hydrogen in ammonia itself, it will have
little of that power when two-thirds of that hydrogen have already
been replaced by sulphonic radicals.

The constitution of the salt in which two-thirds of the base of the
trisodium salt are replaced by mercury next requires attention. It
mio'ht be treated as having the single atom of sodium in the odd
basylous position, that is, united to the nitrogen ; but against this
four objections present themselves. One is that it is quite against
probability that the sodium should hold the imidic relation rather than
the sulphonic. A second is that it is quite as improbable that a
mercury atom should be united half to nitrogen and half to oxygen.

IMIDOSULPHONATES. 1 1 1

while it is usual to find it thus united at once to oxygen and to an
oxyo'enous radical. A third objection is that, whereas in other cases
sulphuryl takes one-andra-half atoms of mercuric oxide to saturate
either of its valencies, it does not do so in this salt unless the sodium
is oiven to one of the sulphuryls of the salt. The fourth objection lies
in the great improbability that sodium in the imidic relation would
resist, as it here does, displacement by hydrogen on contact of the salt
with nitric acid. Avoiding these four difficulties by placing one-
fourth of the mercury with the nitrogen, there remain .the usual
one-and-a-half atoms to unite with half one of the sulphuryls. The
relations of the salt, and its constitution as here developed, require
that its formula should be double that expresed in the lowest terms, in
order to allow of the oxylic mercury being shown apart from the non-
oxylic— Hg:(OHgO-SÜ2)2:N-Hg-N:(S03Na)2.

It becomes now clear that the sodium resists the action of nitric
acid because it is in oxylic relation to sulphuryl, as pointed out in
discussing the constitution of the normal mercury disodium salt ; that
nitric acid removes a fourth of the mercury, replacing it by hydrogen,
because this fourth part is in relation to the nitrogen ; that nitric acid
does not remove the rest of the mercury l^ecause this exists as the oxy-
mercuric group found in oxylic relation with sulphuryl in sulphate
and sulphite, also then resisting the action of nitric acid ; and, lastly,
it becomes clear how it is that the oxymerciiric hydrogen imidosul-
phonate has quite consistently a constitution different from that of the
oxymercuric sodium salt, and how the one salt is formed from the
other. (Cf. Divers and ShimidzAi on Mercurij Sulphites, J. Coll. Sei.
1, 101).

In fact, the constitution of the oxymercuric hydrogen imidosul-
phonate follows obviously from the production of the salt by (acid)
mercuric nitrate in excels. From what we actually can observe in thç

112

DIVEES AND HAGA.

c;ise of the normal mercury dipotassium salt we know that nitric acid
of itself should act as represented by this equation —

Hg(OHgOS02)2NHgN(S03Na)j+2HN03 =

-Hg(OHgOS02)2NH + HN(S03Na)2 + Hg(N03)2

— replacing the imidic mercury by hydrogen, but not touching the
oxy mercuric group, in conformity with its inability in other cases
(sulphites and sulphates) to do so when the group is joined to sulphu-
ryl. In the .absence of mercuric nitrate this reaction is slowly followed
by hydrolysis of the disodium hydrogen salt through the unavoidable
excess of nitric acid, but in presence of mercuric nitrate hydrolysis does
not take place. The mercuric nitrate ünishes the formation, just
formulated, of the oxymercuric hydrogen salt, in the way shown l)y
the equation —
HN(S03Na)2+3Hg(N03)2+'20H2 = HN(S020HgO),Hg + 2NaN03 + 4HN03.

— from which equation it will also be sufficiently clear how the whole
chmo-e can be effected by mercuric nitrate without any addition of
nitric acid.

From the constitution given to this salt may be seen why it can,
so remarkably, be left for days in a nitric-acid solution of mercuric
nitrate without hydrolysing. For hydrolysis can only occur when
some of the sulphonic group becomes acid, or SO3H, and here tlie
nitric acid, especially in pre-sence of much mercuric nitrate, is power-
less to displace the oxymercuric group by hydrogen.

There remains now only to tabulate the three merciu'ic salts,
the constitution of which has been discussed, in order to bring out
their relations quite clearly, and particularly the intermediate relation
of the oxymercuric s<.dium salt.

IMIDOSULPHONATES.

113

Iiiiidosnlphonîite.

Mercuric disodium

Oxymercnric sodium
Oxy mer curie hydrogen.

Hg

Hg

Formula.

N(S03Na)2
■NlSO^Na)^

N(S03Na)2

N(S03ligO)2Hg

HN(S08HgO)aHg

Addendum.

Oxiia))iidosiilp]ionate.'i. — In our last paper, we had to call attention
to the discordance between some of the results of the work done by
otliers aiid by <)urselves on sulphazotised salts. We are glad to be able
to brino- in sup{>ort of the accnracy of certain of our own statements
the te-itimony of one of the other workers, Hr. Dr. Rascliig', who
wr«jte soon after the appearance of the paper on oxyamidosulphonates
tous request wlien next publishing to make known that he now entire-
ly agrees with our account of the decomposition of oxyamidosulpho-
nates bvcnustic alkali (J. Coll. Set., 3, 218), and withdraws his own
statement concerning it, which was based upon qualitative reactions
only.

Jour. Sc. Coll. Vol. VI. PI. I.

m m

m ffl

a H

4- +

M M

>/ V/

¥

U

m

la

m

X

^ BZl
^ 4-

l#^

%

é&

Bâ M

On the Anatomy of Magnoliaceae-

By
Sadahisa Matsuda.

Science College, Imperial University.

With Plates II-V.

Introductory Remarks.

The investigation of the present subject was begun in the
autumn of LSÜO, at the suggestion of Prof. J. Matsumura, and was
continued more than a year. During that time I received much
useful advice from him, and also from Prof. R. Yatabe, to both of
whom I am much indebted. The object of my researches was to find
out what anatomical peculiarities characterize the Magnoliaceas as a
whole ; what distinctive characters are presented by each of the dif-
ferent groups included in it ; and to what extent all the species of
it ma\- be anatomically distinguished from one another.

Tliese questions naturally present themselves, if we remember
that systematic botanists in recent times have not agreed as to the
limits of this family, and have treated its members in different ways.
Some botanists extend its province by including certain tribes not
usually admitted, while others restrict it by omitting triljes coui-
nionlv included. 8iich indefiniteness as to its extent shows that this
natural family is not very natural, and I hope that the study I have
made of its anatomical characters may Ije of some service in remedying
this defect.

IIG

s. MATSUDA.

I examined all the species of Magnoliacete that were aece.ssiljle,
including those placed in other families by some botanists. Sucli being
the case [ find it more convenient to use in my dissertation the term
Magnoliaceœ in its wider sense. It also best answers my ])urpose to
divide this family, as is done by liUerssen' and others, into four tribes ;
namely : —

Maguoliece, Scli izaiidrea',

Jlliciea', TrocJi odciulreœ.

The number of genera included in these tribes does not exceed
fourteen, orj the highest estimation, and that of the sjiecies known at
present ranges Ijetween seventy and eight}'. However, I could ex-
amine only twenty-four species and two varieties. Altlunigh the
number of the species examined is small when compared with that of
all the species known, yet those examined are distributed among ten
genera, which are in their turn distril3uted among the fjur tribes.
Therefore, I am perhaps right in believing that the anatomical
characters of the species I examined represent fiirly those of the
whole family.

With the exception of two dried specimens in the herbarium of
the Science College, which I wa-< allowed to examine, the materials for
my study were mostly obtained in the University lîotanic Garden in
Koishikawa. As I could get no specimens of the main root proceed-
ing immediately from the seedling, but only of younger sec(jndary
roots, I examined these branchlets in order to get some kn(n\ledge of
the anatomical characters of the young root, such as the arrangement
of the xylem-plates, &c.

It will be well, I think, to define at once a few words which
thouiih often convenient to use in describinn- the structure of the
plant-body, are yet somewhat vague in their meaning. " Sclerenchy-

1. Luerssen, Grundzüge der Botanik.

ON THE ANATOMY OF MAGXOLIACE^. HJ

riiatf^is ül)re" is used in the same sense as bast-fibre, and '• sclero-
blast" in that of stone-cell; when either indefinitely is to 1)e
denoted, the expression " sclerenchymatous element" is often used.
Bv "sclerenchymatous sheath" I mean the single mass of sclerenchy-
matous fibres, or the numerous isolated groups of them, which lie at
the external limit of the fibro-vascular bundles and form a rino-
either continuous or interrupted at intervals. I consider this sheath
to be ÎI part of the bundles, and not an independent structure lying
outside them. By " cortex" I mean all the tissues which lie outside
the cambial zone. That part of it which corresponds to the phloem
(including the sclerenchymatous sheath, when this exists), I generally
call the •' inner cortex "; and all the parts outside this, the "outer"
or "external cortex." In the case of very young roots 1 mean by
"cortex" all the tissue lying outside the endodermis.

In the following pages the anatomical characters of each genus
will be first described, and then com])ared. It might a,])|)ear sufficient
to have compared at once the characters of the different generu
without giving a- special description of each genus, but this would
haA'e led to confusion, as well as to the omission of many interesting
isohitcd facts.

Anatomical Characters.

Under the headings. Stem, Pclioh', Blade, and Boot, the points of
structure peculiar to each are descrilied, while under the first are
given also those points common to all four.

Tribe I. Trochodendreae.

This tribe consists of three genera, species of each of which I
examined.

113 s. MATSUDA.

Euplelaea.

E. polijaudra, Siel), et Zucc, is tlie only sjiecie.s of this genus
which I examined. It is a small tree, found in many parts of Japan,
in which the aromatic property so common in Mngnoliacea^ is entirely
absent.

Stem. — The epidermal cells present no peculiarity ; the cuticle is
not well developed. There is found on the e])idermis a number of
lenticels, which appear to the naked eye as white sjiecks. Cork is
developed inunediately beneath the epidermis, its cells being of
com])aratively large size, while those forming the other tissues are
generally very small. The hypodermn is represented by n layer of
somewhîit thick-walled parenchymatous cells bene;ith the cork.
Sclerenchymatous elements of any kind nre totally al)sent in the
outer cortex ; but there are found in it many sacs, each of which
contnins an aggregate of crystals of calcium oxîdate (PI. IT, Fig. 2).

The sclerenchymatous ring which accompanies the fibro-vasculnr
bundles is well developed, but is interrn|)ted at the points where the
large medullary rays run radially througli the ])hloëm (PI. II, Fig. 1).
Sclerenchymatous elements are absent in the inner phlorm ; btif
a few scleroblasts -find their wny into those jxn-tions of the ])hlo('m
rays which run near the sclerenchymatous ring (PI. TT. Fig. 1. .s).
In .specimens collected while the cambial zone is in activity, tliere is
seldom seen any marked distinction between that zone and the xyleni,
gradual transition taking place from lignified cells to uidignified ones.
(PI. II, Fig. 1). In the phloem |)ortion sieve-tubes are distinctly to
be seen, especially in longitudinal sections. The xylem contains
vessels, tracheïds, fibres, and wood-|)arenchyiua. The vessels general-
ly have fibrous markings on their walls. In those portions of the
xylem whicli border on the pith and constitute the medullary sheath
there aiv found, in yonng s])eciinens, grouj)s of elongated cells with

ON THE ANATOMY OF MAGNOLIACE^. ]19

thin uiiliiiiiified walls. Each group siirroniuls the first -formed spiral
vessels and is destined to he liunitied in the course of time. Primary
medullary rays usually consist of tw(^ or three radial rows of cells ;
hesides these there are found numerous small rays consisting of a
single row of cells.

The ])ith C(^mmonly consists of cells with thick, lignified, ])itted
walls, hut sometimes there remains in the central portion (^f the pith a
grou]) of cells with urdignified walls.

Petiole. — On the epidermis a few hairs are sometimes present.
They especially grow in the groove whicli runs longitudinally along
the u])per surface of tlie petiole. The hy])oderma is present. Tlie
(il)i-o-vascular hundles form a ring enclosing the pith. Sclerenchy-
matous fihres are well develo])ed and constitute a continuous sheath
around the bundles ; they are unlignified in the basal portion of the
petiole. The phloem ])ortion is here and there crossed by the
medullary rays, and the portion of the ray which bridges the ])hloëm
is made u\) of scleroblasts. unlike that of the stem the pith is formed
of thin-walled parenchyma.

B](t(h'. — Its structure is generally com])act. The stomata do not
])resent any peculiai-itv. Hairs grow on the midrib and their base is
often made up of several cells. The cuticle of the midrib forms
a number of longitudinal ridges, which j)resent in a cross section
cuticular indentations (PI. II, Fig. 3). Directly under the e])idermis
of the lower side of the blade there is found a layer of cells
which forms the hypoderma. The ai-rangement of the fibro-vascular
bundles of the midrib is not completely circular, but shows a
discontinuous portion turned towards the u[)per surface of the blade.
Aofg'reo-ates of crystals ave also met with in the cells of the cortical
portion of the midrib.

Pool. — In the older root the o-eneral structure does not differ

120

s. MATSUDA.

iniicli from tliat of the .stem, except thnt pith is ahsent. The scleren-
clivmntous ring ec^iitaiiis hoth sliort and elongated elements. In the
voiinu' root, hairs are copiously found on the epidermis. The arrange-
ment of xvlem-plates seems to 1)elong to the diarch type (PI. IT,
Fiii'. 4, A). As the root becomes a little older the two xylem-plates
wliich WYO o]^])osite in tlieir ])osition hecome united, thus forming an
ellijitical mass of wo<^d (PL IT. Fig. 4, 1)). The enchjdermis remains
cellulose ;»s long as it exists.

Cercidiphyilum.

Of this geiuis T examined a single species, C. /(iponicinih Sieh, et
Zucc. Tlie ])lant is found in tlie mountainous parts of this comilry.
Tt attains a great height ;ind yields a valuable tindx'r. Ft is not
aromatic though tlie o])]>osite statement is made by some authors.'

,S'^';//. — The tissue elements of this ]>lant are very small in size ;
the ('(mipactness of texture of the wood is owing to this fact.
The epidermis is early shed, so that except in very young
shoots it is compleielv absent. The cork is very compact like
the other tissues. The hypodermal layer is present undei* the cork.
Cells containing prism;)tic crystals of calcium oxalate are abundantly
found l)oth in the outer c(-)rtex and in the ])hlo('m, and are es])ecially
munerous in the ])hloë,m i-ays (PL FF, I^ig. 5). 'I'he occurrence of
these crvstals is so common that I fourid them in vari(nis specimens
collected in different seasons.

The groups of sclerenchymatous fibres are well developed and
form bands alternating with the soft bast ; but there does not exist a
continuous sheath of sclerencliyma enclosing fhe fibro- vascular bundles.
Sieve- plates are seen iti the phloem, though not very distinctly. The

1. Bâillon, Natura! Histori/ of I'laiit.f, (translated from thcFrench).
En<j,-lt'r and Pi-antl, T'ii' iKiJiirliclii'ii Pilctn-cii/diiiilii'ii.

ON THE ANATOMY OP MAUNOLIACE^. ]^2l

xyJem contains traclieae, übres, and wood-parenchyma. I)ordered pits
are found in the walls of the vessels ; they ap[)ear in ;i surface view
like a slit encircled by a halo, and not like an encircled spot as
commonly seen in the wood of Fitnis. Most of the medullary rays
consist of single rows of cells.

The parenchymatous cells of the pith have thick lignified walls,
even in the young specimen, and contain an ample quantity of starch.

Petiole. — The epidermis consists of a single layer of cells, and the
cuticle is not well developed. The hypoderma is represented by a
layer of thick-walled parenchymatous cells. There is present in the
extei-nal cortex no sclerenchymatous element, but deeper in this region
there lies a zone of tissue presenting an irregular appearance, due
to the distortion of parencliymatous cells (IM. II, Fig. (), a). The
fibro-vascular bundles are arranged in a circle, as in the stem, and
enclose the pith in the centre (PI. II, Fig. (!). The bundles are encir-
cled by a well-developed sclerenchymatous sheath, which is inter-
rupted by parenchymatous ceils only at a few points (PI. If, b'ig. (i).
The filjres are unlignified in the basal portion of the petiole.
Prismatic crystals are found in the phloem. The pith parenchvma
unlike tlraf of the stem has unlignified walls.

Blade. — Its general structure, as also the form of its stomata,
has no iK'Culiaritv. There are se\eral main-ribs instead of a sin^de
midril). and their hljnj-vascular bundles jti'esent a semi-circular
arrangement, while tinsse of the petiole ai-e circularly arranged. A
grouj) of thick- walled cells is found within that poi-ti(jn of the
epidermis, which lies on the u])pei' side of the main-ribs. Aggregates
of crystaJs are found in the cortical region of the main rib. The
CLiticidar ridges ar(3 not so conspicuous as in the midrib ol hliiplehva.

liooi. — The structure of the older I'oot is similar to that (jf the
stem. Crystals are copiously found here also in the phloem.

122

s. MATöUDA.

Sclerenchymatous fibres are also well developed. In tlie young
root the endodermis is distinctly seen, and the cells that constitute it
remain cellulose. The xylem-plates present the tetrarch arrangement,
but sometimes also the triarch.

Trochodendron.

To the present genus belongs the single species, T. anilioides^
Sieb, et Zucc. It is n tree destitute of any aromatic property. It is
grown in many parts of Japan. Articles of various shapes are inade
from its wood by means of the turning-lathe. Its bark yields
bird-lime.

Stem. — The epidermis is of the usual structure. The cuticle is
very well developed, and stride, which are perpendicular to the cuticular
surface, and coincident with the l)Oundary lines between the epidermal
cells, are distinctly seen. Cork is present immediately beneath the
epidermis. The hypoderma is represented by a layer of somewhat
thick-walled, closely arranged parenchymatous cells. The portion of
the outer cortex which lies within the hyp<jderma consists of a loose
tissue with manv interstices, and trichoblasts are here found in great
abundance. They are of peculiar shape and are characteristic of the
present species (PI. Ill, Fig. i)) j many-armed tricholjlasts are found
in a, few other genera of Magnoliacea', such as MiKjiKjlia and MichciiUj
but these are nuich simpler in form.

The sclerenchymatous sheath is well developed and almost
uniiiten-ui)tedly encircles the fibro-vascular bundles. liesides these
fibres the sheath contains short sclerenchymatous cells or sclenjljlasts.
A great number of oil drops is found botli in the external cortex and
in the phloem. Witli the exception of a few spiral trachea' found in
the primary w<j<jd and of the pai'encliyma forming the medullary rays,
the whole xylem is made up of tracheïds. The walls oï the latter

ox THE ANATOMY OF MAGNOLIACE^. j^^;^

present 1)ordered j)its of various forms, nn^re numerous on their
radial than on their tangential surface ; and round pits are mostly
present on the tangential surface, while elliptical forms predominate
on the radial surface. Tlie primary medullary rays are very broad,
consisting of several rows of cells.

The parenchymatous cells of the pith have thick lignified walls,
which are pitted, and there is no trichoblast, though the ojjposite is
stated to be the case by certain writers.^

Petiole. — The general structure is similar to that of the stem. A
great many trichoblasts are found in the external cortex. The
fibro-vascular bundles are so arranged as to present a somewhat
semi-lunar form with its concave side turned to the upper side of the
petiole (PI. Ill, Fig. 7). The bundles are rather scattered in the basal
portion of the petiole, but in the other portions they are more closely
united. There exists at the external limit of the phloem the scleren-
chyuiLitous sheath, but it consists of unlignified fibres.

Blade. — The contour of the epidermal cells is not wavy here as
in the leaves of many other plants. The cuticle at the entrance
of the stoma is elevated, and presents a cup-like appearance (PI. Ill,
Fig. 10). The j^alisade-parenchyma of the upper side of the
blade consists of three or four layers of cells. Trichoblasts are met
with in the region where the structure of the blade is loose, so that a
number of intercellular spaces are left. The tibro-vascular bundles of
the midrib are semi-circularly arranged, and a group of sclerenchy-
mntous fibres is found in the cortical region lying on the l)undJes.
The cuticalar ridges of tlic midrib are wanting'.

lioot. — The general structure of the (^Ider root does not nmch
ditfer from that of the stem. Trichoblasts are abundantly found in
roots a little older, but their form is much simpler than that of those

1 Enyk'i- und I'mutl, I)ic uatHilu-lwn I'tldinfiifamilien.

224 s. MATSUDA.

in the stem ov leaf, and a[)|)n)u<jhes that of tlie triche )l)lastfs existinfj;'
in the external cortex of some species of M(((jNoli(i. The hy})odernia
is al)sent. In the cortex of tlie young root there is found a nundjer
of cells with thick walls, sHghtly lignified (PI. IT[, Fig. <S. f-.s).
Some of these cells are short and others are elongated. T'li^y are not
the vouno- sta2'e of the sclerenchymatous elements found in
the older root, and are destined to be soon shed off with the paren-
chymatous cells among which they are scattered. The true tricho-
blasts come into existence at a much later time, while the sclerenchy-
matous fibres do not appear at all in the roc^t, or only in s(jmewhat
old roots. The arrangement of the xylem-plates is of the diarcli type
(PI. Ill, Fig. 8). The endodermis and pericambium are not well
marked in the present species (PI. Ill, F'ig. 8).

Tribe II. lUicieae.

Two genera, JUiciioii and Drimijs, were examined in this tribe.

Illlcium.

In this genus I examined, two species ; namely, I. reliijiosdDi,
Sieb, et Zucc, and /. Tasliiwi, Max. The former is a small tree with
aromatic odour, and widely distributed in Jap:in. 7. Tasliiroi was
discovered a few years ago in Iliukiu or Loochoo by a Jaj)anese
botanist, Y. Tashiro, whose name the plant bears, and 1 examined
only a dried specimen of it.

Sleiii. — The epidermis has no peculiarity and lias no appendages.
The cuticle is very well developed, and stria', perpendicular to the
cuticular surface, and corresponding to the limits of the epidermal
cells, are distinctly seen, as in Trochodciulruii. Tn stems a little older
cork is developed immediately beneath the epidei-mis. In the outer
cortex there exists no layer of cells that can represent the hypoderma.

ox THE ANATOMY OP MAGNOLIACE^E. ]^95

nil the cells of tliis reo-ion bei no; of simihir fonn and, in most eases,
liavino- ])ittecl thick walls. 'I'here is not found aniono- them any kind
of idioblasts, whicli abound in several other genera of the same
family.

The phloem portion of the fibro-vascular bundles is almost
destitute of hard bast, and only at the external limit of the bundles
are there freund a few scattered sclerenchymatous fibres, which repre-
sent the sclerencliymatous sheath (PI. Til, Fig. 11, .s7). The soft bast
certainly contains sieve-tubes, but I was not able to detect them
clearly. Oil-globules are often met with in the phloem region, and
are a j^robable cause (^f the characteristic odour wliich is given out
when a branch of the ])lant is bruised. In the external cortex of I.
'J'(ts]iirfii certain cells are met with which seem to l)e a receptacle for
an oily substance (PJ. 111, Fig. 12, A) ; but in /. reliijioswn the
existence of such cells can not be clearly detected, thoug-h some cells
of the external cortex may be suspected to be of that nature. In the
xylein there are found trachea^, fibres, and wood-parenchyma. The
medullary rays are very narrow, generally consisting of one or two
rows of cells. In /. irligiosiim the demarcation of the annual rings is
generally not very clear, owing to the fjict that the xylem-elements
formed at the end of autumn of the preceding year are similar in size
to tliose wliich are formed at the beginning of the next spring.

The pith is surrounded by a medullary sheath, in which spiral
tracheae are found. The parenchymatous cells of the pith have thick
walls which are often ])itted. They are lignified in old specimens of
7. rclifiiosinn ; and in 7. Tasliiroi this is the case even in youn»'
s])eeimens. According to PrantP sclerenchymatous cells are found in
the pith of Illiciea?, but I do not find them in my specimens.

I'cfiolf. — The fibro-vascular bundles of the petiole do not form a

1. E:iiJ'l<^r uml PiMnM, />/'' initHrliclwit PtiKtr.i'iif'aiiiiUcn.

J 96 s. MATSUDA.

ring enclosing the pith, as they do in the stem ; but constitute a
somewhat crescent-shaped oToup witli its conr^ave side turned to the
upper surface of the petiole. The xylem ])<)i-tion is incompletely
lignified as is proved hy the action of reno-eiits. In tlie phloëin no
sclerenchymatous fibres are present. The bundles are partly
enclosed by a row of parenchymatous cells containing starch-grains,
which perhaps represent a Inmdle sheath. In all other res])ects the
structure of the petiole is similar to that of the stem.

Blade. — The cuticle is well developed. The fibro- vascular
bundles of the midrib are semi-circularly arranged, and a fcAv scleren-
chvmatoiis fibres accompany the bundles. The cells forming the
palisade-parenchyma lying at the midrib have their height diminished
and become almost round. Resin sacs are not found at least in
7. yrJiijiimim.

Hoot. — An old root almost resembles the stem in general
striK'tiu-e. Tn the young root the arrauii'ement of the xylem-])lates is of
a diarch ty])e. The cells that constihite the endodermis remain cellu-
lose. Oil -drops are scattered about in the cortex of the young root.
l^o sclerenchymatous fibre is seen in the phlorm, even in old speci-
mens.

Drimys.
T examined onlv a dried specimt^n of one s])ecies of this genus ;
namely, J), diprfala, ¥r. M.. which had r-ome from New South Wales,
and T can only give a few points about its antitomical characters,
since I have only a verv im])erfect knowledge of them.

Tn general structure this species resembles /. rcliijiosiim^ but
there exist some decided structural ditferenres between them. Thus
in />. dipftala a great nu7nber of resin-sacs are present in the outer
oortex (PI. TTT, Fig. 12, B) and even in the phlo('m ; a well develop-
ed sclerenchymatous ring accompanies the fibro-vascular bundles, and

ON THE ANATOMY OF MAGNOLTAOE^. jQ^

the xylem consists exclusively of traeheids. The List oh;trncter is
only found in tlu' present o-enus and in 'ivochnilcndron nmono- Mao--
iioliacete. The pnrenchyniatous cells of tlie pith are lignified even in a
young specimen, as in 7. Tasliimi, hut T have not here met with anv
scjerenchyniatous cells scattered aJ)ont. (as they are stated to l)e hv
l^-antl). Tlie le;if is almost sessile, and the til)ro-\'asculai' Ixnidles oi'
the very shoi-t jietiole present a semi-circular arrangement, as in
llJiriimi. The bundles are similarly arranged in the midrib. A
immher of resin-sacs are found in the mesophyll. In the epidermal
cells of the leaf, crystals are often met with, which are prohal)lv
calcium oxalate.

Tribe III. Schizandreae.

This trihe consists of the following two genera :

Kadsura and Schizandra.

The two genera are so closely related in anatomical characters
as to make it convenient to describe them togethei-. Of Kadsuin T
examined K. japonira, L., and of Scliizavdm I examined S. nufnt
Max.. and S. cliiiifnsis, I>ail. The three are small climhing shrubs.
found in se\'eral parts of Japan. When a cut portion of these
plants is put in water it yields a large quantitv of mucilage, which in
the case of 7v. japnuicn is S(^metimes used for dressing the hair. Tlu;
two species of Schizandra give out a sweet odour \vhen any j^oi-tion of
them is bruised, hut K. japonica gives out little or no odour, when
similarly treated. T'he berries of S. eliinensis are said to he edible.

Stem. — The epidermis consists of a single layer of cells, and the
cuticle is not well developed. The hypodei-ma, is not represented.
The cork is well develo])ed directiv under the epidermis and is of the
usual type. Tn the cortical parenchyma there are scattered about a
few cells wliich contain oil globules (PI. TV, Fig. 14. rn) ; these are pro-

128

s. MATSUDA.

b;il)lv the s(^nrce of tlie cbarneteristic odour which the plants o'ive ont
wlien bruised. These cells are conspicuous in Schiiniuhri l)y tlieii* size,
beinu' much larii-er than the cells of the surroundino- parenchymatous
«•ells, but" in 7v. japoidca the oil-cells are scarcely lars^er than the
surrounding cells, and are. besides, seldom met with. The cells of the
(•ortic;d pareiK'byma are sometimes pitted in /\'. jdpnuica (PI. TV. Fig.
14).

Sclerenchvmatous fibres are generally found at the external
limit of the phloem, constituting the sclerenchvmatous slieath, but
they are not much crowded, either forming only a thin layer (jr else
small grou])s (PI, FY, Fig. lo. .s7). Some of these fibres are septate,
as may be seen in K. jd/ioHicd (V\. TV, Fig. '2\). Scattered about
in the ])hl«H'm (and als(^ in the external cortex) tliei-e are found
peculiar cells with tliick lignified walls and a narnnv lumen (IM. IX,
Fig. '20). Some of these cells are elongated like sclerenchyuiatous
fibres, others are very short, simply round in form, or else provided
Avith arms, which they ])ush out freely among the surrounding cells.
The outer layer of theii' thickened wall contains a great number of
granules (^f calcium oxalate, which present an angular configurati(Mi,
without the definite form of true crystals. These peculiar elements are
structurally nothing else than sclerenchymatous filtres or scleroblasts.
liolding a number of grannies imbedded close to one another in their
walls, and for convenience' sake I will hei-eaftei' call them "crystal-
bearing sclerenchymatous elements." I do not think tliat there exists
any essential difference between them and the so-called spicular cells
of WelwiUchia mirahil/Sj which liave been described by several authors,
and are stated to he of tl>e same nature as the crystal -bearing fibres
found in Amiicnria. &c. Ff I am right, the interesting fact is
supplied of two gi'oups of plants which arc remote from eacli other in
systematic jtositiou coiitaiiiiiiL;- a nci'v similar elciuent in their

ON THE AXATOMY OF MAUNOLIACE^ j^^C)

ti.ssiie.s. In the jthloeiii of kadsiini and Schl:tui(lra tliere are found
nianv Jarge intercellular spaces, which are apt to be mistaken for some
kind of laroe ducts, but are really passages of lysigenetic origin.
They are on all sides bounded — and the boundaries nre often verv
irregular — by distorted and broken cells, and in sections cut from
specimens preserved in alcohol, they ai-e often found filled with a homo-
geneous, structureless sid)stance. The}^ seem to serve as reservoirs for
the nnicilage which ul)onnds in these plants, and wliich is probalilv
derived from tlie disorganization of the suriounding cells. They are
mo?t conspicuous in Kadsiini. The xylem consists of tracheae, filn-es,
and wood-parenchyma. The vessels are large, their diameter Ijeino-
several times greater than that of the surroundincj- tissue-elements,
especially in Kadsuni. The walls of the vessels are distinctlv sculptured
with bordered pits. The medullary rays mostly consist of a sinole-
row of cells.

The pith consists of unlignified parenchymatous cells, amontj-
which may be found a few sclerenchymatous cells in Scliiidndra (PI.
IV, Fig. 15), and a few crystal-bearing ones in Kadmm.

Vet'uAi'. — Unlike those of the stem the fibro-vasc;ilar bundles of
the j)etiole are so arranged as to present a somewliat semi-lunar form
with its concavity turned to the upper side of the [letiole (PI. IV,
Fig. 1(S). In a section cut from the upper or middle })ortion of
the petiole, these bundles lie in a few definite groups, as is
shown in the figure just referred to, (where three distinct groups are
seen) ; but in the basal portion they are more looseh* arranged.
There is not present either in the phloem or in its neighbourhood, the
common form of sclerencliymatous fibre or scleroblast ; l)ut scattered
about in the parenchymatous tissue surrounding the bundles there
are found the ci'vstal-bearing sclerenchymatous elements (V\. IV, Fiu-.
ly, 6'.s). In the case of Kadstira a single row of parenchymatous

130

s. MATSUDA.

cells which contain a (juantity of starch-grains, j)artly encloses the
bundles and faintly represents a biindle-sheath (PI. IV, Fig. 18, i ;
PI. IV, Fig. 19, ï). In Schizandra some parenchymatous cells
lying near the bundles contain crystals of calcium oxalate. These
crystals are found either singly in a cell or as an aggregate. In the
former case they attain a considerable size and are (3ctahednd ov
prismatic in form, but in the latter case their exact form is
indeterminate. Such crystals are not met witli in the stem. In the
existence of mucilage-reservoirs, as well as in other points not
specified here, the structure of the petiole agrees with that of the
stem.

Blade. — The stomata are of the usual form. Cuticular ridges
are distinctly seen on the epidermis of the midrib. The arrange-
ment of the fibro-vascular bundles in the midrib is similar to
that in the petiole. Mucilage canals are found both in the
midrib and in other parts of the blade. In the latter place they
generally accompany the veins sent off from the midrib. Crystal-
bearing sclerenchymatous elements are found both in the midrib
and in the veins of Kadsum japonica, A\'hile they are rarely met with
in the leaf of SchiziUidra nvjva. On the contrary crystals both aggre-
gated and solitary are cojnously found in the midrib and veins of
Schizandra fiiyra, while they are rare in the leaf-blades of Kadsura.
Crystals forming aggregates are also found in the epidermal cells of
Sehizandm (PI. IV, Fig. IG).

lloiil. — The (Tystal-bearing scJerenclniiiarous cells, the cells con-
taining oil-globules, and sc^me other })eculiar structures, which are
found both in the stem and leaf, are also met with in the root (Pi.
l\'. Fig. 11, I)). However, the sclerenchymatous sheath, which is
constantly found in the stem, is absent even in somewhat old roots.
The intercellular passages which serve as nmcilage-reservoirs are found

ox THE ANATOMY OF MAGNOLIACEtK. x;-J1

in the roots of K. japonica and S. clu'ncnsis, but in a somewhat old
root of S. niyra, 1 found the passages not yet developed, and but
little mucilage present. Thus it is seen that the formation of
the passages has a close relation to the production of mucilage, a fact
which seems to favour the view that the nmcilage is derived from
the disorganization of the pre-existing tissue. As to the number ot
xylem-plates in the young root, K. japonica and S. nigra agree, both
presenting the diarch arrangement ; but in S. chinensis the triarch,
as well as the diarch one, may be seen (PI. IV, Fig. 17). The pith
is absent in old specimens.

Tribe IV. Magnolieae.

Under this tribe I examined the following three genera :

Magnolia, Michelia, and Liriodendron.

The anat<3niical elements of these genera may Ije described to-
gether, as they present nothing characteristic enough to distinguish
the genera from each other. Only one sj^ecies is known to Ijeloug
to Liriodendron, and this I examined. Four species of Michelia were
examined, and ten species and two varieties of Magnolia. All of
these plants are trees and generally attain a great height. Aromatic
properties are prevalent among them. The names of the species
examined are as follows :

Magnolia stellata, Miq.

M. parrißora, Sieb, et Zucc.

These two are ornamental trees.
M. Kolws, I). C.

Also ornamental. Its wood is used in cabinet work.
M. hijpoleuca, Sieb, et Zucc.

It is one of the most useful trees, its soft wood being ex-

j;-^9 S. MATSUDA.

tensively used for various purposes. Suiall articles uf

furniture, chopping blocks, &c., are made fro7n it. It is

also used for making pencils. The charcoal obtained

from it is used for polishing lacquered ware. Also the

tree is ornamental.
il/, mlici folia, Miq.
M. ohovata, Thunb.
M. ohovata, Thunb. var. (commonly known as Katisliiu-

mokuren),
M. conspicita, Salisb.

31. coHspicua, Salisb. var. parpurescens, Max.
J/, pmnila, Andr.
J\L Jl'aisoni, Hook. fil.
M. tjrandißora, L.

The last seven of the above plants are ornamental exotic

trees.
Michclia comprcsm, L.

This is an ornamental tree. Small articles of furniture

are sometimes made from its wood. It is grown in the

hotter parts of this country.
il/. Cliampaca, L.
il/, lomjifolia, Blume,
il/, fnscata, Blume.

These three are ornamental exotic trees ; il/. Cltaiiipaca is

said to be of some medicinal value.
Liriodendwn hdipifcra^ L.

An exotic tree.
Stem. — The epidermis is of the usual structure with the cuticle
w^ell developed in many species. Epidermal hairs are found in all
species of Michclia and in some of Magnolia, {e. (j., il/, (jrandißora, par vi-

ox I'HE AXATOMY OF MAGXOLTAOE^E. J 33

ß.orcC) ; but not in Lirioderulron. Development of cork seems to take
place in these genera immediately under the epidermis. The hypoderma
is generally represented by a layer of thick-walled parenchymatous
cells which are packed very closely (PL V., Fig. 22, hp). The cells
constituting the hypoderma are in some cases transformed into
scleroblasts, as is seen in Magnolia (frandifl ova ^ hijpoleuca (PI. Y, Fig. 28,
///)), piiniila, imrvißora^ &c. Resin-sacs are scattered about in the ex-
ternal cortex. Each consists of a cell having a form similar to that of
the surrounding parenchymatous cells, but distinguished from these by
the nature of its contents, the thickness of its wall, and also in most
cases bv its size. These sacs are conspicuous in Magnolia and Lirioden-
dron, ])ut hardly so in Miclielia. In many species they are found in the
phloem, and even in the pith. The contents of these sacs are probably
the chief source of the aroma possessed by the species of these genera.
Like that of the resin-sacs the occurrence, either singly or in groups,
of scleroblasts in the external cortex is universal in these genera,
These cells are, however, very rare in some species, e. g., Magnolia
pa rri flora. Some of them are many armed and may properly be
called trichoblasts.

- The sclerenchymatous fibres are greatly develo])ed, and not only
form a sfroug sheath to the fibro-vascular bundles, l)ut arc found
mixed among the soft bast. A few scleroblasts are also found with
the sclerenchymatous fibres that lie at the external limit of the phloem,
and with them constitute an almost continuous sheath. The phloem
portion consists of the usual elements. The xylem contains the
trachea3, filn'ous elements, and wood-parenchyma. The vessels present
bordered ])its in their walls. Tlie medullary rays vary in breadth,
some consisting of three or four rows of cells, and others of a single
row. Sclerenchymatous elements are generally absent in the phloem
ravs.

134

s. MATSUDA.

The pith consists of iinh'onified cells, and is traversed in places by
a kind of horizontal septa. This structure is known as " diaphragms "
(PI. Y, Fig. 27, dm'), and is universally found in tlie species included
in the present tribe. In some species, cjj., ]\fafiH(ih'(t (jrmnliflora, it is
very conspicuous and distinctly visible to the naked eye as streaks
traversing the pith. A diaphragm consists of a horizontally stretched
mass of scleroblasts, whicli, when highly developed, occupies an almost
entire cross section of the pith, though it is inconsiderable in thick-
ness; and this mass is continuous at several points with the wood-
parenchvma that lies at the inner limit of the xylem. In Liriodcndron
the greater part of a diaphragm consists of cells with thick, ])itted,
but not lignified walls, while the remaining small ])orti(m is made u])
of scleroblasts. The diaphragms of Müijnolia Inipolcuea consist of
scleroblasts having a very irregular shape. Those of MaiptoUa par-
r i 11 ara iuid 31. sal ici fol id are not so well developed as those of other
sjiecies.

rctioh'. — Tlie cuticle is strongly developed in Michelia, but not

in Liriudcndron. In Mü(juolia it is well developed in some species, l)ut

less so in others. Epidermal hairs are abundantly found in Michelin,

l)ut in Liriodoidrou they are almost absent. In M(ujuoJi(( many species

have the hairs well developed, while some are destitute of them. In

DffKjHoIia jiarvifJora and a variety of MmjuoJin ohorala (KdH.diiu-inolii-

rcii) there are found, in addition to the kind of hairs common to other

species, smaller haiis which are sometimes l)ranched (PI. V, Tig. '24,

cli). The hairs are generally cuticularised, but in MdtjNolia Kohus, in

which a few are found, they remain cellulose. Resin-sacs occur in

the outer cortex of all the species, and even in the ])hloëm of some.

The scleroblasts, which are often many-armed, are generally found in

the external cortex ; but in the single ca.-e of TÀriodcndron they are

almost absent. The number of hbro-vasculnr l)uiidles which entei'

ox THE ANATOMY OF MAGNOLIACE^.

IP.O

tlie peliole is variable, in some species exceeding twenty, in others
l)eing miK'li fewer. These l)und]es are g-eiierally isolated from one
another in the basal portion of the petiole, but in its u])per portion they
become gradually united so as to form a ring enclosing the ])ith.
This arrangement is usually very regular, but sometimes several liundlcs
stand isolated outside the main ring, and disturl) the regularity of the
ai-rangement. The sclerenchymatous fibres are very well deve!o])ed
and constitute the bundle sheath. They are generally unlignified
at the b:isal portion of the petiole, but in Liriodendron some of these
fibres are lignified eNen in the basal portion. The diaphragms .are
present in the species examinech In several cases, however, tiiey are
only faintly represented by a few sclerol)lasts found in the pith; while
in others (c. <)., Miclirlia fuscata), a bro.ad mass of sclerenchymatous
cells constitutes the diaphragm.

Blade. — With the exception of IJriodciidron the cuticle is general-
ly well-developed. In MidieUn and Maipiolia epidermal hairs are
generally met with, especially on the lower surface of the leaf; wliilc
in Liriodendron many epidernud cells of the lower side of the leaf are
provided eacli with a little ])rotuberance, which is really an im])erfect
hair. In many cases the hair is not an elongation from a single
epidermal cell, but several cells take part in ibrming its l):isal portion,
as will be seen in referring to the figures, 25 and '2(1 (PI, V). The
stomata are generally elliptical in form, but in MofiHolio (jntnditlorit
they are rather I'oundish, the curvature of the guard-cells being very
great. In Magnolia jnimila the cuticle is much I'aised at the entrance
of the stomata, and presents a cu])-like appearance, reminding us of
the stomata of TrocJiodemlrox (PI. V, Fig. oO). lu MaijiioJia (jrandifJora
there is found between the epidermis and the palisade-parencliyma a
single layer of parenchymatous cells, which do not contain chlorophyll
grains (PI. \\ Fig. :^9). This layer extends to tlie verv margin of

136

s. MATSÜDA.

the lenf-bhide. and some of the cells constituting it become sclerotic
as this region is ajiproached, and, together with the sclerenchymatoiis
fibres that abound there, form a very strong margin to the blade.
The blade of JSIdijuoJia (jniHdijIora is also itself very thick, the palisade-
])arenchyma consisting of several layers of cells, but in MaijuoJui
mlicifoh'a, which has very thin leaves, there is only a single layer of
cells constituting the palisade parenchyma. Although there are the few
just noticed peculiarities to be detected in certain species, the gener«al
structure of the blade is similar in the three genera of ]\fa<inolie(v.
The fibro- vascular bundles are circularly arranged in the midrib, as
they :dso are in the petiole. Resin-sacs are found in the leaf-blades
of all the s])ecies examined. The (K'currence of stone-cells is also
very general. MiKjHolid piniiiJa contains somewhat el(^ngated scleren-
chymatous cells in the hy])odermal region of the lower side of the
midrib. Red ])atclies ai'e often met with in the n.esophyll of some
species of MtKiHoli/i, c |/., il/, ohovata, M. cou^]yieu(i, and M. coiisjricua
^'ar. jivrpinrscrns. These were proved to be tannin.

Boot. —The general structure of the older root does not nnich
deviate from that of the stem. Resin-sacs are ])resent without
exception in the outer cortex. This region also contains a number of
scleroblasts in some species, while it is destitute of them in others,
'I'he sclerenchvmatous fibres are generally developed at the external
limit of the ])hl<x'm. Pith is absent, and its place is occu])ied by a
group of elongated woody fibres. In the young root the number of
xylem-plates is very variable. Thus in Lin'o<lr)i<Jron their arraiigement
is diarch or triarch, but mostly the former. In MicJielia it is triarch or
tetrarch (PI. V, Fig. 3i, A). In Miij)u^Un diarch, triarch, and tetrarch
arrangements are each commc^n ; but in one species, namely, ^hufiinVut
fjraudißoni, the arrangement varies from tlie tetrarch to the heptarch.
General l\', \vlu;n the root becomes a little oldei" the xylem-plates which

ON THE ANATOMY OF MAGNOLIACEil]. l^J

are separate at first, tend t(3 unite and form one body of xylem ;
but in }Iü(jnoli(( (jmuditloya they conti nne separate for a comparatively
long time, so that tliere remains in the central portion a mass of
iinlignirted cells. In KdnsJiin-iuoktircH, (a variety of MiUjnolia oJiocatti),
I saw only a continuous mass of ligniiied cells instead of separate
xylem-plates, even in the youngest s[)eciinens obtainal)le. In general,
the endodermis is distinctly marked off from the surrounding tissues
(PI. A^, Fig. 32, {'), and in most cases it is traceable even in somewhat
old roots. In Michelia compressa and Miujnolia coNspicua, var. pur-
pumscens, somewhat elongated cells with lignified walls appear \evy
early in the cortex (1*1. V, Fig. 32, es), while the xylem-plates still
remain se})arate. The widls of these cells in the latter species are not
uniform in thickness, the part (^f the wall facing the circumference
being not much lignified, while that facing the centre is thickened and
lignified.

The cortex of the older root of MarpioJia Kob)is contains many elongated peculiar cells,
found near the intercellular spaces of schizogenetic origin, which also abound in the root of
this plant.

When the roots of some species of MagnoUeœ, such as Michelia Cliainpaca, Mich, com-
pressa. Magnolia Kohus, &c., are kept in alcohol they impart a I'ed colour to it.

Anatomical Characters of DifTerenl Genera Compared.

In the fores'oino; najj'es the anatomical ch;u"acters (jf each i>'enus
of the present family having been described, it is now possible to
give a comparative view of them. The structural points to be com-
pared will be treated under the same four principal heads as used
in describing each genus : Stem, Petiole, Blade, and lîoot.

Stem.

The characters of the stem require fuller treatment than the rest,
and will be compared under the several structures.

1 . Epidermis. — The epidermis generally consists of a single layer

l'^<i^ s. MATSUDA.

(jf cells. The cuticle is sotnetime.s well developed, as in flliciu}ii,
Driiiiij.s, Troclioilendrou, and many species of Magnolia. Eindermal
h iirs are found only in the two genera, Michelia and Magnolia und in
the latter genus are almost absent in some species.

2. Cork. — This structure presents no peculiarity. As with
m;iny other Dicotyledons, cork is here developed inunediately beneath
the epidermis. In Euptelcea it is conspicuous fnjm the size of its cells.

3. Hypoderma. — This is totally wanting in JlUciuin, Vriiiiij.s,
Kadzvra, and Scliizandra but present in all the other genera. When
present, it consists of a layer of thick-walled and closely packed paren-
chymatous cells (PI. V, Fig. 22, hp), and in some species of Ma(jnolia,
as M. gmndiflom, hjpoleuca (PI. V, Fig. 28, ///>), &c., these cells are
transformed into scleroblasts.

4. Secretory Reservoirs. — Two kinds of these are found, resin- or
oil-sacs and crystal-containing sacs, the latter chiefly found in Eiiptekea
(PI. II, Fig. 2) and Ccrcidiphyllmn (PI. II, Fig. 5). The crystals are
of calcium oxalate in both genera ; 1jut in Euplekm they exist in the
sac as an aggregate and are confined to the outer cortex, while in
Cercidiplryllwii tliey are not aggregated, and the sacs containing them
exist in the phloem, as well as in the outer cortex. In Schizandra
crystal-containing sacs are confined to the leaf (petiole and blade).
Ilesin-sacs universally occur in the external cortex of Ma<jnoliea', and
in many cases are found in the phloem, and sometimes e^'en in the
pith. Cells containing oily substances are also met with in the
extern-.d cortex of Schizandra (PI. IV, Fig. 14, CD). Resin-sacs are
also present in Driuiys and iUicium Tashiroi, but absent in lUiciina
reliyiosiun. Trochodendron is entirely destitute of any kind of reser-
voirs.

5. Scleroblasts and Tricliohlasis (in the external cortex). — The latter
are simply a modified form of the former, lîoth forms of cells are

ON THE ANATOMY Oi' MAGNOLIACEJS. J 39

met witli ill Mngnoliuceiu. Trichoblast« are very abundantly found in
the external cortex of Trocliodemîro'n, and send their arms freely into
the intercellular spaces which also abound in this region (PI. Ill,
Figs. 7 and 9). Their presence is a feature that distinguishes this genus
from others. Scleroblasts are generally found in Maijinjliece, and in
many cases are accompanied by trichoblasts. The external cortex of
Illicinm, (perhaps of Drùiu/s also), Eupteliea, and CercidiphylluDi is
entirely destitute of them. Kadziira and Schizamlra contain in their
outer cortex and less often in their inner one a number of crystal-
bearing sclerenchyniatous cells (V\. IV, Fig. 20), which are nothing
more than scleroblasts with numerous crystals imbedded in their walls.
Some of these cells are elongated and form crystal-bearing sclerenchy-
niatous fibres.

6. Fihro-rascidar Bandies. — The fibro-vascular bundles existing
in the stem of Magnoliacese belong to the so-called collateral bundles,
which are uni\ers;illy foiuid in the stem of phanerogams, so that it is
unnecessary to give an acc(junt of their general features. But there
are certain structural points in the bundles peculiar to different genera
which need to be noticed briefly under the following heads :

(1) Sclcrencliiimatous Sheatli. — This is made u}) of sclerenchy-
niatous fibres often accomjKinied by scleroblasts, and is more or less
developed in all the genern. Howe^^er, in Illicium it is very imper-
fectl}' represented by scattered fibres that are found at the external
limit of the phloem. It is })oorly developed in Kadzura and Sltizandra
(PI. IV, Fig. lo, .s7) ; rather well developed in Eiiptehra and Ccrcidi-
plujlhnii; in 'rrucliodendroN and all the three genera of Jic/^/wZ/ca' most
strongly developed aud, together with a number of scleroblasts exist-
ing there, constitutes an almost continuous ring of sclerenchyina.
Such is also the case in Driiniis.

(!?) Vltloeiii. — The soft portion of the phloem probably contains

140

s. MAT8UDA.

those elements which are usually found in this region of other Dicoty-
ledons, Ijut clear detection of them was generally very difficult,
and the presence of sieve-tuhes was pnjved only in a tew s{)ecies. In
Cciridijihiilhuii and all the three genera of MiUjnoUea: tlie hard-l)ast
or sclerenchymatous fibres are especially well developed, and are found
amonf tlie soft bast. Also in Kadinm and Schizandra sclerenchyma-
tous fibres are found among the soft bast, but these are heavily loaded
with miimte crystals, and transformed into the crystal-l)earing
sclerencliymatous fibres (1^1. IV. Fig. 20). Sacs containing crystals
exist in the phloem, as well as in the external cortex of Ccrcidi-
phiilhiui (PI. n, Fig. 5). In Kadziira and Scliimndra it is a remark-
able feature that there exist in this region large intercellular passages,
which serve as tlie reservoirs of mucilage.

(3) Xiilem. — This portion generally consists of trachete, fibrous
elements, and wood-parenchyma. But T roch ode n Iron and Drinujs are
exceptional in this matter, their wood consisting abnost exclusively of
tracheïds. This [)eculiar structure of the xylem distinguishes these two
srenera from others of the Maunoliaceœ, and at the same time allies
them with Conifera3. But, as stated by Prantl,* their wood may be
distinguished from that of the lattn- f imily by the fact that in C(3nifera3
the cells of the medullary rays hav^e their longer axis horizontal, while
ill Drimys und Trocliodendron moHt of t\m cells of the medullary rays
have their longer axis vertical. lîorclered pits are found in the
tracheïds of 7);7'/»//,s and Trocliodendroii ; in the other genera where
vessels are present, pits of this kind are generally found in their walls.
It may be added that the exact determination of the various elements
that constitute the zylem was not very easy.

(4) McdidJarij llaijs. — Their breadth varies, consisting mostly of a
single row of cells in Illlcliuu, Cereidiphiilhiiii, Scliizafulra, and Kadzura,

* Engler and Prautl, Die natilrllclun Fßanzeiifamüien.

ON THE ANATOMY OF MAGNOLIACE^. 141

hut of several rows in TroclioiIcNdrnu. In Mufinolieœ d\iïeve.nt hrencUlis
are fonnd. In the pliloc'in rays of Evptchva a few scattered sHero-
blasts find their way among soft parenchymatous cells (PI. II, Fig.

1, 4

7. ]*ith — The parenchymatous cells of the pith are completely
lignified in Ccrcidiphijlhnn^ Drimijs, and Twchoilendron, even in young
spefimens. This is also the case in lllichim Tasliiroi, hut in lUicium
relitj'umnn they are unlignified in y(Ming specimens. The young stem
of Eiiptehrn (^ften has a small jiortion of the pith remaining unlignified.
The parenchymatous cells of the pith are unlignified in Kathura and
Scliiranihri, hut a few sclerenchymatous cells are found in it (PI.
IV. Fig. IT)). ^[(KjuoJieiv are distinguished by the existence of
dia])hragms in their ])ifh (PL V, Fig. 27, J»?), the only part of the
pith that liere becomes lignified. As I have stated before, the asserted
existence of stone-cells in the ])ith of JUicicœ is not verified by my
specimens.

Petiole.

The structure of the stem having been comparatively described,
there remains scarcely anything to be said of the petic^le. I will here
only notice a few anatomical ])oints in which the structiu'e of fhe
petiole deviates from that of the stem.

The epidermal hairs are generally more strongly developed in
the petiole than in the stem, as we see in many species of Maijunlia ;
and even Evptehva, which bears no hair on its stem, has a, few on the
petiole. In those s])ecies which contain resin-sacs, sclerol)lasts, and
tri('hol)lasts in their stem, the presence of the same in tlie petiole is
almost certain, as is also the case with crystal -bearing sclerenchymatous
elements. A number of crystals is found in the outer cortex of the
jtctiojc of Sell i Id mini, tliough none is })resent in the stem. The fibro-

1^2 S- MATSUDA.

vnsciilar hiinrlles are in many cnses arranîred as in the stem ; that is,
in tlie cross section of the ]>eti()le, they ]^reNent a circiilnr arrangement
enclosino- the pith in the centre (?J. 11, Fi,i>-. ß). But in the petiole
of llh'chim, J)n'm)i!^, Trochodcndwn, Kadiura, and SchizaNdra, the iihro-
vascnlar linndles have a semi-circnlar arrano-ement with the concav^ity
turned to the npper side of the petiole (PI. IIT, Fia'. 7). The
sclerenchvniatf^ns sheatli of the Inindles is often wanting', as in the
petiole of lUicium, Dn'imj.'^, and Scliizandra. Troeliodfudwn has the
sheath consistino- of nnho^nified fihres, while all the other o^enera have
well developed sclerenchymatons sheatlis (PI. TI, Fii;'. ()). The fihres
forming- these sheaths remain nnlignified at the hasal pt^rtion of tl^e
petiole in all cases, except in the ])etiole of Lin'odpndron. where a few
fihres are found lio-niiied. Wliether this local non-lignification of the
iihres has some pli3^siological meaning or not still remains undecided.
In .T/rvf///o//>o' dia]ih7'agms are found in the pith-])ortion of the petiole
as ill that of the stem.

Blade.

The general structure of the hlade does not deviate from the
dicotyledonous tvpe. 'i'lie up])er portion of it is made up of palisade-
parenchvma, whi<h consists of vertically elongated cells varying in
diffeient species from one to several layers. Wliere these cells lie
di recti v over the main rihs. their height is generally diminished and
their f)rm hecomes almost round. The lower portion of the hlade is
generally made up of loose ]>arenchyma ; hut in those ])arts lying
directiv under the main I'ihs there is found a more compact ])ai'en-
chvmatous tissue. Tn some genera, namely, Jlliciinn, Trocliodoidroti,
]\[cu}Noli((. and Michelin, the l)lade has a well developed cuticle. In
Sclri:(Uidreœ the cuticle of the midrih — especially that of the lower
surface— forms fine ridges running longitudinally (PI. TI, Fig. 'î).

ON THE ANATOMY OF MAGNOLIA OETR.

148

Tliese rido-es nre also observable in Eiiptelfra, and less distinctlv in
Ccrcidij)]nilliiiii, but are wanting in Trochodcmlrou whicb with these two
o'enera forms Trocliodendnvr. Epidermal hairs are generally found in
M(«juoJii( and DlicheUa, and also in EupfeJoYi, l)ut in other genera are
almost wanting. In general the stomata do not ])resent any
]>eculiarity, being elliptical in form and a('C(^m])anied by the two guard-
cells, but in Maiinolid (imndiffoni their form is nearly spherical
and the guard-cells are ]iressed outward, so that they rest u])on the
neighbouring epiilermal cells. Both in Maffnolia pvmHa and Twchn-
dt'iidi-oN iinilioidcx. the cuticle is much raised around the entrance of
tlie stomnta. and ])resents a cup-like a])])earance (PI. Ill, Fig. 10 ; PI.
V, Fig. oO). The arrangement (^f the fibro-yascular bundles of the
midril) is circular in Maijnolhuv, semi-circular in fJlicicœ and Selii:nn-
drtw. In all these cases the bundle arrangement of the ])etiole agrees
with that of the midrib. In ('t'lcidifdijiJlinii and Euptekm the Inindles
are circularly arranged in the ])etioIe, but in the former they are seini-
cii'cularly arranged in the midrib, wliile in the latter their circle is
incom])lete in this ])art. F)ut Trochodeudron aralioidt's has the bundles
semi-circularly arranged both in ]ietiole and midrib. Thus in this
res])ect the tliree genera (^f Tmrhodcndnuc do not agree with one another.
I»esin-s:ics are present in the mesophyll, as well as in the midrib of
the examined three genera of Maçiuoliew. They ai'e also jn-esent in
these parts oî DrimnA d.ipctala, but have not been observed in those of
Jllicimn religiosum or of Schizaudrav and Trorhnd(>ndrra'. although in
the former of these the occurrence of mur-ilage canals both in the
mesophyll and the midrib is very common, fhe blade in a few genera
contains crystals. Thus aoföreoates of crvstals are found in the
cortical region of the midrib of Eiiftfehrn and Cercidiphylhim, and in
that of ScliiztiNdrn. In DriDU/s dij>('t(d(( the e])idermal cells contain a
number oi' (Tvstals. As to the existence of scleroblasts. trichoblasts,

144

s. MATSUDA.

îuirl ci'vst;il-beai*ing sclerencliyinntons elements, whnt I have stnterl in
tlie case of the stein is a])p]icab]e here to tlie hlade.

Root.

The root, when it lieconies a little older, presents tlie same
structnre as tiie stem, except that the central portion, which corres-
ponds to the pith of the stem, is occnjned by the xylem. Generally
when sclerohlasts, secretory reservoirs, &c. are found in the stem,
they are also present in the root ; hnt in the case of lUic'nim rcUijio-
swn, sclerenchymatous elements are totally absent in tlie root, thon,o-h
a few fibres are present in the stem. In the yonnii' i'^'^^" I'^dial bundles
with the endodermis arc present in the central portion, and are sur-
rounded by the cortical ])arenchyma. The cells constitiitinii' the
endodermis oenerally remain cellulose. The Casj)ary point is not
clearly seen, l^ioth endodermis and ])ericambium are generally well
marked in the voung root, except in that of Trocliodcndron (PI. Ill,
Fio-, SV In a few cases a number of cells with liquified thick walls
are found in the cortical portion (PI. Ill, Fig. <S, cs ; PI. Y, Fig. o2,
r.s). The arrangement of the xylem-plates varies, tliere being usually
found the diarch, triarch. and tetrardi types, while the number is not
constant even in the same individual. In the single case of J/c^^///o//r<
(intudiffoni the tetrarch to heptarch arrangement is found. In this
resj)ect, therefore, any definite statement can not lie given.

In all cases the numlier of xyleui-plates was detevmined only in young- brauchlets of the
secondary root, nnd not in the uiiin root proceedin»- direct fi'om the seedling, of winch, as I
have already mentioned, T conld olitain no specimens.

Concluding Remarks.

Plaving li'iven a ('om])arative view of the anatomical characters of
different genera of the present family, it remains now to summarise
the results of niv ol)scr\'ations, which are mai id v negative. Thus, I

Tril)e Miupioliea'.

ON THE ANATOMY OF MAONOLlACEit:. 245

have l)eeii uijal)Je to tind any aiiatoiiiicaJ character tliat might «erve
to distiiigiiisli MagiioHaceie as a whole from other dicotyledonous
families. There is certainJy not a single character common to all the
members <jf this family, except such as are common to them and
the members of other dicotyledonous families. Again, I have been
unable to find in most of the species of this family any anatomical
character that might serve to distinguish them (jne from anothei'.
However, I found tliat each of the distinct groups included in the
family is marked out by certain anatomical characters, as the following
synopsis shows.

A. With diaphragms in the pith :

Magnolia^
Michelia,
Liriodeiidron.
V). Without diaphragms in the pith :

I. With crystal-bearing sclerenchymatous elements in the
cortical portion of both stem and ])etiole :

,, , . , [ Tribe ^chizandreœ.
cicluzandm.

II. Witliout crystal -bearing sclerenchymatous elements in

the coi'tical portion of stem and petiole :

1 . A few scattered sclerenchymatous fibres present in

the cortex of tlie stem ; though totally absent in

that of the petiole, and that of the midril) of the

blade. Resin-sacs either indistinct or wantinii' :

lUiclum. Tribe llUcieœ.

'1. Sclerenchymatous sheath well de^'eloped, and

resin-sacs present both in stem and leaf:

IJriiin/s. 'JVibe Illicieœ.

0. Sclerenchymatous fibres well developed in the

14G ö. MAT8UDA.

cortical portion of botli stem and petiole, (iii
a .single case only the libres in the petiole remain-
ing unlignified). Resin-sacs wanting :
a. With a great number of triclioljlasts in the
outer cortex, as well as in the mesophyll ;
wood almost exchisively consisting of
tracheïds :

Twchodendroii. Tribe T roch ode ndrea\
h. Cells containing single prismatic crystals
abundantly found in tlie cortex of the stem :
Cercidiplujllwn. Tribe Trockodcndreœ,
c. Cells containino- an aoorefi'ate of minute
crystals often present in the outer cortex of
the stem ; each aggregate presenting a stel-
late appearance :

Enptckm. Trilje TrocJiodendmi'.
As will be seen from the above synopsis, the four tribes of
Magnoliacea3 may be distinguished from one another by certain
anatomical characters more or less peculiar to each. Of these four
triljes Mcujnolieo' and Scliizundrea' are well defined in tlieir characters
and may be easily distinguished not only from etich other, l)ut from
either of the other tribes. Altln^ugh tinsse two tribes are included in
the same family by most botanists, yet there does not exist any
anatomical character which seems to 1)e sufficient to connect them.
It is true that the secretory reservoirs found in tlie cortex of Manual (ctr,
are faintly represented in Schi'uiiulrcü' by reservoirs quite different in
forju; and that the diaphragms in the pith of Magnolieœ are said to be
represented in Schizandreœ by a few sclerencliyniatous cells.* But I
hesitate to consider these points as sufficient connecting links,

* Bâillon, Natural History of Plants (Translated from the French).

ON THE ANATOMY OF MAGNOLlACEiE. ^47

since there exist in other anatomical characters of these tribes such
marked diiferences as those which I have pointed ont. Thns, then
from a ])nrely anatomical point of view, Magnoliece and Schizmidrece
may be taken without any impropriety to be two distinct groups.

In JUicieœ the only two genera, lllicium and Diiiiiiis^ whicli I
examined, are markedly different in some points, though similar in
general structure. On the one hand, the semicircular arrangement
of the fibro- vascular bundles of the petiole allies this tribe t(3 Scliizan-
dreœ, and cm the other hand, the existence of resin-sacs in Drinujs, and
probably in some species of lllicium, allies it to Magnolieœ. Still the
tribe lUicicœ is widely separated from these two tribes b}" the absence of
certain anatomical characters possessed by one or the other of them :
for instance, both the mucilage-canals and crystal-bearing sclerenchy-
matous elements of the Scliizandreœ, and the diaphragms and scleroblasts
which are found respectively in the pith and in the outer cortex of
Maijnolieœ, are wanting. Tluis, here again is a tribe which may be
anatomically considered as a distinct group.

The wood in Driniij.s is almost exclusively made up of tracheïds,
and the leaf-petiole has its tibro-vascular l^undles arranged in a, semi-
circle, points which ally this genus to Trocliodeudron in another
tribe. Thus lllicic<v and 'irochodcndrcœ might seem to l)e C(jnnected
throufjh these two genera, were it not that the very characters that
ally TrochodendroJL to JUicieœ, widely separate this genus from Eiiptekea
and Cercidipliyllinn, the other two genera of Trochodoidnui'. Further,
these other two genera- are not only but slightly related to each other,
but are also destitute of any anatomical characters that may serve to
connect them with the members of the other tribes. Such being
the case, it is evident that the tribe TrocJiodcndreœ is very indefinite
in its anatomical characters, and that on these grounds Ave ought to
place Trochudcndron in the JlUciea', and establish a new tribe, Eupieleii'^

14.S

s. MA.TSUDA.

which would include Euptelœa tiiid Cercidiphylhtm, the remaining two
genera of Trochodendrea'.

Summing up, the Maijttoliücea may be split up into the following
four groups :

First group, identical with Magtioliece.

Second group, identical witli Scliizandreœ.

Third group, consisting of Trochodendroti and the genera of
lllicicœ.

Fourth group, consisting of Euptelœa and Cercidiphylluiu.
Now, it is worth noticing that the four groups into which
Mao-noliacecT3 might he divided as the result of a purely anatomical
study, correspond in the main to its four tril)es, the distinction be-
tween which is based on the external characters ; and we should not
perhaps be wrong in making the general statement that resemblance
in the external characters of certain plants indicates that there also
exists resemblance in their internal or anatomical structure.

As we have just seen llUcimn and Driuiijs present some marked
differences in their anatomical characters, as do also to some extent
Etrptclœa, Cercidipliylluiii, and TrocJiodcndiwt, which are the three genera
constituting T roch ode lulrav ; but Magnolia, Miclielia, and Einodendron,
the three genera belonging to MaynoUea are so similar in their internal
structure as not to be distinguisliable from one another, and this is also
the case with the two genera of Schiiandreer^ m\n\e\y, Schi :a h dra and
Kadmna. From this we see that anatomical structure is not always
to be relied up(m in distinguishing different genera of the same tribe or
family. However, we must remember that Sclriiafidra and Kadsura have
differences so slight in their generic characters* that lîaillon comliines
tiiem into «jne genus. t Again, Maynolia, Michelio, and Eiriodendron

* Generic characters are generally founded on external peculiarities, and not on in-
ternal ones.

t Bâillon, Tlie Natural Uisturij of Plants, (Translated froui the French).

ox THE ANATOMY OF MAGXOLIACEyE. ^49

are so related in tlieir external eliaracters that there are not wanting-
cases in wliich some species of one of the three genera is often ])laced
in either of the other two. J If we except the case of sncli closely
related «genera, it is probahly the rule that any two genera of the same
tribe or family present anatomic:d eliaracters of such difference as to
distinguish them from each other.

If it is difficult in some cases to distinguish certain genera of
the same tribe or family from one another by anatomical characters
alone, much more would it be so to distinguish in this way species
included in the same genus. Such is actually the case in the ])resent
family, and so it w<Mdd perhaps be in other families.

I Tvish to express here my obligations to Professor Edward
Divers for his kindness in looking through the present article and in
suggesting many improvements in Eiiglisli.

X Thus, ^lafinolia ]nt)iiilii, Andr., JAriodciidron coco. Lour, and Urindeiidnm lilifcra, Liun,
are synonymous; a\so MicJu'lia fu'^cata, B\\\u\e, Magnolia fm^cata, Audi:, and Liriodciulnni F'kjo,
D. C, are synonymous — The Journal of the Linnean Societij, Vol. XXIII, Xo. 150.

Explanation of Plates.
List of Reference Letters.

(Those not found here are explained where they occur.)

ac. aggregate of crystals.

c. cambial zone.

Gp. cortical parenchyma.

cr. crystals.

c.y. crystal-bearing sclerenchymatous element.

en. cuticle.

(///(. diaphragm.

e. endodermis.

ell. epidermal hair.

ej). epidemnis.

es. elongated sclerenchymatous cells.

If. guard cells,

h)>. hypoderma.

/. intercellular space.

U. jinner limit of xylem.

/,•. cork.

;//. cells constituting medullary ray.

)ii(L pith.

vip. passage containing mucilage.

inf)' . passage containing mucilage in process of formation.

p. parenchymatous cells.

pc. pericambium.

j)]i. phloem.

pin. cell constituting phloem ray.

re. sac or reservoir containing crystals.

rn. sac or resei'Aoir containing oily oi- resinous substance.

s. stone-cells or scleroblasts.

EXPLANATION OF PLATES.

sq. group of sclerenchyraatous fibres.

si. sclerenchyraatous fibres or l)ast fibres.

ss. sclerenchyraatous sheath.

t. trichoblasts.

X. xylera.

^. one of the cells tliat form a vow representmg a bundle-slieath

PLATE IL

Plate II.

Fig. 1. — Cross section of phloem and cambial zone of Enptehea poh/amlra . x'285.

w. a zone of newly formed wood, where lignification is incomplete.
Fig. 2. — Longitudinal radial section of outer cortex of Kuptehea jxjli/andra, showing

three crystal-containing sacs. x355.
Fig. 3. — Cross section of epidermis of midrib of leaf of Euptebea polijiuulnt,

presenting cutieular ridges in cross section. X355.
Fig. 4. — Cross section of young root of Euptehea jxjli/andra : A, younger; B, a

little older stage. x200.
Fig. o. — Longitudinal radial section of a portion of phloum of Cercidii>lti/Uti)ii

Japoiiicum, showing crystal-containing sacs ; / indicates the side nearer to

the centre of the stem ; /•, the side farther from the centre. X2ÜÜ.
Fig. 6. — Cross section of the middle portion of petiole of CercidiphijUum. japonicum ;

a, (I, a, mark the zone where distorted cells are found. X45.

Jour. Sc. Coll. Vol. VI. PI. II.

J^ig. 3.

/^'rff. L<

S. Mathilda, del.

Lith. (É Imp. the Seislubiinsli,

Plate HI.

Fig. 7. — Cross section of the middle portion of petiole of Trochodenilro)i /u ulioides.

X45.
Fig. 8. — Cross section of young root of Trodiodendruii araliuides ; endodermis and

pcricambinm not well marked, x 2ÜÜ.
Fig. \). — A trichoblast (separated by maceration) from outer cortex of Trochodeii-

dnui (inilinidc's. XloO.
Fig. lu. — Cross section of a stoma of Trochodendion (indioides ; cs, cup-sliaped

cuticular elevation at the entrance of the stoma. X445.
Fig. 11. — Cross section of phloem and a portion of xylem of IliiciiDii reli(jiosum.

X2b5.
Fig. 12. — A, cross section of a small portion of outer cortex of [lliciiiin, Tashiroi,

showing a resin sac ; B, that of Di'uiujs. dipetdla ; (only dried specimens

were examined). X20Ü.

Jour. Sc. Coll. Vol. VI. PI. III.

s. Matinda, del.

Lith. d- Imp. the Seishihunsna.

PLATE IV.

Plaie IV.

Fig. 13,^ — Cross section o{ owier GOïtex o( Kadzu ru jajjoniat. xllO.

Fio. 1-i. — A, secretory reservoir iu longitudinal tangential section of outer cortex of

Schizandra niiira ; B, tliat of S. ohineitsis ; G, that of K. Jajxniica ; D,

that of voot oî K. /(ij)oiiica. Ail xl30.
FiG. 15. — Longitudinal section of pith of .S'67t/c(//wi^/y< nù/ra. X45.
FiG. 16. — Superficial view of an epidermal cell of Schizuiulra nii/ra containing an

aggregate of crystals. X445.
Fig. 17. — A, cross section of young root of S. vhiiienaia, presenting the triarch

arrangement of xylem-plates ; B, that of a little older one of the same.

X200.
Fig. 18. — Cross section of the middle portion of petiole of Kadzura j<ipo)iica. X45.
Fig. 19. — Cross section of a part of the fibro-vascular bundles of petiole of Kadzura

jajJO)iica. X 200.
Fig. 20. — Various forms of crystal-bearing sclerenchymatous elements ; A and B,

from stem of K.jajjonica ; C and D, from its petiole. X200.
Fig. 21. — Septate bast-fibre from A". Jc/^johü-ö. x70.

Jour. Sc. Coll. Vol. VI. PI, IV.

s Ihlmdii. tirl.

1 ilh, .1 Imp Hl' S,i.llU.un^lut

PLATE V.

Plate V.

Fig. 22. — Cross section of outer cories. oî Maijnol in ohorata. xl80.

Fig. 23. — Lougituclinal tangential section of a small portion of outer cortex of

Michelin Champaca showing resin sacs. xl30.
Fig. 24. — Epidermal hairs of the smaller kind found in Mnt/nnlia nhnvatti, var,

(Kanshu-moJcurenJ. X 355.
Fig. 25. — The basal portion of an epidermal hair from the lower side of the leaf

of Mfu/nolia consjncua, var. pitrpuresceus. X355.
Fig. 2G. — Superficial view of a portion of epidermis of Mni/imlia Watfioni ; hs, basal

portion of an epidermal hair cut crosswise. x355.
Fig. 27. — A, pith of Michelia compressn in longitudinal section showing " diaphragms,"

(xl5); B, the same in cross section, and more magnified (x70); C, in

longitudinal section, and magnified as in B.
Fig. 28. — Outer cortex of MafpioJin liypoleuca, showing hypoderma ; A, cross section

of younger stage ; B, cross section of a little older stage ; C, longitudinal

radial section of the same stage as B. Xl30.
Fig. 29. — Cross section of the leaf of Mt/f/nolia (irtindißora. I, a layer of cells lying

directly below the epidermis of the upper side of the leaf. xl30.
Fig. 30. — Stoma of Mafiuolia pumila in cross section ; cs, a cup-shaped cuticular

4 elevation at the entrance of the stoma. x445.
Fig. 31. — Stoma of Marpwlia (jraudiflora in crosss ection ; guard cells lying directly

upon epidermal cells. x355.
Fig. 32. — A, cross section of young root of MinjnnJin (;n)ii.prt'Sfi/i ; B, longitudinal

section of a small portion of the same, showing an elongated scleren-

chymatous cell. x200.

Jour. Sc. Coll. Vol. VI. PI. V.

^tlH^

Lilh il linij Ihr HHshtlmmlM.

m

m

I

fï

*

m

BJ3

•fê

•fè

-tJ-

■tf

/N

1 .

^

¥

-b

-b

J!

H

--

- j-

Ä

31

B

e

x^

w

fî

m

Tic

m

T

9d

0

Mi

^

m

Researches on the Multiplication

of Elliptic Functions.

By

R. Fujisawa.

Jacobi, in one of the Suites des notices sur les fonctions eUiptirpies, gives,
without cleinonstration, remarkable expressions of sn 2v, sn 3?^, sn 4:V
and sn 5u in terms of tlie differential coefficients of \/.r-(l — ,r-xl — A-.r-)
and yU~^"X —h-'V-) ^.jj^gj^ y^r[i\^ respect to x\ whereby x stands for

sn?/.* Namely, writing

we have

sn Au =

x^

d{x^)

dm

sn Su =

- x^

d{x-)-
(PA '
d{x'f

1 d^A

sn 4u =

1

;T-'

2.3 f/(j^^f

1 r/-'i^

1

rf^l^ dB 1 rr^i3 '

2 dix'f

2

c?(a;^)2 d(:x') 2.8 c?(i--f

1 fPB

1

^«B 1 dm 1 f?*B

a?^-

2.;-^ r/(.r2)B

2.8

f?(,r^f 2 d{x-f 2.8.4 f/(.r2)^

sn 'ju —

1 d'A

1

f/M 1 d'A 1 rfM •

2.8 £?.(a;7

2.8

fZOr-^f 2 fZ(^2^2 2.8.4 (/(.r^'}'

* Crelle's Journal, Bd. IV, pp. 185-193, and Jacobi's gesammelte Werke, Bd. T, pp. 2G6-275.

152 R. FUJI s A WA.

Jncobi adds " hi loi (jcucral de la compomtion iJc cfs expressions est
aisée à saisir,'' and iVirther remarks that we sliaJl have aiialoofoiis
lorinula' l^y usiiio', instead of the ditterentia] coefficients of A and B,
those of

and

V.?'-'(l - -r-^Xl - F-.r') ' /(±^ ./•-)( I - /.-'■') •

x^

The o^eneral form of these expressions is, however, by no means easy
to infer from the ])articular cases just f^iven, and I have tried to trace
among- the writings of Jacohi, the steps wliich might have led him to
these expressions, but without success.

The present memoir is divided into two ])arts. In the first part,
the miiltip]ication-formn]a' of elliptic functions are derived from Abel's
theorem for the elliptic integral of the first kind. It will be seen that
one of the results arrived at is the general formula in question. It
appears, however, liighly improbable th.at flacobi obtained liis formulae
in this way.

In the ])aper just alluded to, pJacobi gives, also without demon-
stration, the partial differential equation satisfied by the numerators and
denominator of the nudtiplication-formula?. This partial differential
e(|uation has since been obtained by lîetti,* Cayley,** Briot et
r)()n(|uet,'j' and others ; but the final results to ])e obtained by
applying it (o the actual evahiation of the numerical constants involved
in the multiplication-formuke, has not, to my knowledge, hitherto
been developed with much completeness or success.

In the second part, Jacobi's partial differential equation is derived
in a manner which is most probably the one followed by Jacobi

* Betti, Annali di Mateniatica, Vol. IV, p. 32.

** Cayley, Cambridge and Dublin Mathematical Journal, Vol. II, pp. 250-26G.

t Briot et Bouquet, Theorie des Fonctions Elliptiques, p. 529.

:MUi;riPLicA'riox of elliptic functions. 153

liiuLself, und tlieii applied to tlie iiivestigiiticju of tlie multiplication-
foi'inuJa'. The ])i-esent paper i« thus composed of two parts, each of
them having referenœ to the tlieory of mnltiphcation hut otherwise
unconnected.

Throughout fhe paper, \ have adopted the notation of Fund.
Nora, the only exception l)eing tViat I write ^1, fi.,, 0.^, H instead of ft
and II, wherehv 1 follow dacobi in his lectures.*

Part First.

Denote the fundamental elliptic irrationality by s and the corres-
ponding Riemann's surface by T. Let S stand for an algeljraic func-
tion of i, one- valued on tlie surface 7', and of the order </, having for
its zero- points

Of, {.--'m I -s^}, fi = \,'2, 3, q,

and for its intinity-points

-'■ IC" i '^"f» î^ = 1, '2, 8, (J,

then iVbcl's theoi'eni is expi'essed by

(1.) y ^^ - y ^ = 0.

Xow .S is necessarily of tlie Inrni

i" + ^ ■s-
U

where l\ (J, 11 are integral functions of ï of the degree <[, <[-- and ([

* Jacobi, Gesaumielte Werke, Bd. I, p. 501.

154

R. FUJIKAWA.

respectively. Tlie iiinneratov 7' + V s- imist be .so (leteriniiied tli;it it
VMtiishes ill q jK)int^! '\i mikI. besides, in niiother set oi' q ])(>iiits

e'v ICr I -6v\, > = 1,'^, a, q.

It will 1)6 convenient to write ^v+p instend of s;, iind then /' + ^^s
vanishes in 2q ])oints

On |.2V 1 -vK /^ = 1, '^, 3, ... (7, q+\, ... 'i^.
Equation (1.) then takes the form

Ai =2'/ .7^

(2.) 2 ^ = 0.

We now take for the fundamental irnitionalitv Iviemann's form

and write

f7?

so that

Sil // .

Two cases are to be distin<iuished accordini»' as n is odd or e^■en.

AVhen // is odd, put n = 2in+ 1=^:2']—], and lei one of the 'Jq jioints
in which J'+(Js vani.shes, coincide with the point { >!' | o } :>nd the
l'ciiiaininu' (2]— 1) points with the p(jint | ,2- | ,s}. E(piation (2.) then
becomes

MULTIPLICATTOX OF ELLIPTIC FUNCTIONS.

155

(3)

-^- + n = 0 , n odtl,

0 s

and, writing P + Qs in full,

we must huve, denoting differentiation with respect to z by D,

K+«i:+ «2:-+ ••+ 'A«+i c^+i + ^v^ + öi('?:) +...+Ö»-! (^r-')=o,

«1 + «22.? + . . . + «,„^1(7». + 1 ).f" + h,Ds + h,D{sz) + . . . + 5,„_iD(s^"' -1) = 0,

(4).

/;„ £>-'".'? + b^D-'\sz)+...

Since «o? ''^n ? ^i? ^ ••• ^^^ ^^^ ''^^ vanish, the determinant obtained

by eliminatinof a^,, «,, , h,„ h, ... must vanish, that is.

(^^)

1 f «'2

1 f Z^

5,

f^2

.9,r,

1 , 2,?, (;;i + 1 )z"\ Ds, Z)(.s.?}, Z)(,s-.2-),

.jD(.s,ï"'~»)

= 0.

Let the expansion of this determinant according to the elements of
the first row be written,

156

R. FU.JTSAWA.

(6) [P„+ a: + . . . +p,„+ir"'^^] + [a+ Qi:+ • ■ +9.-ir^M''^=o.

Theii Ç2ui-{-'2) roots of tlie erjuntion

are C, and r repeated (2m + 1) time.s. Thus we ohtain the followirig
identity :

(7) {P,+P,Z + ... + P„,,,Z''^-^'}'^{Q, + Q,Z + ... + Q,,_,Z^>'''}\^^^

^PUAZ-:){Z-zr^\

Herein putting Z = 0, Z — \^ Z = ^7j- successively and reducing, we
obtain

(8)

V:

Vi-r {i-z) '

Pn

Vl-A-C {l-'k-z) 2

p '

Po + Pi+...+P^^l^

Pmfl

P,

jre+i

In extracting squnre root, strictly speaking, we have to ])refix the
double sign ± ; but by taking some particular value of w or by putting
/,•=() in the final results to be hereafter obtained, it comes out that we

have to take the + sign.

§ -

Let us now investigate the êX])ressions which occur on the right-
hand side of equation (<S). For this purpose, put

MlMVni'LI<A'l'K)X OF EliLIPTFC FU\("I'K>NS.

157

CM

A =

I)'" l^^ ]y"-S- 7)ml2_^.,m-l

n'"'s, J )-"'.s.';-, 7)-"'.s.?"'-i

Inrtlier let the cxpiinsion of A arconliii^" to tlic clcinctits of the lirst
rt)\v lie wriflcii :

(10.) A ^ D"'+Ks . Ai + ])"'^h.y . As + . . . + D"'+i.s^?'"-i . A,

Xow

.) J\ =

1 , 2z, . . .{m + 1 )z"\ Ds, Dfi:, Dsf"-'^

{m + 1 ) ! , D'" ^^s D"'+hz, 7^'«+i.s .'»-1

D2"'.<?, J)5"'.<?r, 7)2'".S,:''

= A

, ~2 «)« j-îrt+l

»•^j '* ) ■^

, -ir, m:"'-\(ni+ \):

})i ! , (»1+ 1)!

-(/^/+iy.Ai

^, /:-, . .

z

,S

1,-Jr,..

. . . /;/

^■m -

-1

D.s

-(?;/ + l)!Ao

;;/ ! , 7)"'.s
'2:, 'm.f"^\ Bft-

-(w + niA,

lôS

1.'. FUJrSAWA.

Writiiiu'. for slioi'tncss,

1! -2! ...

m ! = m II

wo hnvo

- y2 „wj -.?H+1

■^ ) - > •< ) -

1,2", m.f"'\{}n + \)f"

;;/ !! .^"'+^ ;

îilso

Z, .?-, . . . .■ "^ a;

/// ! , 7)'" .S'

I . 1 . . . . 1 , .s
1. 2, ... w, :7).s-

;;^ ! , fl)'"!^
^ (;;/ — ] ) ! ! [ ■.■"'7)"' .s- — VI:'" -'^ly ^s- + , . . -f ( ■

)'"/// 1 .s-

imilnrlv

\,\i.-,...iiiv"'-\ Dkv
m ! , 7)"'x:-

wi ! , 7>"^r

- (///-!)!! ■•"'+i7)"'.s' ,

= {m—\)\\ :'"'+V7>"'.v; ,

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

159

yin „ „m— 1

m ! , D"\sz"'-'^)

{m—l)ll ^™+iD'^(s^"' 2)

Hen

ce

(12.) P,=^m !! z"'+'\a-{)ii+\) TW^Yai
Aoain

+ D"\s. A., + . . . + D"'sz"''-. A

.]

(13.) P,„+i = (-l)'"+^

},z, z^,... z'", s, sz,

. sz

l,'2z,... mz'"-"'^, Ds, Dsz, Dsz'"''^

m\,D"'s, D"\sz, ...D"'sz"''^
B"^+h,D"'+hz,...D'''^hz"'^^

D-"'s, D'"'sz ...D-"'sz"'-'^

= (-1)'«+1 ;;i!! A .

Suiwtituting the values of 1\ and P,„+i just found in the tirsf uf

2in+l

e(juation.s ((S.) and dividing by z ^ , we get

(14.)

V: = (-i)'«+V2

_A--(/M+ 1)|/J''/— YAl + Z)'^^A,,+ ... + D''^s;^•™-2. A,

A

The expression f(K- A given in ([).) may be greatly sinipJitied.
J>y an easy reduction, we hud

(15.)
A =

D"'+\S {m+ \)D"'s, {,ic + l)D"'-i.s, {{)n+ \)iii. . .^ \D-s

W'-^h, {m + 2)£>'«+\s, (VM + 2)u/i + 1 )D"\s, { {m + 2)(??i + 1) . . . 4 [ D=^s

£»2^s, '2mD-"''h, 'Imi^m - 1 )L>=^"'--'i-, { 2m{2ni - 1) . . . (yy^ + 2) [ D"'+\s

160

R. FUJISAWA.

th;it is,

Mm-l) (On A

(16.) A=(-l) '^ ^■'"''

m !!

'2!^^' 3!^^' {m+l)r '

B^o T)i<i D"+-<;

The expression

(/;i+l)p'«— .Ai + £)"'6'.A2+ .. + Z)'"s^"'-lA^j

which occurs on the righthand side of (12.), admits of being simplified
in a simiJar manner ; namely denoting, for a moment, this expression
by a', we liave

A'=(m+1)

D'"^-, D"'.s, D™s.s"'-'

W !!

z

3! '

'2 ! ,î- '
1

4!

D's,.

^ -.D"^-^h, ,_ ^.,.Z)"'ns,

(m + 1)! '(m + 2)!

1 ^ ,s

y;t ! /:

(/yi + '2)!
1

Z)"'+-.s

(2;;i)!

D-"''.ç

Leaving the factor ( — 1)"'+! V^ . ( — 1) ^ .. oat of consideration,

tlie numerator of the righthand side of e(|ualion (S.) becomes

MULTIPLTCATTOX OF ELLIPTIC FUNCTIONS.

KU

2 ! ' 3 ! '

— D^s — — 7)^.<?

8 ! ' 4 ! '

1

(w+1)!

1

(7// + 1)! ''(w + 'i)! "

{^2m)\

-D2-.S

— — D'ls

o !

2 ! ^ '

_1_

4!

D's,

(w+1)! "(7;^+2)r "■

1 D»A

1

(w + 2)!

D'»+"-^.<?

C27?0!

D2'».S

The two determinants may l)e componnded to,2:ether ; thus we obtain

(17.)

i^Kt> -^K-) (^!°"^'(t)

3!

ns.s — — D'.s

1

(w + 2)!

_pM4 25

(m+l)! ''(7» + 2;!

(!2m)\

A .slight]}^ diiferent form miglit l)e üiven to this expression, viz

(18.)

l/^i +

162

E. FUJISAWA.

Let this determinant (omitting the factor f") be called il/, and the
determinant on the righthand side of (IH.) M. 01)serve that i)/ and
A differ from each other only by a mumerical factor. E((iiation (14.)
now assumes the form :

(19.)

Vr =(-ir+^^'

f Ml

M

Consider now

§• 4

p„ + p, + ... + p,

m+l

Avliicli may he written in tlie form of a determinant, viz.

»i=^1H+l

(20.) 2 P>^

1,1, 1, 1, 0, 0, 0

1 Z ?} ;?'"+\ ,S, ,S,e, .S.^'"-^

1 2.? (w + l)/% D^, B^z, Z)ä2'"'-i

(7;^ + 1 )!, 7)'"+^S, D"'+1,S?, -£,m^r\ç,ym-\

Tliis determinant may he reduced in exactly the same manner as J\„
îind then we find

MULTTPLICATIOX OP ELLIPTIC FUNCTIONS.

ig;^

H=»i+1

2P.==A

1,1, 1,

1, 1

„ill ç,?H + l

1,.?, .?^

1,2;?, 'mz"'-\ (;» + l):''

m!, (w + 1)!,«'

No

w

-(w + 1)! Ai

1,1, 1, 1,0

1, ^, ^^ z'^, s

1, 2,?, m.z"'-\ Ds

m ! , D"'s

(m+l;! A,

1,1, 1, 1,0

1, Z, Z^, Z"', fiZ

l,'2z, viz'^-\ Dfiz

m ! , D'^fiz

:w + l)! A,

1,1, 1, 1,0

1, ,?, z\

] , '2 ?, ;;? i"'~\ Z).sr"*~^

wi ! , D"'.s::™-^

1,1, 1, 1,1

1, 2^, mf"-\ (w + 1)-;™

?;i ! , {vi + 1 ) ! ,•-'

(-1)"'+';«!!(1 -;?)»'+» ,

164

R. FU.ITSAWA.

1,1, 1,

1,0

1 r, ,~2 „in o

1,2?, wr"'-\ D^

- - (- ] )"'+\?^^ - 1 )!!(1 -, :')"'+'!)"

1-? '

1,1, 1, 1,0

X, /i, /4 , A. , o..

1,2,?,

.. 7;?-?'

D,s?

?» ! , Z)"'.s,?

( - 1 r+^.7;i - 1)!!(1 - ,?r+iD"'^^

1,1, 1,

1, z, z\

1,0

,,'»-1 T"loV'«— 1

1, 2,?, y;7?'"-\ Z)«

m ! , D'".s'«-i

-(- \y"-^\m- 1)!!(1 - ?/«+iD"

l-.e

He

nee

n— »l+l

(i^ii ^ L_ i - J ;?

l-z

(-l)"'+'w!!(l-.:r+2

7)"

.'J 1

J^m+\

7>"'+2s', I)'"+2s^^ J)'«+2.<5 -'

D^^i; J)^>»<fy r)2»«^~?H-l

Further, we dcclnee witlioiit inueli difîienlfv

MULTIPLICATION OF ELLIPTIC FUN'CTIONS.

165

H=0

1

1

,»»+2

i

i\ \-z '8! V-z

1

3

-l^ o, J. t '

I

(ni+\)\ '(w + i)

('Jy;^)!

-D'"

or

(1=0

m{m~\>

[2'A.) 2 ^A. = (-l) 2 (-l)"'+V2m.)!l(I~,if"

1

1+1

_1_ ,^
•2\ 1-;

D'^

8! 1-
1 jj. s

3\ i-z' 4! i-z

1 ^i;.+i ^^'

[111+ [y. v~z

{in + n \

^ -n-^-M •'^

1 s

1 ^..- ^

(w+1)! i-,^ ' (;yi4-'2)! i-^ ' " (2/;^)! l-z

DeuDtiiig the deteriiiiuaiit on tlie rightbaud side of (--.) by Mo

and renieinI)ei'iiJi'-

\ve get l)y division

P™+i '"^ '' iVf

iSiibsLitiitiijg this in the .-second of equations (.S.). we (obtain

166

K. FUJISAWA.

li=m+\

§• 5.

Let us consider the determinunt

/.2,»+2^ A--"\ A:2'""^ ... 1, 0, 0, ... 0

1, Z, Z\ ... Z"'-^\ S, HZ, ... SZ'"-^

1, 2z, ... {:m+\)z"\Ds, Dsz, ... Dsf"-^

2 P.iJc'

2)«+2-2ti

This determinant is of the sunie form as tlie one in (20.), and we tind
likewise

fi=0

+ D"

sz

^_^^ .A,+ ... +D-,— ^^. A

sz"

]

:(-ir+'y/ii!a-A-2.^y"+2

D"

^ J)m-Ti . ^'^

^SZ,

D

m+l_

SZ"

\-k^z

Z)"'+-',s, D"'^hz, ... Z)"'+2s^'»-i

D2'».ç,

Z)-"'s^, ... D-"\s2'"

whence

(•25.) 2 PA'A'2"'+-'-2M = (_iyH+i^2;;i)!!(l-Ä2^')"

Ln.'^ Lna_l

Z)

D

D"

2! l-A^^' 8! i-Â;^,^'-U/i+l)! i-Ä;-.

8!

B's,

4!

DSs,

■(m + 2)

:D'«^-,S

1

[m+iy ' (m + 2)!

".s, . . ,

7)2'» s

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

167

or

ti=m+l
)i=0

m(m—l)

C20.) 2 iV/^-"'+'^''=(-ir^X-l) ' (2m)!!(l-^^Ä')'

2.A2(n+l

^^D^^

'J! l-A-2^'

1.^. '^

1

8! i-^'^

3! l-A'^' 4! l-A'^^

-.D

TO+1_

1 ..

(;« + 'i)! i — A-^

1

.D

»i+i.

-.D"

1 ,s

>+T)r 1-^2^ ' (7;i + 2)! 1 - A-2-J ' ■ • ■ {'2m)\ 1 - Ä;^^
Cull the dererininarit on the righthand side of (26.) M„ we liave

p -(I-A^j -^ .

SubstitutiDg this in the last of equations (8.) we obtain

C27.)

Vi-A'c = (i-^'-2)'

M '

§. (;.

In case // is even, sny // = 2»?, put m = q - i, and, referring to
§. 1., let one of the 2^/ points in which 7^ + Qs vanishes, coincide with
the point IC I ^}, another point witli |0 | 0)-, and the remaining
(2q-2) ])oints with the point {z \ s\ . Then

(28.)

dZ , dz ,,

—^ + II = 0 , n even,

0 s

and

P-\-Qs = [a,z + a.j- +...+ <*,„+i^"'+') + {K+byZ+...+ b„,^,z^"^')s,

168

11. FUJISAWA.

«0 bciiïg' zero. If, as will be convenient, we write

k-C , s

^ -V

then ^^±2i = (rt^ + rt,.+ .. +rt„,i""0 + (l}, + b,z+... + b,„^,^u,
z

and

(29.)

+ ö„,_iD'" («^™-i} =0,

whence

(80.)

1,:, :^...

1,^, ^^...

1 , 'Iz, . . . ni

7\ 0, 0r,

Ô'™, IL, ItZ,

r -\ Du, Duz,

0^m 1
71Z"'^-^

DiLz'"-^

m\,D"'u, D"'uz, ...D"'uz"'-^
D"'+hi, D'^'hiz, ... D"'+htz'"'-'^
D"''^-u, D"'^hiz, ...D"'+-icz"'-''

D'"'-hi, D-"'-hiz, ... D-"'-htz"'-^

= 0

The expansion of this determinant according to the elements of
the first row may he written :

MULTIPLTOATTOX OF ELLIPTIC FUXCTTOXS.

1()1)

(31.) {Pi+p,:+ ..■\-p„„,:"'\ + ia+^w+ •. + a.-ir'n 0 = o-

Hence

Herein putting Z=0, Z = l, Z = -7;2 successively, we oLtnin, after sligh)
reduction,

(88.) ^ ^Yzr^d^zY ^

Vi-A'r(i-/.-.?y" =

^ m+1

PiA-"' + PoA-"'"^+... + P„Hi

Tlie reduction of (he expressions ■which occur on the rig'hthand
side of (38.) runs on the r;anie line as tlie reduction of the corresponding
expressions when n is odd, discnssed somewhat in detail in preceding
sections. We niay therefore at once w'rite down tlie results ;

(34.)

i'„Hi = (-l) ' C27;i-1,!:

\\^ z

2! 0

in\ z

8! ;? ' •••(w+])!^ z

m\ z '(m+1)! z ' '■- {2m-\)\

B"-

170

R. FUJI SA WA.

a = (-l)'"'^^^(2^«-l)!!

1

4!

D's,

fi t

Dl.,

1

(w + 2)!

(y//'

-i 2)™+is ! D'"-<i = D-

+ 1)! '(w + '2)! ' ■■2m -1)1

(36.) 2 PM-(-l)'^'^(2w-l)!!(l-^f"

M = l

^I) -^

^ D^- -^

1 ! ?(l-,~)' 2! 41-;?;

-i-D"' — - —

m! z[\-'z)

1_^. .

L^s -^

1

D"

2! ^,1-;?)!' 3! z{l-z)' ■■■(w + 1)! ^1--?)

1.^. -^

,D"

.1)2"

w! 41-^)'(w + l)! ."(1-^)' ■■■ (2m-l)l z{l-z)

(37.) 2, Pßl--'" --'" = {-}) -' (2w-l)!!(l-Â-Vf»

'd- '

1^._ -^

1! z[\-}r.zy 2\ zil-Pzy
s i _.. .s

yy/! z{l-A-z)

-Id

iz)

1

Z)"

2 ! .:(1 - P^)' H\ zi]- ¥zy ■ ■ ■ 'vi+ 1)! ^1 - F^

1

D

1 .s

7)"'^l

1

D'-

m\ z{\ - l^zf {m-\- ] )! ^(1 - ]^h)' ' ' ' {2m - \)\ z{\ - A-^)

Cîilling the deteriiiinniits on the right-haud sides of equations (84-o7.)

N^, A^ij A^2, iVg in order, and, substituting in (33.), we ol)tain

( J^ — r — lV"-i — ^

(3.S.)

vr

VI -/.-r = (i-H?r

N

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

171

Writing-, as is usutil,

S. 8.

so that

^/z = so?« ,

we have, in virtue of equiition« (3.) and (2(S.),

V'c — — sn mi,

vr

= cunu,

where n is any even or odd integer.

When // is odd, say // = 2m + 1, we have from (16.), (19.), (-o.)
and (27.)

I sn nu = { — \)^z

M '

(89.)

en nu — (1 — ^)-

^ ilf.

M '

where

(40.) ilA

M '

1 ,s 1 .<?

'2! 2 ' 3! ^ '

1 .s

4d'-

1-D'-'

1 ,9

8! ^ ' 4! ;? ' ■•■(m+'2)! ^

1 .s

1 sis

^ T)TO+2 ^ ^ r)2?»

(w+1)! ,r ' (7» + '2)! ^ '■■' i2w)!

172

R. FUJTSAWA.

l^o s

(41.) M,=^

iß~ •'

1

D"

2! 1-;?' 8! 1--' "(m+\)r \-z

1 ^, s

^-D' '

1

D"

3! 1-;?' 4! 1-;?' ■■■iw+2)! 1-

7)"> + ^ '^ J)m+2 ■''

1 .<?

(w+1)! l-;?'(w+'2)! I-;?'" (2w)! 1-.?

(42.; il/, =

iß" •''

l._D. -^

1 .<;

2! 1-ÄV 3! 1-Ä-V ■■■(w+l;! \-V-z

il)" -^

1 D^^

1 ,s

:!^ J^m + 2

3 ! ^ 1 _ /;2^ ' 4 ! 1 - Pz' ■ ■ ■ {m + 2)! 1 - Z--'^

1 .si .s

1 S

(7;i + 1 )! 1 - /r^' {m + 2}! 1 - lih' ' ' " (2??i)' ^ - A'?

(43.) M =

^B%

——D^s

3 !

3!

J_

4!

D's,

1

(w+1)!

1

(w + 2)!

(m+1)! ''(m + 2)! ''

(2;«.)!

D""s

111 case // is even, say fi = 2i]i, we have, in virtue of (SS.)?

(44.)

N '

' sn im = ( — 1)'
en nu = (1 — ^)'

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

173

vvliere

(45.; iVi =

a!
J_

4!

D's,

_1_
4!

1

D%

■•■(w+i)!
5! ' ■••(m + '2)!

[m+lû '(m + 2)! '"•('Jw-ljr

(46.) iV.

'-fl ■'

1

Z)2

1! 41-.:;)' '2! ;2a-^)'

——D''"' — -

ml z[i-~z)

i.ß. •'

'-D^ •'

1

D

m+l

2! nl--^)' 'àl -ni-'?)' ■■■(w+1)! ^1-^)

l.Z)- '

A _2)™+^ -

J)2n

ml z{i - z)' [m + 1 )! z{l-zy '" C2m - 1)! z{ i-z)

(47.) N, =

1

D

1 .Z)._ '^

^D"

1 !^( 1 - A:-'^)' 2 ! ,^( 1 - Ä2?)' ■ • • m\ z{l- Pz)

\d^. '

1

■D

1

D"

2 ! ^(1 - A-2^)' -öl z{\- k-z)' ■ • • {m+ 1)! ^(1 - A-2

i.D- '

1 s

1

D-

»il z{l-k'z)' {i)i+\y z{i-k'zy " (2';/i-Jj! z{l-k^z)

(4S.) iV

1 2)^ J-D^^

1 !

^ I z

2 !

3! z

nil z

^ Jjm + l '*''

(m+l)! ^

-^D"

^ £)m + l "^

i^ J)2fft-1 ^

(m+ly! - ' '" i'^m-^y z

It will be observed that the loriuulie of ^wnn for // = 2, o, 4, 5,
are those given by Jacobi in the Memoir referred to, which appeared

1 74 ^- FUJISAWA.

on the opening page of the present paper. I was not aware of the
existence of Jacobi's formulae when I first found the formula for
snmt which was, in fact, then obtained in a sUghtly different form
as furnished by (17.), and which is perhaps in some respects preferable
to the one here given. I have, however, given it its present shape,
in order that, for the particular values of //, it may exactly coincide
with the formula} given by the illustrious mathematician whose
memory is sacred to every student of the theory of elliptic functions.

§. 9.

Before proceeding further with the reduction of the multiplica-
tion-formulas, it is necessary to give a few formula? relating to the
differentiation of composite functions, to which frequent reference will
subsequently be made.

Let u be a function of y and y a function of x. It is required to
find the n"' differential coefficient of a with respect to x. By actual
differentiation, we find

du _ du dt/
dx dy dx '

dPu
dx

I _ dit d-y d-n. / di/ \-
'' ~ ~dy W' "^ 111/' \dx) '

(Pa _ dt(_ d^ d-u dy d-y d^it / dy V

dx' ~ dy Ix'^'dj/'l^'dx''^ ~dJf \dx) '

dx' dy dx' "^ dy' \ ^ dx dx' "^ ^ V dx') S "^ *' dy' \W) W

.dhi/dyV
■^ d,/\dxj'

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

175

d^u _ du d'^ij d''n ( ^ dij d'^ij . d^ji dhj
dx-' dy dx^ dif ( dx dx^ dx^ dx^

d!^ii/dij\'dhj d'u/d//\

\ dx ) dx^ dy^ \ dx ) '

dy* \ dx / dx^ dy^

and, generally,

d"iù du d'il .^^'■'S'a- I d'"'>''Y ,'^"'W

^^^^ -d^ ^ 7hj^'-^df^'^---^dV^'-^---^l^^'^-''^dr^'^'

where Aj, Ag, V,^ denote a« yet unknown l'uiK^tions which contain

only the differential coefficients of // with respect to x. A few of the
initial and end coefficients may at (jnce be written down :

Y —

d^
dx'

du d"-h/ nhi—l) dh/ d" hi ,

LttAj CLJu

2 ! dx' dx"~
ln{n-l)...{n ^ + 1) ^^ 2 ^j 2

+

('¥)

dx '' dx ''

n(>i— l)...Oi— -IJ-+1) If-in

(50) /

'^ • (^)

, dx '-^

, (/i even)

A..

;iO?. — 1 )(yt - 2) / r?// \ "--^ rZ'V/

/ dy Y~'' d !/
\dx J Tbâ

+

n{ii— 1 )(y^ — -IXn—'^) ( dy

(djiX'Vdj/\
\dx ) \dx-)

_ n{n.~\) /dyy-'-d'y
"—1 o I I ,7 ., I .7 ..2

^A.

/djiY~'d^

dî^-

17(5 1^- FüJISAWA.

To find X,., observe that A'., being a function of tlie differential
coefficients of y with respect to x only, is independent of u as a
function of y. Hence, putting u = y, y'r-y" successively, we obtain

/ d"y _ ^

'^ = 2yX, + 2X,,

'^ = ry'-'X.+ rir- 1)//-%+ ... + rlX,,

\ '^ = ny^^-'X,+ n{n~l)y--'X,+ +n! X„

To obtain A^j, X.,,..., we may solve these equations as was done
by Ijertnind* ; but it is shorter to proceed as follows: — Put n = e^'-^,
then

c-^"^^ = /Ai + /=^X+. ..+/%+. .. + /%,,

and, on the other hand,

d"c^'' _ . d"y '/? d"y^ )? d"y^ /" fZ"/y"

~dx^ ~ ^'dr^"^ TT dx" ^ 3 1 ~dx^'^'"'^ nl dx" '

Multiplying the last two equations together and ecpiating the coefficients
of the like powers of A., w^e obtain

* Bertraud, Traité de Calcul Différeutial et de Calcul Integral, T. I, p. 139.

MULTIPLTCATTOX OF ELLIPTIC FUNCTIONS. ^77

^2 ^2 ! f?.r" ^ d.r" '

(51)

'" ~ v! c/^r" (/•-!}! 1! cZj-" "'*(/•- 2)! 2! fZ.r" ••

+(-ir^

l!(r-l)! f/.r" '

»~ 71 1 d:r'' («-1)!1! ^;r""i (n-2)!2! dx"

\ ^^ ^ l!(7i-l)!fZ:r»'

It may be worth while to notice that the expression on the opposite

side of A', is identically zero for ;dl values of r oreater than n.

From the mode of derivation it is evident that A,, contains

1 d'^dr)
differential coefficients of?/ but not ?/ itself. Hence, if — j — '^^ be

./ ./ ' r ! dx"

broken u]) into two parts such that one part contains all the terms
independent of explicit // and the other part, terms having y as a factor,
then

1 d"(u')
X,. = that part of -^j j^ which is independent of explicit y

1 d'KiD
and that ])art of -^ "77 n which contains explicit // together with

y cny'-') y' d^y"^^) , , -.y, f ^^y

(r-l)!l! dx'' "^ (r-2)!2! dx"" "^ ^ ' \\{r-\)\ dx"
is identically zero. Now

V ! dx"

rocffioieiit of /(" in \i/-\-Ji^ + -rr ^,+ ■ (
(■ dx 2 ! dx- )

178

R. FUJTSAWA.

Hence tliat part of ^-j f^ wliidi is indcpeiKlent of ex])li('it // is

r ! clx
equal to llie coefficient of //" in

that if^, in

TT I TÎJ'^^TÏ?^

^]/(-r + /0-/(.r)} ,

^vliere y=fi-c)-

Thus we ohtsiin

(52, .X, = ^[(Ay|/(.,. + /„-/(.,)|]

Tin's form of A',, has been ohtained in a different manner by U. ^leyer.*
liertrnnd oives tlie followinii' fcjrm of A',, which is sul)stantiall_y the
same as (ol) :**

r ! ax''

where a is to he regarded as constant during ditlerentiation and is
afterwards to l)e re]^laced hv //. Again m^dcing use of the form of
A',, given hy (.51), Ave get

r/.r" ~ ,6l r! (If fl.r'' -^ ^Jr—]}\ (If dx"

/y _J__^ ^7^11 _ . _iv,-i«i^'^
■^ -^ ;e^,(v-2)! r7//- rjx" '"'^^ ^ -^ (Ifdx'''

* Glrnnert's Archiv d<'v IMathouiatik, Bd. IX.
** Loe. cit. p. 1 10.

MULTTPLTCATTOX OF ELLIPTIC FUXC'J'IONS. J 79

whicli nuTecs \\\\\\ flie form of -7-- o-iven l)y R. iro|)i)0.*

Put u = y'\ then —. — = /^. //'"''. where />,. denotes tlie continued
(///' J ' ■

product of /• (|unntities p, yj — l, |) — :2, ...(/> — / + 1). ■\rnltij)lyin<i- e<|u:i-

tions (51) by pjf'~\ /)(/>— 1)//'"",. . . in order und tiddinu', we obtain

For /) = -^, we hîive

;• ! 2y— 1 ax" 2/? — 1 dx" )

d"ir
If ?/ be n rntionnl iuteofrnl function of .r of the tjn'rd deoTee, -r^

,-, -^ dx"

v:iriishes for nil values of r for whicli 'ir < //. In this case, denotino- l)y
/ eitlier — - or the intei>er next nbove -7— according- as // is a inulli])le
of P) or not. we have

,r., d\/Tj_ _ j^^^j^i, .-,j!^ jr!_ iV .

^^^ dx" ~ 2-'\n\f ?/"--• r ' /! 2/-1 dx"^'-
Aofain

+ {-\r\ 1.3. ...(2r-3)(2/yr'-A;+. ..+(-!)". 1.3. ...(2/?-3)X,| ,

* Crelk-'s Journal, UJ. XXXIII.

],S0 R. FUJISAWA.

niid if// be ;i ratioiml iiite^Tal fnnotioii of the tliird degree, -y^ and
all the higher differential coefficients vanish, so tliat

: coef. of h- i„ 4 j Ä 4^^ + ^ ;,= i!* + I /,' 4^ }' ,

r ! ( d.r 'J d.r (j d.r )

Ji-r

X„_, - coef. of //' m --— -r -y;. + TT ^' -xj- + IT '^'

or

UT — r>r>of (^f 7)' 111 I ,, _ ,„

(/i — r)! ( dx '2 c?u^'- 6 djf

wlienee follows that A",,., vanislies for all values of r for which

2»
8

2»
r > :lu — '2t\ that is, r)/- ^ i^//. Denoting the integral ]>art of—- by /,

we have

+ (--lrl.a...(2;/-2r-3)(2/yyX„_,+ ... + 1.8....(2«--3)X„| ,
wliere

"-'- ■ |(» - 2;-)! r! V f7.r / V '2 r/.r"-^ /

■^C»-2r+l}!(r-2)ll! V7/7y/ \2d.r'0 \(]d.r)

{n - 2r + 2;! (r - 4)! 2! V dx ) \ 2 dx'') \ G f/./'V "^ ' ' ' ) '

+

the last term beins"

1 (ävS:~'u\ d'iiy

(,,_.;j;.)!(J_). \dx) V«', ./W

MULTIPLICATION OF ELLIPTIC FUNCTIONS. ]^<^]^

(^n- ^-^) ! 1 ! (^) ! V di' J V -i dj^'J \ Ü d.r'J '

accordiiJii' a8 r is even or odd.
Siiiiihirly

+ (-lr.l.8....(2«-t^/--lX%)'-X„_+...+ 1.8. ...C2»,-1)X„| ,
where A„_,. has the same signification as in (57).

Let / = 6- = ,^1 -,?)(! -Ä:^^) = ^-(l + ^V+A-V ,

Â- ^ - l-2(l+^•v+3^■v,

Now write

(59) -:^D^s = ]^^S,,,

then by (57) and (5<S)

((31) S„ = (-1À1.8. ...(2»-2/-3)(2/)%,_,+ ...

+ i-\y.\.3. ...ßn-'2r--S)(!2/yZ„_,.+ ... + L3. .. (2//.-3)Z„ ,

l<;2 ^- FUJISAWA.

((32) B,, = i-iy.VS, ...C2n-2i-l)C2/yZ,,_,+ ...

+ (-lr.l.3....(2;i-2r-lX'2/)%,_,+ ... + 1.8. ...(2,i-l)Z„ ,
where

fn-2rfr /n-2r-l/r-2/ / /t-2r-2/")-4r2
/f'0\ ry _ yi 72 ■ ./l 72 ./s J ./ 1 Ji J-i 1

\K^ö) z/„_, (,i_2?-)!/-!'^(/i-2r+l)!(r-2)!l!"^(«-2/-+2)!(r-4)!2'. ■" '
the lust term iDeing

accordinii' as /■ is even or odd.

Observe that both .S„ and //„ are rational integral functions of z of the
degree 2/<.

§ K'-

The determinant-expressions of § <S, simple as they niav a])pear,
are not convenient for introducing .S„ and Yi',, of the last section. It
is desirable to derive the multiplication-formulce in forms slightly
different from those given in § 8.

Consider first the case n odd. Referring to § '1^ we might Iku'c

p
taken yi} instead of /'+''^As' and then we shajl ha\'e in [)lace of (5)

the following determinant :

((U)

MULTIPLICATION OF ELLIPTIC PUXCTIOXS.
1

183

c

"2

f-m+l

6'

■■ 6

z

z'

^m+l

s '

S

1 Z z- 2™+^

1, 2z, ... (w- 1)2™- 2 D— , D— , D— , ... D- —

s s s s

1 ^ - 2 /.y?i

(;«" 1)! , D™-i— , D"'-i— , D™^^— , ... D"^ ^—
' s s s s

1 ,- ^2 .,»1+1

.9 .9 S S

m+l

2)m+l_ J)in+\JL- Dm+lf_ J)in+\ ^

S ' S ' 5 ' ' " S

1 „ ,2 ^,m+l
jyim±_ JJ2m_ JJ^mjL^ D^m^

S ' .9 ' S ' '" S

= 0

Now put

(65)

iJ

2)m-l_î_ D^-l^ ... i)»-i:^

.9 S S

1 ~ p,?«+l

D"'J_, D«_, ... D"'i —
■s s s

1

ym+l

Ty2m__ JJ2m '^ jyim ^

.9 ' S ' ■■■ S

nnd let the expau.sion of this determinant îiccording to the elements
of the first row be written :

s s s

We find

a/C ^

2»«+! 0

Vi-C (1-^) ^- =

i^

VI -^'r (i-A-^) '' =

u±l j^^p,H2^/^^^2,»_,_ ^O^

■''^m+2

1<S4

li. FUJISAWA.

und thence, // being odd,

«36)

/ e

n 71/- '

dn;m = {1-khf'^^,

sn //?^

en mi

where

(67) M'

s '
1 !^s '

1! s '

9, ! -^ c '

m'. -s
^ n»t+i *

___7)»"_!_ JJm+1 ^ jyin

m\ s ' {m+\)l s ' '" {2711)1 s

(68) Ml'

— D-

i! 5

i ! s ml s

'2! .9 ' •••(m+1)! .s-

m! s ' {7)1+ \)l s ' ■■■ {'2711)1 s

(69) Mg' =

(l_^)"')i, (1-^)™, (l-^)™-\

0,

J_

s

1 I .s

lis' 2 ! .9 '

7111 S

___L__r)m+l_L

(w+ 1)! s

(m-\)\^ s 'ml s'{m+\)r .s'-'(2w)! .s

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

185

(70) ii/;=

ii-r-zT+\ r-{i-r~zr, Ä-\i-Ä^.^r-s ... a-"

0,

]_

s '■

-1 ±d'-

s ' lis'

lis' ■!'. s'

. 1 D»-I

ml s

1 D..-.1 ±D- 1 1

(7;t— 1)! s ' m! s ' {iit+ i}\

D"

1 _jr)2,„_L

(li/yij:

Ob.serve that the expression of sn mi ns giyen hy (f 7) and ((iS) is
that which has been indicated by Jacobi. It wiJJ, liowever, be more
convenient to write i)// as follows :

(71) in; =

ym+l
•' >

0,

J_

s '

— Z"

..(-1)"

-1 ^dI

5 ' 1! s'

1 ! s ' t2 ! s '

«il S

■•(;y;+L)! 6'

1 T.. 1

C2;;i)! .s

We may liere point ont that the comparison of the preceding
formula' with the corres])onding expressions of § 8, gives some elegant
theorems in determinants. Take for example il/ and M' . We find
easily

where C denotes a constant wliich may be a function of m. To
determine C we may take some particular values of m. For ;// = 1,
we see at once that

•2!

Bh = -.s^

1, dK

I)

-s- ' '2

-D-

186

E. FUJISAWA.

For ?// = 2, we hnve

s s 2! ,s

S s 2' s

.s- 2! s àl s

= s'

SS s 2! ,9 s àl s

±D^^1 J-D^l ±i)J
2r s' 3!^ s' 4!^ s

S 21 s' s àl s' s 4! .s

1 1 „.1

,ç àl s s 21 ,s

1 \-

Multiply the first row by D— and subtract the product from tlie se-
cond row, and then, mu]ti])ly the first column by I) — and subtract
the product froui the secoiid column ; and, observing

l^Dh = -s{d—)\2sW^ . A.D2l_,2 1 j).l
à I \ s / s2! s als

( s à I s \ 2 ! s / ) 4 ! s

M'e 2 et

^Pi, 4.D'1

1!

s ' à

-Ld=1 J_d,1 Xo.i

•J ! s ' 3 ! « ' 4 ! .5

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

187

Thus we find C = (-1)'" and thence

Çrl)

^^'''' ■■■i^irf

{»i + 1

,D"'^\9,

(2«i)!

D-'^s

/ 1 \m^,'2iii + l

1 ^d1

s ' lis

lis' 2 ! s

^ — _D"'+i2

■■■ {m+l)\ s

which identity may nhso be proved directly in the manner exempli hed
by the particular case m =^^2.

§ 11-

Consider next the case // even. Referring' to § 6, if we take
\-Q instead of i'+C^As, we sh-all have in place of (oO) the following

determinant

188

(73)

1 f r2

-l> s- > ^ > • • •

1, Z, 7}, ...

R.

FUJISAWA.

f

f'i

fm~\

^

s

0 >

0

«

z

7^

„m-l

.<? '

.s '

z'^-\

7)^,

D

;?2

.5

.S' '

D

p,m-r\

~ ~2 ~TO + 1

{m- 1) ! , D""-^— , jD™-^— , D"'-^ ^

S .s

D"

„7tt+l

J)m+l^ jr)m+lil 2)'"+'-^

^2

. D^

= 0,

say Ço+Çi:+9,rH...+a,-ir-^+A4r+p.^+...+p,„,,-^-o.

Now

Vc;^^

a

p.

i+1

Hid thence, // heing even,

PiP"' + P^k^"''-+... + P.

Pto+1

?rt+l

— N'
sniin = ( — 1)'" z- -iTfT-,

(74) ; cn?w = (1-;?)" 2

dn nu =(1-F^r" ^,

MULTIPLICATION OF ELLIPl'IC FUNCTIONS.

189

WlU'fC

(75) N':

lis'

— D^ —
'2! s

à ! ,s

m\ s ' (w+J)l ,s '

1

D'" —

(w+l)! s

jyn-i^

C2m-1}! s

0,

(7()) iv;=

II s

— I D'« l —
(?;i-l)! s

m 1 s

(;«— 1) ! .s 7/t ! .s

^ 7)2'» 1 jL

('2w-i)! s

(1-^)'", (1-^)"

(77) iy;:

T'
1 ! s '

2! s

?;t 1 ,9

1 z

{m — 1 ) ! -s ' 7?i 1

1

TTT^^

^Am — 1 ;

(1 - ^zj", ¥ (1 - 'k-zY'-\

— D —
1 ! s '

(78) N',-

1 ! .9

J_i)2A

2! S

?;i ! ,9

(m+D! s

J)m-l_^ * 2)'» ^

)??t— I) ! s ' ';;i ! 6- '

1 z

(2w-l)! .9

190

R. FüJISAWA.

N', N'.,,N'snv,i\ also lie written ns follow«

(79) N'--

' S '

' ' 1! ^ S '

^TO-2 — 1—7)2 _

' 'J ! .9 '

(-1rs -^D'«-^,

I ! ,9 '

'2 ! ,s- '

8 ! .9 '

7)w+l_

w!'^ s ' (w+1)! .9 '

1

D

,. 1 1 !

(w— J)! .s

7;i ! .9

1 Z)-+i —

(w+1

1

(2m- 1)1

TT-

.. 1

<80) 2\r;=3

0, (1-2)'",

-;2^ 0,

S '
1 ! .9 '

(m-D! s

^ ^ ' (w-i)! .9 ' ;;/! 6" ' ('iw-l)!

X)2'«-i_

(81) n;--

0,

(l-A-22)'«, P(l-/.2^)™-i,

0,

.9 '

.9 '

1 ! .9 '

(-l)'"+\

Z)"*~^— 7)'« _

{m— 1) ! s ' 7^1 ! ,9

(7;i— J)! .9

•;;? ! .9

1

i2;»-l)!

D2

-il

§ 1-^-
We ure iiow in ;i position to introduce U„ and <S„ into the niulii-
plication-formukiß. In some cases, we might have introduced ^'„
rather tlian II,, ; hut, for the .sake of uniformity, we ha\^e chosen those

MULTIPLICATION OF ELLIPTIC FUNCTIONS.

191

forms wliicli ;ire ex]!rf;8.siL)le Ijy 11,,. On introducing B,,, it ^^ï\\ be
seen that there comes out ;i certain power of.s- as a factor in the numera-
tor as well as in the denominator, which may lie cancelJed ayainst
each othei', leaving" rational integral functions of -y/z ^ sn // both in
the ntunerator and the denoniiiiator. The final results are:

11 odd, /// = -— ,

(cS'i) i=,ii lilt =(-l)"'y^^

1, ±\--.\-l^z, m-zA-K'zf, ...C2.1-,ï.l-A:2^)"'+'
0, 1, B„ ... B,„

1, Bi,

B„

B.

-^^;;t-l> -^^m»

1 >

i?o

(8;i) en na ^\/l —■<^

1, -±y.\-1^z, !:l.zl-hrzf, ...(-2.^.1 -Â-?)"'+i

0, 1, Bi ... B,„

1, iv'i. Bo, ... -R,„a

■^Art-l) J^tio

B„i+i,

JrCq,,

(«4) dn nil == V 1 — A'-^i

1, -~±A'z.\~z, {-2.Pz.l-zf, ...{-±Ji^:A-zr''

0, 1, Bi, ... B„t

1, il'i, iV.^, ... B„,^y

B,,

B,,,,

B„

B.,

(Ö5)

(küjoni.

1, By, ...B„,

B„ Bo, ...B,„+,

'ri

Ë. FÜJISAVVA.

11 t;\eii, ///

n-\

'2 '

(«(3) bii v/,« = (— l)'"±s

0, i,...Ä.-i
J, iA, ... 24

2^«;-], -fV„(, . . Il 2,11-1

(87) cil ;/,/^

0, 1, -2.^'.1-Ä-.:-, ... (-2..~.1-A2:

(^.\~,A-Jr,r, K-1, B„„

■^Hm-l

(SS) du lut —

a 1, -±k^z.\-z, ... {-±]^-.]-ïr

h 0, 1, ... B.,^_,

(2.1-^.1-A'V', iC-i, i^.,

-^'s,»-]

(8U) tleiiom. =

1, 1, /A, •••2^.-1

2.\—z.]—l -;:, Bi, B», ... ii'„j

I9r}

MUL'lTI'nrCA'ITOX OP ELLIPTFC PLTXCTIOXS,

Part Second.

§ lo.

To avoid ronfn.sion, we shall adopt onœ for all the followiii!
notation :

Fi^lloAvinD- .T:ipnhi ill lii»j lûofn..,-»,. ,,r^ w...;*^^*

E BEAT A.

_2 T^2

Pa ore lî)o, h' ne 10, for -^ read -^ .
„ , line II, for ^ read ^.

194, lines 8 and 11,/^/" ^k' ànnu read — rp dn /m.

t)iii

rho

U\-lr(^'~ '''^'^'

hence

(90) Ç!L+o,,"-'^„j;l+o^,j,..^-^o.

'I" ak du dk

()l)serve that the same ditlerentinl e(juation is satisfied l)v 'h, ^2:iiid H,.

* JacoM's o-<:'saniuielto Werke, Bd. T, pp. 501, 511, and 512.
** Ditto p. 259

t Pitto p. 260,

192

lt. i'ÜJISAWA.

(8(3) Sil mc =1—1 f^-ls

n even, ;// = — p-,

0, l,..-i?,„-i

1, Äi, ... i^„.

0,

1, - ii.,c. 1 - A:-a-, . . . (- 2.^. 1 - A-2~)^

(±i_^.l_A^,y«, ^V i?,«+i, ...i?2. 1

Mur/innjcA'rrox of elliptic fctnctions. 193

Part Second.

§ 18.

To avoid oonfiision, we sliall ndopf once for 'ill the following
notntioM :

X — finir, ç=\/Ji fin)/, ^ = sn-//..

Following Jncoln in his lectnre><, we write*

ß (,,)^i +22l-"lV' g'' cos '^, Uu)= ^2 ^^ ' ^ cos^^^^ ^1^'^",

n = l

»=i

nnd tlien

//7 ^^..w/)

1 T ^///6)

VF^"""^(./)'

Put fo=— loo; (^= ^ , then

l)nt

hence

(JH _ -- d-h u (IK (W
(/(o /i- r//r K dio du '

,,,-r(f)

■ ^ c//^^ r/A- dit dk

0])serve tliat tlio 8:nne ditferentinJ e(|u;ition is s;iti8He(l hy ^^, ^aiind fly

* .Tacohi's ^rosainmelte Werke, Bd. T, pp. 501, 511, and 512.
** Ditto p. -^59.

t Pitto p. 260,

n)4

R. FUJISAWA.

E(|nation (90) may also be written in the form

(91, ^Z^^/rfloH^V

dii'-

\ ai(. / dk da dk

SI mi

1^, ^ +(î?!i^.y+2,,,.l!2sLf „ '^ + 2;,./,- '^ = 0

"^ du- \ du / dk du dk

Siibtractirjo; the former eqna,ti(^n from the latter and I'enhieinii' ,. ,
•^ ^ ' ■ n{u)

Ijy ç^, we (xq\

(92) '^ + f'Ä + 2 S '^^- + Mr- '^ „ I ^^ + llir ''^ = 0.
du- \ da / ( du dk ) (///

dk

N

ow

-%//• sn m

ß,(n

u) /k_
A' y h'

ä{nu) ' y k

cnnu = 77= — r, VÄ- (In;/// =

ä[m()

0{nu) '

mnltiplying the nnmerators and denominators on the ri^lit-linnd side

by 's:!^,

VFsn }iu

wliere

V

(98)

/^(////)^"^-H0)

/^i(;///)/^'''-\0)

7.=

t),{nu)6-'-\0)
ß,{uu)ß"'-\0)

' ä"\u) ' ' •■ ~ fi"\u)

Put //// instead of » in eqnation (91), tlms : —

MULTIPLICATION OP ELLIPTIC FUNCTIONS. j (j^

XoAV

log ß(ni/) = log V + n- loo- fliif) - {),^--})\oa ß{()) .
Siihstitntiiig ill (i'l), we g'et

f//r \ ail / ( r/?^ a/,- ) du

r//,- '( (hl- (Ik )

Au-:nn. ^-^ = Z(i/) , 6 (0) = J- ;

au 'y -

ainoreiitiMtino-, ;nifl oi)servini>- -yy- = — ^r^^ — 7 ^^'^ ohf-nn

(Ik kk

(05) ^-lo^%)^,^.^..jIog^^_^.^„.^^

Siihstitntinf)- tliis in (i)4),

r/?r \ dii / ( r/?/ dk ) dir

dk

or

+ 2nn-k'^'^+ }i\)r- - 1 )/.-2sn^// V= 0.

Iiitrofliiciiio' ^-^-^/k^^^)f :is tlio new indepeiiflent vnriable, and
o]).ser\inu- (9!2),

* .Tacolii's «^esauiuiplte Werko, Btl. T, pp. 1!>8 and 235.

|9(i •>' FU.HSAWA.

/r]-\-rPV fZ-c (TV dV

\(hi / fh- (In- d: dk

Since

er|iiation (97) rnkes the form

+ '-^"'/'''-^ + ''\'''- 1 )--' V-^ 0

This eqnatic^ii is also satisfied by T',, I .,, and 13.

With J'K'oln, we in:iy ])iit /,■ + -^- = a and then equation (Î)S)
heeonies

^ r/ç- r/ç (la

in wliicli forni, tlie p-irtial differential equation was o-iven for tlie first
time hy Jaeohi.

The al)ove demonstration ofJaeobi's partial differential equation
is substantially the same as the one given by lU'iot and lîouquet* in
their Avell-known treatise on elliptic functions, the only difference
being that, in their demonstration, tlie intermediary steps are conducted
bv means of Weierstrass' function .//.

lîciot and lîonqiiet «jive also the following- forms of the pai'tial
differential e(juati()n : —

* JjOO. cit. p. 529.

MULTIPLICATION OF liLLIP'l'iC FUNCITOMd.

197

/, 1-27,= „ ,\<r-v , , ., ,/i--2i-' , ,, ,\<n'

hen

wiierc

■^/ - V T

cii u and

1

Va

^ dn ^^.

§ II-

\\\- si Uli I liaxe, by .suitabJy (Icteriiiining the constaiilsf,

(10-2)

\ V = D[.r),

A{0) - n,
B{0)= 1,

AO) = 1,

' 7j = Va- .<Vi - '^-Vi - /^j^^ A{x-), Aio) ^ II,

(108)

F2 = V ^ ^''''"'■''

ü(0)= 1,

CIO}- 1,
AO)- 1,

11 odd,

II, even,

wliere J, 7», ( ', V ;ire nitioinil inte,ii;r:il I'lind ions ol .r.

lt)3 R. FÜJISAWA.

By division, we obtain

n odd, n even,

xA (x-) x\/i — x-'s/ 1 — A '-x'-A {X-)
sn nii = — ï^ — s—, Sil nu =

(104) cil nu =

^X-x^B^x^)

D{x')

Vi-ÂV-^(7(x-)

du na = j^^^s , du na =

D{x^)

B{.r)
D{x')

G{x')

A.'-') ' D^x'-)'

AVlieii n is odd, ./, />, C, i> are ail of the degree n^—\ iu x. and
when /< is even, A is of the degree /r — 4 wliile ./, />, C' are all of the
degree /r in x. Moreover, A, B, C, 1) are rational integral functions
of Ä;^ whose coefficients are integral numbers. The coefficients of the
highest power oï x in A, B, C, D are respectively

or

(-1)2 A; 2 , Ä; 2 , h'^ , (-1)2m A- 2 ,

(-1) ^ n k ^ , A-2 , A-- , (~1)'^ A--",

according as //, is odd or exeii.
Write

A - i'A,„,x'"^ ^ :i'/i„„(i-.-V' =- :i'.i„„(i-AVr,

(1U5)

^ = ^' C,„,r^"' ^- -■ C.„X1-.^'^)"' -^ ^' C,„x\-Pxr,

then, we liave the well-known relations*

* Compare the work of Briofc et Bouquet already referred to, or Baebr, Sur h's funiin/cs
pour la multiplication des fonctiona elliptiques de hi pn'ini'^Te exjièce, Uruuert's Archiv (h'r
Mathematik, Bd. XXXVI, pp. 125-176.

MULTIPLICATION OF ELLIPTIC FUNCTIONS. ^99

D,„ik) = Ä""A.(y ), DIM = ^-.«(y )•

Observe that A.2,„, i>2,„, C'o,„, 1A,,„, Jl,,,, 1>2„„ C'l„„ 1>2,„ are integral functions
of ¥ of the degree at niost equal to vi.

§ 15.

Consider iirät the case where n is an odd number, and put
n2__ 1 = 4^^ sq.

Introducing the new variable ç = ^/W x in

Sn 7/7/ =

we get

(107) -y/k snna -

where «am? ^^2m '^I'e rational integral functions of « = ^ • + — .

We may suppose A and i> to be arranged according to the
powers of a, thus : —

A ^ lA„,x"'^ - ycuj''" ^ ^E,jjy\
(108)

200 ^^- FUJIKAWA.

From the well known relations

(109) A{x, k) = {- l)'^'D(Jy, äVä^ B{x, ;,) = (- 1)V^^J^, Ak'^x'p,

which may also be written in the form

(llU) A{a, ç)==(- \)^'d(^x, jY', D{a, ç) = (- iy^A(^a, 4^ç^

we see that A and 1) are of the same degree in a and that, moreo\ er,

(111) E„,(ç) = ( - \)^H„(iy'^, H„,{ç) - ( - l)''^V«(y)>-^^.

AVhen the multiplicatcrr // is required to be put in evidence, we
shall write A\_n~\ and lJ\_n'\. Xoav the terms containing the highest
power of a in J[3], J[o], J[7] are -2^-^«, 'A-^V/, -2''^'^y', and those
in D[3], D[5], l)[7] are '2\-V., -i^çi«./, •Ji^i.s^.«. j>^ ,,^^.^11^ ^f the relation

(112) A[n + 2]A[n-2] = iyp]A%N]-(\--aç'+^')A\2]D-lnl

which is easily deducible from the addition-equation, we conclude by
applying mathematical induction that, generally, the terms involving
the highest power of a in A^nl, i)[//] are

(118) E,/j:' = {-\) ^ 2^ç "a", H,//.'J = 2^ç^"f/',

where /„ + /i„ = «^— 1.

To find À,„ /A„ we deduce from (11^^)

/„+2+'<„-2 = 2//,,+ 2 - 2>i2_'2/„,
which may be written

MULTIPLICATION OP ELLIPTIC FUXCTIOXS. 201

or, ]uitting for a moment Wn — ir + ^j — '/'ii >

ÇV2+%''n+^ . = 0.

Observing </'i=-^, <r's=—'ö'' we find

ç''„= coef. of ,«" in -^ ^y:j-^ "" ^~^^ ^ "2

Hence,

9?2 »til ,, ^,2 »2zl 71

(114) ;,-| + (_l)2|._l, ^,,^_|-_(_1)2|.,

that is.

or

1 1

'^« = VC" + 1)-1 ' /A, = ^"("--l).
according as -—^ — is odd or even.

This premised, we now h^ubstitnte D=^^H,/£'' in (99), whirli may
1)e written in the form

(i.^'-^r=:r — dr — 1 )«ç -j^ + 2ura-—- = ( 1 + ç" -y^— 'J(7r — 1 )?'-î^

+ «\^r-l),-^F+8n^|^,
and obtnin

(115) ç^-^^0f—\)^^^+2n'qH,=^O,

(116) ç^ '^^^ - Or- 1 ),-'^ + 2u\q - l)7f,_i=(l + ç^)^^ - 2Cn^- 1 )ç^'^'

+7?,v-i;ç^H ,

202 R- FUJISAWA.

and, generally,

(117) ç^^^^(^n'-\)-^Ê^ + %,^niH„ = U„, , m = q-2, q-3, ... 0 ,

where U„, = (1 + ^-^1^' - '2(n— ^)^"-^^ + »'0^' " D'^'^.u+i

It will be seen that equation (115) is satisfied by ç*^"; indeed
we might have determined Un by means of tliis equation, for if

the operator g-^j- be denoted by ^, then the diiferential equation may

be written

the general solution of which is

where C, C" arearbitrnry constants ; and, as H,j can contain only even
powers off, C' = 0, or 6"' = 0, according as —^r— is odd or even.

Let us now investigate e([uation (iK))« ^or this purpose, it
will be convenient to distinguish two cases according as — -^— is even

or odd. When —2— is even, ^„ = —;?(;?-!) and iJ^ = 2pf^"<""i>, further

rJ^H rlTT

(^ + n^ - %n^ - 1 )^~' + n\n'' - l)^m,^

=2^--2|(«,_2)(»,- l>i(».+ l)fä"("-^> 2+(/, -\)n{n + 1)0? + '2),-"("-i>+2[ ,
so that equation (IKi) now assumes the form

= 2^-2|(;^--2)(n-l);^(H+ L,-'"^"--i)-2+(/i- l);?,(^i + l)('y^ + 2;c^''(''-"+2[ .

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 903

Xow, q being equal to

The complementury function is thus

but, as ^fi(ii — ^) is odd and H^_^ contains only even powers of ç, C" = 0.
Ao-ain, a jmrticular integral is -2^'(?i-l>?.$^"<"-i>-2-2^-^;?(» + ])ç^"("-i^+-',
which, together with the complenienturv function just found, gives

where P^ is an as-yet undetermined function of n.
When —^ is odd, we find likewise

r>y virtue of (HI),

E^_,= r, e^"('^ ;'.)-i_2p-=';,(,i + l)f^"("-i)-^_'2"-«^;?,- l)»f^"(''+i>+i, ^i^ even,

^^_i=2"-\'n-l)».?*"<"-i^-='+2^-^';K'«+l)?-"^"~^^+^-^i çi"'"+3>-\ '-^ odd.
We may put the multiplicator n in evidence by writing

and then e(ju;ition (112) may be written

whence, equating the coefficients of the second highest power of a,

»1+2 n—2 )i— 2 »1+2 r~ n —12 p « -12

»I n

204

R. FUJISAWA.

If — ^^— lie odd, so that -^^ is even, then the term containing

the lowest power of ^

»1+2 Ji-2 „, n-2 9 „IP

in R ^, , is 2^'«+2r,r-" + ^

'7)1+2 '/«-2-1

Equating the coefficients of the lowest power of ^ (that is ç"'-"-) on
the two sides of equation (119), we get

jt+2 9,, ., „ (n-l)(ra+5) ai — ^

r, ^ 2^^'""^'"-^=2 ^ , ^ odd,
or, writing // instead of // + 2,

» (»-3)()t + 3) g 9/« 1\ 7? — 1

/^ _2 — 4 _2?"~'^ = 2^ -^ -

even.

Again, when — ^ is even, so that — ^=^ is odd, the term con-
taining the highest power of ç

H +2 H-2 n— 2 ., , „

in i?„ E , is 2?'"+2ril"- + "--,

'ln + 2 '/n-2-1 ^ '

»1—2 »1+2 11 + 2 .,

-4(l+ç^)[^A 1' „ _22^,,. + ,W-. + 4^
</n '/»I ~ i

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 205

Eqiintiiig- the coefficieijts of the highest power of c (that is ç"'+''+c^ on
the two sides of equation (Hi)), we obtain

ri = '2'^'"~^^"-^=2~^~, '-^ odd,
or, writing n instead of /^ + !2,

n (n-3)(n+3) ., . _ . ^ \

Thus we find, whether is odd or even,

Li

(120) /; = 2^'"~^ = 2"^('^"~^^

and thence, when — -^— is even,

(121) H^_i= -2''-^(;i- l)7ic^"^'^-i>-2-2^-^».(// + l)ç-^"("-i^+2 + 2^-2f^'*("^'«,

(122) £ _i= 2^-2ç^"^"-^'--^-2^-^;H/«+l)ç*'""+i'-^-2^'-^(//-l)/?ç^"("-^2)+i^

and, when —5— is odd,

(124) ^,^_i = 2^-='(;i- l)«ç5"(«-i)-3+2''-^;i0i+ 1)^-"^""^^"^-'^*"^^^^"^"+^^^^

Consider next -/f^_2) whereby we suppose —3— to be even. H^_2
satisfies diiferentiid equation (117)

where

ü,,_2=-2^^(/i-l)H(>r-;i-4)(/^2_vi.-6)f^"(''-l^-*-2^-^/A^M"-2X/^-y)r^^^^^

- 2^^''';?<?/^ + 1 )(nH /i^4)(;?2+ y;-(3)f^""^-i)+*+ 2^-*yA(;i + 3)(7t'+ 8;f^2)ç^"<"+=*)-2

+ 2''-*/i('i-3)(M-- 3yi - 2)ç'"'"+=*^+2.

206 ^- FlJJiSAWA.

The complementary function is

aiicl, a>s H^-2 can not contain irrational powers of c, we must hîive
C'=0, C"=0. Hence, —ç— being even,

(125) H^-s^ 2^\;i- l)7j(H--?i-6)e*»'"-^^-^ + 2^^^7r-2)0i2-9)ç^""'"^^

+ 2^~hiin + 1 )(;r + ?i - G)e^"("-^'+^- 2'^-^vK« + 8)ç*"'"+^>-2

-2^^M?i-8)c^"'"+^'+2

and thence,

(126) E,j_2 = - 2^-^;i(7i - 8)ç="("-^'-=*- 2^^?i(m + 8)e^''("-3)+i

+ 2''-7yi(« + l)(n2 + n- 6)ç^"'"+"-^ + 2'"~%n^- 2){n''- 9)ç^«("+i'-i
+ 2*-7(;i - 1 )n{ii^ - ;i - Q)$Mn+i)+^^

When is odd, we find likewise (or, more sim])ly, by writing — //

in place of n in (125) and (12()),)

(127) H,_ = -2''-')i{n- ;il)ç-"("-3'--2_2î^5„^^^+ 3^^5«(«-3)+2

+ 2^^7?.(7J + 1 )(7r + », - 6)çi«("+^>-^ + ^2^-'^^uP-2){n-- 9)ç-^"f"+i'
+ 2^^??(« - 1 ){n^—n — 6)ç^"("+i^+^

(128) E^_2 = - 2P-^7i(M - 1 ){n^- n - 6)ç*"<«-i^5 _ 2^- 6^,^2 _ 2)^,^2 _ (j^ -^«(»-d-i

- 2^-^w(n - l){n^+n - 6)e^"<"-^'+H 2*-^n(n + 8)c*"<"+^^^

+ 2''-^7î.(??.-3)l^*"^"^^'+\

Next, consider i^y s which satisfies the differential equation
where

MULTIPLICATION OF ELLIPTIC FUNCTIONS. -JOT

When — ;t— is t'Aen,

Ug_^ = '2"~ \ii — \)n()i - 'S){n + '2)( /r - //, - «)( ir -n-l 0)ç'"^" "^^'^

+ '2"-'\ii, — l)n{n - S^Sn + '2){n' + '2;r'- 21 /r - 42;; + G( )lç'"("-i>-2
+ 2f-»»(;?^ + \)(n + 8X3» - 2)(;/,*- 2;/.- - 2 1 ;/- + 42yt + GO)ç^'''"-i'+-
+ 2P"'-'//.(;? + 1 )[it + H){)i ~ 'Dili- + II - H]{ii- + It - l0)ç^"("-i>+6

- 2''-'n{n + 3)iii + 4:)(:ii - I )(;;2 + 8;i - G)|^"( "+«>-*
-2'^^'';^^(;;''-l9;r + 98)ç^"^"+='^

-2-''-''yt(;j- oK/? -4K;? + 1 ){ir- -yii - 0)ç'"(»-3)+4.

A particular integral may easily be foiiiicl. I'lie cum[)leiiieiitary
f unction is /gÇ-"'""^', where f^ is a function of//. Hence we get, — ^j—
being even,

am H,_,= f;ç^"<"-«

_ 2P-i^^-s-in{ii - 1 )(if" - u - 8)(;r - n - lUlç '"'"-^'^ ^
-2^i'V<M-8)(;?'' + 2;i^-21;r-42y/ + (U))ç^"f"-i)--

- 2»'-i"yM;A + 8)(»*- 2;/.^- 2br + 42;f + G0)ç''"<"^^*+2
-2''-^"3-^;K« + !)(''-+ '^-8K'^' + 'i-^0)ç^"^"-i>+«
+ 2^-«;?(yi + 3)(«2 + 3;^ - 6)ç^"("+^^-*

+ 2»-\n'- 19»H 98)?^"'"+^^

+ 2^-«w(w- 3)(7?-- 8?^ - 6)ç-^"("+^^+\

(180) ^^-^3= 2"-'VVi-3Hy^'-87/.-6)l^^"<"-^-''-'
+ 2''-\y/*- l<.)y?- + 98)p("-''-^
+ 2'^-"y?V/ + 8)(/r + 3n - 6)|^*"('^-3)+3
_ 2P-^'K-^-ht{it + \)(n-+ii-8)(ii- + ii- 10)ç-'«<"+i)-7

- 2P-'^''n{ii + 'à)[ii^-2n'''-2\ii- + 4-211 + OU)ç'"^"+i'-^

- 2^--i";/(yt- 8)(M* + 2yr^- 2br- 42/i + 60)ç''""'+i^-^
-2"-^"8-^;K»- l){ii--n-3)[ir-ii— lO)ç^"('^-'^>+'o

When —:t— is odd, we get, l)_v writing — // instead of// in (1-Î'),

208 ^- FUJISAWA.

(131) iî,-3= 2''-VH-3)('i'-3'i-6)e^"(''-«'-*

+ 2^«».0i + 3)(n2 + 3« - 6)c*"<"-=''+''

_ 2P-103-' ;i(?i + 1 )(»- + »' - 8)(;r + ^/ - 1 0)e^"*"+i^

- 2''-^'n(,n + 3)(?i*- 2;i^ + 2l;r + 42;/ + G0)çirt(n+i)-2

- 2^-i";i(« - 3)(n*+ 2>i^- 21»-- 42;/. + GO)^'"^""^^^^

_ 2^-103 -l;,(;i _ l)(;/2_ Vi _ 8)(;t2- »t - 1 0)ç ^"<"+l^+8

and thence, by virtue of (111),

(132) ^,_3=-r3^*''^'-^'>-^

+ 2P-^^n{)i - 3)(u* + 2;/^ - 21;r - 42;i + 60)e^"<"-i'-»
+ 2^^%{n + 3)(«* - 'M + 21 ;r + 42;* + 60)ç^"<"-^'+i
+ 2^-i''3-ifi(« + 1 )(«- + ;i - 8)(;i2 + n - lO)|^^^"("-i^+5
-2^^;<?i + 3)(;r+ 3//-6)e^""'+=""'
-2^'-8^;i*-l9;iH 98)ç*"("+="-^
- 2^-^;K;i - 3){)i^ - 3;i - 6)ç^"<'^+=*>+^

n'— 9

Equating the coefficients of a ^ on the two sides of (11<*^), we
obtain

n+2 n— 2 n+2 n— 2 » + 2 «—2 ?i+2 »i— 2

^Vi+2^'7n-2 - 3 + -^^„+2 - l^'/«-2 - :i + ^q„+2-2^<ln-2 - 1 + ^'/m+2 - 3^'y«-2

/ >i »I n H \

-4(l + fo(2HÄ-2 + Ä,,,_.H,,._,) + Hf'H,,,,H,,.-i-

When —-y- is even, the lowest power of ^ in the above ecjuation is
^,i>-3rt-2^ Equating the coefficients of this power of ? on the two sides,
we get"r3 2''"-2= 2^(^'"-^>, whence

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 209

»+2 iJ^±^_, n-l

' 3 = ^ * , — T— even

or, writing n in place of // + 2,

n»-l

n — 1

/; = 2^-" = 2^^^-«), ^ odd.

n

In like manner, we may shew that /^ has the same value in the ease
where \^ is even. Thus, whether // he odd or even,

(133) r^ = 22(' -^J.

In determining Hq^ix, every time 8/^+1 is of the form /', where r

n

denotes an integer, there comes in the term r^ç-"^"*'\ Indeed all the
terms of H imd E may readily he expressed in terms of /'. On the
other hand, E^ may be determined hy means of (11<S), as has been
actually done in the cases /z = l, o ; but when p, is a large number,
this becomes very laborious. Now in the series

r r r r r r t'

r, = 2^'^ /; = 2'^''-^\ ri =^ 22(''-='>,

so that most probably /;, = Q^n^i-i^). ]^^^^ j i^^^^g j-,q^ ^j-j^,g f.^^^ succeeded
in proving this generally.

For a given particular value of //, the constants E may be
determined in a diiferent manner. For example, take the ca^e n = d.

Sin 9x = œ{9-l20x^ + 4:32x*-51Qx^+2rDQx^).
Again E,^_e contains tlie term /;ç*"<" "-i j^^d iî,,_i„ the term Al""^"'^'.
If we put A:=0, a becomes infinite but in the same manner as -j- .
Thus sin 9a; contains the term f\x^^ whence follows /g = 256.
Obviously /w=l.

910 ^- FUJI SA WA.

The elliptic fnnction.s of vii for ;/ = 2, 3. 4, 5, H. 7. 8 have been
calculated by Jiaelir and others* by the primitive method of succes-
sively applying the îiddition-eqnati<jn, Ihning found the values

9

of r, sn 9?/ may be calculated by the above method without much
difficulty.

§ 16-

As regards sn nv, n being odd, the analysis contained in the
preceding section, whereby the variables are taken to be ^W sn ?/ and a,
leaves nothing to be desired ; yet for the other functions, this is not
the case, and it is better to have as the variables sn ii and h. For the
sake of uniformity, tlierefore, we shall once more investigate Hnnii, but
this time consider it as a function of sn u and /.-.

Changing the variable from f to x, equation (1)(S) takes the form

(I.I- ax

Cl/i

and the equation is satisfied by the numerators and denominator of

V/'" sn nil, Vp en //?/, -yp= dn

nv.

The numerator of sn ////, that is, xA(x-), satisfies the differential
equation

* Se(; the paper of Baelu* tih'oady referred to, and Proooedinsj-s of the Royal Society of
London, Vol. XXXIII (1882) pp. 480-489.

MUL'J'IPLICATIOX OF ELLIPTIC FUNCTIONS. ^ll

dV

a85) |l-a + AV- + ZV['^+{U^;?2._i)^,2_i),_2^,,2_i^y;,2.

y3L

' doc

(il:

We m:iv sujjpose tlie iiiimerator of sii //// to be aiTanged nccordinijf
'to the powers of /r nnd write

(136) .r^Cr^) = 2] Pojr'".

Then, denotino- the o])er:itor x^r- by 'V, we hnve

dx

d'P.

(137) ^ 4- {»\^m + 1) - /y^jPa. = ( i^m - 3)»^ - -i/^^/y + â')P,,„^^

For 7^? = (). we have

and, if we })iit

then

r/2p
a./'-

Po = 2/3.-r^^+\

^9,(2r+l)2r = -^?^_,o,2_^2r-l)2)

And, since ,3',, = «-, /I. = (— 1)'^ . , , -; thu.s we

obtfiin the well-known series

(138) P„ = 2 ,_ i,iH»!=iM-"=i?r^I)l ,.« .

For 1]}:= 1, we have

Since the lowest powcn- of .r in Pg i^^* 3, we write

212

R. FU.TISAWA.

(139)

then

P2 = llrr^''^'>

2r&r+\)rr+{rw'-{2r'- 1) ) Yr^,

= {n^ - 2n\2r - 1) + (2/- - l)-)ßr-i - (»' - 2'- + 3)^»- - 2r + 2)ß,_^
We find

Tr

n{n^—}) _ ??0''^— l)o;t

8!

, r2= ^ ; . ^2(2^1^-8), r3= -

7!

3(37i2-5),

Generally,

2r(2'>- + 1 )r, + {^)f- - (2r - 1 ?)rr-i

\4>-'-\()r+r))n}

^ ^ (2r— 1)!

.+(2r-3)(2r-l) )

/-;•(/v^2-(2;■-l) ) (;i2_(2>--3)2)

^ _ ,_,Mn^-^l),_X^^
(27'— J) !

I ((r-l)»2-(2;--3))

= ^_l),.^K.^--l)...0.--(2>--5mn^-(2.-3)^^^

))

or.

+(5,.^-(2.-i)^x-ir-"<"''''^,!!';;|''-'"(.--i)

((»•-l)»'-(2,--3)),

= _(5«»-(2,-i)', Qv-.-(-ir-"^"'-'^„':^-f'-"'''(.-i)

((,-l)„2_C2,— 3))1;

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 213

whence follows

(140) ., = (-ir"'"'-^"^-;';,/;f-'^'-^'%-(».-(2>-i)).

For ?H = 2, we h;ive

'^ + i^M^-ö^-)F, = {bn^-'ln'ä-\-Ö^)P,-x\n'-ö){n'-&-l)P^.

Since the lowest power of a; in l\ is 5, we write

(141) P, = 2«r^""^';

r=2

then, 2K2>-+ l)o, + (9H2-('2r-l)Vr-i

= ( (2r-l)2-7i2(4;--7) ) }'r-i-0^'-2>-+ 3)(7i2-2/- + 2) jvo ,

whence we find

, _ n{nl-\){n?-^)

».(/r — l)(n- — 9) 2 KN

(142) ^ '•

8, = M>^'-\X|^'--^)2(247y<«-ta25n^-l3757/^H 28625).

Thus the general law is not ohvions as in the case of r, imd it seems
to be impracticable to proceed further in this way.
Reverting to the formulae

since, by virtue of (109),

214 ^- FUJISAWA.

we need onlv detennine either A or D. Let U8 take- D and apply

n—l

(134). We obtain V^^-l, A=0, 1)4^,=(-1) 2 n¥", and, generally,

(143) (2m + lXi^;» + 2)A.«+2+4w|(n^-7;0^''-m|A«. + '2^i^Atl-Ä:^)'^

When VI ^p,

(1 44) D,„, = D,», 0 ( 1 + A-'" ) + A™, 2 (A-- + A""^-) + ■ • ■ + ^2«. 2. (A'" + /.''"--n + . . . ,

the last term heing A,».™ A'"' or A,„,,»-i (A"'"' + A™^') Mccording as /» is
even or odd. Substituting this in (113), we obtain

{'2m + 1 ){2m + 2]{ D,,,^,, ,{] + Ä^-+^) + !)„„,,, , (A- + A'^-^) + . . .

+ Am4 2,2r(A'' + A-"'--^+)+...}

+ 4//t (;r-;;^) |A,«,o (A'+ A'™'') + A».,2(A' + A:-"')+ ...

+ A., 2r( A "■+'-+ /.-'"--'• + ')+...}

- 4;;r I A,,,, „ ( 1 + Ä-'") + D,„, . (A - + A -'"-2) + . . + D„„, 0,. ( A -' + A-'" -"1 + , . }
+ 2ir |I)2„,,,(*+2;;iA;-''0 + A„,,2(2A- + ('im-'2)A2''--)+ ..

+ A», .r (2rA^'- + (2w--2y)A-'«- -"•) + . . . [

- 2n- { A-«, Ü i* + •2;;iA-"'--) + D,„^ , (2 A* + (2;;i - M"") +...

+ A,«,2.i^''A''-'-+(2^«-2r)A2--2'+2) + ...}
+ (n' - 2 ;;i. + 1 )(^r - 2m + 2) ] Do„,^., „ ( A" + A-'" ) + D,„_,, ., ( A '^ + A -™ -^ ) + . . .

+ Am-2, 2. lA-'-^ -+ A-'"-2'-) + . . . }
= 0.

Hence

{2m + 1 )[2m + 2)D.,„, ^0.0= 4y;rD2m, 0 , -^^2,.. n = < '.

(2w+1X2;m + 2A«.2,2 = -4(yr-y/rA,»,2

(2w + l)(27;i + 2) A,«+2, 4 = - 4 1 {fn - 1 )n- - ui' \ A«. 2 - 4i2;/- - //r ) Am, 4 >

- [n''-2m + l)(;/,2- 2;« + 2)A,.-2. 2 .
( 2y/i + 1 X2m + 2)D2,„+o. e = - 4 ^ {m - 2)n- - m^ \ A,„, 4 — 4(3;/- - ^/r^i).,,,,, e ,

-{}v'-2m+ \){n'-2m^2)D.„, .,,, ,

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 215

and, generally,

(145) (2w+l)(2wi + 2)Z)2,„+2,2r = -M{m-r+l)ir-m'\D,„,^,,_,

- 4(;vr - vi')D,„^ ,, - («^ - 2m + 1 ){n' - 2m + 2)D„„_,^ ,^-2 ,

the la«t coefïicient being given by

(2w+l)(2m + 2)Z)2,„+2.m+i = -8|^^^^i^-i';r|A..,m-i

or

(2;;i+lX2w+2)D2„+2,^ = -2\{m + 2W-2m'\D,„,^,„_,

-^{^n'-m^D„,^„,-{n'-2m+l)i)i'-2m+2)D,^

accordino- as m is odd or even.
By means of (145), we find

(146)

2>n-'2, w-2 >

6!

7^V-l)(7^'-4)
8!

A = -''^''' tl ^^{^Oi'-'àW + k'j + ain'-^m^,

^^_ 7^;^^ 1)(M^^^4)(^/^ '')).^2{4{n'-l6W + k')+ l5(M^-4)(^^ + A«) [ ,
Dj2= _»'(^''^^X»^-^)(»'-9)3,g4(,„2_i(3)(,,2_25)(^2^^.io)

+ .S(?i2- 16X47»2_ 185)(Ä;H Ä;«)+ I5(45?i*-5G9M2+ 1544)Ä''[ ,
Dj^= »'0^'-n^^'-4)(^'''-^)32|ö4(M^-l6)(7^^-25)(;^^-36)(A.-HA:^^)

+ 1(3(^^2- 16)(34yi^-982;r + 3300)(Ä,'»+ A-^")
+ (1549;^'-43925?iH 357196m-- 8l5040)(Ä;''+A-8)}.

216 R- FUJISAWA.

Again equation (143) may be written in the form

(147) ('2wi+ 2)(2wi + 3)Ä;2D,p_2„,_2+ '2;i2A-(l - k'f^^^

■\-{n'-2m){u'-'^m+l)D,,_,^^, = 0 .
Now, m being less than p,

(14o) -i^ép-Sni = -^4p-2ot, 2p— Swi ("- "T "^ ) "T" -^4p-2m, 2p— 2?»+2 ("' ^ '"' + «''"■'^ )+...

_L n ( l.23J—2m+2r I I.2i>— 2r\ r

T^ -^4p-2m, 2î)-2»i+2r [f^ T^ "• ,> T^ • • • >

the last term being

-^4J'— 2;ft, 2p-în ^ 01' •^4/)-2m,2;)— jn— 1 ("' "T k " ),

according as m. is even or odd.

From (147) and (148), we obtain

(2?n + 2)(2?;i+8)D4^._2„j_2,2p-2m-2 + |''"-('^''«+ l)'}A?>-2m,2p-2.« = 0 ,

(2;;i + 2)(27;i + 8)D^,_2^_2, 2;,-2m + I ön^ - (2;;^ + 1)^ [ D^p^s,«, 2p-2,«+2
+ {(4w+ l);A2_(2m + l)-[D4p_2m,2p-2,»

+ (;i2-2w)(;i2_0;;^_J_ljX>^^2«i + 2,2i,-2,»r2 = 0,
{1m + 2){2m + 3)D,,^2,n-2, 2,-2.+2 + I ^hl'-{:2m +\f\ D,,_2,«, 2;,-2,« + 4

+ {{-^111—3)1^ — {2m. + If [ 1)4^.-2™, 2;>-2'«+2

4-(»2-2w)(?i2_2^;^+l)2)4p_^,„+2,2?>-2»,+4 = 0,

generally,

(149) (2?^t+2)(2;;i+3)Z)4p_2,,_2,2p_2,„+2r-2

+ I (4r +\)}v'-{2m+lf\ D,^^^^ 2j>-2m+2r

+ { (4/M - 4/- + 5)^2 _ ('2wi + 1 )2 [ 19,^,^2,,^_ 2;^2,«+2.-2

+ (n2-2w)(7t2_'2m+l)Z)4p_2„,+2,2?J-2m + 2r = 0,

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 217

and, lastly,

{Im + 2)(27?i + 3)D4;^2,n-2, 2p-m-2 + (2WI + l)(n2— 2wi - 1)-Di„_2m, 2?>-m
+ {(2m+ 5)7^2- (2?;i+ l)n ^.p-2,«, 2p-,H-2

+ («2— 2w)(7?2-2?» + l)D4„_2m+2,2;.-«, == 0 ,

or

{%n + 2)(2wi + 3)D,,_2„^2, 2;.-«-i + 2 1 (27;^. + ^)n^ - ßm +}y-} D,^,^^ ,,^^_,

+ (?i2— 2??i)(7l2— 27?i+l)D4p_2,„+2,2j^^ + l = 0 ,

according as m is even or odd.

From the above equations, we find

(150)

A^i2 = (-1) ' 13 ; ^ ^Ä;^^^^M(^^— 25)(».^-49)(;^^-81)(7?.^-121)(1 + A:^^)

+ 6(n2_25)(H2_49)(7?.2-81)(6?r-ll)(A=^+P')

- 3(l927';i«- 330687^''- 962«*

+ 1033308»2_ 1819125XÄ'' + A*^)

- 4(3046??« - 38037;/« + 327997?.*

+ 7086477^2- 1299375)Ä«} ,

■^4p— 10

_(_1) 2 /^^^^ ^'11^ — J_A"2p-io|(,^2_25)(,;2_49)(^j2_8i)(i^7,io)

+ 5(7i2_25X?i2-49)(57?2-9)(A-+ A'«)
- 2(24771«+ 325?i*- 137577?H 23625)(A'' + /-") } ,

A^8 = (-1) ' ^ ,j ; ^F^-M(»— ^)(v^^-25X^^^-49)(l + A'«)

+ 4(7i2- 9X7i2_ 25X47?2- 7)(F + A")

-6(7i''+857i*-67l7i2+945)Ä*},

218 K- FUJISAWA.

D,^, = ( - 1) V ''('''- ^ ) k'^^^{{n'- 9)(1 4- k') + 2{2u'- 3)¥ \ ,

D^-, = -(-l)V^K^^^^ ^^F^-\l + F) ,

D,p = (-1)"2 7^ A;2^.

The first five coefficients were given by Jacobi himself. Tlie first
and the last six coefficients of all the four functions have been found
by Baehr by a different method.

§ 17.

Let us now consider the rational integral functions of .t'\ B, C
which enter into the numerators of en nn and dn nit, n beino- odd.
From the well-known relations

(151) B{x,k) = c(J^,k^};^^x\ C{x,k) = B{j^,l>jk

(152) B (kx, -^ = G {X, k), C (kx, -j) = B {x, k),
we deduce

(153) B (kx, j) = B (-^ , k\ k'" x'",

2?) rpip

whence

(154) B,,_,„,{k) = B,^ (j^ ¥^, B,Jk) = i?,,-.2.(y) F^

MULTIPLICATION OF ELLIPTIC FUNCTIONS. ^19

Hence, if we put

IK = 2p

then

so that we need only determine B^,,, and this only for the initial
values of?», in view of (154).

Now (134) may easily be modified in such a manner that the
resulting equation is satisfied by B. We find

(155) |l-(H-A-V+A:V}-^+ I [{2n'-l)k'-S]x~2{n'-2)k^x^\'^

whence follows

(156) (2wi+l)(2wi + 2)02^+2+1 [7r-(2m + l)2J +im{n'-vi)Ji'}B,^

dB,

elk

+ 2?^2A-(l-Ä-2)-^ + (?i2_2w)(n2-2m+l)Ä-2i5,,„_2 = 0

Further we find, for m^p,

(2w+l)(2w + 2)ß2.+2,o +{'^'-(2w+l)2]ß2„,,o - 0,

■^2m,0 = 0 , ?i— 1 <C 27)1 ^ 2,p,

(2wi + l)(277i+2)J52„,+2.2 + {5n'-{2vi+\y-}B,„,^,

+ ém{7i^-m)B,^^, +(??2-2m)(^/2-2w+l)ß2m-2,o = 0 ,
(2m + l)(2w + 2)52^+2, 4 + [9n2-(2w+ \)'}B,„^^,

+ 4[(w-l)n^-m2]S2.,2 +(n^-2m)(«2_2m + l)B2,„-2,2 = 0,
generally.

220 ^- FUJISAWA.

(175) (2m+l)(2m+2)B,,+2.2r + {{4r + l)n'—{2m+lf}B,^^,,

+ 4 [(w -?•+ 1 :«--?».-] B2,„,5,,_2

+ in- — 2m)y - 2m + l)-B2m-2, 2r-2 = 0 ,
and, lastly,

B2mr2m = 0, 7ÏI ^ p .

By means of the above equations, we find

(158)
•ßo = 1> B^ = —

-^10

2! '

2 "I

B, = - '-^^[(«2_9)(7?2_25) + 6mV-9)F+8??V-4)7.*],

B, = '-^^ [(7?.2-9X7i2-25)(;r-49) + 127r(7i2-9)(7i2-25)^2

+ 471 V - 4)(1 5n2 - 107)^,-^ 32w2(n2 - 4)(7i2 - 9)Ä-''] ,
(7?.— l^X7r-9) ^„2_25)(^^2_49)(^^2_^l)_,_207i\;i2_25)(,;i2_49)^2

+ 127/2(7i2_4X297i2-329)Ä-H 327iV-4)(14??2_89)Z«

+ 1287^2(7^2-4)(7^2-lß)A■«],

^^^ — ^1^ — ^'[(^^2-25X';i'-49)(7i2-8l)(7?,-'-121)

+ 307iV-25XV-49X?^'- 81)Ä-

+ 47?2(5937i«- 1708271*+ 1795 177*2 -482708)Ä'^

+ 87iV-4X5757i*-1011l7*2+44276)Ä«

+ 1927i2(7i2-4X«'- 16Xl5?i2_ 89)Ä;^

+ 512712(7*2- 4)(7*2- 16X^2-25)^:1°].

B,, =

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 221

§ 1«.

Consider next the ca«e where n i.s even. To begin with, take
sn nil. Here D is exactly of the same form as in the case where )i is
odd, so that we may restrict ourselves to the consideration of A alone.

Now A is of the degree y^^ — 4 = 4^ say, and

(159) A {X, k) - (- 1)'^' A {-^, k\ k-^ x'",
and, therefore,

(160) A^^,„, = (- 1)"^ A,,, k'^"\

m - 22)

In consequence of (104) and (134), A = ^ A,„^x^"' satisfies the

m = U

differential equation

rPA flA

(161) |^-(l + Ä;V+^'-^'}-Tl+ 1'-^+ [(2n2-5)^•2-5]^2-•2(«2_4)^V^-p

dx^ " dx

dA
dk

dA
+ 2n^k{l - k'')x-^ + («2-4) {(1 + k^)x + {)v'-S)k''x^]A = 0 ,

whence follows,

(1()2) ('2/;t + '2)('27yi + 8)^i2,„+2+ {n^-4:{ni+l)-+ [(4;/i+ l)n--4.{iii+ \f\ k-}A.„,
+ 2n^k{l-k^)^^ + {n'-2m-'2){n'-'2m-l)k^A^,,_^ = 0 .

By virtue of (1()0), we need only determine the first half of the
coefficients A^^. Again, A.^m is of the form

A,„. = A,,,a^ + J^n + A,,,,,A^'+^r''^-')+-+A,,,,,,A^>^'+^^^^^^^^^^^ ,

the last term being ^s™,»«^'" or ^2,», «i-i (^"''^ + ^"'^^), according as ;;« is
even or odd. Substituting this in (162), we get

222

R. FUJISAWA.

(2w + 2)(2m+3M2,„+2.o+ in2-4(w+ 1)2^2^.0 = 0

^2,«,o-( i) (2w+l)!

(2m + 2)(2;;i + 8M2^+2,2 + {5«2_4(?;z+l)2}^2^,2

+ I (4wi + 1 )?r - 40;i + 1)- } ^ 2«, o

+ («2-'2?7i-2)(?t^-2?;i-l).42„,_2,o = 0 ,

(2w + 2)C2w + 8)^o„,+2,, + |9?i2-4(w+ 1)'M2^.4

+ {ti" -2m- 2)(7t' - 2vi - 1) A 2,„_.2, 2 = 0,

(2m+2)(2m+3)^2,»+2,6 + { VSn'-^m + lf]A,,,, ,
+ {(4w-7>i2-4(w+l)^}^2,.,4

+ Cm2_2w-2)(»--2w-1M2«.-2,4 = 0'

generally,

(168) C2m + 2)(2;?i + 8M2„,+2, 2. + { (4>' + 1)'^' - Um +\f\A .,,„, ,,

-f {(4w-4/-+5)n2-4(w+ l)'i.42„,o,_2

+ (;r-2w-2)(«2-2wi-lM2,„-2,2r-2 = 0,
and, lastly,

Clin + 2){2m + 8)^ 2^+2, m+i + '^ i ^^^ + 8)yr - 4(;/i + 1)- [ A o„,, ,„_i

+ {n- — 2m — 2)(h- — 2m — 1 )^ 2»i-2, m-\ = 0 ,

or

C2m + 2X27/t + 8M 2,H+2, ™ + I C-^''^ + 1)'^' - -l^"^ + 1 )' i ^ -',«, m
+ I (2/;t + 5);r - 4(;;i + If } ^ .2»,. «.-2

+ ('/i'^ — 2wi — 2)(n- — 2?;i- l)zi2,„_2,m-2 = 0 ,

according as m is odd or even.

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 22S

By means of the above e(|uatioris, -we tiiid

(104)
Aa = )i, A,= ^-^-| — (I + Â-j,

n{nr-4)

A^ = ^ [^ . ' KM--lÜX>r-8G)[0r--ü4Xl+/v-') + 4f4^r-l8X^- + A'^)]

-0(/i'^+lüÜM*-21i4«H4752)/l*},

^io = - '-^^jir^ {(;r-l6XH'-86K-(34)[0i'MOOXl+^-'")+5(5;i2-l(5XA-+A-«)]
-2('247M«-882/i*'-7-2l02/i* + 661il2yr-l86.S0U0X/^HA*'')},

'^12 = ^^^7^7^^ |(/r'-l6X^i--36)0^^-64)(M2-lOOXvr-l44Xl + F^)

+ 6(«-- l(5):;r-86X'r-64);;r- IOOXOm-- 19XA-+ P')
+ (-57öl;i^'^+170472^i«-4yOi40»*^-22744752;i*

+ 193986816^2- 3837 54240XA,H A;')
+ 4( -3046yt^« + 75579;i«-260532m«- 5901554«*

4- 48907368;r- 93498440)^'* } .

§ 19.

Lastly we consider the numerators of en }tu and dn nn, ii being
even. In this (;ase. ])Ut //"' = À p.

As in the case where n is odd, C...JA) = BoJ-r-jk'-'" by (10(5), so

224 ^- FUJISAWA.

that we may here al«o restrict ourselves to the consideration of B.
Moreover, since in this case

(165) B {X, k) = B (J^, k^ k'^ x'^,
it follows,

(166) B,^,„, = B,,,, k'^-^'\

and we need only determine the first half of the coefficients i^g,,,.
Now B satisfies the diiferential equation

(167) {1-(1 + A:V + AV['^.+ I [('27i^-l)Ä.-2-ll^-2(H2_l)^-2^-3}^

+ 2;i2Ä'(l-A-2)^+ {n\n''-l)k:'x^ + n-\ B =0,

whence we deduce

(168) (2w + l)(2w + 2)B2,„+,+ { {n^-é7ii') + 4:m0v'-m)k^B,r„,

clh:
Substituting

•^2»» = B2n, 0 + -^2/», 2 A"' + • • • + -^^2/n, 2r A-'' + . . . + B.2,n^ -2,11 ^'"\

m ^p,

in (168), we get

(:2m+ l)(27;i + 2)B.,,„+2,o +in'-^m')B,,,^o = 0 ,

B.2m,o = 0, /i<c2w<2^,

(2w+ I),2;;i + 2)ß2«f2.2 +{5n''-^m')B,^,2

+ 4m{n'-vi)B,^^, +{n--2m+\){n^-'2m + 2)B,,,.,,, = 0,

MULTIPLICATION OF ELLIPTIC FUNCTIONS. 225

(2w+ l)(2m + 2)ß,,„+.,, , + (9u'-4m')B,„^ ,

+ 4 { {m — 1 ',n- — m- \ B,,,,,^ ., + (>r — "2 m + 1 )(;r — '2m + 2)ß2w-2. 2 = 0 >

geDeralJy,

(169) {2m+ l)(2m + 2)ü.„,.,,,,r + [(4/ + l)yr-4w-]52„,,2,

+ (;r-2m+l)(;i2-2w+2)£2,«-2,2r-2 = 0 ,
and, lastly,

(2w+ lX27;i + 2}5o„,+2.2«+2 + {{^m+l)}r-4:m'}B,„^,^

+ (n--2m+l){n?-2m + '2)B,„,^._^._^_^ = 0 ,

By means of the above equations, we find

(170)

Bq = 1, -Dg = oT'

B, = |^[(/i2-4) + 2(;i^-l)^-2],

ij^ = !!Î!^Ili)[(,j2_l6)(«2_36) + ]2(;i2-l)(7i2-lG)A2+

+ 4(w-- l)(l5?r- r)l)k' + 32(11'- l)(;r- 9)A«] ,

2^^^^ = _ ^^^^j^ [.;r-lG)(u2-36)(/r-(34) + 2ü(;r-l):«--l6;(«2-86U-2

+ 12(71^- l)(;r-9X29/r- 104)Ä;*
+ 64(7v^ - 1 )(?J- - 9)(7 7i2 _ 22)/.«
+ r28(/r- l)(/i2_ y)(H2_ 16)A«],

2'2Q R. FUJISAWA.

9 / O J \

Bj., = ^'"^'^'^^7 [(/r- 10)(yr-86)(;r-G4)(?r- 100)

+ 80(;r- l)(;r- l6)(;t- - 80)0^- - 64)^'2

+ 40^2- l)(598u« - 14305;iH 1 1 3972^^2 - 257760)Ä;*

+ 40(7^2 _ l)(n2_9)( Il 5,^4 _i 288«- + '2968)Ä:«

+ 2880(M-- l)i7i'-'à){}v'- I6){7v'-3)k^

+ 5 1 2{n^ - 1 )(;r - 9)(;i2 _ 1 e)^,i- - 25)A-i" ] .

Imperial University, Tokio, July 189 o.

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On the Process of Gastrulation
in Chelonia.

(Contributions to the Embryology of Reptilia, IV.)

By

K. IVIilSukuri, Ph. D., Rigakuhakushi,
Professor of Zoology, College of Science, Imperial University,
Tokyo, Japan.

With Plates Vr.—VHf.

The sea-turtle, Clielonia caouana, UVu//. deposits its eg^s
on almost every suitable stretch of sandy beacli in the southern lialf
of Japan during tlie summer months of the year. During tlie breed-
inof season of this animal in 181*1, I Avas enabled, bv the liberaHtv of
the University authorities, to visit Sagara in the province of Tötömi.
with my assistant, Mr. T. Tsuchida, for tlie pur])ose of collecting
materials for the study of its develoj^ment. With the assistance of
several kind friends, we made arrangements to luive reported to us
every deposit of eggs that might he made along about fifteen miles of
sandy beach in that region, and we thus succeeded in getting hold of
several good deposits. As encli of tliese contained over one hundred
Ggg^ — 1-1 i"^ the least, and 145, the largest number in one deposit in
my exjierience, but 16!) has been i-ej^orted in one case— we had command
of over one thousand eggs in all, and as we opened eggs from each
deposit at certain intervals of time, we were able to secure unusually fine
series of embryos, gaps in one series ])eing often filled up by members

228 ^ MITSUKURI

from other.s. This success was in a larsre measure due to Mr. Tsuchida,
whose zeal and perseverance never flagged, even under most trying
circumstances ; and I would here express my deep indebtedness to
him. My thanks are also due to Viscount Tanuma, Mr. Y. Mura-
kami, the Mayor of Sagara, the Chief Officer of the Sagara Police
Station, and several other gentlemen who assisted us in various ways
and showed us much kindness durino- our stay. Messrs. T. Ogfasa-
wara and K. Niwa of Shizuoka were also kind enough to furnish me
with much useful information.

Various observation made by us on the breeding habits of the
sea-turtle together with similar facts which I have ascertained in
other species, I hope to embody, at some future time, in a separate
paper. A short preliminary account of these observations is already
published in the Zoological Magazine (Japanese) Vol. III., No. 35.
I will only remark here for the benefit of those who may attempt a
similar study, that Chelonian eggs can be transported with safety for
some hours immediately following their deposition, but after that,
their removal is apt to bring on death and decomposition. This
seems to be due to the circumstance that the white at the upper pole
is rapidly absorbed, the blastoderm becomes adherent to the shell
membrane, and a large fluid cavity is produced directly beneath the
developing embryo. In this condition, slight jarring seems to disturb
the delicate arrangements and to cause death. After thirty days or
so, Avheu tlie foetal membranes have become definitely established, the
eggs can again be moved with impunity.

The embryos of C h el o nia caouana, thus obtained, together
with those of Trionyx japonicus and Clemmys japonica
which I already possess or can get in almost any desired stage, afford
a good basis for the comparative study of reptilian development, and I
intend to use them for this purpose, as I have previously used those

ON THE PKOCESS OF WASTRULATION IN CHELONIA. ^29

of the two Jast-named species. MeanwhiJe I have discovered that when
C h el o nia caouana deposits its eggs, they are in a far less advanced
condition than those of 1'rionyx or Clemmys and thus enable us
to elucidate many points in the much discussed process of gastrula-
tion in the Amniota. The present contribution embodies the results
of my own study on this point and, it is hoped, will throw light on
some phases of this vexed question.

Preparation and Preservation of the Embryos.

Young embryos were in nearly all cases preserved in Kleinen-
berg's picro-sulphuric acid. Very advanced embryos were placed,
partly in that fluid, and partly in corrosive sublimate. In removing
blastoderms from eggs within one or two days of their deposition, at
which age there is not yet any large subgerminal cavity in the yolk, the
shell was removed and as much of the white as possible. The whole
egg was then placed with the blastoderm uppermost in a deep vessel
and covered with picro-suJphuric acid. The spot where the blastoderm
was to be found was generally marked with a hair since the thin layer
of the white necessarily left over it coagulates in the preserving fluid
and hides it entirely from view. Proceeding in this way the pre-
serving fluid will be found after three or four hours to have penetrated
to the blastoderm and acted on it as also on the upper strata of the
yolk. Incisions at right angles were now made with a sharp knife
on three sides of the blast(jderm, leaving the fourth side and the two

corners uncut, as shown in the accompanying diagram.

It was then found that a little manipulation with forceps

i)v scalpel easily separates the superficial coagulated white

from the blastoderm beneath it. If we then cut the
corners, the sheet of the white will roll up of itself towards the uncut
side, leaving fully exposed the blastoderm which bei)ig already hardened

230 ^'- MITSUKURI.

can then be removed with gi'eit eise. The bhistoderni thus removed
was o-enerally left in a relatively hirge quantity of tlie preserving fluid
lor some hours longer. In more advanced embryos tlie position of
the blastoderm under the shell is easily told in chelonian eggs by the
change of colour in the shell. In all the species I have examined, a
white patch a]jpears in the shell over the embryo, and increases in
extent with the growth of the embryo, or more strictly speaking.
pari jiassK with the disappearance of the white over tlie embryo ; so
that, roughly speaking, the size of the patch is a very good indication
oi the size of the embryo beneath. In these stages the embryo is
firmly adherent t<3 the inside of the shell, with a large subgerminal
fluid cavity in the yolk Ijeneath it, which can be easily pierced
tlirougli the shell and the blastoderm with the ])oint of one blade of the
line scissors. liy thus piercing the cavity and cutting round just
inside the edo-e of the white patch through both shell and blastoderm,
tlie embryo is removed, firmly adhering to the cut piece ; the latter
can tlieii be turned o\"er, exposing the ventral surface of the embryo,
and the preserving fluid be poured over it, using the cut-piece as
a veritable Avatch-glass. After half an hour or so, the blastoderm can
be easily separated fr<3m the shell and placed in a larger quantity of
the preserving fluid. This method has the great merit of keeping
every part of the blastoderm stretched in its natural condition, and
also of milking it possible to remove a Inrge number of embryos in
an incredibly short space of time.

When the embryo is very much advanced and the allantois has
spread itself entirely beneath the sliell, it becomes a serious (question
how to remove the shell without much injury to the fœtal membranes,
especially as the shell is leathery, and not brittle as in some
other reptiles. In this and similar ciises I carefully scrape tlie shell at
one small spot with a knife, until it becomes ((uite thin, and then apply

ox THE PKOOESS OF OA^'IKULATION IX CHELOXIA. 2'dl

to that spot some picro-sulphiiric acid, which reDiovcs calcareous
matter. I scrape again with the knife and again apply the acid. I
repeat this process, ah\ays using great care, until enough of the shell
is worn off to expose a very small patch of the allantoic surface, some-
times not larger than the eye of a needle. However small the opening-
may he, the acid is able to penetrate through it and harden the tissues
for some space around it. The opening may then be enlarged a little,
with perfect safety to the |)arts l)eneath. The acid is then applied
again, a still large area is hardened, and the opening is accordingly
made still larger. At length the opening becomes large enough to
allow of the removal of the entire shell without injury to the mem-
branes. In removing the shell, it is advisable to use the broad,
blunt-pointed forceps and insert them tangentially between the shell
and the fretal membranes. With a little practice, it becomes com-
paratively easy to obtain in this manner embryos with the foetal
membranes perfect, except for the yellow patch where the picro-
sulphuric acid was first applied.

As to staining, imbedding, îind cutting sections, there is nothing
special to communicate. I generally use borax-carmine for staining.
For imbedding, celloidin-paraffin is used.

The methods just described have been used in the case of Chelonia
caouana and in those of other species with equal success.

Description of the stages ofGastrulation in
Chelonia Caouana.

The first stage to which [ wish to call attention is represented in
Figs. 1 and \a. PI. VI. It was taken out of an egg which had been
deposited <jnly a few hours before. We notice tirst the oval -shaped
embryonic shield somewhat elongated in the antero-posterior direction.
At the posterior end of this, and for the most part, lying outside it,

232 K- MITSUKURI.

there is a second much smaller, irregularly circular white patch.
This is the structure called the "Primitivplatte" (the primitive
plate) by Will* (No. 1<S); and in later stages, when cells added on from
the subjacent yolk form an accumulation, is the " P r i m i t i v k n o t e n "
(the primitive knob) of Mehnert (No. <S). These names will be adopted
and used interchangeably in this paper, for it is after all difficult fo
distinguish when the state of the plate ends and that of the knob
begins. One stage of it is also called the • ' S i c h e 1 " by Will (No. 1 8).
In the dorsal view (Fig. 1) there is already in the middle of this area
a large, transversely elongated opening leading into a spacious cavity,
For convenience in description, I anticipate my conclusions by stating
here that I consider this cavity to be the archenteron and its
dorsal opening to be the blastopore. In the ventral view (Fig. la),
the appropriateness of calling the above area a " knob " is clearly
seen, for it is a thick accumnuilation of cells projecting much more
than the adjacent parts into the yolk. It is important above all to
notice that the archenteron at this stage is not open on the
ventral side, although it can be seen from that side through the
cell-mass in the specimen figured.

Fig. 9, PI. VIII. is a median longitudinal section through another
embryo of the same stage as that represented in Figs. 1 and la. In
this the yolk was left intact on the ventral face, so that it represents
no doubt a more nearly normal state of things than if the yolk had
been removed as in Fig. la. The blastoderm has already extended
itself over a wide area and, with the exception of the primitive
plate, is divided throughout into two layers : (a) the superficial layer
(the e pi blast of authors, the blast ophor of Van Beneden), and

* Also in No. 21. 1 regret that this paper came to my hauds only after the present
contriljutiou was nearly in shape to be given to the printer. I could not therefore make as
much use of it as I should have liked to do.

ON THE PROCESS OF GASTRULATION IN CHELONIA g 33

(h) the lower layer (the cœnogenetic hypoblast of Hubrecht,
the parader m of Knpffer, and of Mehnert, the lecitophor of Yun
Beneden.) The superficial layer, which T shall call the epi blast,
forms a distinct membrane and is composed of columnar cells in the
region of the embryonic shield, but changes gradually into low
cells in the parts outside the shield. The lower layer is composed of
irregular, amoeboid-shaped cells and does not probably form a con-
tinuous membrane. I hope to show in the sequel that this layer
ought to be regarded as only a part of the hypoblast, and might be
called the cœnogenetic hypoblast, after Hubrecht (No. 5 ) . In
the region of the primitive plate, there is a different state of
things. Instead of having two distinct layers, this area shows a thick
accumulation of cells. It is composed for the most part of an
irregular network of cells with tolerably wide meshes between, so that
it is not a compact mass. In the middle of this accumulation, there
is seen the i n v a o- i n a t i o n - c a v i t y — the a r c h e n t e r o n — leadino' at
first downwards but soon forwards and ending blindly. The roof of
this cavity shows distinctly a columnar arrangement of cells, and
becomes continuous with the epiblast at the anterior lip of the blasto-
pore. On the floor, as well as for some distance behind the blasto-
pore, so long as we are in the region of the primitive plate,
we see no columnar arrangement : the general network of the mass
extends up to the surface. There is no sharp line of demarcation
between the cell ii lar mass of the \) v i m i t i v e plate a nd the subjacent
bed of the yolk. The latter is divided into especially fine globules at
the boimdary line, and we can clearly see many cells arising at this
]»]ace and adding themselves to the primitive knob. That the
nuclei of these cells are the descendants of the segmentation nucleus,
there can be no reasonable doubt ; in fact I would have this
addition of new cells considered simply as the continuation of the process

234 K. MITSUEÜRI.

of segrnentation.* As the figure shows, the primitive plate be-
co'nes continuous with both the epiblast and the lower layer at its own
periphery. I am thus unable to find any independent sheet of cells
which lies below the primitive plate, and with which alone the
lower layer of the surrounding- parts becomes continuous, as Wencke-
hich (No. 15, Fig. 1) and Mehnert (No. 8, Figs. '20 and 21) and Will
(No. 21, Fig. 49) have found. On this point Virchow's observations
(No. 14) seem to be similar to mine ; Will also sees no such inde] »en-
dent sheet of cells in earlier stages (N(\ 19 Fig. 1, No. 21). I am
also unable to find such a sharp line of demarcation between the
shield and the primitive plate as is given in Will's Fig. 1 (No.
19f) : the primitive plate passes gradually into the epiblast both
anteriorly and posteriorly.

The yolk globules in this series of sections are generally spheroidal
with a uniform yellow tint. Generally speaking, they are markedly
fine immediately below the blastoderm, and become larger farther
below (See Fig. 9) : of their especially small size iinderneatVi the
primitive knob, I have already spoken. I agree with Virchow
(No. 14, )). 67) in thinking that the dark granules which Mehnert
describes in the yolk-globules (Xo. 8, Figs. 20 and 21) are artificial
productions made in the course of preparing sections.

I regret that I was unable to obtain the oTowino- edo-e of the
blastoderm which \\ould no doubt present interesting phases of growth
as observed by Virchow (No. 14) and Duval (No. 2). I am therefore
luiable to thro\N' much light on many questions bearing on cells found
in the yolk. I may however mention that in the series from which
Fig. 9 is taken, two kinds of cells are found imbedded in the
volk. Theii' nature is not donr to me. Tlic inorc iiumon~»us kind T

* Will appears to be of the same opinion (No. 21).

t This sharp deiuarention is insisted on still more strongly in Will's recent paper {Xo. 21).

ON 'I'HE PROCESS OF GASTRULATION IN CHELONIA. 235

have represented in Fig«. 10 and 12. Fig. 10 is part of a section
like Fig. 9 from a more lateral part of the blastoderm than tliat in the
latter figure. Beginning from the upper surface, we can easily recognise
the epiblast and the lower layer of cells lying immediately below.
Under these two layers, there is a rather thick stratum of spherical
yolk-globules. AYe then come to a crowd of cells which are the cells
in question. Some of these are large and full of yolk-granules ; while
others are smaller and formed of vacuolated protoplasm. The size of
their nuclei is tolerably unitbrm — being about .016 mm. in length.
In Fig. V2h are shown more distinctly three of these cells from
another region. One of them is full of yolk-granules, as is also the
unusually large cell shown in Fig. I2a. Another is partly full, with
an area of granular protoplasm around the nucleus. The third, of
which only one-half is seen — having no doubt been cut in two in the
process of microtomizing — has no yolk-globules, but is formed of
vacuolated protoplasm. Below this stratum of cells there is a layer
of closely packed fine granules which represents some liquid coagu-
lated in the course of hardening. Below this, we come to the thick
bed of yolk. The globules are here larger than in the upper layer.
The conclusion seems to me almost inevitable that the cells above
described take up and digest yolk globules and that the stratum of
liquid on the edge of which they are found is produced as the
result of their digestion. This liquid stratum has probably a genetic
relation with the large subgerminal liquid cavity found belovv the
blastoderm a day or two later in the course of development. So
much seems tolerably clear ; but whether the cells have for their sole end
the digestion and preparation of yolk globules for the nutrition of
other cells, or whether they themselves are to form s(jme integral
parts of the growing embryo, I am unable to decide.

Cells of the second kind found in the yolk never occur together

236

K. MITSUKURI.

in laro-e numbers l^ut are scattered, at least near the surface, in-
ditterently through the yolk-substance. At different points in the yolk
we find unusually large nuclei surrounded by a comparatively small
amount of protoplasm (Fig. 11« and h). Sometimes there is only
one nucleus (Fig. lib) and then it is very large. The one repre-
sented in Fig. ll/> measures .04 x .032 mm. Quite iis often, the
nuclei occur in a group of t^^o or three, closely adherent to one
another (Fig. 11a). These cells are no doubt what are called
" Merocyten " l^y Virchow (No. 14). What their nature is, whether
they stand in some genetic relation to other kinds of cells or are of a
nature siii generis, I am unable to say. I have thought it just possible
from the frequency with which two or three nuclei are found together,
that they are cells dividing by amitosis and possibly undergoing dis-
integration (Flemnn'ng [No. 4] and Ziegler [No. 20]).

Let us consider for a moment how such a stage as that described
has been reached. What I am inclined to think as probable is as
follows : — When the process of segmentation has gone on lor some
time, the blastoderm separates itself into two layers, the superficial
epiblast and the lower layer. Tliis takes place throughout the blasto-
derm with the exception of the primitive plate.* Here
cells not only remain undifferentiated but with the addition of cells
from the subjacent bed of yolk form a mass which protrudes into the
yolk — the primitive kn(j]j. In the middle of this region, an
invagination soon appears, which is at first shallow and is directed
straight downwards. I have two specimens of this stage but have
not figured them because tlie blastoderms having Ijeen peeled off from
the yolk to which it is adherent at this stage, the lower part of it is

* And proljably also of the growing- edge of the blastoderm, hut of this part I aui not now,

speaking 1 am gratified to find that what is given above as pro)3able is now verified by

Will by direct observation (See No. 21, Figs. 35 & 36).

ON THE PROCESS OF GASTRULATION IN CHELONTA. 237

probably not complete. But as to the above ])oint there is not room
lor mneli doubt. The specimens are very much like Fig. 1 of Will
(No. 19) with one exception, stated above, viz : that the epiblast of
the shield is continuous with the primitive plate and not separate
as in Will's figure. One peculiarity of this stage is that both the
anterior and the posterior wall of the invagination shows faintly the
columnar arrangement as seen in Will's figure. Later on, this feature
is confined to the anterior or dorsal wall (Figs. 9, 13, 14). After going
straight downwards some distance the invagination cavity takes a for-
ward horizontal direction and reaches the conditiim shown in Fig. 1>.
At the anterior lip of the blastopore, the columnar cells are recognizable
very early, and the epiblast is here reflected downwards to l)ecoine con-
tinuous with the anterior or dorsal wall of the invagination. In that
part of the primitive ])late placed behind the invagination the cell-
mass remains undifierentiated for a long time, there l)eing later estab-
lished in this place the rudimentary yolk-plug, as was minutely describ-
ed in the joint paper of Ishikawa and myself* on the germinal layers of
Trionyx. Robinson and Assheton (No. 10) object to our idea of
considering the structure in question as the yolk-plug. In the course
of tliis paper, I hope to show that the presence of the yolk-plug at
this place is an important feature in homologising the gastrulation of
the Sauropsida with that of Amphibia. I may add that several
authors, as Van I^eneden (Xo. 13), Wenckebach (No. 15), and Will
(Nos. 18 and 21) recognise the yolk-])lug in this place.

I shall next describe how the invagination-cavity, as described
above, comes to open below ;uid l)ecomes united with the large subger-
minal cavity in the yolk. This process has, ,so far as I am aware,

* Coatribiition I. I shall refer to the papers in the present series of Contrilnitions by their
nuuiliers in order of publication. See the list at the end of the present article.

288

K. MITSUKURI.

never been treated with the fullness which its importance deserves. A
careful study of this process has given me results which, I venture to
think, are of the greatest importance in discussing the problem of
gastrulation in the secondary meroblastic egg.

The surface views, Figs. 2-5, and the sections. Figs. 13-17 are
introduced to illustrate this process. Figs. 2 and 2a are of the
stage nearest to that represented in Fig. 1. In the dorsal view (Fig.
2), the dorsal opening of the invagination- cavity has now become a
narrow crescent-shaped slit with the concavity directed forwards.*
In the ventral view, the primitive knob has become larger. Viewed
with a low power, the surface of the knob is tolerably smooth, although
the figure represents it perhaps as a little too much so. The longi-
tudinal section (Fig. 18) of this embryo shows distinctly that the depth
of the primitive knob has grown greater in this stage than in that of
Fig. 9. The invagination-cavity has extended itself much deeper and
shows distinctly two limbs, one vertical and one horizontal. The roof
of the cavity which is as before continuous with the epiblast, shows
a distinctly columnar arrangement which is, however, gradually lost
both anteriorly and superiorly. In these directions it merges gradual-
ly into an irregular network of cells which is in turn continuous
with the lower layer of the embryonic shield. As was the case in
the former stage, there is again below the primitive knob, no in-
dependent sheet of cells continuous with the lower layer of the shield,
as described by Wenckebach or Mehnert. On the contrary, this and
the succeeding figures (Figs. 14-17) give the impression that the
lower layer of the embryonic shield extends below the epiblast

* Will (No. 21. p. Ii7) says: " Dieselbe {i.e. die Urmundspalte) tritt zuerst im vorderen
Abschnitt der Primitivplatte auf, und hat zunächst die Form einer Sic hei- rinne, nach
Schwund der Sichelhörner aber einer rundlichen Delle" That is, his figures 8 and 9 are
less advanced than his figures 4 and 10 so far as the shape of the blastopore is concerned. If
the first two figures named are comparable to my figures 2, 3, 4, and the latter figures (his
figures 4 and 10) to my figure 1, I can not but think that Will is mistaken in his views.

ON THE PROCESS OF GASTRULITION IN CHELONIA.

239

right up to the angle where the epi blast is reflected downwards at
the dorsal lip of the blastopore, and that the primitive knob has been
capped on to it from below, although now irrevocably fused with it
by a protoplasmic network. The floor of the cavity shows two
distinct divisions. In the posterior part (the vertical part in the
section) there is a compact mass of cells which have evidently been
proliferated from the floor of the cavity. This is the posterior median
part of the commencing peristomal mesoblast. In the anterior half of
floor, the vacuolated network ]jcomes very near the cavity, being
separated from it only by a thin sheet of cells.

In the next stage (Figs. 8 and ,3a), we notice one striking
change in the ventral surface view of tlie embryo. While the top of
the primitive knob (spoken of with its ventral surface as uppermost, see
Fig. 3 a) is comparatively smooth as in the former stnge, its base
has assumed a lioney-combed structure and this structure is spreading
itself over the ventral surface of the embryonic shield.

Fig. 14 is a longitudinal section near tlie median line of this
embryo. Compared with Fig. 13, the primitive knob has a longer
antero-posterior extension and it will be seen that this increase is due
almost entirely to the growth of the .'interior hnlf. The forward edge
of this half is gradually encroaching on the ventral surface of the em-
bryonic sliield (cf. Fig. 3 a) and is thus giving the primitive knoh ever
greater extension. Wenckebach (No. 15), AVill (Nos. 18 & 19), and
Mehnert (No. 8) agree in thinking that the forwai'd growth of the
priinitive knob takes place by its front growing edge insinuating itself
between the epiblast and the lower layer of the shield, and quit« indepen-
dently of these two sheets of cells.* My sections do not allow me to

* In his latest paper (No. 21), Will admits that where gastruLitiou ia completed ]}j the
formation of the Kopffortsatz, the " primary " and " secondary " endoderm cells cannot be clearly
distinguished and that the former may grow ))y addition of the cells of the latter formed in situ
(p. 48).

240

K. MITSUKURI.

come to the same conclusion, us a glance at Figs. 18 and 14 will show.
Both the surface-views (Figs, ^a ct i^eq.) and the sections give us even an
impression that the primitive knoh is spreading itself under the lower
layer of the embryonic shield. In the parts where the primitive knob
has once established itself, we can, however, no longer distinguish
cells that have come from the primitive knob from those of the lower
layer of the shield : they are indistinguishably fused. The invagina-
tion cavity at this stage (Fig. 14) has much greater longitudinal
extension than in that of Fig. 13. I can discover neither at this nor
at any subsequent stage a j^osteriorly-directed limb of the invagina-
tion-cavity, such as is described by Wenckebach (Xo. 15) in his Fig. 3.

There is nothing special to say of the nwf of the invagination
cavity, except that the points described in the previous stage are all
more pronounced in this one. In the floor, there are some im])ortant
changes. In the ])osterior half, where the mass of the peristomal
mesoblast, grown much more compact, is easily recognisable, there is
not much that is new. l)ut in the anterior half of the floor,
the wall of the invagination cavity is no longer so sharply
defined as before, and some meshes of the cellular network
in the primitive knob even open into the invagination-
cavity, so that we can here, already in this stage, pass
by a labyrinth of intercellular passages from the in-
vagination-cavity to the subgerminal yolk -cavity. It
should be specially noted that the anterior and of the invagination
cavity is distinct and does not share in the dissolution of the anterior
part of the floor.

With the growth of the embryo, the changes in progress between
the stao-e of Fi«:. 13 and that of Fio-, 14 become more and more
pronounced. The primitive knob grows forwards moi'e and more on
the ventral surface of the shield, so that its antero -posterior diameter

ox THE PROCESS OF GAS'J'RULATIOX IX CHELOXIA. 241

is ever getting longer (Fig^. 14, 15, 16 & 17). In the anterior part
of the lloor of the invagination-ca^■ity whicli was already losing its
sharp definition in Fig. 14, tlie disrii]»tion lias proceeded une step
farther in Fig. 15. In this figure, not only this part of the fioor is
giving away, but the network of cells lying underneath it, and
between it and the subeferniinal volk cavitv, has been laroeh' absorbed.
In Fig. 16, the process of breaking through is seen to be complete,
and the invagination-cavity has now a clear opening below. I think
it almost certain that such a clear and comparatively large opening-
has been produced by the running together of se^'eral small openings,
such as we see in Figs. 14 and 15, which put the meshes of the cell-
network in communicati(Hi with the invagination -cavit}'. In fact, in
Fig. 16 we can still see several such openings in the floor of the
cavity in that part of the network situated behind the large anterior
opening and in front of the compact peristomal mesoblast mass. Com-
parison with Fig. 17 makes it probable that this part of the cell-
network is to be eventually absorbed, for the single large opening-
extends in the latter back almost to the peristomal mesoblast. It
should also be noticed in Fig. 16 that the extreme anterior end of the
invagination-cavity is clearly recognisable and does not particii>ate in
the breaking through, which seems to be confined to the floor. We
should therefore remember that although the anterior end may n<jt be
recognisable in later stages (e.g. Fig. 17), it is the floor which is
open below. The surface views of the stage at which the invagina-
tion cavity has just o])ened below are given in Figs, o his, 4, and 4«.
There is considerable difference in the appearance of the two embryos
Avhich I am not able to explain. I drew them just as they appeared
under the microscope. I am rather inclined to think that Fig. 4(a
represents a more normal appearance, if we are to judge from the
succeeding stages, although I am unable to detect anything unusual

242 K. MITSUKURl.

in the sections of the other embryo (Fig. 3 his), Fig. 16 in fact being
one of them.

From the fict« given above, the conclusion is reached that the
in vagination-cavity comes into communication with the
subgerminal yolk-cavity by the absorption of the most
anterior part of its floor as well as of the cell-network
lying underneath this part. In this view, I find myself in
agreement with Mehnert (No. 8, p. 411) who says : — " Wenn der
Einstülpungssack etwa die halbe Länge des Embryonalschildes
erreicht hat, schwindet indem vordersten Abschnitte seine untere
Wand und das mit derselben innig verwachsene Paraderm, so dass
durch diesen Vorüfany- eine freie Communication zwischen der
Einstülpungshöhle und der Subgerminalhöhle gebildet wird " I
w^ould only remark that this disappearance does not take place
suddenly, as Mehnert's words might ]30ssibly lead one to suppose.
As my sections show, it is already begun as early as in the stage
given in Fig. 14. Wenckebach's Fig. 4 (Xo. 15 p. 60) is very much
like my Fig. 16, except for the differences already specified. Although
his views are not given in detail, I think, they are probably similar
to mine. Will's views are essentially like mine ; only he insists on
the greater forward and lateral extension of the invagination-cavity
before it opens below. This is especially the case with the tortoise.
He (Xo. 10, p. 191-2) says: "Aus diesen Stadien geht nun die
wichtige Thatsache hervor, dass auch dei' Urdarm der Schildkröte noch
in seiner ganzen Ausdehnung hohl ist und dass seine Ausdehnung
absolut und relativ diejenige des Gecko noch übertrifft. Während
derselbe beim Gecko diejvorderen und seitlichen Ränder des Schildes
nie vollständig' erreicht, nimmt derselbe bei der Schildkröte stets
die ganze Fläche des Schildes ein. Der Durchbruch des Urdarms
erfolgt auch hier ganz ebenso wie beim Gecko, so dass die Fig.

ON THE PROCESS OF GASTßULATIOX IX CHELONlA. 243

7 meiner oben zitierten Mittheilung (No. 1<S of my list. Same as
Fig. 176 of No. :21) auch geeignet ist, die ^"erhältni8."se hei der
Schildkröte zu illustriren. Es treten zunächst einige wenige isolirte
Durchbrechungen der untern Urdarmwand (nebst dem unter derselljen
wegziehenden Dotterblatt) ein; indem sodann beständig neue Lücken
auftreten, die alten sich aber vergrüssern gelangt man zu Stadien, Ijei
den von der gesammten unterer Urdarmwand nur noch ein unregel-
massiges, bei den verschiedenen Embryonen verschieden gestaltes
System von Netzbalken erhalten geblieben ist. Schliesslich kommen
auch diese letzten Reste zum Schwunde, wodurch dann das bisherige
Urdarmlumen mit dem subembryonalen Kaum zusanunenfiiesst."
After seeing his Figs, bba and h (No. 21) it seems no longer possible
to doubt the great anterior extension of the archenteron in Gecko,
bef<jre it breaks open. As t(j this point in Chelonia I shall reserve my
judgment until his prijmised full paper on the tort(3ise appears.

Van l^eueden (No. 13) describes in Mammalia two kinds uf
openings by whicli the cliorda-canal comes to open below into the
blastoderm cavity, viz: — (1) an anterior transverse slit, and ('2) several
openings which soon run together into a. single posterior longitudinal
slit. From what has been stated aljo\e. J need hardly say tliat 1 do
not tind any such differentiation of openings in Chelonia.

The changes that follow on the ijreaking through of the
invagination-cavity can best be seen in the surface views. Figs.
5 and oa — (S and 8c( are introduced to illustrate this point, ^\'e
have seen how the primitive knob, at first c<jnfined to a small
accumulation of cells at the posteri(jr edge of the embryonic shield
(Fig. y), gradually spreads itself anteriorly until it comes to occu])y
quite a considerable area on the ventral surface of the embryonic
shield (Figs. 13- IG), when the invagination cavity breaks through be-

244 ^- MITSUKÜEI.

low (Fig«. 4rt and 16). The part that has been covered by the cells
of the primitive knob can be very plainly distinguished on u ventral
view of the blastoderm, showing mostly a trabecular network (Fig.
Aa). This gradual spreading of the cells from the primitive knob
(jver the ventral surface of the embryonic shield is continued long
after the breaking through of the invagination-cavity. Figs. 5 and
ba are only a little advanced on Figs. 4 and 4a. The area of the
network is not yet very large, ])ut in the stage next introduced
(Figs. () and 6a) it has expanded itself considerably. A great change
is now noticeable : a, circidar area at its centre shows no longer a
network but presents a smooth compact surface. This is produced by
a continuation forwards of the process by which cells in the roof of the
archenteric-cavity, beginning at the dorsal lip of the blastopore, have
gradual 1\- assumed the columnar shape and formed themselves into a
compact sheet (Figs. 13-18)

In Figs. 7 and 7a the spreading of the part derived from the
primitive knob has gone one step further. Not only is the area
occupied by the network larger but the compact part in the centre is
considerably enlarged by its extension anteriorly in the median line.
There is also anotlier noteworthy new feature : at tlie anterior end of
the median compact area, there is a slight transverse ridge. This is
the com m e n c i n g head- f o Id. The last stage in which I was able
to dete(.'t traces of the network is shown in Figs. 8 and 8a. At the
front end of the embryonic shield, a patch of the network could Ije
faintly traced. I tliink it almost certain that the part derived from
the j)riniitive knob does not extend itself much beyond the area of the
embryonic shield, and that it gradually thins itself out and ends by
becoudng simply continuous with the primitive lower la^er at or near
the periphery of this area (Figs. 17 & 18). Witli the exception of
the patch above-mentioned, the ventral surface of tlie embryonic shield

ON THE PROCESS OF GASTRULATIOX IN CHELONIA. ^45

presents now n smootli compact appearance. Tlie head-fold and the
chorda -groove have ah'eady Ijecome conspicuous.

As the head-fold, formed well within the edg^ of the emhryonic
shield, marks the anterior end of tlie emhryo, and therefore of the
archenteron or the adidt alimentary canal exclusive of the stomod^eum ;
as the primitive kn<:>h marks the posterior end of the emhryo ; and
as the lateral Iwdy-wall is formed from the lateral folds, also arisen
within tlie embryonic shield, we are justified in coming to the very
important conclusion that the body of the future embryo and
consequently the definitive alimentary canal is formed
entirely within the area covered ventrally by cells derived
from the primitive knob. This speaks in fli vor of the assump-
tion that the invagination cavity is the archenteron and gives rise
to the fiitare alimentary canal. I sliall discuss farther on how we
ought to regard the breaking through of the invagination-cavity and
the gradual sj)reading of the cells of the primitive knob over the
ventral face of the embryonic shield.

The reason wliy the advancing edge of the primitive knob is
marked by a zone of network is probably, T tliiid^, that sucli a
structure allows free and easy access of the nutritive liquid of the
yolk to the deeper parts of the tissue.

The network sucli as is here described, has been seen many
times before. For instance, Ishikawa and 1 noticed it in a
Tri onyx blastoderm (Fig. l/> of Contrilj. I) without knowing its
significance. Again, Fig. 10. C^ontrib. Ill represents the same
thing in cross-section in an embryo of Clem my s . Mehnert (No. 8)
gives beautiful ilhistrations of stages showing the network, in his
Figs. 4-13. He, however, gives an explanation of it which is at
utter variance with the one given above, for according- to him, it is
concerned with the process of the mesoblast formation. He states

246 K- MITSUKURI.

thnt ill the anterior part of the embryonic shield, the dorsal roof of
the archenteric cavity divides itself into two layers : (1) a lower c^ne
consisting of a single layer of low cells representing the definitive
hypoblast, and (2) an upper one consisting of stellate branched cells
representing the " Rumpf-mesoblast " (his Figs. 22 & 23). In the
course of this separation, the dorsal roof which is nt first composed
of compact columnar cells becomes permeated by vacuoles, aiid he
says tliat " das im Flächenbilde eruirte Xetz der Ausdruck fiu' die aus
dem Verbände des Urdarm-epithelhofes (sel. oberer Ürdai-mwand)
losgelösten Mesodermstränge war, welche sieh im Furcluingsspalte
centrifua-al zwischen Ektoderm und Paraderm weiter vorschieben "
(]^. 434). He thus calls the ai'e:i of the network with the central
compact ])nrt the '' Rumpfmesodermhof." Moreover he makes thi.s
[)r<jcess of the mesoblast formation begin at the cnminl end and proceed
backwards. He also says thnt "'die periphere Ausbreitung des Meso-
dermhofes nicht im proportionalen Verhältnisse znr Grösse der Area
embryonnlis (sei. Euibryoniilschild) steht " (p. 434j. Mehnert's views
C!in not be reconciled with mine: one of us is wrong. Except as to
the single ])oint that the network grows centrifugally, I am obliged to
differ from him in almost every particular. This network has in my
opinion nothing to do with the process of the mesoblast formation.
My views on the latter process have already been given in great detail
in two former papers (Contrib. I. & III.) and I do not intend to
go into them again in this paper. The network is simply the surface
expression of cells from the primitive knob spreading themselves over
the ventral fnce of the embryonic shield. In what light we ought to
regard this pi-ocess I shall discuss farther on. But whatever it is, it
does not begin at the cranial end and proceed backwards. In obt:iin-
ing materials for the ])resent investigation, I opened on consecutive
days a certain number of eggs from tlie same deposits, and observed

ON THE PROCESS OF ÜASTRULATION IN CHELONIA. 247

the ]H*ogres.s made durino- the interval of time hetween the two
successive acts of takiii": oat, making' of course due allowance for
fast or slowly developing ego-s. Fig. 1 and Figs. 4-<S with some
intermediate staofes, whicli I ha\'e not introduced liere, helono- to one
of the series obtained in tliis mariner, and tliese show, conclusively
so my mind at least, that the area of the network spreads itself gradual-
ly underneath the eml3ryonic shield from the spot where the
invagi nation-cavity first breaks through, towards the periphery of the
thield i.e. from the posterior primitive knob anteriorly over the
embryonir shield. It is not the network that is gradually encroaching
on tlie central compact area, as Mehnert assumes, Init just the reverse;
for the central compact area is formed out of the area of the network.
The series, Figs, ôa — Ha, shows also that the area of the network
increases with the age of the embryonic shield. I can not therefore
accept Mehnert's explanation of the appearance of the network on
the xentral face of the blastoderm.

As I said just now, I do not ))i"opose to go into the mesoblast-
formation again in tliis paper. I woidd merely remark that in tlie
stao-e corresponding to Fig. 17, I already see the establishment of the
chorda-hypoblast and the stretch of the epithelium on eacli side of it
which becomes transforjned into the gastral mesoblast. (Compare
Fig. 11, Contrib. TIL).

There are two other points on which I wish to make some
remarks.

The first of these is in regard to the position of the primitive
knob relatively to the embryonic shield. In Fig. 1, the primitive
knob lies for the most ])art outside of the embryonic shield, only
about one-third of its antero-posterior extension being within the
shield. In Fios. 2 and 3 it is al)out one-half, and in Fiiz'. 4. entirely
within the shield. This is no doubt brought about by the gradual

248

K. MITSÜKURI.

exteii.sion of the epiblastic area composed of columnar cells. In later
stages, (Figs. 6, 7, 8), the mass of the peristomal mesoblast no doubt
helps in causing opacity in the posterior region. From tlie stage of Fig.
6 on, the embryonic shield, which has hitherto passed gradually into the
surrounding jjarts, becomes sharply marked off from the circumjacent
transparent area, in which it is eccentrically placed, and becomes ap-
parently diminished in size.

The -second point on which I wish to touch is as to the dorsal
opening of the invagination cavity, in the earliest stage 1 ])ossess
(referred to on p. 236-7 ), it is a squarish pit rather elongated in the
antero-posterior diameter. In Fig. 1, it is a wide open cavity
elongated transversely. In Fig. 2, it is a crescent-shaped transverse
slit, no longer gaping, and with its concavity turned forwards. The
same can be said of Figs. 3 and 4. In Fig. 5, the l)]astopore is
nearly straight across. It has a slight notch in the median line open
backwards. In Fig. 6, the ends of the slit-like opening have turned
backwards so that now the concavity faces backwards. In further
growth, the backward curvature becomes greater and greiiter, until it
becomes a h«3rse-shoe shaped slit, as can be seen in figures contained in
the former Contributions. Of the significance of this change of shape I
shall speak later on.* The yolk-plug, which can be traced more or
less clearly from the first, becomes very distinct as the backward
curvature becomes greater, and sticks out between the two limlis of
the horse-shoe.

T'he yolk-[)lug in Fig. S is very peculiar in that it has a groove
in the median line. The cross sections of this reofion also show it to be

* In his latest paper. Will (No. 21) seems to consider the enclosure of the primitive knol)
within the em1>ryonic shield as intimately connected with the change of the shape of the blasto-
pore-opeuing. Both are, according to him, due to the forward growth of the yolk-plug and the
consequent shoving forwards of the Wastopore-opening (see p. 127 et seq.). I find myself
imable to accept his views.

ox THE PROCESS OF GASTRULATION IX CHELOXIA. 249

ji deep fiwsiire cutting the yulk-plug into liuJves. I am unable to give
any ex[)Janation of thi« groove, whicli is very unusual, this being
the sole instance among hundreds of chelonian embryos that have
passed through my hands. 1 think it may probably be teratological.
Kuptfer (No. G, Taf. IV. Fig. 40 /. & y.) gives two figures of
Coluber that are strikingly like this.

To sum up the facts of Gastriilation as above described : —

1. When segmentation has gone on for some time,
there is established in the blastoderm two layers: (<<) the
superficial epiblast composed of columnar cells, and (/))
the lower layer composed of irregular stellate cells and
probably not forming a complete sheet.

2. This separation into two layers takes place in all
parts of the blastoderm v\^ith the exception of a small
area at the posterior end of the future embryo. Here
not only is there no differentiation of layers but a thick
knob consisting of a network of cells is produced by the
accession of cells from the subjacent bed of yolk. The
mass can not be said to belong to either of the two
layers above named. This is the Primitive Plate or Prim-
itive Knob.

o. In the middle* of the Primitive Knob, an in-
vagination cavity is produced, which at first goes straight
downwards but soon takes a forward horizontal course.
This is the Jnvagination-Cavity or the Archenteron. its
dors;il opening is the Blastopore. The in vagination-ca vity

* Will (X'u. 21) is uo doubt quite correct in printing out that the invagination -cavity
begins much nearer the anterior than the posterior end of the primitive plate. In front of it
there is only the future anterior or dorsal lip of the cavity.

ç)50 K. MITSUKURI.

extends itself gradually forward«, pan passu with the
anterior enlargement of the Primitive Knol).

4. The roof of thi« invagination-ca^■ity which be-
comes continuous with the epiblast of the embryonic
shield at the anterior lip of the blastopore, assumes a
colunmar arrangement, the process beginning at the
posterior end and proceeding gradually forwards. Out of
the median part of it is established the Chorda dorsalis,
and from a certain stretch of columnar epithelium on each
side of it is developed the gastral mesoblast.

5. The floor of the invagination-c;ivity is divided
into two parts: — ((/) the posterior which proliferates the
peristomal mesoblast, and (h) the anterior which losing
deiiniteness is finally absorbed, together with the whole
thickness of the cell network placed beneath it, ih ns putting
tlie invaginaiion-cacit ij in commun ication with the large suIj-
germinal cacittj in ttte yolk.

6. The primitive knob which was gradually spread-
ing itself over the ventral surfjice of the embryonic shield
before the breaking through of the in vagination-cavity
continues to do so after that event. It spreads from the
spot where the in vagination -ca vity first broke through
away towards the periphery of the shield. Its advance
in later stages is marked by a zone of cell -net work
with a compact central area. AVhen the whole of the
ventral surface of the embryonic shield has been
covered, the process sto])s. The cell-network afterwards
changes into compact cellular sheets.

7. The head-fold is formed some distance behind
the anterior edge of the embryonic shield.

ON THE PROCESS OF GASTRULATION IN CHELONIA. 251

8. The future embryo and consequently the defini-
tive ulimentary canal is formed entirely within the
area covered ventrally by the part derived from the
primitive knob.

Putting the results in another way, they may be summed up as
follows : —

From the epibla.st of the embryonic shield, the
EPiBLAST and ITS DERIVATIVES of the future animal is
derived. In the region of the primitive plate and its
anterior enlargement are produced the invagination-
CAVITY (the Archenteron), the yolk-plug, the chorda,
the MESOBLAST (botli peristomal nnd gastral), and the
DEFINITIVE HYPOBLAST and ITS DERIVATIVES. The primi-
tive lower layer forms the wall of the yolk-sac, and
contributes to the future animal only in so far as some of
its cells are unrecognisably incorporated with the cells
of the primitive knob, when the latter spreads itself
over the ^^ e n t r a 1 sur fa ce of the e m b r y o n i c shield.

On the last point, I find myself at variance with Wenckebach
who makes the cœnogenetic hypoblast take part in the formation of
the anterior ])art of the embryo-body, "namentlich an dem cranialen
Wachsthimi von Chorda und gastralem Mesoderm (No. 15, p. 76).

Theoretical Considerations.

If we represent the chelonian egg dia grammatically in the light
of the facts described in the preceding pages, we shall obtain something
like that given in Woodcut I.

25â

K. MITSUKURI.

Woodcut I.

Invagination Cavity ( Archente ron)
Yolk-Plng

Head-fold

X

Diagram of a Chelonian Egg.

A — B represents the embryonic shield with the enclosed primi-
tive knob. Within the shield is established the whole of the future
embryo. A—X—B is the yolk-bag with the large subgerminal cavity
filled with nutritive liquid. The invagination -cavity which has ex-
tended forwards pari passu with the anterior extension of the primitive

ON THE PROCESS OF GASTRULATION IN CHELONIA.

253

Woodcut II.

knob has by tlie absorption of the anterior part of its floor (indicated
by dotted lines) been put in communication with the subgerminal
cavity in the yolk. The anterior end of the invagination-cavity is
clearly recognisable at the time of the breaking through ; it becomes
invisible for a time after that event, but is soon marked out again by
the commencing head-fold. Tlie tliick part of the hypoblast (marked
with slant lines) is intended to show the extent to which cells from
the primitive knob spread themselves. The structures l^ehind the
invagination cavity — the yolk-plug, the peristomal mesoblast — have
been fully described in ('Ontribs. I. & III.

When we compare
this diagram with tlie
well-known one of an
amphibian egg (Wood-
cut II.) given by Hert-
wig, their similarity be-
co7nes very striking.
.p.Tist-Mesoi.i. The structures dorsal to
the line Z— Yin the am-
phibian egg can be
identified,^ part for part,
on the emljryonic shield of the chelonian egg. In homologizing
these two eggs a great deal depends upon the view we take as
to the nature of the invagination-cavity, the breaking through
of the same, and the large subgerminal cavity into which the
the invagination cavity opens. I have already made mention of the
assumption that the invagination cavity gives rise to the definitive
alimentary canal (p. 245). The following considerations will make
my views clear and I trust, justify them at the same time : — There
is in the chelonian egg a large yolk reservoir A (Woodcut III. 1).

Y.^lk-Pluff

254

K. MITSUKURI.

This, let us suppose, is surrounded by a layer of cells (although in
point of fact the lower pole is not enclosed until a much later
period), except at the point C where there is a mass of cells in
which both the epiblast and the layer surrounding the yolk are
merged. This is the Primitive Plate or Knob. Tn this knob,
there arises an invagination, B (Woodcut III. 2) which grows forwards
together with the anterior elongation of the primitive knob. Assume

Woodcut III.

ON THE PROCESS OP GÂSTRULA.TION IN CHELONIA. 255

for the present tlie invagination-cavity (B) to be the Arche n teron .
Then the yolk reservoir A must from the nature of the
thing be an appendag-e of the in vagi nation-cavity B.
Hut owing to its enormous size compared witli B, the bodily invagina-
tion of tlie yolk is out of the tpiestion. It forms the most conspicuous
part of the egg from tlie first, and begins to surround itself with a
cell-layer long before the invaginntion-cavity B makes its appear-
ance even. Hence, A can only secondarily come into connection with
B. This ha])pens by the anterior part of the floor of B flaring out,
so to speak, into a funnel-shaped opening. This is the mean-
ing of the breaking through of the invagination-cavity. The spread-
ing of the cells derived from the ])rimitive knob over the ventral
surface of the embryonic shield after the breaking through of the
invao'ination-cavitv mav be regarded as the gradual Iv thinnino- wall of
the funnel-shaj^ed opening making itself continuous with tlie cell-layer
surrounding the yolk (Woodcut III, o-longitudin;d section, 4-cross
section). When the alimentary canal is formed definitely, later on in
the course of development, the wall of the funnel-shaped opening of
the invagination cavity is again tucked in as the splanchnopleura.

The above course of reasoninu* explains all the events accompany-
ing the invagination and thus justifies the assumption that the in-
vagination-cavity (B) \fi the Archen ter on corresponding to the part
marked as such in the amphibian egg (Woodcut II.) and the whole
yolk-bag must be regarded simply as a part of its ventral wall that
has become bulged out on account of the enormous accumulation of
nutritive matter within it. The jiresence of a large subgerminal
cavity in the yolk filled with a nutritive li(piid is a physiological
accident, so to speak. I agree with Van Beneden, Keibel, and
Wenckebach in regarding it as intercellular space in the yolk.
It is a cavity arisen solely from physiological necessity

256

K. MITSUKURI.

and having a comparatively insignificant morpholo-
gical value. Although the whole yolk-sac should he regnrded as a
diverticulum of the archenteron and although it has a definite morpho-
logical value, it is a matter (^f comparative indifférence, so far as
morphology is concerned, whether its inside is filled with cells
charged with yolk-granules, or with free yolk-spheres, or with a
nutritive liquid or with a mixture of all three. Looked at in this
light, the chelonian egg is nothing hut the ani])hibian egg, with an
enormous ventral saccular appendage surcharged with nutritive
matter.* The whole yolk-sac (Woodcut I. J-X—B) must not,
however, he looked on as strictly homologous with the part of the
amphibian egg ventral to the line Z~ Y. For, in the latter, the
epiblast of that part becomes the ventral abdominal wall of
the future animal, while in Chelonia the epiblast of the yolk-bag
becomes later n part of the serous envelope, — the ventral a b d o m i n a 1
wall of the embryo being formed within the embryonic
shield above the yolk-sac, and the yolk-sac with the enclosing sheet
of hypoblast and mesoblast cells migrating within the body of the
embryo. When it has done so, nobody has any difficulty
in accepting it as an appendage of the alimentary
canal which has for its function the storage of nutri-
tive matter. My contention is that as such it should be
looked on from the first. The archenteron is at first so utterly
insignificant in size compared with the yolk-sac that the true na-
ture of the latter is obscured : none the less the yolk sac is a mere
appendage of the archenteron. This < view makes it necessary to regard
the primitive lower layer enclosing the yolk-sac as a part of the

* It will 1)6 seeu that f urtlier considei-ation has made me modify in some details my views
as set forth in the preliminary notice sent to the Aiuitomhcher Anzeiger, and pulilished in that
journal, Nos. 12 & 13, 1893..

ox Tue process OE UASTRULATIOX IX CHELOXIA. 257

hypoblast. That it arises before the invagination of the archen-
teron can be explained by the principle of precocious segrega-
tion, as has been pointed out by Hubrecht (No. 5). The name
"cœnogenetic hypoblast" which he applies to this layer, seems
therefore very appropriate as the part derived Ijy invagination may be
called the "palingenetic hypoblast."

I think the objection on the part of Robinson and Asshetoii
(No. 10) that the yolk-})lug can not be present at the spot designated
by Ishikawa.and myself is fully answered by comparing the two
diagrams (Woodcuts I. & II.). The yolk-plug can not only pro-
perly be found at this spot but its presence here is one of the signi-
ficant, although not the essential, features in homologizing it with the
amphibian egg. This is an example of those cases where a secondary
characteristic is of great service in identification.

According to the views set forth above, the enormous accumula-
tion of yolk has profoundly affected the course of development in the
chelonian agg, especially in the precocious develo})ment of a part of
the hypoblast, and in the rapid spreading of the blastoderm over the
surface of the egg. There is left, however, in the centre of
the blastoderm a certain amount of raw undiiferentiated
materials in the shape of the primitive ])late or knob
in order to go with it through certain developmental processes of
palingenetic character : — the invagination of the archenteron with the
consequent establishment of the cliorda-hypoblast, the peristomal and
gastral mesoblast, the yolk-plug, and the definitive hypoblast. The
changes in the shape of the blastopore from a crescent with its con-
cavity turned anteriorly to that of a horse-shoe with its two limbs
directed backwards and enclosing the yolk-plug between them nnist
be looked on as the remnant of that process by which the epiblast
gradually encloses the endoderm cells in the amphibian ovum or the

'25S

K. MITöUKUßl.

Woodcut IV.

yoJk in the Elaismobranch egg. If we make a companion diagram
to the well-known series given by Ballour (Comp. Embryol. vol. II.

Fig. 175) it will be like Woodcut
IV. The upper pole of the
egg in Keptilia is capped
by a small patch where
nearly all the changes
which in Amphibia are
gone through by the whole
Qgg, are performed. Ac-
cordingly, the enclosure of the
yolk by the blastoderm in the
chelonian egg is of a very difterent
nature from the enclosure of the
yolk in the Elasmobranch, for while the former is a simple growth of
the edge of the l)lastoderm, and of cœnogenetic character, the latter is a
]jart of the process of invagination and of palingenetic character.
That the yolk in Chelonia is not completely enclosed till the embryo
has made much progress is due to its large size, and may be regarded
as of quite secondary significance. I thus find myself obliged to put
aside tlie yolk-l)l;istopore of ])alfonras no longer tenable in Sauropsida.
After what has been given above, I need hardly say that I accept
the views of Rabl (No. 9) as to the loss and acquisition of the yolk in
vertebrate eggs several times in the course of the phyletic development.
All the facts given Jibove tend to pro^e that Chelonia possesses a
secondary meroblastic ovum in contrast to the primary meroblastic
ovum of the »Selachians.

My views overlap more or less those of previous writers, such as
Wenckebach (No. 15), Will (Nos. 18, 1!) & iM), Mehnert (No. 8)
and Rabl (No. *J). It would, however, be a tedious and useless task

ON THE PROCESS OF GASTRULATION IN CHELONIA. 259

to go over the writings of these authors and point out wherein we
agree or differ. The reader acquainted with the hterature will be
able to do this for himself. The points which I want specially to
emphasize are however as follows : —

1. The pklmitive; plate or knob is raw-material left at the
centre of tue blastoderm, by means of which certain palingenetic
processes are gone tlirough.

'J. The INVAGINATION-CAVITY is the APCHENTERON, and givcs rise
to the alimentary canal and the organs derived from it exclusive of
the proctodti3um and the stomodseum.

o. The YOLK-SAC must be regarded as a ventral appendage or
diverticulum of the AiiCHENTERON in which nutritive matter is stored
in solid or liquid form.

4. ÜAving to the enormous size of the yolk-sac, it and the
AKCHEXTEKON are formed separately from each other, and come only
secondarily into connection.

Having considered the chelonian egg in its relations with that
of Ichthyopsida let us now see how it compares with the avian or
mammalian egg.

If the process by which the blastopore in Chelonia has assumed

a horse-shoe shape (W oodcut V. A) continues on after the state A

is reached, as actually happens in Amphibia (See Figs.
Woodcut V.
^^.^^ 18 & ID, Ao. 10), the lateral lips will coalesce* and

I there Avill result the primitive streak of the avian egg
(]>). In cases where the lips have not quite coalesced,
we .should expect to find the ^^olk-plug sticking out

* I am gratified to find this verified within the group of Re'ptilia. Will (No. 21) has found
in Gecko that the lips of the blastopore approach each other very closely and form a primitive
streak.

260 ^- MITSUKURI.

between them, and such is actually the case as seen in Figs. 15
and 32 of Duval (No. 3). The annexed woodcut will also explain
why the posterior limit of the primitive streak is not as sharply
defined as the anterior. This view makes it plain that the
homologue of the primitive streak in Chelonia is the lips of the
blastopore which are, however, still so wide apart from each other
that the name " streak " is hardly applicable to it. It should be
noted that the primitive plate or knob is not the homologue of the
primitive streak. Tlie latter has potentially in it not only that but
a great deal more. It is in fact a mass of raw undifferentiated
material from which various structures are produced. This view also
makes it evident that as the |)rimitive streak is almost the first feature
visible in the development of the avian blastoderm, a great many
changes of palingenetic character observed in the chelonia n eg^ before
the establishment of what corresponds to the primitive streak, are
necessarily skipped over in Aves, which are therefore not very good
subjects in which to study the process of gastrulation. The removal
of the primitive streak to tlie centre of tlie blastoderm must also be
explained in the way I have indicated abovein the case of Chelonia.
Comparison of the reptilian ovum with the mammalian seems
easier. The facts given in this paper agree, with the exception of
some minor details, very closely with those communicated in \ an
Beneden's preliminary notice (No. 13). It seems to me that the
primitive streak and Heiîsen's knob together corresj^ond to the
primitive plate of Chelonia, and the " Kopffortsatz '" to the forward
growth of the primitive pJate. I can not, however, accept A an
Beneden's theory of '' Lecit()})hor " and "• Ijlasto])hor." Exactly what
I cannot accept lies in the emphasized words of the following qucjta-
tion: — " Wenn diese Auseinandersetzuno'en richtio* sind, "vvie ich es
glaube, so ist es klar dass das sogenannte zweiblätterige Stadium

ON THE PROCESS OP GASTRULATION IN CHELONIA. 261

der Säiigethiere der Gastrulution d. h. der Eiastiilpung, die man von
der Epibolie auseinanderhalten muss, vorangeht, und das s die ZAvei
Schichten respektiv dem Ektoderm und dem Entoderm
des Amphioxus nicht entsprechen. Dieser Schhiss geht
schön daraus hervor, dass nicht allein die Organe des Epiblastes,
sondern auch die Chorda und der ganze Mesohlast aus der äussern
Schicht sich bilden." According to my views, the epiljlast of the
Am ni Ota is liomologous with the epiljlast of Amphioxus. The
difficulty which keeps Van P>eneden from accepting this idea lies in this,
that not having for comparison the comparatively simple story of the
reptilian development he has reckoned as epiblast what corresponds to
the primitive plate of Reptilia. If he had recognised the structure
which, as 1 have sliown above, can not be said to belong to either layer
and then considered the lower layer as precociously developed hypoblast,
the conclusion would h:ive been inevitable that the outer layer corres-
ponds to the epil)last of Amphioxus . Keibel (Xo. 7) has also shown
to what contradiction Van Beneden's theory of the " Blastophor " and
" Lecitophor " leads. I think, however, I have now removed the
second objection of Keibel:—" Dazu kommt dann ferner, dass uns
Van r>eneden den Beweis dafür durchaus schuldig gebhei^en, dass nun
wirklich die untere Schicht des zweischichtigen Säugethierkeimes und
die Keimhöhle desselben mit der Bildung des definitiven Darms der
Säuo-er nichts zu thun hat." I think, the fact that the whole yolk-
sac with the subgerminal cavity within it does not form in Chelonia
any permanent part of the alimentary canal, makes it highly probable
that the same is also true of the homologous structure in Mammalia.
As I have more than once stated above, I accept Hubrecht's view of
precocious segregation. In many respects my views are very much
like his, but I do not think, he makes a clear distinction between the
Archenteron and the yolk-sac. Nor do I know from personal

262 K. MITSUKURI.

obsei'vation whether such a distinction is possible in Mammalia. I am
only inclined to think that, since the reptilian and mammalian eggs
are alike in so many points, what is tree in the former as regards the
development of the alimentary canal will in the main be found true also
in the latter. I can also find inChelonia nothing corresponding to
his " proto-chordal plate." As to whether there is such an annular zone
of hypoblast as he describes which gives rise to the mesoblast I wish to
express no opinion. That tlie '' FÎ umpfmesoblast " arises entirelv
within the embryonic shield from the materials derived from the
primitive knob I hope to have made at least probable in the preceding-
pages, but whether some temporary mesoblastic structures of the
embryo may not arise in Rep til ia from such an annular zone as he
describes, I am not in a position eitlier to stffirm or to denv.

ox THE PROCESS OF GASTRULATIOX IX CHELOXIA. 263

Postscript.

The foreo'oino- article was nearly finished in January of this year.
I made an extract of it in the early part of tliat month and sent it to
the Afiafoinisdier Anzeiijcr as a preliminary notice.* As I was giving
linal t(Ki('hes to tlie article I received from Dr. Ludwig Will an article
of his own entitled '' Die Anhuie der Keimblatt er heim (rech) " (Zool.
Jahrhilrlier ; Ahth. f. Anat. v. Oui., \ I Baml 1 Heft). As I men-
tion in a previous page, I was not under the circumstances able to
make full use of Dr. Will's paper, but inserted remarks on it mostly
in footnotes. The foreo'oino- article has since then ])een Ivin«- readv
for the press, but its publication was greatly delayed, owing to various
extraneous circumstances. Wlien it was at last to be put in the prin-
ter's hands, I received a second article by Dr. Will : '"'Die Anlage der
Keimhläfter hei der menoquiNi,schen Snmjifschildkröte" {ZooJ. JuhrJnlcher :
Ahth. f. Anat. u. Ont., V] Band, 3 u. 4 Heft). As it is too late to go
over my article again in the light of the facts brought out by Dr.
Will, I have decided to add here as a j)ostscript a few remarks on
Dr. Wills two papers, as well as on some other articles whicli have
appeared recently.

Will's observations on the two species, Platydacty 1 us face-
tan us, Sdireih. iiud Cistudo lutaria, Gesn. coincide throughout.
They, I am glad to see, agree also in many essential points with the
results I have brought out in this and pre\ious contributions. There
are however, several points on which we diifer and some of these, it
must l)e confessed, are by no means insignificant.

1 . According to Will, a stage in which a sickle is present precedes
the establishment of the primitive plate in both the species. Since receiv-

* Published in Anat. Anz., VIII Jahrg., No. 1^1 13.

264

K. MITSUKURI.

ing- Ins second article, I have again gone through the chelonian emhryos
in my possession in order to examine this point. Tn Chelonia
caonana, the two youngest embryos whicli I possess (i-eferred t(^ on p.
ioG) are not prol):iblv much older than that corresponding to Will's figs.
1 and 13 (II Art.) but neither in the sketches I liad made of surface views,
nor in the sections, was I able to detect any structure resembling the
sickle. In T r i o n }' x , I vva snot mcore successful . lUit in C 1 e m m y s
japonica, some eml^i-yos which 1 had taken out of the ovi(hict
siiowed a structure wln<'h on surface views looked very much like a
sickle.

Tlie annexed tigure (Fig. A.) represents one of these in which the
sickle is seen to extend to the sides more than in the others. This stage is
7nor(' ^d^'arl(vd than that in which Will figures a sickle, (Fig. 1, If

Art.) inasmuch as the
invagination cavity has
a I read V l)roken through
below. On cutting sec-
tions of this embryo, the
sickle was found to be
due to an ac(-umulation
;:,-: (^f the lower Layer cells

continuous with the pri-
mitive plate. (See Fig.
B.). The epiblast is
'^^ " shar})ly marked off from

1^ this mass, so that it can

: not be regarded as a part

of the primitive plate — at
ï'ig- A. least not in this stage.

Ventral View of a Clemmys Embryo taken from the Oviduct. A ffpv })eC(^mino' familiar

C5

ON THE PROCESS OF GASTRULATION IN CHELONIA.

265

Post

> Ant.

Edge of the Sickle.
Fig. B.
■ Fig. B. Posterior part of a. longitudinal section of the Embryo given in Fig. A.

with the îippearance of the sickle in this series of sections, I was able to
detect the same structure persisting in the sections of some older embryos
of CJemmys. There i.^ ;i great deal of variation in the degree of
development to which this structure attains in different individu;ds as
well as on the two sides of the same individual. It seems to disappear
entirely later. As the mesoblast develops afterwards quite independent-
ly of this, it is not what Will calls Kuplfer's sickle. For tlie present,
I think, it corresponds probably to the sickle (Koller's si(.'kle) which
Will describes in the earliest stao-e, althouu'h there are some features of
it which I do not yet quite ccmiprehend and which may fin;dly esta-
bUsh its ditference from Koller's sickle.

Aj'arr from the structure which I have described, I can detect
nothinii' comparable to Koller's sickle in mv materials,

2. Will makes out a sharp line of demarkation between the
ectoblast and entoblast at the edge of the primitive plate (1 Art. Figs.
43, 44, and others ; II Art. Figs, lo a. and />.). Since reading Will's
second article, I have again carefully gone over the sections of my
earliest stages, but J am unable to make (nit sucli a line at all. As
this line is figured in AVill's papers as pei-sisting to quite late stages,
I aui surprised that I do not see it at least in some of my sections, if
it really exists.

,'). Perhaps the most serifjus jxnnt of ditference in the observations
of Will and of myself is in regard to the extension of the invagination
cavity, before it breaks through below. In Cistudo, Will states that

2g6 E. MITSUKURI.

the invagination cavity becomes exactly co-extensive with the epi-
blastic embryonic shield, before that act takes phice (II Art. figs. 6fo,
7h, 8h). In Platy dactyl us it is said to be only slightly less. In
Chelonia caouana, which I have studied, the invagination cavity
breaks through, as I have stated in the foregoing article, when it is
quite small compared with the epiblastic shield. Will accounts for this
discrepancy by assuming that Cistudo and Platydactylus on
the one hiind and Chelonia on the other are really different in this
respect. (7/ Jrt., Nach sell rift. Also in a note " U. d. iTüstndation
r. Cistudo a. Chelonia,'' Anal. Anz., Ill J Jahnj., No. ltl/19).
In Tri onyx, I possess several embryos which are like Fig. 11). of
Contrib. I. or Fig. (3 of the foregoing article, so that I think I am
justified in concluding that Trionyx is like Chelonia in this
respect. In Clemmys there seems to be individual variations as
to- this point. For instance, if we comj^are Fig. A. in this postscript
with that given in Fig. 1 of my Contril). Ill, Ave find that in the
latter, the inviigination cavity must have advanced farther forwards,
nearer the anterior end of the embryonic shield than the former. iSo
that it is an actual fact that there are variations in different species
ov within tVie same species in the proportion of the invagination cavity
to the shield. For the j)resent, I am therefore willing to accept
Will's assumption as the correct explanation of the disagreement be-
tween his statements and mine. And yet I can not help having some
doubts lurking in my mind that his Figs. 6/>, 7/>, and <S/> (II Art.) are
expressions of something other than the breaking through of the in-
vagination cavity. That in Cistudo the invagination cavity be-
comes both in length and breadth exactly coextensive
with the embryonic shield — not one whit more or
less — seems to me very extraordinary. The figures Will copies
from Clarke do not certainly show the laterjil extension of the in-

ox THE PROCESS OF GASTliULATION IX CHELOXIA. 267

VîigiDîition cavity to be equal to tlie width of the shield. In this
connection we must renieniher jinother fact which Will Irinos out in
another place and which I belie\e myself able to corroborate, that
" die gesammte dorsal Lh-darmwand zur Bildung der ClKU'da und des
gastralen Mesoderms aufgelrraucht wird" (II Art. p. (il:^).* If the
invagination cavity becomes, as Will maintains, really coextensive
witii the embryonic shield, it follows from the above-mentioned fact
that the lower layer covering the entire ventral surface of the embryo-
nic shield is used up for the above-mentioned j)urpose and the gut-hypo-
blast (Darm-Entoblast) must come fnjm outside the shield. Ikit this
can not be reconciled with the fact whicli I have brought out in the
foregoing pa])er and of which there can not be any doubt, that the
gut-hypoblast comes from the cells derived from the primitive plate
and arises within the embryonic shield. It can not be urged
that there are actual differences in this respect between Cistudo
which Will has studied and C h el o nia which I have studied, for his
Fig. 1) (II Art.) is very much like my Figs. 6a or 7a in the fore-
going article, and shows, beyond a shadow of doubt, that in Cistudo as
well as Chelonia the gut-hypoblast arises within the embryonic shield.
These considerations force me to suspect that possibly there is no
great difterence in the actual facts between Cistudo and Chelonia
in this matter.

4. As I have stated in the foregoing article, I was unable to
detect in Chelonia any differentiation of the primary and secondary
endoderm such as Will and several others describe.

5. In one place Will does me injustice. On p. 587 (II Art.)
he says : '• Die Hedeutung dieses Flächenbildes, von dem ich in
Holzschnitt Fig. 7. (my Fig. '2, Contrib. III.) eine einfache Skizze

* I am iudeliteil to Dr. "W^ill for poiuting ont the inaccuracy of uiy exj^ression on this point
undor Heading- 4 of my proliuiinary notice f J Hr(^ Aiiz., VIII. Jahr(j., So. 12J13.).

268

K. MITöUKURl.

gebe, konnte von iinserm Autor (i.e. by Mitsukuri) nicht erkannt und
auch nicht interpretirt werden, weil demselben damals die ähnlichen
Oberflächen bilder vom Gecko noch niclit bekannt waren, die allein
dieses vereinzelt stehende BiJd deutbar machten. Wir erkennen in
die Skizze zwei in ihrem hintern Absclmitt nahezu parallel der Mittel-
linie verlautende Linien, welche vorn plötzlich stark divergiren. Ich
kann dieselben nur als die Insertionsgrenzen des gastralen Mesoderms
ansehen." That I was aware of the significance of my figure referred
to above is shown by the following words in my Contrib. Ill (p. 4G).
"This inward extension* of the gut-hypoblast is probably the
cause of the grooves converging posteriorly into the single median
chorda groove seen in tlie surface view Figs 2 and o(^" If these words
are read in connection with what precedes and follows, I think, it will
be plain that I had in my mind the significance of this figure to which
Will refers above. I am, however, willing to admit that Will has
made this point very clear and liis tig. E. (II Art. p. 586 or tig. 4, I
Art. p. 94) is certainly a very suggestive one.

6. I may |)erhaps lie allowed to make remark on a part of
Will's observation on C i s t u d o. In his second article (p. 542) he says :
" Während der Embryonalschild bisher noch vollkonmieu im Kiveau
der iibrio'en Keimscheibe lao-, tritt dieselbe auf diesem Entwicklnno-s-
(Stadium zuerst als deutliche, wohl umschriebene Erhebung von herz-
förmiger Gestalt aus der Keimscheibenoljertiäche hervor. Dement-
sj)recliend macht sich diese A\^ölbung an der Dotterseite (Fig. 'M).)
durch eine leichte Concavität bemerkbar." In another place (p. 568),
he is surprised that Mehnert's euibryos are not more vaulted or bulged
out. When Ishikawa and I first luidertook the study of Tri onyx, we
used to open the shell and try to cut the blastoderm out as is usuaJly

* Perhaps the words I used were not. entirely happy. If I had said the "inward uiove-
uient," it would have expressed uiy meaning more clearly.

ox THE PROCESS OF GASTRULATION IX CHELONIA. 269

done in taking ont cliick-embryos. At tliat time, we used to find
Tri onyx embryos vaulted dorsally, just as Will describes in the
({notation given above. Since adopting the method given in the
foregoing article of preserving embryos stretched in tlieir natural con-
dition, I have never found the shields vaulted in this manner at any
stage : they were always on a level with tlie rest of the blastoderm.

7. In a note entitled " On Mcsoblast Formation in Gecko,"
(Anat. A)iz.) No. 12 u. 13, 1893), 1 ventured to criticize Will's views
on the mesoblast formation of Reptiles. Will replies to my criticism
in the postscript to his second article, and :iIso in a note " Zur Frage
nach der Entstehung des gastralen Mesoderms bei Reptilien "' in the
Anafoinisclicr Anzeiijcr Xo. 20, 1803, wliich has just come to my hand.
I must refer the i-eader to Will's original pa])ers as well as to the above
note for his views. Suffice it to say here that AYill considers the
gastral mesoblast to be formed by a fold which arises in the outer wall
of the archenteron and grows towards the median line, thus cutting
otf tlie dorsal portion of the archenteron from its lower main portion.
The wall of the small dorsal portion thus cut otf is said to become the
mesoblast. This is put forth in (^p[)osition to tlie N'iew which was first
pro])oniided l)y Hertwig and which appears true to me, viz : that the
mesoblast is formed from two diverticula of the archenteron arising
directly on each side of the chorda.

Will considers that Fig. 23 of my Contrib. III. which shows a.
distinct diverticulum on each side of the chorda can not be held to
prove the " Divei-tikelbildung " as it comes from an old embryo which
has the chorda already cut otf in the middle dorsal region. In his own
words* : '" Hier sielit man thatsachlich rechts und links neben der
Chorda ein kurzes Divertikel, von dem die solide Mesoblastmasse
ausgeht, jedoch lässt sich an einem solchen i^ild aus dem Ende des

* The note above referred to. Anat. Anz. No. 20. p. 681.

270 ^- MITSUKUEI.

ganzen Processes natürlich nicht melir erschliessen, oh es sich nm
echte Divertikelbildiino- oder nm Unterwuclisung' von Seiten der
Urdarnifiilten handelt, oh das Divertikel das l^rimare nnd die solide
Mesoblastmasse das Secundäre ist, oder uintrekehrt." Now, the fact is
familiar to every embryologist that at a given stage of development a
structure, one part of which is already finished may show at another
])(3rtion of its length only the commencing phases of the process of fn-ma-
tion, so that one can see in one and the same specimen the whole ])rocess
from the beginning to the end. Such seems to me to be the case with
the mesoblast in the CI em my s embryo from which my figure 23 is
taken. The fact tliat the mesoblast formation is complete and the
chorda is cut off in the middle dorsal region, does not necessarily vitiate
what is seen in the head region : here the ])rocess of the mesoblast
formation is in a less advanced phase, and if a diverticulum is seen
there, it is highly probable that a diverticulum is a feature of the
mesoblast formation. That there is no such distinct diverticulum seen
earlier in the middle dorsal region is because the epi blast presses closely
down, and there is no space for the diverticula to curve upwards to
any large extent as in the head region. I think, I have sufficiently
demonstrated in my Contrib. Ill, that the diverticulum in the head
region corresponds to that part of the primitive hypoblast in the dorsal
region which Will calls the " Zwischenplatte," and that this 7nust
therefore be regarded as a shallow diverticulum.

Will also objects to my views on the following grounds : " P)ei
der Auffiissungder Zwischenplatte als ein gestrecktes Divertikel miisste
der solide Teil des gastralen Mesoderms (war. in Fig. 1 ]>.) nicht an
dem Rande der Z\vischen])latte inserirt, s(jndern aus der Mitte dei-
letzteren hervorgewuchert sein." (Anat. Anz. No. 20, l<Si)3. p. 6<S])
Again "Wäre die Zwisclieni)latte ein abgeflachtes Mesodermdivertikel,
•so musste aus iVir sowohl der somatische wie die splanchnische Mesobkist

ox THE PROCESS OF ÖASTRULATION IN CHELONTA. 2T1

licrvoro-eheii." (/^/*?V) The first of these ohjections occnrred to me,
while writino- mv (^ontril). III. If one examines tlie cross-sections of
Amphioxns ns o-iven, for instance, in Hatschek's Taf. IX. (Sfiidioi ti.
Entiv. (J. Jiiipliio.rus. Arh. a. d. Zool. Inst, Wien. Bd. TV. See also
Hertwig's Lehrlnich fio-. 72), we shall find that the mesoblnst ])ouch
sprenfls ventrnlly :tnd laterally not from what «corresponds to the
npex of the earlier diverticulum, but from its outer or lateral wall.
The same thinü' takes place in Reptiles, Although I did not express
this distinctly, it was present in my mind, as a reference to the middle
of p. 41 (Contrib. III.) will show. These two objections on the part
of Will are, I think, fully answered by this consideration.

Will again says : " Die Urdarmfalte würde bei der Mitsukuri'schen
Auffassung überhaupt belanglos für die Mesodermbildung und deshalb
unverständlich sein " (loc. cit. p. 681), I do not quite see the force
of this objection. A fold is ueeded to m.ark the outer limit <^f the
di\ erticulum,* and when tlie diverticulum is finally to be cut off from
the main portion of the archenteron, it takes place by this fold advan-
cino- towards the median chorda. I described this inw^ard moveiuent
of the fold! in my contrib. III. (pp. 42 and 4() ; also Figs. lG-17).
It is this last ])hase of tlie mesoblast formation which Will ein]>hasizes
above all others, and on which lie builds what he considers to be a new
theory of the mesoblast formation (I Art. p. 102). Even in his own
views tlie ]iart of the mesoblast which is formed by " Septenbildung "
is only a smidl ])roximal ])ortion near the chorda, for the part wgr in
his fig. 1 B. C. D. (Anat. Anz. No. 20, 1893) is according to himself
not formed by " Septenbildung" but proliferated from the archenteric

* In a sent(,'nce siuiilar to the above, in my note " On the Mesoljlast Formation in Gecko "
(Auat. Anz. Xo. 12-13, 1893) the word "mark" is by a most unfortunate oversight in proof-
reading misprinted " snack " — a mistake which makes my sentence Avell nigh incomprehensible.

t I admit that I used then the expression " gut-hypoblast " — instend of the word " fold "
whicli I ought to have adopted. But a reference to figs. 16-17 will show that a« a, matter of
fact I had observed a fold.

272 ^- ^nTSTiKURi.

wnlJ. This course of reasonînof reduces Will's views practically to the
same thing as mine as given in Cnntrib. IIT. with the exception of the
single ]ioint that I consider tlie *' Zwischenplatte " as a flattened
diverticulum, while he does not. I have already urged above the
reasons for my views, so that I will not again go into them. I
must refer the reader to it as well as to my C<^ntrib. III. Notwith-
standing that A¥ill says, I have fallen into a fundamental error in con-
founding " Septenbildung " and " Divertikelbildung," I still think,
I was not without reason, when I snid in my note (Anat. An:. No,
12 and 13, 1893. p. 4o4). that ■' * ->^ ::- whether the ])resence of the
fold is emphasized or the diverticulum is pointed ont as the essential
feature does not alter the facts of the case much. Will's objection to
Hertwig's theory may therefore be only an apparent one." The
difference between " Septenbildung " and " Divertikelbildung " which
Will points out is exactly like that lietween the process of budding and
of division. It is not possible to draw a hard and fast line in one
case as in the other.

Finally I would like to add that while Will and myself agree as to
the essential features of the reptilian development, the above discussion
shows that on nifiny minor points we must for the jn-esent " agree to
disagree," (as I lieard the late Prof. Balfour i-emark on a similar occa-
sion), until fresh observations bring out new facts and enable us to
settle these vexing points.

I have very recently received througli the kindness of the
author, Keibel's " Studien zur Entwicklungsgeschichte des Schweines."
(Morphologische Arbeiten. III). It would perhaps be going out of
my way too far to offer any extensive remarks on this article in-
teresting though it is to me. The foregoing paper shows that, like
himself, I divide the gastrulation into two phases, but these two

ox THE PROCESS OF GASTRULATIOX IN CHELONIA. 273

phases are different in his case and mine. My own views are (1) tliat
the cœnogenetic hypoblast is formed by precocious segregation, and
('2) that the definitive hypoblast is produced Ijy the formation of the
invagination cavity which gives rise to the definitive alimentary tract
as well as to the chorda and the gastral mesoblast. According to
Keibel, '* In der ersten dieser riastrulationsphasen wird l)ei den
Säugern das lùatoderm des Darmes und des Dottersacks gebildet, in
der zweiten Mesoderm und Chorda " (loc. eil, p. 10<S). That is, the
second I )hase which corresponds to the formatitjn of the invagination
cavity in Chelonia gives rise simply to the mesoblast and chorda and
has nothing' to do with the formation of the alimentary canal. On
the latter structure he says, quitting from an earlier work. '' * * * so
habe ich doch wohl festo-estellet, ' dass.' so saü'te ich damals. ' wir das
Homologon des Urdarmes unter der zweiten Schicht des zw ei blättrigen
Säuü'ethierkeimes zu suchen haben. Doch wurde demselben nicht die
ganze Höhle des Bläschens entsprechen, sondern nur ein ideeller
Spaltraum zwischen der unteren Keimschicht und dem Inhalt des
bläschenförmigen Keimes, welchen Inhalt ich dem Dotter homolo«iisi-
ren möchte. Die untere Keimschicht des zweiblättrio-en Säufyethier-
keimes entspricht aber nicht dem gesammten ürdarmepithel des
Amphioxus, sondern nur den Theil desselben, weiche z(un definitiven
Darm werden. * * *." ( j)p. 1 — '2) Lwotf* has also come to a
somewhat similai' conclusion. I would not like to be understood as
opposing this view in a dogmatic spirit. On tlie contrary, I think,
there are several points which seem to favour such an interpretation.
P^or instance, when the invagination cavity breaks open below in
Reptiles, tlie dorsal wall, which alone remains, gives rise only to the
chorda and the gastral mesoblast, as Will points out, and if we looked

* Basilius Lwoff : — Ü. d. Kciuiljliittorliilduu"- hei don Wirbeltieren. Biologischem Central-
blatt, XIII. Band. No. :i Sf 3.

274

K. MITSUKURI.

simply lit such u figure as Fig. la of the foregoing article, we might
naturally come to the conclusion that the invagination ca\'ity gives
rise only to the gastral mesoblast and the chorda, and has nothing to do
with the formation of the definitive alimentary tract. But we should
always remember îi fact which I hope to have proved conclusively in
the foregcji ng article that t he 1 j o 1 1 o m o f t h e i n v a g i n a t i o n c a v i t y
has been removed. When the bottom is still present, we may

Fig. c.

Diagrauiiiiatic CruBS-section of the
Invagination-Canity ill Chelonia. ^^S- ^■

Diagrammatic Cross-section of the
Archenteron of Amphioxus. {■■iftcj-
Göttc, from Born. Ergehu. d. Anat. u.
Entw. I Bd. p. 49a.)

represent the cross-section of the invagination cavity as in Fig. C. The
dark portion is the part that is removed. AVhen we comjjare it witli
a homologous section of Amphioxus (Fig, D.) we are struck with their
similarity and I think, we are justified in concluding that the invagi-
nation cavity of the reptilian ovum is homologous with the archenteron
of Amphioxus, and has potentially present in it not only the chorda
and the mesoblast but also the definitive alimentary tract. The re-
m<jval of the bottom, or that part whicli represents the definitive hypo-
blast, must in any case be regarded as a secondary process, and can
not stand in the way of homologizing the two structures. These
considerations, together with the reasons which I have brought out in
the foreofoini»' article, incline me more t(jwards those views which \ have
set forth in tlie preceding pages than to those of Keibel and of Fwotf.

ON THE PROCESS OF GASTRULATION IN CHELONIA. 275

On another point I would like to say a few words. Referring to
the structure in Mammalia which \^an Beneden homologizes with the
yolk-plug in Amphibia, Keibel says : — ^' Der Dotterpfropf hat bei den
Ami)hibien keine grosse morj)hologische Bedeutung, er ist ein Ent-
wicklungs h i n d e r n i s s, er hat kein Funkti • )n. Warum sollte gerade
diese Bildung so zäh festgehalten werden, während doch so vieles
Andere, das von ungleich grösserer Bedeutung ist, undeutlich wird
und verschwindet." As I have insisted on the presence of the yolk-
plug in Iveptilia, ever since Ishikawa and I tirst discovered it. I may
perhtips give my own view on Keiljel's objection. I have in a
previous ]jage tried to explain the chang.^ of shape in the blastopore
in Reptilia as repeating that process by wliich the lower half of the
Amphibian egg becomes enclosed by tlie epiblast. Taking this in
connection with the presence of the yolk-plug, I think that wh.at is
inherited is not simply the yolk-plug but the whole process of the
epiboiic invagination which is gone through in the region of the
primitive plate. This is certainly important enough to persist for a
long time.

Tokyo, Oct. 1893.

276 K. MITSUKURI.

Contributions to the Embryology of Reptilia.

I. K. MiTsuKURi and C. Ishikawa : — üu the Formation of the Germinal Layers in
Chelouia. Quart. Jour. Micr. !Sc. Vol. 27. Also Jour. Sc. Coll. Tokyo, Vol. I.
]>t. S.
II. K. MiTsuKURi : — On the Fœtal Membranes of Chelouia. Jour. Sc. Coll. Tokyo.
Vol. IV. pt. 1.
III. K. MiTsuKURi : — Farther Studies on the Formation of the Germinal Layers in
Chelonia. Jour. Sc. Coll. Vol. V. pt. 1.

List of References.

No. 1. Agassiz, L. and Clark: — Contributions to the Natural History of North

America. Vol. II, Pt. 3, Embryology of the Turtle.
No. 2. Duval, M. : — Etudes histologiques et morphologiques sur les Annexes des

Embryons d'Oiseau. Jour, de l'Anat. et de la Physiol., A'.V. 1884.
No. 3. DuvAL, M. : — De la Formation du Blastoderme dans l'Oeuf d'Oiseau.

Ami. d. Sei. nat. 6 ser. T. 18.
No. 4. Flejiming, W. : — Ueber Theilung und Kernformen bei Leucocyten und

ueber deren Attractionssphären. Arch. f. Mikro. Aiiat. Bd. 37.
No. o. Hubrecht, A. A. W. : — The Development of the Germinal Layers of Sorcx

vulgaris. Quart. Jour. Micros. Sei. Vol. XXXI.
No. 0. KuPFFER, C. :— Die Gastrulation an der meroblastischen Eiern der

Wirbelthiere und die Bedeutung des Primitivstreifs. Arch. j. Aiiat. u.

Fhys., Anat. Abth. Jhnj. 1882.
No. 7. Keibel, f. : — Zur Entwicklungsgeschichte der Chorda bei iSäugern (Meer-

schweincheu u. Kaninchen) Arch. f. Anat. v. Fhys., Aiiat. Abth, 1889.
No, 8. Mehnert, E. : — Gastrulation und Keimblätterbildung der Emys lutaria

taurica. Morph. Arbeiten lierausy. von Dr. (iustar Schwalbe, o Bd. 1 Heft,

1892.
No. 9. Rabl. C. : — Theorie des Mesoderms. Morph. Jahrb., 15 Bd. 1889.
No. 10. KoBiNsoN, A. and Assueton, E. : — The Formation and Fate of the Primitive

Streak, with Observations on the Archenteron and Germinal Layers of Rana

temporaria. Quart. Jour. Micros. Sei. Vol. XXXII.

ON THE PROCESS OF GASTEÜLATION IN CHELONIA. 277

No. 11, Sarasin, C. F. : — Keifung und Farclmug der Eeptilieueier. Aib. a. d.

Zool. — Zoot. Inst, zu Würzhimj 6. Bd. 1883.
No. 12. Strahl, H. : — Die Dottersackswand und der Parablastder Eidechse. Zeit.

f. Wiss. Zool. Bd. 45.
No. 18. Van Beneden, E. : — Kemarks made in tlie discussion following the reading

of Kabl's paper on the Mesoderm.' Anat. Anz. 1888 p. 675 — Demonstra-
tion of the plates illustrating ihis investigations on the Formation of the

Germinal Layers, the Chorda-Canal and Gastrulation in Mammalia.

Anat. Anz. 1888, pp. 709-714.
No. 14. ViRCHOw, H. : — Das Dotterorgan der Wirbelthiere I. Zeit./. Wi-ss. Zuol. 53

Bd. Siipplewtmt. II. Arch.f. Mikros. Anat. 40 Bd. 1 Heft. 1892.
No. 15, Wenckebach, K. F. : — Der Gastrulationsprozess bei Lacerta agilis. Anat.

Anz. 1891 Nos. 2. ii. 3.
No. 1(5. Weluon, W. F. R. : — -Note on the Early Development of Lacerta muralis.

Quart. Jour. Micros. Sei. Vol. XXIII. 1883.
No. 17. Whitman, C. 0. : — A Eare Form of the Blastoderm of the Chick and its

Bearings on the Question of the Formation of tlie Vertebrate Embryo.

Quart. Jour. Micros. Sei. Vol. XXIII 1883.
No, 18. Will, L. :• — -Entwicklungsgeschichte der Gecko. Biol. Centralbl. X Bd,

Nos. 19 & 20, 1890.
No. 19. Will, L. : — Zur Kenntniss der Schildkröten-Gastrula. Biol. Centralbl. XII

Bd., No. 6, 1892.
No. 20. ZiEGLER, H. E. : — Die biologische Bedeutung der amitotischen (directcn)

Kerntheilung in Thierreich. Biol. Centralbl. II Bd., No. 12 & 13.
No. 21. Will, L. : — Beiträge zur Entwicklungsgeschichte der Reptilien. Die

Anlage deri Keimblätter beim Gecko (Platydactylus facetanus, Schreib.)

Zoöl. Jahrbücher, i Abth. f. Anat. v. Ont., VI. Bd. I Heft.

PLATE VI.

Plate VI.

Fig. 1. — Dorsal view of an embryo of Chelonia caouana a few hours after its deposi-
tion. Zeiss aaX2 (Bia)

Fig. 1«. — Ventral view of the same. aaX2

Fig. 2. — Dorsal veiw of an embryo of Chelonia caouana \\ days after its deposition.
The lateral parts of the embryonic shield are not represented. Zeiss
aaX2 (Dla)

Fir,. 2(;. — Ventral view of the same. aaX2

Fig. 8. — Dorsal view of an embryo of Chelonia canuana 1^ days after its deposition.
Only a small part around the primitive plate is represented. aax2 (C2)

Fig. 3«. — Ventral view of the same. aax2

Fig, 3. /»/.v.— Dorsal view of an embryo of Chelonia caouana about 2 days old. Only
a small part around the primitive plate is represented. aaX2 (O?)

Fig. 'Sa, bis. — Ventral view of the same. aax2

Fig. 4. — ^Dorsal view of an embryo of Chelu)iia caouana 1\ days after its deposition.
Only a small part around the primitive plate is represented. aa=2. (B4a)

Fig. 4(/. — Ventral view of the same. aaX2.

Jour. Sc. Coll. Vol. VI. PI. VI.

Fig. 4

Fig. 4a

Fig.

Fig. 2

Fig. 8a h\%.

r

Fig. la

K M,l,„l,uri J- A-, .Va,„„),„„„, ,w.

Fig. 2a

Fig. 3a

Fig. 3

Fig. 3 bis.

Ulh. i Imp. Tlu .Seishibuviha

PLATE VIL

Plate VU.

Fig. 5. — Dorsal view of an embryo of Chelonia caniKnin 3| days after its deposition.

aaX'i (B5a)

Fi(j. 5a. — Ventral view of the same. aaX2
Fig. 6, — Dorsal view of an embryo of (Jhelouia camiaiia o^ days after its deposition.

aaX2 (B7a)

Fig. 6t/. — Ventral view of the same. aaXü
Fig. 7. — Dorsal view of an embryo of Chelonia anxtana o\ days after its deposition.

aaX2 (B7h)

Fig. la. — Ventral view of the same, aa x'2
Fig. b. — Dorsal view of an embryo of Chelonia caoitana 1\ days after its deposition.

aaX2 (B9a)

Fig. 8r/. — Ventral view of the same. aaX2.

Jour. Se. Coll. Vol. VI. PI. Vil

K. Milsiikiiri * K. Nasuhar; dti

Lith. tV linp. The SeisiUbiimha

PLATE VIII.

Plate VIII.

Fig. 9. — Longitudiiuil section near the median line of an embryo of tlie same lot
and stage as that represented in Figs. 1 & la. CCx2 (Bib, 41. 2c. last-2)

Fig. 10. — Part of a longitudinal section of the same series as Fig. 9. From more
lateral parts. DD x 4 (Bib, long. 2, 51., last-2)

Fig. 11 a k h. — Two merocytes from the same series as Fig. 9. DDx5

fvi\U f :T" l"»g- 2, 51. last— 2\
(.I^l'^lh. long. 2.51. 7.S ;

Fig. 12 a k h. — Cells of the same kind as those represented in the middle stratum

of Fig. 10. DDX4 (Bib.l-S-^^.-^-^)

Fig. 13. — Longitudinal section near the median line of the embryo represented in

Figs. 2 & 2a. C0X2 (D la, 31. 2c. 5s.)

Fig. 14. — Longitudinal section near the median line of the embryo represented in

Figs. 3 & 3a. CCx2 (C2. long, i, il. 2c. 7s.)

Fig. 15.— Longitudinal section near the median line of an embryo 1 day older than

Figs. 1 k la and 1 day younger than Fig. 4 k 4a. CCX2 (B3. loug. 1. 41. last)
Fig. 10. — Longitudinal section near the median line of the embryo represented in

Fig. 8 his. and da. his. CCx2 (O? long. 1. 21. 2c. 2s.)

Fi(i. 17.— Longitudinal section near the median line of an embryo slightly younger

than Figs. G & 6«. CCx2 (B6. 31. lOs.)

Fig. 18. — Longitudinal section near the median line of the embryo represented in

Figs. 7 and 76. BBx2 (B7b. 31. 2c. is.)

m?:

■-;"i'H!P'.j'.^;;;;'fP!;n;'niiKi'',«<>miiin«iiiiiiiMi|ii|(r((f[^(fj« y..,^ ,/

Jour. Se. Coll. Vol. VI. PI. VIII.

,^._pi;!;^jrnii.aWrr;'Ä'^'y''

■•;-:• '.t;^;.;^**;,.

■•:.",rr^',',ÏÏ. ...

!iWl«iî"fc

f;!.'ff.'iWf»[iiigÄ;i«i

:#%.•

Note on the Eyes of Cardium Muticum Beevel
K. Kishinouye, Riéakushi.

Zoologist to tlie Department of Agrienltnre and Commerce.

Witli Plate IX.

The eyes of the genus Card! um have been studied by many
authors; but. so far as I know, no one has yet found such conspicuous
eyes as I describe below in the molhiscs of this genus. The following
note may therefore ])rove interesting to investigators of the molluscan
eye.

The mnntle-edges, both right and left, of Cardium muticum are
beset witli dnrk brow^n. almost black pigment. They unite at the
posterior end of the shell and form a triangular pigmented area, sur-
rounding the siphon;« 1 openings. Over this area the right and left
valves of the shell do not meet closely but leave a rathei" wide slit. In
this triangular, pigmented area w^e find a, great many tentacles, ar-
ranged in many irregular rows round the siphonal openings.

The tentacles are of \'arious sizes : those on the peripliery of the
pigmented area are larger and longer; tliose towards the siphonal
openings, shortei- and more sleîider till at the margin of these
they are reduced to mere fringes (fig. 1). Clenerally speaking the
larger and longer tentacles about 100 in number bear the eyes. They
are bent away from the siphonal openings, and eacli of them has a
longitudinal band of black pigment on the siphonal side, 7. e. on the

280 K. KISHINOUYE.

side exposed to the li^ht.

The eye can easily: he found without a microscope as a black spot
on the si|)honal side of the tip of a tentacle, opposite to the position of
the eye of Cnniium efhile, which is '' on the shell side of the mantle." *
Along the upper, posterior f side of the eye, the epithelium is raised
into a triangular screen (figs. '2. 6, 7, 9, .s.). At one time I thought
that this screen is the tip of the tentacle, but a closer examination
showed that it is raised secondarily, probably io ])rotect the eye, and
that the true tip is occupied by the eye.

In the eye i^f Cnrdium muticum we find all the essential ])arts
of an eye. Its structure resembles on the whole that of the eyes of
Fecten. Spondyiius^ and Cardivm edide ; but ditlers in S(jme essential
points, since it is ditHcult to find the liomology of some constituent parts.

The cornea (fig. 2. <;.) consists of thin ])a veinent cells, while that
of other molluscs consistSigenerally of columnar cells. The general
epithelium ber-omes gradually thin as it ap])roaches the lens and forms
the thin cornea over it. The corneal cells are colourless, transparent,
and polygonal in outline. The convexity of the cornea is great, its
external surface being almost hemispherical.

The lens (fig. 2, /.) is large and consists of a great number of
cells. Instead of the usual biconvex, flat form the lens of Cardium
imdicwn is ovoid. Its longer axis is parallel to the o])tic axis and
its broader end is directed below. It is more or less constricted at
the middle' ]])art. The cells composing the lens are large, colourless
and compressed in the direction of the optic axis, the degree of
compression being greater nearer the cornea. In the median lon-

* Patten— Eyes of Molluscs and Arthropods. Mittheil, aus der Zool. Station zu Neapel.
188G.

t For the sake of convenience, the word upper is used to designate the distal end of the
tentacle, and the word anterior that side of the tentacle turned towards the sijihonal openings.
'I'he words lower and posterior naturally indicate the opposites of the above worda.

NOTE ON THE EYES OF CAKUrUM MUTIOUM Ueev^. 281

rritudiiial section, we lind that the cells of the lens are arrauf^ed ior the
most part in two lonuitiKlinal eoJumu.s. The cells are nucleated,
havins' nuclei near the external surface of the lens.

The n'luia (üg. 2, rcl.) is in contact with, and directly below, the
lens. It is very sim])le in structure, consisting of colourless, columnar
cells arranged in one layer. These cells have rods (tig. 2. ro.)
which are not clearly fjund in the eye of Cdrdiimi cdule (Patten,*
J:îutschli **). The rods are directed away ironi the retino})horae and are
separated from them by a jtseudoniembrane. They seem to be hollow, as
they appear as rings in cross section of some well ])reserved specimens.
They are longest at the optic axis and gradually diminish in length as
the distance from the optic axis increases. They are stained homogene-
ously and rather deeply. As regards the retinophorse, the nuclei are
deeply stained, while the protoplasm is very faintly stained. I could
not find the ganglionic-cell layer, although it is stated to be ])resent in
the eyes oï Pect en and Canlimn edulc (Patten, Biitschli).

lîelow the retina and continuous with it at the circinnference,
there is a layer of fiat cells (tig. 2. c//.). These cells, are small and thin
at the juncture with the retina, so that it is difticult to tind out tliis
connection in well develo])ed eyes. 1 shall name tliis layer as the
choroid, as it seems to ht. liomolog<jus in position to tiie pigment epithe-
lium of the choroid of the \ertebrates. If the ta])etum (argentea of
Patten) of Peclcn and Curdimii e.dide is cellulai' in orign, as it is stated
to l)e, tVie cjioroid oï Cardimii )tnificinii is j)robably homologous with it.

Below the chorijid there is a membranous layei". It is the iapctuvi
(fig. 2. t.). It consists of many thin, shining layers, stained deeply
and homogeneously by colouring solutions. 1 cannot tind any sign of

* Patten— /oc. cit.

** Biitschli — Notiz zur Morphologie des .Vllgo^^ der Muscholu. (Zoologisches Jahresljoricht
für 1886).

232 K. KISIIINOUYE.

celliiliir atriictiire, tboiiu'h iiccordiu^- t(j J 'at ten (he like-uamed part of
Pecten and Canliinii edah is waid to liave been ])roduc('d from cell
Invers. The rapetuni covers entirely the lower surface of the clioroid,
except at the spots through wliich tlie branches oi (he <jptic nerve
enter. If the taj)etuni of I'ccten and Cordimil cdule is cellular in oriu'in.
there is in these iiujUtiscs iio organ homologous with the ta])etnni of
Cardium midicum. Moreover, the tapetuni of I'ccten and Cardiinii cdtdc
is said to be found below the layer of retinophora\ while that oï Car-
dram muticum is beh^w the choroid.

Lastly there conies a layer of pigmented cells (tig. z^, p.). 'Lliis in-
vests all the external surface of the above described parts of the eye
below the cornea, leaving only a small round area o^er the upper
hemispherical portion of the lens or the pupil only. The pigmented
layer as a whole is cu|)-shaped or rather urn-shaped. It consists of
fiat,. polygonal cells arranged in one layer. The cells at the neck of the
urn are smaller and thicker than those elsewhere. ihe ]>it>'ment is
black and serves to absorb rays of light which fall obliquely upon the
retina. In Pecten the greater part of this function is fulfilled by the
iris, and its red pigment layer, probably homologous with the pigment
layer of Ccwdimn niuticmii, aljsorbs rays of light fr(3m the lower
side alone. As the eyes of Canliuni innticum are destitute of the
iris, the pigment layer is well developed.

The eyes are innervated from the viscero-parietal ganglia. A nerve
(tig. 2, n.) runs through the central axis of each tentacle. It is
divided into two branches where it touches tlie eye. One branch
(fig. 2, /i " ) passes through the ])igment layer near the optic axis and
spreads between the choroid and the tapetum, while the other branch
(fig. 2, n ' ) passes through the pigment layer at tlie level of the retina
from the shell-side and seems to innervate the retina.

As the numerous eyes of an adult individual are sometimes found

NOTE OX THE EYES OF CARDIUM MUTICUM Ikeve. 2SS

ill dirtereiit stages of dei'dojninni. we ••an study their formation
from their sections. It seems to me that the eyes of CardiiDii develoj)
ill two wavs. Althoiiuii we must regard botli as more or less al)ljrevial-
ed, vet we can ilistiuguish one as //(/' iiioir ahhnciaicd proo'ss from the
(jther ///<• l('ss aJihrcnated. If an investigali«j]i into their development
could be made in immature spe<-imens. mcjre satisfactory resiiUs would
doubtless be obtained, unless they are as in the young oï Veclcii in wliicli
Patten was disa])j)oint€d to lind any abbreviated development of tlie
eye. I shall describe the less ahhrerlaled process of development tirst.

Tiiere first forms on the tip of a tentacle (tig. .'>) an invagination
the mouth of wliich then closes (tig. 4). Thus a solid mass of cells is
produced, with a slight conca\ity at the place where the mouth of the
invaoination at tirst opened, and this is turned towards the siphonal
opening. it consists of cells in many irregular rows, while tiie general
epithelium consists of columnar cells in one row. A ner\e running
aloDii' the central axis of the tentacle touclies the lower surface of the

Ci

(•ell mass and there divides anteroposteriorly into two branches.

In the upjter jtart of the cell mass, a s])herical j)ortion becomes
differentiated and sei)aratcd from the rest by a basement membrane
(tig. 5). This spherical portion is the rudiment of the lens, which is
therefore epiblastic in origin, not mesoblastic as in the case of the eye
of Pecten, according to Patten. The cells constituting the rudimentary
lens are large and some of them have nucleoli. At this stage,
although the lens is differentiated and separate from the surrounding
cell mass, it is still imbedded in ihe epiblastic thickening.

In the next stage (fig. 6) the epiblastic thickening surrounding the
lens is obser\ed to be separated into two parts — an up|X3r and a lower.
The upper i)art is continuous with the general e])ithe!ium and consists
of fiat cells in (jne row. It is the cornea. The lower part is cup-
shaped and is n«3t connected with the general e})ithelium. It consists

^)^4 ^- k:t«hinouye.

of cells in about two rows. It is the retiuji. Un the upper side of
the base of the eye there is foi'iiied ;t Jarge hollow. This makes the
eye stand out and at the snine time pushes it towards the siphonal
opening. Thus the hollow di^■ides the tip of a tentîjcle into two — a
prominent eye and a somewhnt triMUguhn- screen behind it. The
hollow corresponds in position -with the original mouth of the in-
vagination for the eye. Pigment is produced in mesoblast cells
surrounding the retinjil portion. These pigmented cells form the pig-
ment layer. A lumen is secondarily produced between the lens and
the retina.

'I'he lens grows l)y the multiplication of cells in the direction of
the optic axis und assumes the shape of an ellipsoid, and consequently
the lumen between the lens and the retina distjppears (fig. 7). The
peculiar arrangement of cells in the lens begins in this stnge.

• Late in de\elopment, the retinal portion is divided into two layers,
the retina ])rr)per and the choroid. Tliesc two layers are continu-
ous with each otlier at the cii'cnmfereiK'e. Soon after the separation
of the choroid from the retina, the tapetum is formed below the
choroitl. probalily by the secretion of the cells wliieVi constitute the
latter. I cannot corroborate the view tliat the tapetum is formed of
modified cell layers, tor even in these early stages 1 cannot find any
thing of a cellular nature in it. Afterwards rods are ])r(^duced from
the retinal cells.

T]ie wore ahhrcnatrd process of development (see ]>. 2^1^) is
as follows :

At the top of a tentacle the epithelium becomes thickened and
forms a little knob within (tig. 8). The little knob is next cut off from
the epithelium (fig. 9). In this stage a hollow is produced behind tin;
little knob and thus the triangular screen (fig. 9. s.) is formed. The
little knob cut off is spherical in form jind consists of a lew, large

XOTE Oy THE EYES OF OARDTUM MUTICUM Reeve. 285

cells. Tt enlarges by the division of the ceils and assumes an ellijî-
soidnl form (tig. 10). Cells forming the lower part of the ellipsoidal
mass become small by division. These smaller cells are not clearly
distinguished from larger cells, as there are many cells of infermediate
size. J^ater, liowever, the smaller cells are separated from the larger
ones and form the retinal portion, while the larger become the lens.
The later stages «^f dcveloj)ment are quite like those in the first process.

In tlie two jyrocess of development, botli the lens and tlie retina
are produced from the epiblast. The process of their formation is
|)robably abbreviated. Originally thev were perha))s produced l)y two
separate invaginations as in the case of the Vertebrata, one invagina-
tion for the retina and one for the lens. Tlie invagination for tlu;
I'etina must ha\e l>een the first to Ix^ closed and cut off from the
epithelium ; it formed a hollow sphere, the upper wall of which
became the retina and the lower wall the choroid. The invagination
ior the lens was next formed and cut off in its turn from the epithe-
liuu). and had its lumen obliterated.

The eyes of CardiiDii differ from those of Vecleir chiefly by the
presence of the choroid between thci retina and the tapetum and by the
mesoblastic origin of the pigment layer ; but I am inclined to think
that i n Pecten the choroid disappeared after secreting the tapetum, and
that the red pigment layer is mesoblastic in origin, and not directly
in connection with the retina.

* It is hard far mo to accept Patten's ohsorvation that tlie lens of Pecten is mesoblastic in
oiigin.

PLATE IX.

Explanation of Figures.

List of Abbreviations.

c.

Cornea.

ch.

Choroid

1.

Lens.

n, 11^ n'-. Optic nerve and its l^ranclies.
p. l^ig'uient layer.

ret. Metina.

ro. Rod.

s. Triangular screen.

t. TaiJetum

i'^i'^- 1. Iiilialoiit siplioii ofCV//7//i/»/ )iiiiticiiiii, iniigmûed. One longitudinal half uf

it is c\it away.
Fig. 'I. Median sagittal section of a tally developed eye (constructed from two

sections).
Fios. 3-7, S(3nii-diagrammatic representations to illustrate the less abbreviated

process of the development of the eye (sagittal sections).
J^'i'ts. b-10. Semi-diagrammatic representations to illustrate the more abbreviated

process of the development of the eye (longitudinal sections).

Jour. Sc. Coll. Vol. VI PI. IX.

/o

/C.Xô.^hùiou,ye /Vf f.

Note on the Cœlomic Cavity of the Spider.

by
K. Kishinouye, Riè^kushi.

Zoologist to the Department of Agriculture and Commerce.

With Plate X.

The species investigated belong to the genera, Lycom and
Agaleiui.

My olyject in undertaking tliis investigation was to ascertain
whetlier the stercoral pocket is mesoderinic in origin as I described in
my former paper* or ectodermic and a part of the proctoda^um, as is
generally believed.

When the embryo has reached the stage in which there are nine
segments in the ventral plate, i.e., seven segments between the cephalic
and caudal lobes, the mesoderm is divided into as many segments,
while a ]3air of the cœlomic cavities appears almost simultaneously in
each of the first five segments, exclusive of the cephalic lobe (Fig. 1).**
As at this stage the segment of the cheliceroe is not yet formed,
the cœlomic cavities now formed belong to the segments of the pedi-
pal]ii and the four ambiTlatory appendages. The rudiments of appen-

* On the Development of Araneina. This Journal, vol. IV.
** As the segmentation of the ventral plate seems to take place simultaneously with the
division of the mesoderm into somites, I will hereafter for the sake of brevity, simply describe
the formation of a segment and will not mention every time the formation of the correspond-
ngc mesodermic somite.

28g K. KISHINOU^E

dao-es make their appearance in these five segments as small round
knobs. The cœlomic cavities of the same segments are just below the
evaginations for the appendages.

In the next stage the segment of the chelicerie is cut off from the
cephalic lobe, and three new segments are added between the last seg-
ment and the caudal lobe (Fig. 2). By this time the cephalothoracic
appendages have become much elongated, and the mesoderm has spread
into them. The cœlomic cavity extends to the distal end of each
appendage ; in fact the larger part of the cavity is now found within
the appendage. It spreads a little at the base of tlie appendage, so
that three portions may be distinguished in it in a cross section of
the embryo (Fig. 5) as Schimkewitsch* observed — one portion in
the appendage, a second extending a little in the dorsal direction,
and the third extending a little ventrally. The last two are the
horns of the basal enlargement. In the abdominal region, the pro-
visional appendages are not yet formed ; but the mesodermic somites
develop rapidly, and in each of the first and the second abdominal
segments a pair of cœlomic cavities is produced. Thus seven pairs of
the cœlomic cavities are now found.

Subsequently two more new segments are added between the last
formed segment and the caudal lobe. A pair of the cœlomic cavities
is formed in this stage in each of the following segments: the cephalic
lobe, the segment of the chelicerae, and the third to the seventh
abdominal segments (Fig. 4). These seven newly formed pairs of
cielomic cavities together with the seven pairs already existing make
in all fourteen pairs, the caudal lobe alone being now devoid of any.
In tlie cephalic lobe the mesoderm is not divided into two lateral
parts, therefore the two cœlomic cavities, right and left, are separated

* Étude sur le Uéveloppement des Araigaées. Arch, de Biologi . Touie XT. 1887.

NOTE OX THE CŒLOMIC CAVITY OF THE SPIDER. 289

by the inedijin partition of mesodermic ceJJs ; while the right nnd loft
cœlomic cavities of the other segments are separated by the yolk
(Fig8. 3, 5).

The cephalothoracic appendages, with the co'lomic cavities in
them, elongate very much, and bend towards the ventral median line
(Fig. 5). The mesodermic somite and also the co^lomic cavities of
the first abdominal segment have developed little since their formation.
Korschelt and Heider state in their text book that in this seoinent a
pair of the provisional appendages is formed as in the four succeeding
seginents. Though a pair of slight elevations is found in this segment,
tliey are very much lower than the ])rovisional appendages of the suc-
ceeding segments, and moreover they are chiefly of the ectodermic
thickening (Fig. 4). I am therefore inclined not to call them pro-
visional appendages. In each of the second to fifth abdominal seg-
ments, a pair of provisional appendages appears. They are short
elevations of the ectoderm, into which the cœlomic cavity enters, as in
the case of the cephalothoracic appendages. The C(Elomic cavities of
these segments develop in the dorsal direction : hence in cross
sections of the embryo these abdominal somites differ from the thoracic
segments in having a shorter branch of the cœlom in the apj^endao-e,
and a longer one towards the dorsal side (Fig. ,5). The ectodermic
cells covering the mesoderm somites are always high and columnar
in shape, and are easily distinguished from the cells of other parts.

Soon afterwards one more new segment is cut off from the
caudal lobe, and in its mesodermic moieties a pair of cœlomic cavities
appears. At this stage, the ventral plate attains the greatest antero-
posterior extension round the egg, so that the cephalic and the caudal
lobes almost touch each other (Figs, o, 6). The caudal lobe is raised
a little above the general surface of the egg. I stated erroneously in
my first paper that the mesoderm of the caudal lobe is split to form

290 ^^- KISHIXOUYE.

an unpaired cavity in thi.s atage. In the abdomen the growth of the
mesodermic .somites, except thîit of the first abdominal segment, is
enormous, extending rapidly towards the dorsal median line. Thus
in the abdomen, the dorsal portion of the cœlomic cavities develops
rapidly, while their ventral portion as well as the portion Avhich enters
into the appendage remains only slightly developed. In the céphalo-
thorax, on the contrary, the portion of the cœlomic cavities which
enters into the appendage develops rapidly, while their ventral and
dorsal portions remain undeveloped.

The reversion of the embryo now begins, and when the process
advances a little, the two nerve cords and the appendages of both
sides of the ^'entral j^late, «n* the two lateral divisions of the germinal
band, begin to separate from each other. This lateral extension of the
ventral plate, together with the rapid growth (jf the dorsal portion of
the mesodermic somites in the abdomen, causes the dorsum of the
embryo to elongate longitudinally. At this stage the mesoderm of
the céphalothorax shows no noteworthy changes. The C(L'lomic
cavities of the cephalic lolje develop anteriorly, or towards the dorsal
region (Tigs. 7, 8). The first abdominal segment begins to de-
generate ; its mesodermic moieties and ganglia may however be seen
with some ditliculty. In the second to fifth abdominal segments the
mesodermic moieties develop greatly towards the dorsal median line,
so that they nearly meet each other at that line (Fig. 7). The
ectoderm covering these mesodermic somites is elevated a little, and
forms the s(j- called tergal portion of the abdominal segment. Locy*
illustrates his paper with a figure (PI. II. Fig. D) in which a pair of
terga may be seen before the second a1jd(jminal segment ; Ijut he gives
no description of them. I myself am unable to find them, lie says

* Uu the Development of Agalena Dtevia. Bull. Mus. Comp. Zoul., vol. Xll. 1886.

NOTE <")N THE CtELOMIC CAVITY OF THE SPIDER. 291

also that tit firwt " the only dor.sal element« developed were the live
pah'8 belonging to the abdominal somite« («econd to «ixtli), but during
this stage the dorsal elements of the limb-bearing somites begin a more
rapid growth." But according to my observation this is not the case,
the dorsal elements of the limb-bearing somites remain undeveloped as
in the pre\ious stage.

Between two consecutive terga there is a furrow. This furrow
was described by Schimkewitsch* as ha\ ing no relation ^^ ith the meso-
dermic somites. He says that it is due to the niesodermic somites
fusing together, before they develop dorsally to cover the dorsal sur-
face. He says moreover that the number of these furrows never
correspcjnds with that of the somites. My observations as given above
do not corroborate these statements.

The last three abdominal segments (sixth to eighth) gradually
deii'cnerate and their cœlomic cavities seem to fuse tooether into one
pair. The pair of the cd'lomic cavities thus formed by fusion is pushed
into the protuberance of the tail as the process of reversion proceeds
(Fig. 7). I find tliat fusion of the cœlomic cavities does Mot take
place before this stage though Schimkewitsch says that it does in the
céphalothorax and in the alxlomen and I also erroneously stated that it
occurs in the thorax before tliis stage.

A cavity is produced in the mesoderm of the tail lobe. It is
iüi])aired (Figs. 7, (S). The unpaired cavity tlius made cannot be con-
ceived otherwise than as a homologue of the ccelomic cavity. Though
the cavity is certainly not formed by ;in invagination, I thought that
the cells in the tail lobe might be produced by the proliferation of the
ectoderm. But I found that the cells enclosing the unpaired cavity
are the remnant of the mesoderm cells which gave rise to the meso-

* loc. cit.

292 K- KISHINOUYE.

dermic somite.s of many preceding segments and that they are entirely
separate from the ectoderm. Pi-evious authors who have studied the
development of the spider, overlooked this cavity in the mesoderm of
the caudal lobe, and observing the stnge at which the unpaired cavity
communicates with the proctodaBum consider the former as a portion
of the latter. If the cells enclosing the unpaired cavity are ectodermic
in origin, the numerous mesoderm cells in the caudal lobe must disap-
pear all at once, as there are no cells in the lobe except those surround-
ing the last fused pair of cœlomic cavities. But the disappearance of
many cells at once is cjuite impossible.

As the process of reversion proceeds still further, each half of
the pair of the f.'celomic cavities in the cephalic lobe is divided into two
portions — one at the side of the stomodaeum (Figs. 9, 10, 1 ceci, h),
the other below the anterior border of the semicircular groove of the
brain (Figs. 9, 10, i cœl. a). The former disappears soon afterwards
but the latter elongates towards the dorsal median line (Fig. 11), and
the mesodermic walls of the cavities of two sides meeting nt the median
line fuse together, leaving, however, a canal between them. This canal
is the aorta.

The portion of the egg which is not covered l)y the ventral plate
or the abdominal terga is characterised by the absence of the meso-
derm cells and by the presence of the secondary endoderm cells or the
fat cells instead. The secondary endoderm cells are directly under
the ectoderm. Most of them are enclosed between the walls of the
mesodermic moieties of the abdominal somites and becojne the blood
corpuscles.

The cœlomic cavities in the segments of the chelicer£e and the
pedipalpi degenerate and disappear. The greater portion of the cœlo-
mic cavities of the four ambulatory appendages degenerates, the meso-
derm cells forming their wall becoming gradually changed into muscles.

NOTE OX THE CŒLOMIC CAVITY OF THE SPIDER. £93

Their proximal and outer portions remain distinct at the base of each
leg (Fig. 9). The cœlomic cavity of the first ambulatory appendao-e
communicates with the exterior by means of a duct which is produced
by an ectodermic invagination. The first abdominal segment disap-
pears entirely.

The mesodermic moieties of the second to fifth abdominal se»--

o

ments, or the segQients bearing the provisional appendages, meet
one another at the dorsal median line and form there the wall of the
heart. The wall of the cd'lomic cavity of the second abdominal seg-
ment meets that of the cephalic lobe (Figs. 9, 10). The forma-
tion of tlie dorsal circulatory system, in which some thoracic somites
do not take part, resembles greatly that of Limulus.* As a lateral slit
or ostium is made where two consecutive somites meet, the number
of the slits in the adult heart shows approximately the number of the
segments which took part in the formation of the heart.

The sixth to eighth abdominal somites are entirely degenerated,
their mesoderm cells are disintegrated and fill the caudal lobe at the
sides of its unpaire 1 cœlomic cavity (Figs. 9, 10). The latter becomes
wide, and over its posterior end, the ectoderm is slightly invaginated
(Fig. 9). The invagination is the rudiment of the proctodeum, so
that the unpaired cavit}^ of the caudal lobe is produced independently
of the proctodeum.

In the next stage in which the embryo assumes the ventral
flexure and the constriction between the céphalothorax and the abdo-
men appears, the cœlomic cavities undergo great changes. In the
céphalothorax, they all disappear, except the small portions at the
outer bases of the first to third ambulatory legs. These remnants
fuse together and form the coxal gland (Fig. 12). The lumen or the

* Kishiuouye — On the Development of Limulus longispina. This Journal, vol, V.

294

K. KISHINOUYE.

cœlomic cavity of tlie gland is so small at this stage that the gland
seems almost solid. In the ahdomen also, all the C(Blomic r-avities
disappear except the unpaired one in the caudal lobe, which, inexpli-
cable as it may seem, remains as the stercoral pocket, as was described
and figured in my former paper. At this stage the mesoderm cells .at
the intersegmental portions grow inward into the yolk and form the
dissepiments, these, being specially well developed between the second
and the third, the third and the fourth, the fifth and the sixth
abdominal segments (Fig. 12).

Those portions of the embryo which were destitute of the meso-
derm in the stage of Fig. Î) now dwindk^, according as they are
encroached on either liy the ^-entrai jilate (^r by the aljdominal terga in
their growth: in the céphalothorax, the cephalic lobe is bent towards
the dorsum and its lateral margins fnse with tliose of the ventral plate
of the thorax, Avhile in the ventral jwrtion of both the céphalothorax
and the abdomen, the two lateral cords of the nervous system with
the underlying mesoderm meet each other at the ventral median line.

PLATE X.

Explanation of Figures,

Reference Letters.

ahd.

xrg.

abdominal segment.

jiedi}!.

pedipnlpi.

br.

brain.

proct.

proctodjenin.

ch.

cheli'eree.

.<?('«(. f/r.

semicircular groove.

c.l.

caudal lobe.

St. p.

stercoral pocket.

cœl.

coelomic cavities.

.^toni.

stomodfenm.

cox.

!/'■

coxal gland.

Via. 1. Side view of an embryo at tlie stage in wliicli the first rufliuients of append-
ages liave appeared.

Fig. 2. Side view of an embryo at the stage in which all the cephalothoracic
appendages have appeared.

Fio. H. Dorsal view of an embryo at the stage of the maximum ventral fiexnre.

Fio. 4. Sagittal section of an embryo at the stage of Fig. 3.

Fig. 5. Cross section of an embryo at the stage of Fig. 3.

Fig. (!. A portion of the median sagittal section of an embryo at the stage of
Fig. 3.

Fig. 7. Oblique side view of an embryo at the stage of reversion.

Fig. 8. Oblique sagittal section of an embryo at the stage of Fig. 7.

Fig. 9. Side view of an embryo at the stage in which the process of reversion lias
greatly advanced.

Fig. 10. Oblique sagittal (about median) section of an embryo at the stage of
Fig. 9.

Fig. 11. Frontal section of the cephalic region of an embryo at the stage of Fig. 9.

Fi(i. 12. Side view of an embryo at the end of the process of reversion.

Jour. Sc. Coll. Vol. VI PI. X.

pediß

abd. seq

-10 coel.
n coel.

15 coel.^^st. p: ^^^^ '' "''^- 1 ''oei: a.

proct .

K. Ki^hhionye del.

Lit h. d' Imp. The Tokio-Seishihumka.

studies of Reproductive Elements.

II. Noctiluca miliaris, Sur. ; its Division
and Spore-formation.'"'

By

C. Ishikawa, Ph. D., Rigakushi, Rigakufiakushi.

Pi-ofessoi- of Zoology, Agiicnltnral College, Imperial University.

With Plates XI-XIV.

A. DESCRIPTIVE PART.

a. Division.

The individuals of the nenn« jSToctilncn which are preparing to
divide can easily be distinguished from the others by their spherical
form, caused by the absenœ of tlie peristome and of teeth, and by the
diminislied size of the tenta<"le (Figs. 1 and 2) ; but the Stahorgan is
always to be seen as a narrow line of cytoplasm, extending backwards
from the nucleus and marking the line of division. The tentacle ap-
pears to be drawn into the body of the animal, as Cienkoivski/ (6,
p. 50) first observed, and not thrown off, as Robin ( 6, p. 1064) is

* The materials upon which the following* observations Avere carried out were in part collect-
ed by myself from different parts of the sea coast near Tokyo, for instance, in Numadsu, Zushi,
Tateyama-in-Boshiu, etc, and in part from the Misaki Marine Biolo<,àcal Station of the Science
College of the University, where I worked for a fortnij^ht during- the summer of 1890, for a
week during- the winter of 1892-93 and again for a week at the end of August this year. For
all these opportunities and for the free use of the library of the Zoological Laboratory of the
Science College, I express my warmest thanks to the authorities of the College, and especially
to Professor Mitsukuri.

298 C. ISHIKAWA.

inclined to think. My figures (1 and 2) show the stages of its with-
drawal.

The central protoplasm, which in ordinary conditions takes a
more or less elongated form, then becomes concentrated into a star-
shaped mass around the nucleus. Close to the nucleus, part of it
differentiates itself from the rest by coarser granulation. It is usually
spherical in shape (Figs. 1 and 2), but sometimes elliptical (Fig. 31.).
This cytoplasmic mass represents the kinetic centre of division, as will
be later shown, and corresponds to the sphère attractive of van Beneden,
or Arclioplasma of Boveri. In many cases this archoplasmic mass is
formed before the complete disappearance of the peristome, teeth,
etc., as is shown in Fig. 31. The relative position of the archoplasm
and the nucleus does not seem to be always the same j but both
bodies lie in the direction of the Stahorgan, as will be seen in Figs. 1,
2, and 31. In Figs. 1 and 31 the archoplasm lies behind the nu-
cleus, whereas in Figs. 2 and 3 it lies in front of it. In living
animals the nucleus at this stage appears as a transparent vesicle, and
the archoplasm is very distinct, owing to coarse granulation.

The archoplasm now stretches itself in a direction more or less
in the vertical plane passing through the Stahorgan, and forms a
relatively large spindle (Fig. 3.). It undoubtedly corresponds to the
central spindle of Hermann (20, p. 580). This spindle gradually
elongates and assumes a position longitudinal to the line of division,
and divides the nucleus (Fig. 4.). The central mass becomes at the
same time separated into two portions, each concentrating around its
archoplasm. A narrow line of cytoplasm now appears also in front
of the central mass in the direction of the Stahorgan. The spindle
jjarts at the middle, dividing the nucleus into two daughter nuclei.
These (the daughter nuclei) at first lie in the line of division, but
soon change their positions and come to lie right and left of it

STUDIES OF REPRODUCTIVE ELEMENTS: IL ^99

(Figs. 5-0). Which of the two nuclei goes into the one half and
which into the other, is not easy to ascertain, as a narrow line of
cytoplasm, as above stated, now stretches also in front of the nucleus
nearly in the same manner as does the Staborgan behind, so that the
fore and hind ends are difficult to distinguish. But in all ascertained
cases, the front nucleus o-oes into the rio^ht side and the hinder one
into the left. All the while the archoplasm remains close to the
nucleus. The line of division gradually extends both ways towards
opposite surfaces of the body, and the animal assumes the well-known
biscuit-shape. Meanwhile, the two protoplasmic masses with their
respective nucleus and archoplasm shift their positions and come to lie
on each .side of the line of division, nearly opposite to each other (Fig.
9) ; so that division of the animal now appears to take place in the
longitudinal plane. The relative position of the nucleus and the
archoplasm does not seem to remain the same during the earlier stages
of the division, (Figs. 5-9), but at later stages the former lies farther
away from the dividing line (Figs. 10-12).

The time during which all these changes take place, from the
stage shown in Fig. 3 to the stage of Fig. 9, varies greatly in
different individuals ; but from observations upon fourteen animals
kept in a moist chamber at the Misaki station, during the latter
part of August 1893, the time required varies from six to eight
hours.

The division of the body proceeds, until only a small connecting
bridge of cytoplasm is left (Figs. 10 and 11). During this stage, tenta-
cles usually grow on both halves, which swim together like the
Siamese twins. Such a couple, with rather short tentacles, is shown
in Fig. 11. That shown in Fig. 12 has tentacles nmch more developed.
This figure is drawn from a living specimen, so that the nuclei (//)
appear quite transparent, while both the archoplasms (a) are seen as

300

C. ISHIKAWA.

dark granular mas.ses close to the nuclei. The protoplasmic connec-
tion (/>. c) is reduced to a narrow bridge.

The changes which take place in the nucleus as well as details
ot the nuclear division will be spoken of in connection wi(h the pro-
cess of spore-fonnation.

/). Spore-forniatiun.

S[)ore-formation is always preceded by tlie concentration of the
central plasm into a rather small area, and liy the disappetu-ance of
all other structures, such as the tentacle, flagellum, teeth, etc. The
individual ap]'e:u's, therefore, as about to divide, except in not possess-
inf^' mouth and StahorijaH. Its central plasm is also raised a little
above the general surface of the body, and forms a low knob-like ele-
vation, as has been observed both by Cienhjirskii ( 89, p. 54) and,
by Hohin (6, p. 10()9). Just as in division, we find here a spherical
concentration of the granular cytoplasm about the nucleus, which is also
quite transparent in living specimens. Thus, Fig. 1 o, which is drawn
from an individual in this stage, shows the nucleus (/<) as a spherical
vesicle, while the archoplasm («) appears as a granular mass. The
division of the nucleus is also preceded here by that of the archo-
plasm, and we thus obtain a stage, represented in Tig. 14, which
exactly corresponds to the stage of di\'ision represented by Fig. 4.
In what direction the spindle lies in this case, it is not, however, pos-
sible to tell, as the external signs of the bilateral nature of the animal
can not all be recognised. The division proceeds still further, till
the stage is reached represented in Fig. 15 or 1(), the latter of which
is drawn from a living specimen. We sometimes meet with a sort of
partial segmentation of the entire body at this stage, the plane of
division cutting the connecting axis of the two nuclei, as has been
observed by Gienkowshj (7, p. 134). Fig. 17 represents a stage with
four nuclei, on the side of each of which is seen a. large archoplasmic

STUDIES OF REPRODUCTIVE ELEMENTS: II. ^Q\

spiiidle, sliowiiig thus their further division. The cytoplasm sur-
rounding each of the nuclei and archoplusnis is connected with the
others hy a fine network, like the protoplasm in the segmentation of
Leptodora eggs obserxed l)y W'ci.sfnanii. and by myself ( 37, Taf. II,
Fig. t^O). Fig. 1<S gi\es a side view of a four-nucleated stage, in
wliich the nucleus on the right side is already divided into two
daughter nuclei, while that on the left is in the spindle form. Fig. 19
represents a stage immediately succeeding this. Here, in this in-
dividual, is seen a pai'tial line of division of the entire body, marking
its surface into two hemisjjheres. A further stage is represented Ijy
Fig. '20, where we see six nuclei preparing to divide. Of these two
belong to one half and four to the other, showing thus that the
division of the buds does not occ.-ur quite regularly even at this early
stage. The first line of division, although now very laint, is still
clearly to be recognised. In the living condition the body of the
Noctiluca had been imperfectly divided into two portions at the
part where the nuclei lie. This is shown more distinctly in Fig. 21,
where ten nuclei are to be found. Of these six belong to one half
and four belong to the other. The partial line of division of the body
is also ])lainly to be seen in this specimen. Fig. 22 represents a
further stage of budding, viewed laterally. All the nuclei visible are
represented in this ligure, and their number is fifty-one, showing
again the irregularity of division. A further stage of sporulation is
seen in Figs. 2o-30. Figs. 23-26 are drawn from sections of nearly
ripe spores, while those represented by Figs. 27-30 are from prepara-
tions of entire spores, drawn under different magnifying powers. In
Figs. 25, 2G (the spore on the right hand side of the figure), and
27-30, the spore cells are quite separated from each other, and they
have all got a long flagellum springing out from the basal third of
the cell, where the archoplasm lies.

502

C. ISHIKAWA.

Nuclear Division.

In a short j)a]>er ^ sent to the Nahirforscli enden Gesellschaft zit
Freiburg im Breisgau, I treated at some length of the structure of the nu-
cleus in the resting stage, as well as of the peculiar manner in which
the division of the Noctiluca-nucleus takes place. As the present
account covers the same ground, some of the facts described in that
paper are necessarily repeated in this one. Nevertheless I do not
hesitate to go again over the same subject here, for two reasons,
namely : (a) many new facts have become known, and the nature of
some of the d(3ul)tfnl phenomena been ascertained since then ; (h) the
papers appeal to t^^•o ditferent sets of readers.

1. Xuclens in resting stage: — The living nucleus appears quite
transparent and homogeneous, as all other observers of Noctiluca
have long ago described. It is covered by a rather thick membrane,
whose contour is often seen as double. When treated with reagents,
such as acetic acid, osmic acid, chromic acid, etc., a number of chroma-
tic bodies can e;isily be distinguished in it. These elements appear as
strino-s of deeply staining bodies which often take an S -form. Each
strino- consists, in well prepared specimens, of a luunber of disc-shaped
microsomes arranged one after the other like the chains of mammalian
red blood-corpuscles. Tliese strings lie either singly or, more often,
two or four united together. In the paper named above, I sup-
posed the single string to be primary, and the compounds of two
or four to be secondary, probably produced by division of the single
string. In accordance with this supposition, I designated the
number of chromosomes as ten. In many cases this number seems
to hold true, as Figs. o3, o4, 40-47 show ; but in other cases, such

1 This pajjer was written for the l'\'xii<ch7-iffen couiuieuioratiug Prof. Weisuiann's 60th l.irth-
day which occurs on the I7th January, 1894.

STUDIES OF REPRODUCTIVE ELEMENTS: II. 3()3

as Figs. 35 or 36, the number of chromosomes counts greatly above
this, and I am not, at present, able to tell exactly their num-
ber.

2. Fropluisis : — Figs. 31, 33, and 34 are taken from preparations
of nuclei which are just preparing to divide. In Fig. 31 the chromo-
somes are not very distinctly to be seen, owing to the deep staining
with hematoxylin. We see, however, on the left hand side a
curved double line undoubtedly representing the segmentation of one
of the chromosomes. Tliis is very beautifully seen in preparations
represented by Figs. 33 and 34, both of which are stained with a
combination of acid-fuchsin and methylen-blue. In hoth. nuclei the
chromosomes are seen to be composed of a double row of minute
microsomes, the number of which in a single chromosome is not
with certainty to be determined, since the entire length of a chromo-
some is very rarely to be made out. In a few cases, however, I count-
ed them to be ten or twelve in a single row, as wdll be seen in the V-
shaped chromosome, lying a little to the left, in the nucleus represented
by Fig. 33. A nearly equal number will be counted, though not so
clearly, in one of the two chromosomes at the right hand of the same
nucleus. Similar states of chromosomes are seen in Figs. 44-1:7,
drawn from preparations of s])ore-forming individuals in about the
same stage as that represented by Fig. 18. In Fig. 44 the chromo-
somes, except a few, do not seem to lie in any definite order, but to be
scattered more or less irreo-ularlv in the nucleus: but in Fig-s. 33, 34 and
45, they lie more or less parallel with each other, their long axis being in
the direction of the archoplasm found just external to the nucleus.
In Fig. 46 the above position of the chromosomes is not so clearly
expressed as in the figures just mentioned, although the archoplasm can
be distinctly seen on one side of the nucleus. This, however, is only
an optical illusion caused by the position of the archoplasm which does

304 C. ISHIKAWA.

not lie in the plane of the paper, but a little beneath it. 15y focussing
the tube, however, we can distinctly observe the chromosomes as
radiating from the pole where the archoplasm lies. A close examina-
tion of the chromosomes, represented l)y Figs. 44-47, of the spore-
forming individuals, and those. Figs. 33 and 34, of tlie nuclei just
liefore division, seems to show that there is some marked difference
between the two. While tlie cliromosomes of the nuclei of the divid-
ing individuîds are represented by a double row of microsomes, those
of the nuclei of the spore-forming individuals appear to consist of
four rows (comp. Figs. 47 and oo). This difference is undoubtedly
due to the division and the spore-formation. /// divi^iion ilic nucleus
lias to dividr onlij once, and lience tlie cliromostwies require onlij
once to divide, while in the spore-formation dirisions of tlie nuclei t(d;e
place rapidlii one <fter tlie other, and two dirisions tedcc place iduioM
simultaneouslij. The w^ay in which two divisions take place simultane-
ously can be made out from Figs. 44 and 47, where some of the
chromosomes are seen at their ends. The microsomes liecome thicker
at their periphery, and form a row of ]'ings. E:ich (-hromatin-i'ing or,
perhaps more correctly, the microsome-ring becomes thickened at
four ])laces and the interspaces between these thickenings break uj)
and thus form four daughter microsomes. Soinctimes the thickening
does not take place simultaneously at four places nnd the ring does
not break \\\) into four at once, as will be seen in a ring at \\\^ right
hand side of Fig. 47, where it is thickened only at one point. An-
other ring at the left hand side is broken at one place, and appeiu's
C-shaped. A ring represented on the i-ight hand side of the nucleus
Fio-, 44, shows a verv interestino; staiiv. This rino- consists of two
thick curves connected together by narrow cementing substance — the
Z/////i-thread. The ends of these curves are again thickened, showing
that the division of the ring into four daughter microsomes does

STUDIES OF REPRODUCTIVE ELEMENTS : IT. 3Q5

not in reality take place simultaneously, but by a succession of two
divisions rapidly following one after the other.

3. Spindle stage : — In my paper for Prof. Weismann's birth-day
celebration I have given a figure of a nucleus and an archo])Jasm in
the latter of which could be observed a (;lear space next the nucleus.
This space was in apjmrent communication with the interior of the
nucleus. Since then I have been enabled to find a number of fibres
in this s)»:ice (Fig. 40), but the existence of any communication be-
tween tliese fibres and the interior of the nucleus can not with
certainty be made out. The archoplasm becomes elongated and foims
a very Itu'ge spindle. This corresponds undoubtedly with the central
spindle of HeniKuni, as stated above, but will be called the arcik^-
PLASMIC SPINDLE, in distinction from other spindles îdi-eady de-
scribed in karyokinetic divisions of other animals and whose origin is
still under discussion; while the fibres of this spindle will be
called the central eibres, for reasons stated below. At first the
archoplasmic spindle lies tangentially on the surface of the nucleus,
but soon assumes a curved figure, depressing at the same time the
nuclear wall. In consequence of tliis the nucleus takes a broad
C-shaped figin-e, in the concavity of which the archoplasmic spindle
lies. A notion of these stages may be gathered from Figs. 3, 21, and
24. The chromosomes wliich, at the stages of Figs. 45, 4G, 4(S, &c.,
are seen radiating from a siiigle point, become with the division
of the archoplasm separated into two masses, one attached to each end
of the spindle. The archoplasmic spindle elongates and the nucleus
assumes the form represented by Figs. 20, 35, 36, 49, and 52.
Fig. 35, prepared from specimens killed with Flemminijs fluid and
stained with Böhmers hematoxylin, represents the division of tlie
nucleus in a dividing individual. The nucleus assumes more or
less the shape of a dumbbell, and the archoplasms at both ends

306

C. TSHIKAWA.

of it are irregular in outline. The chromosomes have now
distinctly divided into two portions radinting from the pole of the
nucleus wliere the arclioplasms lie. Eîich chromosome consisting of
a number of microsomes arranged like bends, is bent ii|)on itseli',
as is so usiimI iu this stage of karyokinesis in other ;inim;ils. The
number of tliese chi-omosomes, ns stated above, is very difficult to
ascertain. The most interesting phenomenon in this tigure (o/)) is
the spindle fibres, of which there are two kinds ; those, whicli I call
CENTRAL FiBHES (Figs. 35, 8(j, 37, 4>>, cf.), ^y'"o imbedded in the
concavity found in the wall of the nucleus, and standing in no direct
relation with the nucleoplasm ; and those (Figs. 35, 3^, 37, 4(S,
rf.') which are seen at the side of tlie central fibres, riuming from
the centre of an archoplasm to the bent ends of the cliromosomes,
and which I will call the radiating fibres. Where they join the
nuclear wall, the latter appears (piite indistinct, but whether they
pass through the wall of the nucleus at these points and are dii-ectly
attached to tlie chromosomes can not be made out. In Fig. 37,
which represents a section of a dividing nucleus at about the same
stage as Figs. 35, 36, and 37, a homogeneous mass of nucleoplasm
is found ;it the poles of the nucleus, external to the bent ends of \}\q
chromosomes. In this mass of nucleoplasm, wln'ch is easily distin-
guished from the more granular archoplasm lying outside, ar(i seen
many parallel fibres running in the same direction as the radiating
fibres of the archoplasm represented in Figs. 35, 30, and 4(S, but
unfoitunatelv not to be seen in the series of sections represented by
Fig. 37. We shall come to this again in the part devoted to general
considerations, as further discussion of the subject is beyond the region
of observation.

WIhmi the division has proceeded as far as Fig. 35 or 3(5, the
median poi'tion of the archoplasmic spindle is swollen up a little, as

«TUDIEiS OF REPRODUCTIVE ELEMENTS: II. ^07

u'iJI l)e seen in tliese two finure.s :iiicl also in tiii'. 4. I'liis juirt of tlie
spindle is left behind after the coni[)]ete separation of the nuclei iu
the form of a small diagonal figure (Fig. 39 x). What becomes
of this figure is not ascertained. Fig. oö is drawn from preparations
of a specimen killed with picro-acetic acid and coloured with fuchsin-
methylen 1)1 ue. The archoplasm, and tlie central as well as the
radiatiny fibres are seen as in Fig. 3G, but some of the chromosomes
are here more or less swollen up, owing perha})S to the action of the
acetic acid. Fig. 35 A. is drawn from the same nucleus as that of
Fig. 35 but nt a level lower than that of the archoplasmic spin-
dle. It should be mentioned by the way that the spindle lies always
nearer the surface of tlie cell than to the nucleus. In both these
figures will be seen a number of narrow lines [)assing between the free
ends of the separating chromosomes, and qidte distinct from the
central fibres. These are the Verhimlumjsfäden of German authors,
whose origin is undoubtedly to be found in the ///u'/i- thread of the
nucleus.

fhe difference we (observed, in the chromosomes in the case of
prophasis, betw^eeii the nuclei of the dividing individuals and those of
the spore-forming ones, is also here discernible. Fig. 48 represents an
individual in which the first two buds are just dividing; these are
seen more magnified in Fig. 48 A and B. In both of them the
nucleus i.s much elongated and dumbbell-shaped, with a large
archoplasm at each end. represented as usual by rough granulations
staining very deeply with aniline dyes. Scattered within the
swollen ends of the nuclei are seen many chromosomes, each of
which is distinctly discernible as a double row of minute micro-
.somes ; whereas in the nuclei of dividing individuals at al)out the
same stage, the chromosomes ccmsist of only a single row (compare
this figure with Figs. 35 and 36). In Fig. 48, A. in which the ob-

308

C. ISHIKAWA.

ject is looked at a little from one side, the central filtres (c. f.)
lying in the furrow of the nucleus, and the radiating fibres (r. /.)
proceeding from tlie upper archoplasm, are distinctly visible. It
thus seems that segment ati(^n of the chromosomes in spore-formation
takes place before the previous division of the nucleus is yet at its
end. This is also to be looked upon certainh^ as the result of the
rapid progress of the bud-formation.

4. Anaphasis : — The details of anaphasis can only be observed
in individuals which divide, since in those which foru) sjjores the
nucleus divides successively without the intervention of any resting
stages, as mentioned above; and at the end of the spore-formation the
diminished size of the nucleus makes it very ditficult to observe the
individual chromosomes with any accuracy. I will therefore only
give my obser\ations of the changes, at this stage, in the nuclein
subs<tance of the dividing individuals. Fig. 38 represents one of tiie
nuclei soon after division. The parallel arrangement of the chromo-
somes, their relative position in regard to tlie archoplasm («), and the
geijeral shape of the nucleus are all dis[)layed very pklinly. The
positions of some of the chromosomes at its right lower corner, how-
ever, become more or less irregular in their relations. Fig. 40, which
represents one of the nuclei nearly at the stage of Fig. 7, shows that
the chromosomes have lost their parallel arrangement. The micro-
somes are, however, still very small, represented only ])y small dark-
coloured dots. In Figs. 39-43 the microsomes have become somewhat
swollen up, assuming nearly the characters of those in the resting
nuclei. The microsomes in Fig. 42, however, appear to be a little too
large, owing perhaps to the action of the acetic acid with which the
specimen was treated.

5. The fate of the arcliuplasin : — The archoplasm, as we have
stated, comes to be seen first at the stage a little before the division or the

SlUinrES OF REPRODUCTIVE ELEMENTS: II.

309

spore-fon nation. During the whole process of the division of the
nucleus it remains always closely attached to the outside of the latter,
and in the dividing individual is still to be recognised just before
the separation of tlie individuals, as represented by Fig. 12, and even
for some time after the complete separation, as one of my ])repara-
tions of the animal twelve liours after lias shoTvn very plainly.
After the formation of tlie peristome, teeth, etc. it becomes indistinct.
In the spore-buds the archoplasm remains till the final stage when
the spore is ready to swim olf iVoni the body of the mother animal,
as Figs. '2o-'M), 19-51 show very plainly. In nearly ripe spores,
Figs. 27-30, the arclioplasm lies on that side of the Ixxly which
is attached to the mother cell, and is therefore at the liead end
•)f the spore when it detaches itself from the bod\^ of the mother.
This is more clearly to be seen from sections of nearly ripe spores,
represented by Figs. 2o-2G. I sought after this body in ripe spores
treated wnth acetic-acid -methyl -green, but could not get any satis-
lactory knowledge of its existence. From the position of the archo-
plasm, and from the manner of division in the foregoing stages,
it is clear tliat the free end of the spore corresponds to the equatorial
part of the archoplasmic spindle, while the end attached to the surface
of the mother animal is the pole of the spindle.

'... 0. The cenfrosouie : — In many cases in the centre of the archo-
plasm is seen a small round body, which often colours very deeply
with aniline dyes, especially with such colouring matters as rubin,
iodine-green-methylen-blue, etc., but also with luvmatoxylin, when
this is used in C(jnjunction wûth iron ammonium alum, as given by
Heidenliciu (15, ]). 118-119). In a short note on the conjugation of
Xoctiluca (26, ]>. 3) I stated that at the poles of the two C(nijugat-
ing nuclei, deeply-staining round bodies are found, similar to those
seen in the centre of the archoplasm, and suggested that such bodies

3io

à. ISHIKAWA.

are the centrosouie.s so commonly observed ut the poles of the spindle
of the dividing nuclei of other animals. The ])ody now in question
is seen in Figs. 35, 31), 45, 49, 50, 51, (c). In Fig. 39 it is very
plainly visible in the centre of the upper archoplasm as a small round
body surrounded by a clear space. In the Ljwer archoplasm of the
same figure, it can not be seen so well, owing to its position beneath
the nucleus, but is discernible by focussing the tiil)e. In Fig, 44 two
centrosomes are seen close together in the centre of the archoplasm.
The nucleus to which this archoplasm belongs is, as stated above, in
the condition just before division, \vhile in Fig. 45 it is again seen as
a single body. It is also seen in four cells in Fig. 49, as small dark-
staining dots. This is also the case in Fig. 50, and in the arclio-
plasm on the left hand side of Fig. 52. It is not visible in other
figures, wliile in the archoplasm represented by Fig. 42 and more
clearly by Fig. 48, there is seen a number of small bodies in place
of the centrosomes. These bodies are not always quite spherical like
the ordinary centrosomes, many of them being more or less elong-
ated, and often presenting curved rods like those, described by Her-
niiuin (20, J). 585), in the spermatocyte of Proteu s ; tliey are perhaps to
be looked upon as a group of centrosomes like those, described by M.
Heidenhein (16, p. 54-()8), in the lymphocytes of rabbits. In my
[)revious paper I have given a case where tlie pr<jbable origin of the
centrosome from the nucleus is shown. Since then I have met with
no case similar to that one, but in another, I saw, at the side of a nu-
cleus whi(,*h had just divided, a sm:dl deeply stained body (c), close to
the nuclear wall, (Fig. 38) probably representing the last stage of the
disappearance of the centrosome within the nucleus.

STUDIES OF "REPRODUOTIVE ELEMENTS: IL 3JJ

B. GEN^ERAL CONSIDERATIONS. .

a. Division.

Cienknirshj, in liis second pnper on Noctiluca (8, p. 55), speaks
of its reprodncfion by means of division and says : — '' Sie wnrde am
voMständig'sten von BrifiJifirdI \evfo\(j!;t, dessen Beobachtiini^eii ieli so
wolil an norniîil u'eb-mten wie auch an einofekno-eJten Individuen;
besfätigen konnte." C^icnkoirskij thus believes with Bri(j]\twcll (5, p. 10)
that the division can take place not only in n<n"nial individuals but
also in animals in which the tentacle, mouth, etc. h:ive become lost.
So fîir as I have observed. I can only confirm Jiohin.s observations
(6, p. 1064) that the division takes place only in individiinis in which
the tentacles have dis-ippeared. The plane of division is longitudinal
to the body of the animal, as Hohin nlso states ; but the first
spindle lies in the direction of the longitudinal ])lane, cutting the
animal, therefore, transverseh'. The resulting nuclei separate from
each other ;ind tiually come to lie opposite, as is ;ilso ])ointed out by
Rohin. In the dividing individual, the plane of division of the nucleus
apj^ears to lie in the longitudinal axis of the body from the hrst.

The primary dividing furmw makes its aj)pearance along the
dorsal median line of the continuation of the mouth and the Stdbnrgan,
and extends till the body hîis taken the form of a biscuit; but the coîu-
plete sepnration of tlie body into two parts pn^ceeds, according to my
observations, from the periphery towards the central protoplasmic
mass, which for a long time remains conti mious, as may be seen in
Figs. 10, 11, and 12. Hohin., wlio studied very thoroughly the divi-
sion of Noctiluca,, has arrived, so I learn from Buti^clili (6, p. 1068-
1067), at conclusions which in the main are in accordatice with nu'ne;
the only difference beinii' in the manner of the final division of the

312

C. ISHIKAWA.

individuals, which, according to liim, ;^'oes either from the periphery
towards the central mass or vice versa.

h, Spùre-formation.

The general phases of spore-formation have heen worked over
by previous writers, such as Cienliowskti (7, 8), Fiohin (31), and
others. Tlie spore-forming individuals appear to possess a less
quantity of cell -substance than ordinary individuals, owing, to
the flowing together of the protoplasmic network with the central
mass, as Cienkoirsl:ij ( 7, 8 ) rightly observed. The observations
of the same naturalist of the partial division of the entire body in the
earlier phases of budding, are, as stated above, in accordance with my
own, although the lines of division are not so deej) as C'ien-
Icoii'slxii sliows them in his Fig. 14 (7); hut then division of the body
into four parts is more seldom met with than that into two.

Tl»e two nuclei resulting from the division of the first nucleus
do not usually remain stationary, but soon change their relatixe posi-
tions, and thus tlie second division does not generally take place at
right angles to the first, as will be seen in Figs. 15, 1(), and 48. In
consequence of this, the first four nuclei very seldom lie regularly
on the surface of the cell. The time of the division of the first four
nuclei is also not exactly the same in all of them, ccnisiderable fluc-
tuations l)ein"' observed in this and also in all the succeedinii' stages.
Accordingly, the number of the nuclei, as well as that of the s]X)res.
is not uniformly a multiple of two. as counted by Uohin (6, p. 1069),
but<jiiite irregular. Ci<'nl-<iir.4-if ( 7, p. 1 .^5) and also Bohin (6, p. 1070)
admit, in fact, that the diviyi<m becomes tolerably irregular when the
number of the nuclei becomes large. Tlie number of ripe spores also
varies much according to the size of tlie individual, being less in smalls
er individuals than in larger ones. I have counted in one case more

STUDIES OF REPRODUCTIVE ELEMENTS: II. 31 D

than live-hundred spore« in a very large animal, while in other cases
the number wns little over three-hundred. lUit this also is not al-
ways the case, for the area c<3vered by the spores varies in different
individuals, though in usumI cases it is nearly one-lialf to one-tliird of
the half surface of the cell.

We will now speak about some pnnts of more or less interest in
the phenomena displayed in the karyokinetic division of the cells of
other animals com])ared with those presented in the above described
cell-division of Noctiluca. These points concern the archoplasm,
the C'KXTROSOME, and the spixdle fibres.

The ARcnopj.ASM is, as will have been seen from the in-ecedinf
description, first generally found quite near the nucleus when this is
in pre[.aration to divide, (see Fig-s. 1, 2, 13, 31 etc.). At this time
the nuclear wall is always distinctly to be seen, and the size of the
archoplasm as com])ared witli that of the nucleus is such as to make
its origin from the nucleoplasm very improbable. The nucleus at
this stage and in the living condition, appears more or less homo-
geneous and transparent, wliile the archoplasm, consisting of coarse
granules, is so conspicuous as to be easily mistaken for it. This fact
perhaps explains tlie statements oi'Cienkowshj ( 8, p. 54), who supj)oses
the nucleus to disappear at tlie beginning of the spore-formation in
tlie animal. JUnn (6, p. 1070) lias observed the archoplasm and the
changes it passes through in nuclear division, but seems to have
mistaken it for the nucleus. In dividing individuals, the archoplasm
remains closely attached to the nucleus and for a, long time until
the complete separation of the daughter individuals, as will be
seen in Fig. 12; and in spore-forming animals up to the time
when the spore is ready to swim away from the surface of the

314 C. ISHIKAWA.

body, the most remarkable thing here being the flagelliim whicli
takes its origin from the centre of the archo])]asm, as ah*eady stated.
Strashiinjcr f 35, p. 65) gives many similar cases in the swarm
spores of many pJants, where the cilia spring ont of the hinopliisiii
(archopla^>ii of Ijoveri), or at least from the point where the last
trace of the l-itiophism was found. We shall come back to tliis again
soon.

Looking over the literatui-e of the nuclear diNision of Proto-
zoons, there is, so far as I know, as yet no case mentioned of the
existence of the archoplasm and the centrosonie, exce])t in a short
notice of mine ( 26, ]). o). The gathering of tlie cytoplasm and the
corjical bodies at the poles of the dividing nuc.-lens was ol)served by
Schcirial-of (32, ]). 2^1) in En g I y ])h a. a 1 v eo 1 a ta. Ihit these apjtear
at the op[)(»site poles of the niu-leiis from the beginning, as he very
explicitly savs : — '' Ik'Aor nocli die Al)pl:!ttinig des kugeligen Kerns
stattfindet, sah man das (\ytophisnia an zwei beliebigen entgegen-
gesetzten Stellen, den zukünftigen Polen, sich anhäufen. Kurze Zeit
darauf beginnt die Abj^lattung und man l)einerkt gleichzeitig, dass die
Kernwandung an diesen beiden Stellen in den Kern sich etwas ein-
stülpt, wodurch zwei kleine Dellen gebildet werden. Auf dem Grund
diese Dellen o^e wahrt man einen kleinen homogenen Höcker, dessen
Umrisse, dank der starken Lichtbrechung, deutlicli her\'ortreten.
Besonders scharf treten sie bei abgetödteten Thieren hervor, und er-
scheinen als mattglänzende, gut begrenzte, ellipsoïdale Körper, die
von Farbmitteln nicht im mindesten tingiert werden." He jnstly
compares this body with the polar corpuscles oi' Ed. rafi llencih'it, and
considers the hyaline p-art of the Spirochona nucleus, ol)served by
11. Heiiii-i(j, to be in the same category. According fo J I. Ilcrhriy
(23, p. 15()), the nucleus of S])irochona gemmi])ara consists
of two ])arts, a. larger granular part and a smaller hyaline part,

STUDIES OF REPRODUCTIVE ELEMENTS: 11. 3]^5

separated from each otlier Ijy a transverse line. Stained witli Beule s
enrmine, tlie former colours much more (jiiickly than the latter. The
same state of things was ohserved l)y this author in Lepto-
discus med u soi des (24, p. oll), the only representative of
Cystofl;i gel lata, other than Noctiluca, found hy Hertwig at
Messiiin. The nucleus consists here also of two parts, '' einem fein-
k()rnigen und einem homogenen." " In dem einen Falle war die
homogene Ivernsnbstanz unverändert, dao-eg-en fanden sich in der
feink()riiigen grössere nnd kleinere Körperchen, die sicli in Carmin
dunkler iarl)ten und otfenbar eine bedentendere Dichtigkeit besassen.
In anderen Fall liess sich die Düferenzierunii- in zwei Substanzen
nicht nachweisen " (24, p. 31:?). The first change that occurs in
tlie homogeneous part of the S pi roch on a nucleus before its division
is the appearance of a small central corn (23, p. 161). This he
calls a " nucleolus," which is very probably the free area around the
centros(3me, such as is repreaented in my Fig. lo. Some changes occur
in the granular part also, and the nucleus soon assumes an elongated
form with homogenecjus masses placed at its poles. In this way the
nucleus assumes a form in which five parts are distinctly to be
made out : viz., two homogeneous terminal plates, two striated
portions, and a granular median part, considered Ijy Her(iri(j to cor-
respond with the nuclear plate. There is tlius a remarkable resem-
blance between the homogeneous part of the Spirochona nucleus
and the archo[)lasm (jf Noctiluca, although the aspects of the two
are different, the former looking transparent and the latter granular.
But no one will doubt of this similarity, who conq)ares my Fig.
14 or 52 witli Hertiriifs Fig. 17, a. What transformations the homo-
geneous part of the Spirochona nucleus undergoes during the
division is not quite clear from Herkuig^s descriptions. From what
we get from the Noctiluca nucleus, it seems very proljable that

316 C. ISHIKAWA.

the division commences in this homogeneous part. The median
granular part of the spindle of the Si)irochona nucleus, as is given
in Hertwiijs Fig. 17, l> and c, is not to he seen in tlie Noctiluca
spindle. It is, however, very probable that here the unifurm length
of the chromosomes causes the appearance of a distinct median por-
tion. One great ditference to be observed between the tw^o, is that in
Spirochona and Leptodiscus the archo|)lasm is so rlosely united
with the nucleus that it appears as if it formed a part of it. Whether
we have to consider the tw^o nuclear parts as a single nucleus, or
whether the granular part alone is to be taken as a nucleus is not
quite clear, since no membrane is to be seen around the nucleus, al-
thoutJfh Hertivui believes in the existence of (^ne from anaK^üv with
luiclei of other Protozoons. In J^ e p t o d i s c u s , however. Heriwifj
speaks of the existence of a distinct membrane around the t^\()
portions of the nucleus taken together. In this connection ticii-
wuf says (24/ p. /^l 1-812) : — "Der Kern des Lei)todiscus stimmt
S(;mit vollkommen mit dem der Spirochona gemmipara iiberein.
Wie ich in einer früheren Arbeit gezeigt, habe, besteht der Nucleus
dieses Infusors ebenfalls aus einer feinkörnigen und einer homogenen
Substanz, die sich beide gegen einander mit einer glatten Contour
absetzen und ç^emeinsam von einer zarten Kernmembran umhüllt
werden." Although division of the Leptodiscus nucleus was
not observed by Hertwig, the perfect similarity of its structure with
that of Spirochona, makes it very [)robable that here also the
homogeneous portion plays an active ])art in the cell-division and cor-
responds with the archoplasm of the Noctiluca nucleus.

The occurrence of the kinetic centres — the archoplasms — apart
from the nucleus in Noctiluca, deserves special attention. Hichdnl
Hertwig (25, p. 10(5) in his very interesting lecture on fertilization
and conjugation delivered in lierlin, speaking of the constant occur-

S'l'UDIES OF liEPRODUOTIVE ELEMENTS .-il. 3]^^

rence of the œntrosonie in Metuzou, makes the following remarks
on it as rencards 1 1 le P ro t o z o a : —

" Mit diesen für die Metazoen geltenden Verallgemeinerungen
sind die Beobaclitungen an Protozoen zunächst nicht in Einklang- zu
bringen. Meines Wissens ist nur fiii- Aoctiluca die Anwesenheit
eines vom Kern unabhängigen Centrosomas you Lslrihuni angegeben
worden, und iiuch hier handelt es sich nur um eine V^ennuthune'.
Wo sonst i'rotozoenkerne genauer auf ihre Theilung hin geprüft
worden sind, hat sich herausgestellt, dass die activen Substanzen,
welche die Kerntheilung veranlassen, in Inneren des Kernes liegen, und
als Bestandtheile desselben angesehen werden müssen. Ich habe das
Gesagte für Actinosphierium nachweisen können ; bei Eugly|)ha,
welclie nach Sclicirialiriirs Untersucliunuen in der Kerntheihin^' mit
Aclin(jsi)luerium sehr übereinstimmt, scheird ein gleiches A'erhalten
zu herrschen. Am beweiskräftigsten sind aber die Nebenkerne der
Infusorien, deren Theilung am auffälligsten unter den Protozoen an
die Sporenbildung der Metazoen erinnert."

In Actinosphan-ium cichhorni and in »Spirochona it is
true that the kinetic substance of the nuclear division — the archoplasm
— lies within the nucleus. In the micrtjuuclei of all the Infusorians, as
far as they have been studied by many eminent investigators, there
is as yet no case known where extranuclear kinetic centres are
proved to exist. The state of things appears, however, to be a little
different in Euglypha alveolata, where the Polkörperchen lie in
small concavities of the nuclear membrane at two opposite poles of
the nucleus. Of the origin of the Volkorperchen, ScJiewiakoJt (32, p.
222) says :-

" Verfolgt man aufmerksam seine Entstehung, so kann man die
Vermuthung aussprechen, dass es, wenn auch theil weise, aus dem sich
ditferenzirenden Cytoplasma gebildet sind." It, therefore, appears

31g C. ISHIKAWA.

that the kinetic centre — the l'olkörijcrcfwn — of Englypha is derived
from the cytoplasm and not from the niicJear suhstance, just as in
the case of No(;ti 1 iica . Ihit -while in Noctiluca the archoplasm
appears to remain in the cytijplasm, in Englypha it l)ecomes inter-
mixed witli tlie nucleus, when the division of the latter is completed,
as we see from tlie following words of Sdienuakoff' (32, p. 2o(S) : —
" Vor alien Dingen wird das ehemnJigen Polkörperchen vollkommen in
die Ivernsuijstanz eingezogen; man gewahrt nichts mehr von den linsen-
artigen Hervorstiilpung ; wie es noch in den kurz vorhergehenden
Stadien der Fall war, sondern der Kern erscheint einheitlich und besitzt
eine regelmässige glatte Oberfläche." It tlius a])pears, that the Polkör-
perchen here are formed from the cytoplasm at the beginning of the
nuclear division, and become drawn into the Ixjdy of the nucleus at
the end of it. The Folkörpercheii is therefore not a permanent body,
but has always to be formed anew from a part of the cytoplasm at
the beginning of each nuclear division. As I have no ground to
doubt the accuracy of the very beautiful ol^servations of Sclwirialwf,
I have onlv to conclude that the fate of his PolkorpercJien is not
exactly the same with that of the archoplasm of Noctiluca, nor
do 1 doubt the observations of Herlie'uj and others of the kinetic
centres lyiug within the nucleus. From all this it apjiears that
the above statement of 11. Herlwiij as regards the position of the
kinetic substance in Protozoa, and Metazoa, holds true ; that in
I'rotozoa, this substance lies within the nucleus and that in Meta-
zoa it lies outside it. Jkit it apjjears also that there is considerable
fluctuation within the Protozoa. Of special interest in this con-
nection is the ])Osition of the archoplasm in Noctiluca and in
Le pt od is eus — the two relatives of Cy s to flagellât a : for while
in Leptodiscus this substance lies within the nucleus, in Nocti-
luca it lies outside it. Cystoflagellata thus forms, in this res-

STUDIES OP REPEODUCTIYE ELEME^TS : II. ^^9

])ect, an intermediate staa'e, so to speak, between the Metazoa and
other Prot o z o a . If in E n g I y p li a a I v e o 1 a t a the Polkörperchen is
formed from the nuclear substance and l^ecomes connected with the
cytoplasm during the nuclear division, to be again withdrawn into
the nucleus, it forms in tliat case an interesting connecting hnk between
Leptodiscus and Xoctiliica! But we must confine ourselves to
facts only.

Another interesting tiling about the Noctiluca archoplasm is
its extreme similarity with the Neheuh'rii of vo)t la Valette Si. George
(13, U\ and riatNer (29). Just like the archoplasm ofNoctiluca,
the NerenkerH of these authors is formed by the consolidation of a
part of the cytoplasjn at one side of the nucleus, and wlieu this be-
comes visible the '^ Schleifenbiindel des Kerns ist stets mit der Spitze
gegen jene Verdichtiuig hingerichtet" (Ay l'alette 13, ]). 7). The
])art ])layed by tlie Xemil-ern ap])ears, however, not (juite clear, but
roN la \ (dette St. (leonje considers it very })robable that the spindle
fibres are formed from its elements in lîlatta, since these transform
again to fi^rm fhe Xelteukern (13, ]>. 10). AVhen tliis fact is ])roved,
then tlie parallelism between the Noctilnca archoplasm and the
Nebenkern, of run la Valette St. Geonje will be proved l)ev<^nd all doubt.
It will not be, however, a rash conclusion to consider these two ])odies
as identical, especially when we consider the fate of the Nehenkern in
the foi-matioH of tlie spermatozoa, as will be seen later on. The
close similarity l)etween the two will also be seen by com])aring von la
Valette s F\g. (J on Plate 111 <)f '' /vö7///.vr'.s Festschrift, 18 <S7 '' with
ray Figs. 1, 2, 6, or 7.

Of not less interest tlian the archoplasm is the origin of tlie spindle-
fibres. Two different opinions are here also to be met with as in the.
case of the origin of the kinetic centre. Str((s]nir(jer in his earlier
works (33, 34) as well as in his newest (35, p. 59-62). has shown in

820

C. ISHIKAWA.

inanv vegetable cells the cytoplasmic oriiiin of the spindle fibres.
In Spirogyra poJytojniata a part of the cytoiJasm— the kino-
plasm — differentiates itself in the form of parallel fibres on l^oth
sides of the flattened nucleus, even when the nuclear wall is very plainly
to be seen. Meanwliile the nucleus becomes flatter than before, and
the chromosomes ari-ange themselves in the equatorial plane. The
parallel fibres come now to lie seen distinctly within the nucleus on
both sides of tlie equatorial row of chromosomes. The fibres are.
according to Slnishunjer, the same fibres as were seen outside the;
nucleus and have pnshed into the nucleus through the sieve-like pores
formed in the nuclear wall, and attached themselves to the chromo-
scmies. The nuclear membrane disappears after this, and a perfect
spindle with the ecjuatorial row of chromosomes is formed (34^ p.
11). In the nuclear division of the ])rot()])lasmi(' l]'^tind}>eh'(i of the
embryo-sac of Leucojum a^stivum, the same author describes the
formation <^f the spindle out of the cytoplasm surrounding the nucleus
at the time when the miclear wall still exists. In the ])ollen-
mother-cells of Lilium biill)iferum (3Q. |>. 1 8::}-183). ^Ym.s7rH/y/^/'
describes the formation of the sjundle as like that in Spirogyra.
Here twelve fibres are thrown out from both the Ct'tttrosjilnnrs towards
the centre of the nucleus and push into its interior, and then,
the nuclear wall bec<)niing indistinct, ;i perfect spindle is formed
by the union of the free ends of the fibres at the ec[uator. In
his latest paper (36) StraiyJmrgcr still holds liis views as to the cyto-
plasmic origin of the spindle-fibres for plant cells. On the other hand,
Fßtzwr (28, I». 6o5). Iluhl (30, p. 26\)), Z<irhnrias (39, p. 852 ; 40,
p. o34), (L llniiriii ( 22, p. 16o). and others, believe in the tin-mation
of the spindle fibi-es out of the nuclear substarice. <>. Hcrtwiij «m this
point says (22, ]>. Ki.H):- — " l>ei einzelnen ^lollusken (Ptei-otrachea.
Phyllirhoë) liaben Fol und ich beobachtet, das« die Polspindel im

STUDIES OF JJEPRODUCTIVE ELEMENTS: II. 35!

Innern des Keimbläschen ??, welches hier übrigens von geringer Grösse
isr, angelegt wird, so lange noch die Kerimiembran vorhanden ist.
J)ie Annahme, dass in diesem Fall Protoplasma von aussen iji den
Kernraum hineingedrunuen sei, will mir wenis^stens als eine o-ezwuno--
ene erscheinen." While thus the views of Straslmrger and Hertwig
are quite di Itèrent as to tlie oi-igin of the spindle fibres, the former
deriving tlieni from the cvt(^plasm and the latter from the nucleo-
plasm, the opinions of Flemuiiiuj (12, p. 75), Plainer (29, p. 70),
and othei-s stand midway between, since according t(3 these authors
the eijuatorial part of the spindle is formed from the nncleoj)lasm,
and the polar parts from the cytoplasm. The late investigation
by AiKjust Brauer (4^ of the spermatogenesis of Ascaris megaloce-
phala is very interesting on this point, inasmuch as the origin of the
spindle fibres is different in two varieties of the same species, fn
Ascaris megalocej)ha la uni Valens the centrosome is found in
the micleus, and there forms a small spindle which this authoi- com-
pares with the central spindle of Heriiianii, although the origin of
that is ditferent, as will be shown later on. In Ascaris mes-alo-
cephala bi va lens, on the other hand, the centrosome is first found
in the cytoplasm, closely attached to the nuclear w^all. By the divi-
sion of this is also formed a small spindle lying first tangentially to
the surface of the nucleus. Init the two centrosomes gradual Iv separate
from each other until they come to lie diameti-icallv opposite to each
other. What becomes of the central spindle fibres is not known, but
the author is inclined to think that these are divided in the separa-
tion of the centrosome (4, p. 181).

Of no less interest than the al)ove are the observations oï Herman ii,
the result of whose investigation on the spermatocytes of Sa I am a n-
dra maculata, specially directed to the study of the origin of the
spindle fibres, nnnarkably accoi'ds with that of mine upon the

322 C. ISHIKAWA.

Noctiluca spindle. In the cytoplasm of the spermatocyte ut some
distance from the nucleus, there is formed a small spindle, from the
poles of which, when it has attained a certnin size, a number of fil)res
is seen passing to tlie chromosomes. By growth and change of
position, it gradually attains the ordinary form of the spindle with
equatorial chromosomes. In this full grown spindle there are thus
two sets of fibres : the one continuous iDctween the two centrosomes
and lying axially ; and the other passing from the centrosoines to
the chromosomes. The latter set of fibres therefore lies more or
less like a mantle above the other set, which he calls the central
spindle (20, p. 580). When division of the cell is completed the
fibres of the central spindle return to the protoplasm (?), while the
other fibres collect together to form the archoplasms of the daugh-
ter cells. This description of h[eriii(tnn\s stands thus in beautiful
correspondence with that of the formation of the spindle in Xocti-
luca, given above; but with the difference that in Noctiluca, from
the persistence of the nuclear wall, the centnil spindle does not
lie, strictly speaking, in the axis of the whole system, but in tlie
groove formed on one side of the nucleus by the depression of its wall,
and therefore the mantle-like fibres as well as the chromosomes do not
lie quite around the axial spindle. Tl\is otherwise exact correspon-
dence of the spindle fibres in these two widely different animals is,
beyond all doubt, a very interesting phenomenon, and renders desi-
rable, in my opinion, further investigations into the structure and
formation of different kinds of cells in other animals and perhaps also
in plants.

In this connection I nuist not leave unnoticed the question of the
V('rhindi(N(j!>fa(lfn, of whose origin there are also as many interpretations
as there are questions on other matters. Many investigators, such as
0. Hertiri,! [2\, 22\ Boven {3\ Ed. ran Jkneden and Nejit (\\ and

.STUDIES OF REPRODUCTIVE ELEMENTS: II 393

Brauer (4', consider tlie ycrhiNdiiiKjstadcH to be formed from tlie /?'//?7?-
substance l)y the separation of the chromosomes towards the poles of the
spindle. Accordino- to Stra!ihur(icr (36, p. I8ö), these are formed from
tlie same substance as the kinetic tibres, which enter from the poles
of the spindle and meet in the equatorial plane. This view accords
well witli tliat of Hermann, in so far that the latter author considers
the central spindle as forming the ]'erhinilungsjaden of other investiga-
tors. Tn s{)iteofthe great similarity between the structures of the
spindle in Noctiluca and Salamandra this view oï Hermann's con-
cerning the ^'erhindnngsfäden can not l)e ap]>lied in explanation of the
same structure in the Noctiluca s[)indle, since, as we see from the
above description, the persistence of the nuclear wall in Noctihica
naturally shnts off all possibihty of confounding the libres of the
central spindle with those extending between the free ends of tlie
separating chroiiK^somes. Moreover, the two sets of fibres opticallv
appear quite di itèrent from each other, as will be seen in Fig. 37, (c.f.)
and (/■./.).

Of the origin of the radial tibres in Noctiluca, I can say but
a few words, since the whole pr<)blem still remains very obscure
and requires a thorough study with better methods and optical instru-
ments.' In sections given above, these tibres. whicli are found within
the nucleus and probably attached to the chromosomes, appear to come
into close juxtaposition, but not to be continuous with those without,
i.e., those seen within and without the nucleus appear to be diiferent
from each other, the f )i-mer originating from the nucleoplasm and the
latter from the cyto])lasm, just as Braver thinks concerning the forma-
tion of the spindle fibres of Ascaris megalocepha la bixalens
(4, p. 182-183).

1 . It may be here remarked that our climate is not fitted for the use of Zeiss apochromatic
systems, the dampnoss of the air in the warm season soon blunging- considérai )]<_■ alterations in
the lenses.

;^24 ^- ISHIKAWA.

Last of all, the part played by the archoplasm in the formation of
the cilia in Noctiluca deserves some attention. Stni si mrger in Yns
Histologische Beiträge, Heft IV., gives many similar cases of the forma-
tion of the cilia from the archoplasm (35, p. 62-13:^; 36, p. 184). Ihe
observations of Henking (17, 18), Flat iter l29', and above all oï von la
Valette St. George (13, 14', of the part played by the Nebenkern
in tlie formation uf the Spermatozoons are very interesting in this
(jonijcction, since the Isebenhern of these authors (especially of
voïi la IdU'tle St. (reimje) corresponds, as said above, in all particulars
with mv 'archoplasm "; and it is, therefore, very reasonable a priori to
consider that that part of the cytoplasm which is especially ditferen-
tialed for kinetic functions, is transformed to form a part of the tail of
a spei-matozoon or the flagellum of a Noc ti luca spore.

C. Summa r g

1) 'J'lie division of the animal is preceded by the loss of the
[icristome. teeth, and the tentacle, the last of which is not thrown oif,
as /('(*/*/// is inclined to think, but is withdrawn into the body of the
animal. The moutli and the Staborgaii arc. however, always present
(th>J>i,i).

2) The spore-forming individuals differ from the dividing ones
in not possessing tlie mouth and the Slahorgan in addition to the
organs above mentioned, and by the excessively empty appearance of
the cell interior (Cienkoivskg).

3) The division of the nucleus is always preceded by the con-
centration of part of the cytoplasm in the form of <*i spheri(*al or oval
granular bod}', mostly elose to the nucleus. This is the archoplasm
or the kinetic centre (of division, and corresponds most j)robablv with
the Sebeitkern of roit la I alette SI . (reorge.

4) 111 living animals at the stage of (o), tbe nucleus appears more

Si'UDrivS OF REPRODUCTIVE ELEMENTS: II.

3i>5

or less homogeneous :nid"ti-;insp:ireüt, and is not so distinofly to be
seen as the archo|)l;isiii. Ihit treated with reagents, the chromosomes
come into view distiiictlv.

5) Knch chromosome consists of a row of disc-shaped microsomes
irregularly scattered in the nucle<jplas]n. The number of the chromo-
somes is not clenr. but in most cnses has been counted to lie ten.

6) The chromatin substance of each of the microsome discs
collects at the {>erij)hery and forms a microsome-i'in"".

7) In the nucleus of a dividing animal, each microsome-ring
splits into h;df-rings thus dividing a chromosome in halves, while
in that of the spore-forming animal two successive divisions of a,
microsome-ring take place, so that a single chromosome is directly
divided into four daughter ones.

8) The clu-omosomes collect on that side of the nucleus which is
nearest to the archoplasm, and spread out towards the other j)ole.
The pole where the archoplasm lies thus corresponds to Rabrs Folfeld
and the other pole to his Gengenpol.

9) The archoplasm divides and forms a very large spindle which
lirst lies tangential to the surface of the nucleus. This division of the
archoplasm is succeeded by the separation of the chromosomes into
two groups each attracted (?) by its respective archo])lasm.

10) The archoplasmic spindle thus formed pushes-in the nuclear
wall on which it lies, and the nucleus assumes in consequence a half-
ring form.

11) lîy the separation of the archoplasms, a sj)indle is produced
wliicli in all essential characters appears like the form known as the
" disaster stage," with a large archoplasnn'c mass at each end of the
spindle.

12) The fibres of this spindle are therefore continuous from one
pole to the other and lying outside the nuclear wall become in no way

^'2Q c- ishikaWa.

connected with tlie cliroinosoines. But there is seen ;it this stage
another set of tihres running from the centre of the nrchoplasm to the
polar ends of the chroniosotnes. This structure of the spindle corres-
ponds exactly with that of the spermatocyte of Salamandra macu-
la ta, as investigated hy Henna nu, with only the difference of the
persistence of the nuclear wall in Noctiluca. and tlie necessary
modification in consequence of this tact. The optical appearance
of these two kinds of fil)res is different, just as in Salamandra.

13) 1 besides these two sels of fihres, the I eiiundungsfäden are clear-
Iv to he recognised extendini»- hetween the separating' chromosomes.

14) The central spindle fihres originate from the archoplasm,
the radial fihres probahly from both the cyto- and nucleo-plasms, and
the VerhiinhuKisfäden from the //'/////-substance.

15) In the spore-buds the archoj)lasm is to be seen lying close
to the nucleus u]) to the time of tlie full development of the sj)ore
just l)efore its detachment from the mother animaJ, and a part of it
l)ecomes transformed into the flagellinn, just as iji many vegetable
swarm-spores, as Stra.sbiinßr sliows.

16) In the centre of the archoplasm is generally seen a centro-
some, which often show^s a dumb-bell form. »Sometimes, however,
two centrosomes are found in the archoplasm of the spore-forming
cells. Fn many cases, again, there is found in the centre of the
archoplasm a number of small oval, rod-shaped or curved bodies,
staining exactly like centrosomes, instead of one or two centrosomes.
These may repi-esent the group of centrosomes of Heidenhein.

IT) The origin and the fate of the centrosome are not known.
In a few instances it appears to be fbi-med from the nucleus.

Agkiculti ral Cullegp:.

End of November, 1893.

STUDIES OF KEPRODUCTIVE ELEMENTS: II. 327

List of References.

1. nin Heitedeii, Kd. et Adiilplie Xfi/t. — Nouvelle recherches sur la fecondatioii et la

division mitotiqne chez l'Ascavide mégalocéphale. Bulletins de l'Académie
royale de Belgique, S'' série, tome XIV, no° 8 ; 1887.

2. Herr/h, H. S. — Recherches sur les noyaux de l'Urostyla grandis et de l'Urostyla

intermedia, n. sp. Archives de Biologie, tome IX ; -1889.
8. Boveri, 77t.— Zellen-Stndien, Heft 2. Fischer, Jena ; 1888.
4. Brauer, Au;/. — Zur Kenntniss der Spermatogenese von Ascaris megaloeephala.

Arch. f. mik. Anat. Bd. XLII ; 1893.
6. Hriiihtivt'll, T. — On tlie self-division in Noctiluca. Quart. Journ. Micr. Sei.

Vol V; 1857.

6. Biitschli, (). — Cystoflagellata in Bronn's Klassen und Ordnungen. I Bd. xAhtli.

2; 1889.

7. Cienkowftlaj, L. — Uel)er Schwariuerbildung bei Noctiluca miliaris. Arch. f.

mik. Anat. Bd. VII ; 1871.

8. Cienkowskij, L.-— lieber Noctiluca miliaris Sur. Arch. f. niikr. Anat. Bd. IX ;

1873.

9. Flemminij, H'.— Neue Beiträge zur Kenntniss der Zelle, t. Theil. Arch. f. mikr.

Anat. Bd. XXIX; 1887.

10. Flemming, W. — -Neue Beiträge zur Keinituiss der Zelle, IL Theil. x\rch. f. mikr.

Anat. Bd. XXXVII; 1891.

11. bleimniiiij, W. — Ueber Zellthoilung. Verhaudl. der anat. (Tesellschaft zu

München ; 1891.

12. Fle)iiinin(i, \V . — Zelle. Ërgel)nisse der Anatomie und Entwicklungsgeschichte.

1892.

13. St. <Tt'i)n/t', roll la Irt/^'^/^^.— Spermatologische Beiträge, LI. Mittheilung. Arch.

f. mikr. Anat. Bd. XXVII; 1880.

14. St. (rmrtje, von la Valette. — Zelltheilung und Sameubildinig bei Forftcula

auiicularia. Festschrift für von KöUiker. 1887.

15. tleiileiihein, M. — Kern und Protoplasma. Festschrift fiir \on Kijlliker. 1892.

^'2 H •'• isniKAW'A.

IG. Heidenlu'i)iy M.- — lieber die Centialkurpeignippe in den I.yuipliocyten der
Saugethiere. Verbandl. der anat. Gesellschaft zu Göttiiigen ; 1898.

17. HenkiiKj, TL — Untersuchungen über die ersten Entwicklungsvorgänge in don

Eiern der Insecten. I. Das Ei von Pieris brassicffi L., nebst Beraerlanigen
über Samen und Samenl)ildnngen. Zeit. f. wiss. Zool. Rd. XLTX ; 1890.

18. Henkhvj, H. — Tlie Same. IT. lieber Spermatogenese und deren Beziehung zur

Eientwickluiig bei Pynliocovis apterus L. Zeit. f. wiss. Zool. Bd. LI; 1891.
10. Irlenlnnij, H. — Tiie Same. Iff. Spezielles und Allgemeines. Zeit. f. wiss. Zool.

Bd. LTV; 1893.
•20. Ht'fiiifOiH, F. — Beiträge zur Lebre von der Entstehung der karvokinetisebeii

Spindel. Arch. f. mikr. Anat. Bd. XXXVTI ; 1891.

21. lli'ittn'ii, (). — ^Vergleich der Ei- und Samenbildung bei Nematoden. Arcb. f.

mikr. Anat. Bd. XXXYI ; 1890.

22. Hertii-Uj, <>. — Die Zelle und die Gewebe, Fischer, Jena ; 1892.

28. HertwUf, Jl. — Ueber den Bau und Entwicklung der Spirochona gennnipara.

Jenaisch Zeitschr. f. Naturw. Bd. XI ; 1877.
24. Hcrtirit/, E. — Ueber Leptodiscus medusoides. Jenaiscjie Zeitscbr. f. Naturw.

Bd. XI; 1877.
2;"). Herta-'ui, 11. — Ueber Befruchtung und Conjugation. Verhdl. d. deutsch, zool.

Gesellschaft zu Berlin; 1892.
20. f.sfiikdtvd, (-'. — Vorläuüge Mittlieilungeu über die C'onjugationserschehuuigen

bei den Noctiluceen. Zool. Anzeiger; 1H91.

27. hitikawu, C. — Studies of Beproductive Elements. I. Spermatogenesis, Ovo-

genesis, and Fertilization in Diaptomus sp. This journal Vol. Y; 1891.

28. I'rit'.ii>i\ li'. — -l^nträge zur Lobve vom Bau des Zellkerns und seiner

Tlieiliuigsersclieimingen. Arch. f. mikr. Anat. Bd. XXII; 1883.

29. Fliitner, (t. — Beiträge zur Kenntniss der Zelle und ihrer Theilungsei-scheinuiig-

en. Arch. f. mikr. Anat. Bd. XXXIII; 1889.

80. l'uthl, C— Ueber Zelltheilung. Morph. Jahrb. Bd. X ; 1885.

81. Unhiii, (Jh. — Keclierches s. la i-eproduction gemraipare et fissiparu des Nocti-

hiques. Journ. anat. et physiol. 1878.

82. Schi'n-idknif, IF.-— Ueber die karyokinetische Kerutheilim^- der Euglypha alveo-

lata. Morphologisches Jahrl)uch. Bd. XLll ; 1<S8H.
88. Stidshiinjer, /'.''/. — Ijofiuichtungsvorgang bei den i'hanerogamen Fischer, Jeuîi ;
1884.

STUDIES OF PkEPEODUCTIVE ELEMENTS: II. 3^9

34. Stnisbtinjer, Ed. — Histologische Beiträge, I. Kern nucl Zelltlieihnig im Pflan-

zenreiche. Fischer, Jena ; 1888.

35. Strashuiyei; Ed. — ^Histologische Beitrüge, Heft IV. Schwärmsporen, Gameten,

pflanzliche Spermatozoiden, iind, das Wesen der Befruchtung. Fischer, Jena ;
1892.
3G. Strasbuiyei; Ed. — Zu dem jetzigen Stande der Kern- und Zelltheilungsfragen.
Anat. Anzeiger VIII. Jahrg ; 1893.

37. M'eitiiiuoui, A. and C, Ishihiin/. — Ueber die Bildung der Piichtungskörper bei

tierischen Eiern. I. Ber. d. Naturf. Ges. zu Freiburg i. B. Bd. III ; 1887.

38. Widdeyer, W. — Ueber Karyokincse und ihre Beziehungen zu den Befruchtuugs-

vorgängen. Arch. f. mikr. Anat. Bd. XXXII; 1888.

39. Zacharias. — Ueber die Spermatozoiden. Ann. Bot. Ztg ; 1881.

40. Zacharias. — Beiträge zur Kenntnisse des Zellkerns und die Sexualzellen.

330

C. ISHIKAWA.

Explanation of Plates, XI-XIV.

a — avclioplasm.

a sj; — archoplasmic spiudle.

cf — central fibi'es.

n — nucleus.

rf — radial fi]:)res.

t — tentacle.

.1' — median part of the archoplasmic

spindle left after the division of

the nucleus.

-^ X Z -— Drawn l)y means of camera lucida under Zeiss' objective B with eye-
piece 2 : magnified 60 diameters.

-jjj- X S = Drawn by means of camera lucida under Seibert's objective III with
eye-piece 1 : magnified 120 diameters.

-ji — X Z := Drawn by means of camera lucida under Zeiss' homegeneous immer-
sion Vi2 with eye-piece 1 : magnified 350 diameters.

-Yi — X Z '-=-- Drawn by means of camera lucida under Zeiss' homegeneous immer-

/l2

sion V]2 witli eye-piece 4 : magnified G50 diameters.

Plate XI.

(7)/r/.y/o» of the aiiiiiial.)

Fig. 1. Individual preparing to divide; tentacle (/) much reduced in size ;
archoplasm (a) very plainly visible as a granular spherical body lying close to the
nucleus (n). Killed with Flemming's stronger fluid, and stained with Böhmer's
hiematoxylin.

Fig. 2. Another individual of about the same stage as that of fig. 1 ; in
centre of archoplasm a ceutrosome surrounded by clear space. Similarly treated
as above.

Fig. 3. Central plasm of a dividhig individual with archoplasmic spiudle
{(( s/>) lying close to the nucleus ; chromosomes gathered towards pole of iiucleus

STUDIES OF REPRODUCTIVE ELEMENTS: II. 331

where spindle touches it. Acetic acid methyl-green preparation.

Fig. 4. More advanced stage of division ; nucleus now duinbbell-sliiped
with archoplasms at its poles ; narrow line of protoplasm marking line of divi-
sion of animal seen in front of and behind central protoplasmic mass ; the latter
also constricted into two portions. Acetic acid methyl-green preparation.

Figs. 5-9. Different stages of division, showing various pusitions of archo-
plasm in regard to those of nucleus. All acetic acid methyl-green preparations.

Plate XII,

(Figs. 10, 11, Si 12 represent fiàv'.y/oy; ; Figs. 13-19, .sporc-foriimtion.)

Figs. 10 and 11. Further stages of division; tentacles already at stage
of fig. 11. Killed with Flemming's stronger fluid, and stained with Bohmer's
hematoxylin.

Fig. 12. Portion of dividing individual with narrow protoplasmic connection
{p c) still visible ; tentacles grown much longer. Drawn from a living animal.

Fig. 13. First stage of spore-formation ; archoplasm (a) as a granulated
mass lying close to nucleus {)i) ; dumbbell-shaped clear space within archoplasm,
which represents clear zone around a dividing centrosome. Drawn from living
animal.

Fig. 14. Further stage of spore-formation about the same as that of fig. 4.
Acetic acid methyl-green preparation.

Fig. 15. Still further stage of spore-formation ; division of nucleus now
nearly at an end, parts still connected by narrow bridge. Acetic acid methyl-green
preparation.

Fig. 16. Central plasm and dividing nucleus at stage similar to that of last
figure ; drawn from living animal.

Fig. 17. Four-nucleated stage of spore-formation ; archoplasms belonging to
nuclei in state of division, each forming large spindle. Drawn from acetic acid
methyl-green preparation .

Fig. 18. Side view of stage slightly later than that of tig. 17. One of the
four nuclei, on right-hand of figure already divided into two daughter nuclei. (The
two dumbbell-shaped miclei, not represented in the figure, are also in a similar

332

C. ISHIKAWA.

stage of division to tbat represeiited on the left of the figure). Acetic acid methyl-
green preparation.

Fig. 19. Stage similar to that of fig. 18, sho-\ving line of partial division
of animal. Picro-acetic acid preparation without staining.

Plate XIII.

(Figs. 20-30, represent spore-formation; Figs. 31-37, dichion.)

Fig. 20. Stage of sporulation more advanced than that of fig. 19, showing
also line of partial division of animal. (It will he observed that the number of the
nuclei in each half is not equal, four lying on one side and two on the other).
Acetic acid methyl-green preparation.

Fig. 21. Stage of spore-formation still later than above, showing also line of
partial division, six nuclei lying on one side and four on the other. Acetic acid
methyl-green preparation.

Fig. 22. Stage with fifty-one nuclei seen laterally ; spore-cells on the other
side represented by fainter shading. Acetic acid methyl-green preparation.

Figs. 23-2G. Sections of spore-buds in last stage of division. Killed with
Flemming's stronger fluid and stained with combination of Bohmer's hfematoxylhi
and rubin.

Figs. 27-30. Eipe spores just before detachment from the mother-body.
Killed with Flemming's stronger solution and stained with carbolic acid fuchsin.

Fig. 31. Part of an individual just preparing to divide ; very large archo-
plasm close to nucleus ; tentacle and teeth still visible. Killed with Flemming's
stronger solution and stained with Bohmer's hematoxylin.

Fig. 32. Nucleus of individual preparing to divide. Killed with Boveri's
picro-acetic acid, and stained with acid-fuchsin and methylen-blue.

Figs. 33 & 34. Nuclei of two dividing individuals, in which the longitudinal
division of chromosomes are beautifully to be seen. Killed with Flemming's stronger
fluid, and stained with acid-fnchsin and methylen-blue.

Figs. 35 & 35. A. Nucleus in process of division as seen at two diflerent
levels: fig. 35 at level of archoplasmic spindle, and fig. 35. A. at a lower level ;
centrosome within the archoplasm on the lower side. Killed with F]( inming's strong-
er fluid and stained with Bohmer's hfematoxylin.

STUDIES OF REPRODUCTIVE ELEMENTS: IF. 333

Fig. 36. Stage like the last drawn at the level of avchoplasmic spmdle.
Central- and eadial-fibres beantifnlly seen ; owing to accumulation of dirty matters
over archoplasms, centrosomos not to be recognised. Killed with Fleminiiig's stronger
till id and stained with Bohmer's hirmatoxylin.

Fig. 37. Section of nucleus in division at about same stage as that repre-
sented by fig. 35 or 30 ; radial-fibres in the homogeneous mass of nucleoplasm at
lower pole of nucleus, different apearance from central-fibres. Killed with Flem-
miug's stronger fluid and stained with Bohmer's hnematoxylin.

Plate XIY.

(Figs. 38-43, (liri.siuii; Figs. 44-52, sjmre-formatiim.)

Fig. 38. One of the mielei of a dividing individual just after division ;
chromosomes still radiating from pole where archoplasm lies ; small centrosome
lying close to nucleus. Killed with Flemming's stronger fluid and stained with
Bohmer's htçmatoxylin.

Fig. 39. Stage same as fig. 38. Centrosome surrounded by clear space at
centre of upper archoplasm ; in lower archoplasm hidden below nucleus and there-
fore not well seen ; part of central-spindle in shape of a diagonal figure (.c) between
the separating nuclei. Killed with Boveri's picro-acetic acid solution and stained
with Bohmer's hfematoxylin.

Fig. 40. Another nucleus in anaphasis ; microsomes plainly visible. Killed
with Flemming's stronger fluid and stained with Bohmer's hferaatoxylin.

Fig. 41. Nucleus at stage slightly later than that of fig. 40. Microsomes no
longer in rows, but more or less irregular and thus assuming the position in the
resting nuclei. Killed with Flemming's stronger fluid, and stained with acid-fuchsin
and methylen-blue.

Fig. 42. Similar imcleus treated with Boveri's picro-acetic acid and stained
with acid-fuchsin and methylen-blne ; microsomes very much swollen by action of
acetic acid.

Fig. 43. Nuclei of a dividing individual at about the stage fig. 7 or 8.
Killed with Flemming's stronger fluid and stained with acid-fnchsin and methylen-
blue.

334

C. ISHIKAWA.

Fig. 44. Arclioplasm and nucleus of spore-forming individual at about same
stage as that represented by fig. 13 ; the tAvo centrosomes plahily seen at side
of nucleus within arclioplasm. Killed with Flemming's stronger solution and stained
with acid-fachsin and methylen-blue.

Fig. 45. Similar stage to above. Single centrosome surrounded l)y a clear
space and many fine fibres seen in the arclioplasm next to nucleus ; chromosomes
gathered up at pole of nucleus fachig arclioplasm. Killed with Flemmhig's stronger
fluid, and stained with acid-fuchsin and methylen-blue.

Fig. 46. Similar stage to fig. 45. Here the arclioplasm lies at a lower
level than nucleus. Treated as above.

Fig. 47. Nucleus of another individual at about same stage as above. Many
of the chromosomes showing very plainly longitudinal divisions into four rows of
microsomes. Treated as above.

Fig. 48. Two spore-buds in division ; represented more highly magnified in
fig. 48 A and B. Chromosomes within poles of nuclei longitudinally divided already
at this stage ; many darkly-staining bodies within archoplasms, probably corres-
ponding to group of centrosomes observed by Heidenhein. Killed with Flemming's
stronger fluid, and stained with acid-fuchsin and methylen-blue.

Fig. 49-51. Divisions of spore-buds ; centrosomes in many of them. Killed
with Flemming's stronger fluid, and stained with acid-fuchsin and methylen-blue.

Fig. 52. Abnormal division of spore-bud into three daughter cells. Treated
as above.

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Lith. 4* 7mp. the Tokw-Seiêhibuv.êhi

On the Sero -Amniotic Connection and the
Foetal Membranes in the Chick.

By

S. Hirota. Rigakushi,

Zoological Institute, Imperial University, Japan.

With Plates XV-XYII.

CONTENTS.

Introductory lîemarks.

1 Origin of tlie Sero-Amuiotic Connection.

2 Prolongation ni' tlie Sero-Aiuniotic Connection.

3 Limit of tlie Prolongation.

4 Changes in the Extra-Embryonic Germinal Layers.

5 Pieplacement of Cells in the Sero-Amuiotic Connection.

6 Expansion of the Mesoblastic Comiection,

7 Perforation in the Mesoblastic Connection.

8 Supplement to tlie Preceding Sections.

9 Effects of tlie Sero-Amniotic Connection on the Foetal Membranes.

10 Comparison between the Sero-Amniotic Connection in the Cliick and that

in Chelonia.

11 Methods for the Preparation of the Specimens.

12 Explanation of the Plates,

There is probably no other animn] whose development has so
much been Tn;ide the subject of study not only by students in <'eiieral
but by S[)e('iMl invesrio-nfors ns that of the chick, and, vet. it is a
remarkable far-t that erroneous notions jH-evail at tlie j^resent dav in

3:iS TTTRo'l'A : OX TTTE 8E170- AMNIOTIC CONXECTTOX.

reo'ard to some fundamentnl points in the structure and relations of
tlie foetal memhranes of the chick. It is generally assumed hy writers
on eml)ryol<^n'v that several amniotic f(^lds, I'ising over the emhrvo.
come togethei' in the median line aho\ e the emhryo. t]\v edges of which
entirely coalesce and whose cavities hecome continuous from one side
to the other. The classical scries of diagrams given in Foster and
]>alfour's Elements of Embryology (2nd ed., pp. 28-3.')). and copied
very extensively, is constructed on this supposition. Some authors, as
Schenk,' Köliiker, and Hertwig, have, it is true, remarked that the
cavities of tlie lateral folds are separated hy a longitudinal partition
(n;nned ' Amiiionnaht ' by Köliiker) for some time after tlie amniotic
cavity is closed ; luit they also state that shortly afterwards, the parti-
tion breaks down, so that the cavities of the former lateral folds
l)ecome dh'ectly continuous and the amnion proper entirely isolated
fr(3m the serous envelope. In reality the facts are very different.
While the horse-shoe shaped anterior amniotic fold grows backwards
over the dorsal surface of the embryo, the cavities of the lateral folds are
not in coalescence in the median line, l)ut ai-e from the first separated
from each other by a ]>artition which never breaks down. This
is quite similar to the state of things in Chelonia, recently described
by Mitsukuri,"^ who gave the name of '' Sero-Amniotic Connection"
to this partition. This sero-amniotic connection undergoes in the
cluck some histological changes in the course of the development of
the embryo, but remains as a partition to the last day of the incuba-
tion. This persistence of the sero-amniotic connection exercises, as in
Chelonia, a great and complex influence on the foetal membranes in
later stages. At the suggestion of IVofs. Mitsukuri and Ijima, 1 have

1. S. L. Schenk: Beiträg,-e zur Lehre von Amnion. Arch. i. Mikros. Anat. Bd. 7. 1871.

2. K. Mitsukuri: On the Poetal Meuiliranes of Chelonia. .Tourn. of the Colleo-e of Science,
I nil., l^niv., Ja]ian. Vol. IV., Pt. 1. 1890.

ANT) THE FOETAI, MEMBKAXKS. [X THE ('Hiriv.

:53!)

been engaged in elucidating these points in fhe history of the foetal
inenil)runes of the chick and ha\e hi-ought to light some facts which
appear to me very remarkable. The present article contains the main
results of my investigation.

Before going further, I wish to express my best thanks to my
teacher-, Profs. Dr. Mitsukuri jind Ijima, for their constant encourage-
ment and advice throuHiout this investig-ation.

Dia^. /

25-30 Hours.

1. Orujui of llie Scrit-AmuKAic Connection.

If we examine a chick embryo of 2o-o0 hours (Diag. 1) we
find that the extra-embryonic coelomic cavities (E) have already
Diaé. 3, extended themselves on l)oth sides,

some distance in front of the embryo
itself. These anterior parts of the ex-
tra-embryonic coelomic cavities have
not, however, yet extended themselves
directly in front of the head, and
leave here a sort of Ijay (P), where

40-45 Hours.

Head not represented the hypoblast and tlie ej^blast alone
Diaè. 2. Diag. 4, are as yet present.

When the embryo has become
30-40 hours old (Diag. l>), the an-
terior limbs of the extra-embryonic
coelomic cavities meet in front of the
head, cutting off the ])revious bay
from the region in front. Thus an
area (P), which is formed by the
hypoblast and the epiblast only, has
)>een circumscribed just in front of, and beneath, the head. This is

30-40 Hours.

Longitudinal sectidii

through a line, x y

(Diag. 3). Heart

not reprei-euted.

Ö. — Sinus termimiHs.

P. — Proamnion.

H.— Head.

E. — Extra-embryonic coelomic cavity.

M. — Mesoblastic septum.

F. — Amniotic fold.

340 HIROTA; OX THE SERO-AMXIOTIC COXXECTIOX,

the Proamnion. ' The extra-embryonic coelomic cavities \yhich ha\ e
thus met in front of the Proamnion, do not fuse together for a time,
but have a median mesoblastic septum (M), consisting of two sheets
of cells, each limiting the extra-embryonic coelomic cavity of its own
side. At this stagne the head fold of the amnion is raised alonof a
line which passes through the anterior limit of the proamnion
(F. F.).

When the embryo is 40-45 hours old (Diag. 3), the head is
somewhat sunken down in the yolk sac, and the head fold of the
amnion has grown posteriorly over the anterior part of the head. Thus
the head is received into a cavity which ends blindly in front and is
open posteriorly (Diag. 4). The dotted line of Diag. 3 shows the
limits of this cavity. The mesoblastic septum is now obliterated near
the dnus tenninalis but the backward oTOwth of the amniotic fold has
prolonged it in the posterior direction (M). The proamnion (P),
into which the extra-embryonic coelomic cavities gradually penetrate
from both sides, is now out of sight, concealed beneath the he;id and
the amniotic fold. In Fig. 1, and Figs. 57-51), the surface \iew
and the sections of the mesoblastic septum are represented.

In Chelonia the extra-embryonic coelomic cavities secondarily
insinuate themselves from both sides into the iimniotic fold, which
is at first purely epiblastic (See Figs. 2-4 and Figs. 17-10 of Mitsu-
kuri's article). In the chick the extra-embryonic coelomic cavities,
which are of course bounded by the mesoblast, spread themselves, from
the very beginning of folding, almost coextensively with the epiblast, (jr
in other words, nearly up to the posterior horse-shoe sha|)ed edge, so
tliat from the initial period, the dorsal part of the amniotic fold of the
chick does not contain an extensive area tree from the mesol)last,

1. E. Ravn : Ueber die Mesodermfreie Stelle in der Keimscheibe des Hükaerembryos.
Ai-ch. f. Anat. u. Physiol. (Auat. Abth.). 1886.

AXI) THE F(^K'rAL MEMIiKAXES, IN THE OHICK'. ^.\\

such ;is lias been übser\efl in Chelonia. Hut, still, we see the siinilaritv
betwo^en them in the pî-ocess of folding. From the tiinc when the
amniotic fold ha« C(jme to enclose some of tfie anterior part of the head,
the epiblast seems to grow backwards always just a little in advance of
the mesoblast, forming at the poslerioi- edge a mass of epiblastic cells
in the median line. Thus in Fi^-. 1 and ¥"i<r, {^) we see that a mass
of tlie epiblastic cells is inserted between the two sheets of the meso-
blastic septum. The same state is also o13ser^'ed in Chelonia when the
amniotic fold has advanced considerably over the d<jrsa] part of tlie
emljryo. Conse(|uently the chick differs from Chelonia only by the
fict that the early or solely epiblastic stage of the folding is abbreviat-
ed,— and with res])ect to this point it is highly interesting to notice thai
in the chick the anmiotic fold is raised much later than in Chelonia.

Tlie wedge-shaped e])iblastic cell-mass near the posterior edge,
inserted between the mesoblastic sheets, directly connects the ej)iblastic
layer of the amnion and that of the serous envelope, and thus is
seen at this stage quite the inci[»ient condition of the sero-amniotie
connection. Finally, the sero-aniniotic connection lies in the direct
prolongation of the line ol' tlie mesoblastic septum formed in
front of it.

^. I'rulofKjaliun of the Serd-^iiiiitiutlc CoiuucdDn.

From tlie time when the first trace of the sero-aniniotic connec-
tion has appeared, the wedge-shaped mass of the epiblastic <'ells in the
posterior edge of the head fold (referred to in the previous section) is
enormously thickened in tlie dorso-ventral direction especially ne;ir
tiu; median line. From this place the mass is gradually attlenuated
posiei'o-latei'ally along the horse-shoe sha[»ed edge (Figs. 1— o, 8<S— 4(1
and ()2-()5). Conse(juently along the posterior edge of the amniotic
fold there is t"orme(l a delta-sliaped epiblastic mass, which marks the

342 HIROTA ; ON THE SERO-AMNIOTIC CONNECTION,

posterior end of the .sero-uumiutic connection. The extru-enibryonic
coeJomic cavities of the tw(j sides ure sejKirated by this delta. In a
transverse section throuyfh the delta we see tliat the sero-anmiotic
c(3nnection is a very thick bridg'e between the outer and tiie inner
epi1)Jastic layers of tlie fold, and that the bridge is laterally constricted
from both sides by the niesoblastic slieets, so that the connection is
bi-concave (Fig. 60).

As the posterior edge of the amniotic fold with its epiblastic
delta grows constantly backwards, and the extra-em br}(jnic coelomic
cavities as (-onstantly encroach on the epiblastic mass of the posterior
edge, a sheet of cells (Fig. 90, s. a. c.) connecting the amnion and tlie
serons euxelope is alwaws left between the mesoblastic sheets of the
rwo sides, and thus the sero-amniotic connection or the ' Amnionnaht'
is prolonged backwards, so long as the posterior edge grows back-
wards. This goes on nntil the head fold meets the tail fold of the
amnion. The delta also urows somewhat larder in absolnte size in the
course of the backward progress (Figs. o<S-J:2, (iO, h'2, and (H)), Whil(;
these processes are being carried on, the curvature of the embryo takes
place (Figs. 1-5) and the sero-amniotic connection also l)ecomes curved
at the same time.

The mesoblastic septum, which has Ijeen formed in front of the
sero-amniotic connection, is gradually obHterated, and the extra-
emlji-yonic coelomic cavities of the two sides become continuous in
iront oï the epiblastic sero-anmiotic connection (Figs. 2-4).

Finally, in Chelonia tlie epiblastic delta is observed in the
[)osterioi' edge in comparatively later stages of the folding, l)ut iu
it the delta is not thickened dorso-ventrally so much as in the
chick.

AND TFTK FOETAL ^[EMP.RANES. TX TTTE r'TTTCK. 34^

3. Liniil of till' rroIniKialioH of llii' Si'vo-Amtiioiic
Convfcfio)!.

At the (Mid ol' tlio second day or o;u'iy in tlie third day ol' the
iiiciih;itioii, ;i ne^v amniotic fold arises hehind the tail end as tlie
])i'iinitive t:ùl I'old (Fiii's. '2 and ()1). The process of its folding is
(jiiite similar to that of the first stages (^f the liead fold. Bnt its
horse-shoe shaped edge is dh-ected anteriorly and there is never fonnd
a nv trace of the mesoblastic septum which has been observed in the
head fold. Early in the fourth day, while its cavity is still \'ery
shallow, it becomes fused with the head fold at tlie level of the still
rudimentary right hind limb (Figs, a and 4).

The seeming growth of the tail fold in the anterior direction is
due, in my opinion, not so much to actual advancement in that
direction as to the backward growth of the tail and its consequent
sinking into the yolk sac. If the head fold of the amnion arose a
little earlier or grew backwards a little more rapidly than it actually
does, it would reach the posteri<n* end before any tail fold had risen
and the case would be somewhat similar to that of Chelonia.

When the tail fold approaches the head fold, its anterior epi-
blastic edge becomes thickened into an irregular cell-mass, especiallv
developed in the median line. Thus the elliptic opening which
mai-ks the point of the final closure of the amniotic cavity, or the
' KommunikationsoiFnung ' of Schenk, is surrounded bv a thick
epiblastic ring, anteriorly deltoid and posteriorly irregular (Figs. 3. 4.
41. and 67). By degrees, as the epiblastic ring becomes thickened,
the diameter of tbe opening is reduced until the folds have entirely
coalesced, usually in 75-<Sö hours (Figs. 4, 5, and 42). Fig. 89 is
a transverse secti(jn througli the o])ening, when it lias just closed, and
when the epiblastic laiig is greatly swollen.

;> I ( nmOTA: ox THE SEKO-AMNTOTK' rOXXEPTTOX.

AVe thiLs see that the prolonGratioii of tlic sero-amniotie connec-
tion is st()))[)ed 1iy the closure of the amniotic cavity. Tliis is Ihe
))eri()d of the niaxininni (leve!(^pment of the cpiJilast/'r s('ro-ai]niioiir
(uniiit'i-lioti. At tin's time \t has (Fili". 42) in a dorsal view tlie a]>-
|>e:iran('(' nf ;i slender streak sliolitly cur\'ed, witli a concavity on tlic
riiilit side, and a swollen epihlastic mass at the spot where the am-
niotic 1"olds have coalesced (Fio's. 5 and 42). From this time on,
aithonprh cells in it may mnlti]>l\^ hy division, it does not seem to
enlargv as a whole — ;it any rate, in any oreat degree — but rather
grows less and less, being encroached uyion by mesoblastic cells, in a
manner to be explained liereafter. until by the tenth dav of the
incid^ation il has been entirely I'eplaced bv a xcrDiuhirii Dirsolilasfic
coiiiicclioii.

4. (^hiiiKjcs IN i]ir l\.rtr(i-J'jinhrii()iiii- (lennimil Lai/ns,

Every one who has stmlied the development of the chick well
knows, that from vei'y e;irlv stages the epiblast of the embryo proper is
split up into cell-strata. The up]>ermost layer is built of flattened cells,
while the »ujder layei* or layei's are composed of slightly columnar cells.
The stratification is not confined to the em])ryo j)ro])er, but graduallv
extends ceiitrifugally into the epiblast of the amnion. Fig. DO is
taken h'oni the dorsal part (fn' its exact locality see the explanation of
figures) of a noianal enibrvo of 7.S lioiu's. whose amniotic cavity has
just l)een closed ; and in this figure we see that the stratification has
advjuiced considerably beyond the lateral augle (A) of the amnion,
but that there nre as yet no distinct layers near the sero-amniotic
eoîuiection.

The extra-embryonic piu'ts of the coelomic cavity are at first
lined by a single e])ithelial la \-er of mesoblastic cells. From an early
|)eriod. howe\er, a I'eticiilnr tissue graduallv spi-eads itself outwards

AND THE EOF/I'AL MEMT.llAXES, TX THE ('HTOK'. 3^5

from th(^ body w:>li <>ii tliat side of the amniotic mesoblast which is
tnriied towards the e])il)last in the manner already described by Schenk/
Tliis net-work advances hand in hand with the stratification of the
epiblast, and tlie mesol)Iastic net-work also scarcely reaches the sero-
aniniotic coniiectii^n before the amnion closes np (Fig. 90).

The strati ti cation of the epiblast and the reticulation of the
mesoblast spread firther in later hours of the incubation. At the end
of the fourth day, the mesobListic net-work has already reached the
sero-amnititic connertion and |)assed along both sides of it towards the
serous envelo])e over which it also gradually spreads (Figs. 71 and
7.')). Distinct stratification of the epiblast is also found from the same
jK'riod in the amnion as well as in the serous envelope (see the black
layers in any of Figs. 70-S7). Thus, in Fig. 71, which is of this
stage, the previous fpiflu'lial reih of the coelom on both sides of the
connection are no longer distinct, and their place is taken l)y an
ill-defined reticular tissue which is intimately applied over the com-
pact epiblastic bridge.

These new changes in tlie connection are very important, for they
are steps to the formation of the secondary sero-amniotic connection,
which will be explained in the following sections.

It is necessary to say here a few words about the allantois. 'I'his
organ, which first appeared between the amnion and the serous en-
velope, on the right side of the embryo, about the time wlien the
timnion had closed itself, has in this stage come near the sero-amniotic
connection. As far as it has expanded itself, its vascular mesoblastic
tissue has fused, on its way, with the mesoblastic epithelium of the
serous envelope ; and so the mesoblastic net-work, ^vhich has spread
over the serous envelope, some distance beyond the connection is on

1. Loe. cit.

;^4(; UTIJOTA : ON THE SEKO-AMXTOTrO rONXECTTOX,

tlic right .side directly traceable to that of the allantois (Figs. 5. 6,
and l)n).

-0. Itt'jihiceiiieiit of Cells in t]\c Scrn-Jmfiiotic ('oniicctioN.

T liave mentioned in Section 2, that as the head-fold grows hack-
wards, the epiblastic delta of it« posterior edge is c(mstantly, though
only partially, being removed by the insinuation of the extra-enibryouic
coelomic cavities of the two sides ; and that the remnant of the delta is
prolonged backwards as the ' Amnionnaht ' or the sero-amniotic con-
nection. This process of insinuation prevails from the very beginning
of the formation of the delta, and is continued after the amniotic cavity
is closed. The mesoblastic net- work, having reached both sides of the
connection, begins to pierce across the remnant epiblastic cell-bridge,
so tliat at lenoth the mesoblastic cells of one side meet with those of
the opposite side, as we see in Figs. 72 and 74.

F'rom the sixth day the epiblastic bridge is broken uj) here and
there, beginning with the anterior part of the connection (Figs. 71, 72,
73, 74, and Figs. ()-7). In the posterior part of the connection, where
the amniotic cavity has finally closed, remains of the epiblastic cell-
mass, which there was the last to form, are found for a long time as
irregular patches, as in sections (Figs. 76-7 (S and Fig. 8). Sometimes,
the epiblastic cell-mass appears in sections as isolated ])atches iml)edded
in the mesoblastic tissue, as in Fig. 7<S, but by tracing them in other
secti(His they are found to l^jecome continuous with the (qjil)last either
of the amnion or of the serous envelope, lîy tlie ninth to tlie tenth
day of the incubation, the epiblastic bridge has been (•ompletely
replaced, along the entire length of the connection, by mesoblastic
cells (Figs. 4.S-50).

The epiblast of the amnion and of the serons enveIo])e are now
eiilii'clv discoiitiimoiis. lint the amnion and tlie serous envelope,

AND THE FOETAL MEMBRANES, IN THE CHICK. 347

in both of which the epiblast aud the mesoljlast are by this time
inseparably attached, are stil] comiected along the line corresponding
to the length of the sero-anmiotic connection hy means of insinuated
mesoblastic net-W(jrk. Thus the mewhlaüic or secondanj >iero-am.niotic
connection is formed.

6. Expan,s((jn of the Mesoblastic Connection.

The siin[)Ie replacement of the epil)Iastic Ijridge by the mesoblastic
net- work is not the sole event in the formation of the secondary sero-
amniotic connection. From the nature of the mesoblastic connection
its breadth is of course invariably greater than that of the previous
epiblastic connection. But, (jver and beyond this, the mesoblast of
the amnion and that of the serous envelope begins after the first week
of the incubation to coalesce for a certain distance on both sides of the
connection, so that in ten to twelve days, the connection is very much
widened and assumes the form of a plate, viewed from above. Fi^-s.
4o-50 represent the l^readth (jf the connection at different stao-es, eadi
projected on a straight line. Figs. 7G, 81, and 84 are transverse
sections of the secondary connection, through the approximately middle
level of the connection represented in Figs. 45, 4i), and 50 respe<--
tively.

If we examine sections of embryos in tlieir seventh to tenth dav of
the incubation, the ])rocess of coalescence is unfailingly witnessed. The
plate-like mesoblastic connection seems to be produced by the intimate
contact of mesoblastic cells (Fig. 75) in the narrow space on both
sides of the preexisting connection, similar to the coalescence of the
allantois with the serous envelope (Section 4).

The breadth (jf the secondary connection may, however, vary to
a certain extent in different individuals, and not necessarilv be pi'o-
})ortional to the age of the embryo (Figs. 4o-5()).

34S HIKUTA : OX THE SEllO-AMXIOTIC CONXECTIOÎf,

In some individuals a peculiar triangular sheet is observed at the
anterior end of the connection (Fig. 11). This is due to the following
circumstance. The anterior ])art of the connection (jften obtains a
considerable dorso-ventral extension, and thus produces a triangular
sheet whose base corresponds to its anterior free edge, and whose
sides correspond to the lines along which it joins respectively the
amnion and the serous envelope (Fig«. 11 and 79. Figs, (i and 70).
This triangular sheet does not stand edgewise, but (-(jmes to lie on
its surface and hence is seen from outside, as in Fig. 11.

Thus, the mesoblastic bridge, which is supposed by all writers
on the embryology of the chick U) be absorbed, making the extra-
embryonic coelomic cavity continuous from one side to the other,
is in reality greatly widened and remains to the last day of the
incubation.

7. Verforaiion in the Mcsohlaslic Connection.

From the eleventh day, the plate-like connection begins to be
perforated here and there by numerous round and elliptic ]:)ores, which
]nit the amniotic caA'ity in communication with the space outside the
serous envelope, so that the plate is now changed into a sie^e, and the
epiblast of the amnion and that of tlie serous envelope again become
continuous. This state of the connection is represented in Figs.
ol-5o. Each of these figures is of a view from inside the amnion,
part of which is represented in circular outline (Amn.) ; a tube-
like structure attached to this piece of the amnion is a part of the
alliumen sac which is lined by the serous envelope. How the albu-
men sac has assumed such a shape will be explained fully farther
on. It is represented as tilled with a l^lue mass. The membrane
which separates the amniotic cavity from that of the allîumen sac is
the sero-amniotic connection. It has nuiny ))erf ^rations which con-

AND THE FOETAL MEMBRANES, IN THE CHICK. 349

iiect tlie two cîivitiey, mid appears «ieve-like. Figs. 85-87 are from a
series of sections throiio-h a perforated sero-amniotic connection of the
twelfth day. In Fiir. 85, which is the most anterior represented, the
connection is entire and intact. In Fig. 86 it is very thin. In Fig.
87, there is a perforation.

These pores in the plate-hke connection grow larger in size in
later days, while the niunl^er becomes less by the running together of
some adjacent ones. At length, after the sixteenth or seventeenth
day the two sides oi' the plate are connected only by a few sti-ands
which remind us of the cliordae tendineae of a mammalian heart
(Figs. 54-56).

Thus the epiblast of the annii(jn and that of the serous envelope
again become continuous ; the exti-a-embryonic coelomic cavities of
the two sides are as completely separated from each other throughout
the entire extent of this sero-amniotic connection as in any previous
stage.

6'. SuppU'iiieni to tlic I'recediiKj Sections.

1 will now make some necessary remarks on other points Avhicli
it was convenient to witlihold until I had traced, in the preceding
sections, the main course of the changes in the connection itself.

In the extra -embryonic part, near the sero-amniotic connection at
least, the epil)lastic cells can usually be distinguished from the meso-
blastic cells by the following points. In form, the mesoblastic cells
are flattened in the plane of the layers themselves, wherever the
cells are densely disposed, and are stellate, wherever the number of
cells is very small, wliile the ejnblastic cells are mostly ])erpendiculai-
t<j the phine of the layei's (Figs. 75, 80, 85, and DO). Conse-
(piently the mesoblastic tissue is schistose or reticulated, while the
epiblastic layei's are in more or less regular strata and compact.

350 HIKOTA; ON THE SEKO-AMNIOTIC CONNECTION,

On this a,cc(niiit the two kinds of germinuJ layers are differently
stained in the exti-;i -embryonic region.

These differences are, however, not apparent in all cases. In
later stages, both the layers are intimately united and in some places
they can not l)e satisfactorily distinguished from each other. One
may even come to doubt whether the epiblastic bridge is replaced by
the mesoblastic tissue. Ikit I have many preparations in •which there
can not be any dou1)t on this point. For instance, in the sections
given in Figs. 71 and I'l, taken from the stage rei)reseiited in Fig. 6,
in which the epiblastic bridge has begun to be invaded by the meso-
blast, the one (Fig. 71) has the epi blast bridge still intact, while in
the other (Fig. 72), which comes behind the first, there is distinctly
no trace of the bridofe, althouo-h it must have existed here earlier.
The same state of things is observed in the case represented in Fig. 7
and Figs. 78-74. From these cases we may be certain that the
epiblastic cells have been entirely \\ithdrawn fi'om the prcAious bridge
and that the netted tissue is purely mesoblastic, as I have mentioned
in Section 5.

That, near the posterior edge of the head fold of the amnion, an
epiblastic cell-mass is produced was observed by Schenk in 1(S71.
Of the fate of this mass he says : *' So bin ich geneigt anzunehmen,
dass die Zellen, welche die Verdickuno- ausmachen, l)is zum einem
bestinnnten bleibenden Theile derselben nach und nach in Liipior
Amnii aufgehen, und ihre Zerfallsprodukte scheinen zum guten Theile
an der Embryonalleibe ihre Verwendung zu finden." Th.\< inter-
pretation induced me to look carefully foi' the supposed processes of
dissolution, l)ut I have been unaljle to confirm his idea for the follow-
ing reasons.

I have examined sections <jf nearly three dozen specimens of
embryos from the second to the tenth day of the incubation, and

AXr> 'I'lIE F(M^yrAL membranes, TX 'I'HE (""ITTOK.

:;r)l

hfive liecii ini;il)le to find :i single case like that represented in Fiii'. 4
of Schenk. On the conti'ary, the epililastic cell-mass seems to me to
have ;üwaA> a definite h(Minda,ry on the amniotic side. Often we find
il C(^aguhnn well stained with borax carmine, attached to the ('ell-mass,
but a similar coagulum is fonnd in other spaces, snc^i as the central
nervous canal, the alimentary canal,, and also in the coelomic cavity.
In Fig. (SS we see the cell-mass projecting into the amniotic cavitv.
and in Fig. 7(i the renmant of this projecting mass is I'epresented.
These are, however, extreme instances and in ordinary cases the cell-
înass. as a Abhole, is only slightly sunken into tlie amniotic cavit\-
(Fig. ()6). Such sinking into the amniotic cavity is not to lie wonder-
ed at when we consider the weight of the mass itself.

As I have described in Sections 2 and 5, the epiblastic cell-mass
is being ^constantly foi-ined and constantly removed from the very
beginning of the formation of the connection, and being situated near
the free edge of the head, fold, the product of its dissolution, if it wei-e
actually dissolved, would diffuse wddely through the surrounding
albuminous fluid, and but little'of it be mingled with the amniotic
fluid, which is speedily increased in volume, evidently from othei-
sources.

Finally, from the physiological point of view, it seems to me to
be improbable that the epiblastic cells dissolve nj) for the use of othei"
epiblastic cells of the same surface.

From the preceding discussion, and from the fiict that from an
early stage of folding the deltoid cell-mass is constantlv added to at
the posterior free edge and constantly reduced neai- its anterior apex,
it is probable tliat the cell-mass is used u]) foi- the extension of the
surface of the fold, and not for any otlicr special pui-pose. It is,
besides, a noteworthy fact, that in Chelonia tlie deltoid area is not
thickened dorso- ventral ly -iiid undergoes, of course, no dissolution.

HTT^OTA : ON THE SERO-AMXTOTTr' fOXXEr-TTOX.

0. Effects of the Sero- Amniotic CoiincctiôH on the
Foetal 2Ieui]ir(ines.

About the time when the amniotic cavity is closerl the allantois
appears on the viulit side of the embryo in front of the level of the still
rudimentarv right hind hmb ; and it spreads itself graduully between
tlie serous envelope and the amnion.

The first structure which we notice, and which seems to owe ka
ori""in to tlie presence of the sero-aniniotic connection, is a curious
infolding- «^f the amnion near the connection, given in Diag. 5. It is

Diag. 5.

Yolk.

Transverse section of an embryo thront^h
the alhintois and tlie fold.

Diag. 6.

;Mio Oonneotian.

Tjontjitudinal section throui;-li the
sero-a mniotic connecta* >n.

alw^ays on the right side of the eonnection and is pushed on towards
the left. There takes place no folding before the allantois appears, and

AND THE FOETAT; MEMBRANES, TX TEE rHK'K. 353

fho longitudinal extent of the fold depends on the length of tlie sero-
araniotic connection. The form and extension of such a fold, vary
greatly, however, in different individuals and at different stages.
One instance of great development is given in Fig. 37, and its sections
in Figs. 91-93. The planes of these sections are indicated by dotted
lines in Fig. 37; in wliich the points, A, B, and C, on the dotted line
marked 92, correspond t(^ the points marked hy the same letters in
Fig. 9l^ It is not clear what significance this fold has.

At both extremities of the sero-amniotic connection the amnion
is also slightly folded longitudinally (Diag. 6) ; transverse sections
oi" such folds are represented in Fig. (i(S (at the anterior end) and Fig.
()9 (at the ])Osterior end). Tn these figui'es, above the general am nicotic
ca^ ity there appears a second smaller cavity marked Amu. A reference
to Diag. (i will show that this appearance is due to the folds. How-
ever, these transverse and lonor-itiidiiial folds, beinçf disturbed bv the
oTowth «^f the amnion as well as [)v the chano-es of the sero-amniotic
connection itself, become insignificant after the first week of the
incubation.

These temporary effects upon the amnion are, however, fol-
lowed by others much more important on the allantois and also on
the albumen-sac. The albumen-sac of certain birds, and the yolk
sac of the chick have been studied by Duval ^ and recently by A^irchow,"
but knowledge of the general development of the foetal membranes of
the chick is still left very imperfect. Hence, it may not be superfluous
to give here a l:>rief outline (^f the development of the foetal membi'anes

1. M. Duval : Études histologiques et morphologiques sur les Annexes des Embryons
d'Oiseau. Journ. de L'Anat. et de la Physiol., XX. 1884.

2. H. "^^irchow : Der Dottersack des Huhnes. Internationale Beiträge zur Wissenschaft-
lichen Medizin. Bd. 1. I regret that I know this article only by extracts given in Ergeb-
nisse der Anat. u. Entwickl. (1892 ; Wiesbaden), and Zoologischer Jahresbericht 7.ur 1891
(I89b; Berlin).

3,5 ( HTROTA: OX THE SERO-AINIXIOTTO rOXXEOTIOX,

themselves, in ordei- tliat the influence of the sero-Mmniotic connection
iipc^n them m.'iy he l)ei"ter understood.

Wlien nt the end of the tliird dny or early in the fourth day, the
allantois ap))ears on the rio'ht side of the emhryo. hetween the serous
envelope and the amnion, it is a discoidal sac with circnlar outline.
At the end of the fifth day we can distinü'in'sh in a surface view the
riüht and left allantoic arteries and one larsfe allantoic vein in the
allantois (Fia*. 7). The right artery, which is always i)ifiircate in the
inner limhof the allantois, supplies the posterior part of the allantois.
while tlie vein is divided into two or three main hi-anches in the
outer limh. The left artery is smaller than the otlier two \cssels, and
is destined to supply the anterior part of the ;dlantois.

From this stage the allantoic vein begins to indent the pre-
viously circular maraiu of the allantois at the point where the
\ein ]iasses fr<^m its outer to its inner lind). The obstruction by
the vein restrains and retards the growth of the allantois at this
point, althougli it does not stop it entirely. Contiguous to this
point, therefore, the allantois grows fister than at the ])oint itself
and thus an indentation is produced. With the growth of the allan-
tois, the indentation becomes of great depth in later days, and the
lobe of the allantois which lies in front of it comes gradually to
overla]) the lobe posterior to it, beginning from tlie a])ex of the
indentation (Figs. <S, 9, and 10).

The riglit and left arteries also have a tendency to restrain the
free gr<^wth of the allantois at tlie points Avhere tliey ]iass from the
inner to the outer limb. The eifect is, however, (piite trivial in
comparison with that of the allantoic vein.

It is noteworthy that there is an intimate relation between the
allantoic vein and the sero-amniotic connection. The allantoic vein
always restrains the free growtli of the allantois at a ]ioint a little fo

AND ruE Foetal membranes, in the chu'k.

855

(lie figlit of the coiiijecti<Jii (Figs. S, 1), jiikI 11). Whether the sero-
aiiiniotic connection, towards which th(; alJantois s[)re!ids itself, is able
to persist by the presence of the allantoic vein, or the niargiji of the
allantois is specially indented by that vein on account of tlie presence of
the sero-aniniotic <'onnectioii, is a question which can not be answered
at present.

The overlapping of the allantoic ]ol)es which began at the apex <,)f
the indentation and spread gradually [»eripherally (Fig. 10) progresses
without any hindrance, until it reaches the sero-anuiiotic connec-
tion. As this otfers an impassable

Diag. 7.

A Longitudinal Section.

Diag. 8.

^eia

Allantoic cavity.

A Surface \'ie\v.

Egg in the 'Jth Dii,y of luculjatiou.

obstacle, the allantoic lobes with
their serous envelopes rise alcove
the level of the sero-anmiotic con-
nection, and pass beyond it, pro-
je(.'ting like the eaves of a roof.
Diag. 7, which is a longitudinal
secticjn thrijugh the plane oi' the
overlapping, shows how tlie allan-
tois has risen above tlie sero-
aniniotic connection and passed
lje}ond. There is thus jtroduced,
above the yolk sac and below the
allantois, a sort of cave, at whose
blind end is the sero-anmiotic
connection. Diau-. ,S yives a sur-
face view of these ])oints. The
part marked with lines slanting
to the left shows the area of over-
lapping, wiiile tiie part niai-ked
with lines slantinu' to the rii-ht

35 G HIEOTA ; OX THE SERO- AMNIOTIC CONNECTION,

shows the extent of the cave. The line at which the cave comes in
contact with the amnion represents the sero-amniotic connection.

Tlie two lobes of the allantois that overlap do so with their accom-
panying serous envelope, so that there ought to be sheets of epiblastic
cells between the lobes, witli the mesoblast and the hypoblast lining
them. In point of fict, it becomes impossible to distinguish these
layers, and only a simple septum, whose nature is not clear, divides
the cavities of the two allantoic lobes. The epiblastic sheet is visible
only near the outer limit of the septum (Fig. 82). In future I shall
call this the ' Inter allantoic Septum.'' It has a free edge aloug which
the allantoic vein runs, and is, for the time, somewhat heart-
shaped. At tliis stage other two minor interallantoic septa are
formed by the right and left allantoic arteries in their respective lobes
(Fig. 12). These are much smaller than the primary septum but in
other respects show no essential difference. The amniotic cavity
is now much enlarged and the yolk-sac much reduced. In fact, it
is at this stnge that the epiblastic sero-amniotic l)ridge has become
ahnost entirely replaced by the mesoblastic tissue. Fig. 12 represents
the surface view of the foetal membranes on the ninth day and Fig.
13 the vicinity of the sero-amniotic connection. In this particular
example the amniotic cavity is drawn out towards tlie connectiou,
and the cave (Fig. 13 DABC) is narrower and longer than that in
Diag. 8.

With further growth, the amniotic cavity becomes enlarged, the
yolk and the albumen-sac diminished, the allantois extended, and the
interallantoic septum widened. Through these changes, occun-ing
simultaneously, the foetal membranes reach in later days a degree of
complexity, which well nigh defies description. 1 will only atteiii})t
here to explain the arrangement of the foetal membranes in an embryo
of the tenth day by means of several figures and diagrams which are

AXn THE FOF/l'AT. ^[EMl^RAXES, IX THE CHICK. 357

madt* so us to supplement one imother. Fig. 14 is a surface view of the
membranes when the shell is removed, m which, as usual, tlie allantois
is coloured oreen. Fii>-. 15 iiives the outlines of the memhranes with
the folds in the allantois shaded. These two figures are made exactly
alike so that they may be superposed. Fig. 1() gives the vicinity of the
sero-amniotic connection on an enlarged scale. Flo-. 17 is intended to
give the diagrammatic rejiresentation of the allantois with its folds and
the interallantoic septum. In Fig. 18 the position of the amnion,
the yolk-sac, the albumen-sac, and the sero-amniotic connection is
given in addition. In the last two figures the outer lin^b of the
allantois is represented as extensively removed, so as to show the
inside of the allantoic cavity, and, in addition, the small end of the
egg is figured as nearly cut off from tlie main portion of the egg and
reflected to the left. In all the figures the egg is in one position,
tlie anterior lobe of the allantt^is occupying the top of the figures and
the posterior lobe, overlapped by it, appearing in the lower porti<m.

At this stage, tlie allantoic lobes have not }et closed entirely o\er
the small pole of the egg so that the albumen-sac peeps through for
a small space between them at that end. How the albumen-sac
has thus come to lie ^^ithin the allantoic lobes is easily understood
if we imagine to ourselves the ])rocess explained in Diagrams 7
and <S to continue until the allantoic lohes have extended them-
selves to the small pole of the egg. Within the space thus enclosed b}'
the allantoic lobes, the three structures: the amnion, the yolk-sac, and
the albumen-sac, are disposed, roughly spe-aking, in three stories (Fig.
IN). The amnion with the enclosed embryo proper occupies the upper-
most story at the larger pole of the egg. lîelow it comes the yolk-sac
rallier discoidal in shape, and then the albumen-sac. The perpendicular
.<■//. (Fig. I'S) axial to the three stories does not ('(jincide with the axis
of the egg thi-(jugh its poles, but is incliiied to it as shown in tlie

;>5,S HIROTA: ON THE SERO-AMXIOTIC CONNECTION,

fio-ure. The ulbumeu-sac .separated from the aiiiiiioii foi' the most
[»art hy the intervening yolk-sac, «ends out o\'er the latter a hodow
cave-Jike diverticulum towards the amnion (see Figs. l-S and l<)j,
with which it c<jnies in ct)ntact. The contact plate at the lu^ttoni
of this diverticulum is the sero-amniotic connection (S. A. C,
Fio-. IS and K! : sections in Figs. 80 and <S1). In fact, it is
the presence of the sero-amni(jtic connection that lias j)roduced the
peculiai" cave-like diverticulum. Let us now turn to the allaubjis
itself. '!<) understand the complexities arising in this membrane,
it is necessarv to bear clearly in one's mind the facts whicli liave
been brought ont in tlie preceding pages ; (1) that the allantoic vein
in passing from tlie inner to the outer limb of the allantois causes
an indentation in the margin of the allantois, retarding its growth
at this point, and the lobe of the allantois in front of the indentation
comes t(j overlap rhe lobe posterior to it, the area of overlapping
being continuously increased with age; (2) that a terminal part of the
left allantoic artery and a branch of the right allantoic artery produce
respectively a minor indentation in the margin of the allantois. We
nmst therefore kn<j\v well how these blood\'essels run in the allantois.
This may be learned by com|jaring carefully Fig. 14 and 15. First
notice in Fig. 14 where all the allantoic vessels come out together
Irom the umbilicus (for the exact position of which see also Fig. 17),
tlie allant(3ic vein runs from the umbilicus to the point F (Fig. 15),
along the inner limb ; then from the point F gradually ascends along
the line FE to the outer limb on which it: emerges '<^^t the point
E. The line F E nuist therefore be regarded as that along which
interference witli growth takes ])lace, and the shaded areas F D E, and
EGM DE (Fig. 15, for the present disregard the area ABC) together
represent the area of overla{)ping of the two allantoic lobes, or in
other words the extent of the interallantoic septum. The reason

AND THE FOF.TAL MEMBTIANES, TX THE THTOK.

:;')9

B

wliy this is in two nreas FDE niid ECMDE is that tlic
iiiterallnntoic scptnm Iims heeji ioklcd in two ülonn' the :ixis E D.

A reference to Di;ig. il will
°' ' lielp to make this as well

as some other points clear.
Lettering- in the diao-ram
is as far as possible the
snme ;is in Figs. 15 and 1 7.
Tu A. the rod F E, vvhidi
represents the allantoic vein
in its passage from the in-
ner to the onter limh, is
enfolded l)y a piece of |>a]ier
so that it stands at the head
of a deep fissnre or incision
(MEFÜ). FF FM S re-
presents the anterior allan-
toic limb, ÜFEMT the
E posterior lind), and E M V F
the interallantoic septnm.
in W, the area E M ü F is
folded along the line K D
so that it a])pears now as
1 w<^ areas, FDE and E M U D E. The triangidar area E D K is two
layered, and the corresponding area in Figs. 17 and 1<S is painted
blacls. Tlie line EM corresponds to tlie line ECM in Fig. 15 and
E C Fl in Fig. 1 7. The line F D Ü corres])onds to the line FDM in
Fig. 15 and FDG in Fig. 17. 1 ho|)e 1 have now made clear the
disturbances cansed by the allantoic vein and the ndafioiis of the
interallantoic se])t iim.

f>ßQ HIROTA ; OX THE SERO-AMXIOTIC CONNECTION,

Let US now torn to tlie two arteries which c;uise disturbance in
the allantois. One of them is the terniiiial part of the left allantoic
arterv. This artery, emerging from the umbilicus, runs along the
inner limb towards the right in Fig. 14. It leaves the face repre-
sented in that figure on the right side low down, and running alorig
on the other face, it enters the fio'm-e as^ain at the rio-ht side above. Ft
runs all this while along the inner Hmb and continues to do so until
it reaches tlie point A in Fig. 15. From this point, it begins to rise
along the line A V) and emerges on the outer limb at the [ioint 15.
The terminal portion of the left allantoic artery in thus passing from
the inner to the outer liml) causes the same sort of disturbance as was
made by the allantoic vein, but to a, smaller extent, and produces
I he fold AlUJ (in Figs. 15, K^i, 17, and IS) which is marked on the
outer allantoic limb by the line C B. This is therefore a sectmdary
disturbance of the interallantoic septum.

Another minor septum is produced by a Ijranch of the right
allantoic artery. This artery proceeds from the umbilicus towards the
left in Fig. 14 along the inner limb until it reaches the })oint Z (Fig.
15), giving off branches on its way. At the ]joint Z it begins to leave
the inner limb and rising gradually along line Z Y reaches the (jutcr
limb at the point Y, from which it is distributed on that limb. 44iei-e
is thus produced a. mesentery -like f )ld, Z YS,' of the allantois embrac-
ing the vessel along the line Z Y. (Cf. Diag. 9, A.) A secondary
complication is produced by the fact that tlie surface of the fold is
slightly curved, aiid is two layered in the area W Z Y (Fig. 15).

There are many individual variations in the configuration of
the primary and minor septa, but the egg explained above in full
may be taken as tlie ty])ical one, and all the other cases can be ex-

1. A minor scptniii. ciuisod li.v iinothcr Imiuch of tlie riolit alhintoic artery, which proiliiccs
still further eouii)lic;ition in this septum, is here. t'<>r the sake of simplicity n^'t referred to.

AS\^ THP] FOETAL MEMBRANES. TX THE THTCK. ^Ql

plained as iiioditicutions of this fm-iii. Thus, in Figs. 19, 20, and 21,
from an embryo of tlie eleventh day which correspond respectively to
Fiii's. 14, 15, and 1(5 of the ])recedini>' stage, the terminal portion of the
left allantoic arterv whicli runs alonu' the edofe AB in Fisf. 15 and 17,
runs along the axis on which the interallantoic septum is folded, and
consequently the septum formed by the left artery alone is very
small (ADK Fig. 21). Besides, in this example, the right allantoic
artery, running along the inner allantoic limb, is divided at the
point Z into two branches which, running along the free edges ZY
and ZX, emerge on the outer limb at the points Y and X respec-
ri\ely, and thus a somewhat complicated minor septum ZvSYXZ (see
also fig. 20 his.) is ]Droduced by that artery.

By the eleventh or tAvelfth day the aJlantoic lobes come together
at the vegetable poJe of the egg. Consequently, the albumen-sac
or the ' .s'rtc placentoide ' of Duval is entirely cut off from the sur-
rounding space, as Duval says but contrary to Virchow's observation,
while its cavity is lined l)y an epil)lastic layer which has been con-
tinuous with that of the outer limb of the allantois. The leno^th of all
the allantoic septa is determined by the closure of the albumen sac,
Mud nil come together at that point where the allantoic lol^es finally
close.

It is at this stage that the perforation of the plate-like connection,
before referred to, begins and it is a remarkable fact that for a few days
after, the amniotic fluid soon coagulates in alcohol or in Kleinenberg's
picro-sulphuric acid, just like the fluid in the albumen-sac. This
seems to be due to the presence of albumen which has found its way
through the perforations into the amniotic fluid.

;;g2

HfROTA; OX THE SERO-AMXIOTIC rOXXEOTfOX,

Diaé. 10.

Interall, Septum

Part of a Cross-Section
of the Albumen-Sac.

As fasf :is the albiinien is absorbed the primary interallantoic
septum widens itself inwards at the expense of the inner limbs of the

two allantoic lobes, which progres-
sively coalesce with ench other as the
albumen diminishes nnd the albu-
men-sac contracts in volume. A
reference to Diag. 10 will make
what is meant clear. Suppoae the
outer limit of the albumen-sac to be
the line 1-1. It is bounded by the
two lobes of the allantois, and the area
witliin which they overlap (a b) is the internllantoic septum. If now
the albumen-sac decreased in diameter to the line 2-2, the inter-
allantoic septum would then be widened to the point c, by the coales-
cence of the inner limlîs of the two allantoic lobes. Thus the decrease
of albumen will make the width of the interallantoic septum increase
and the diameter of the albumen-sac grow smaller. Hence, the
line along which the interallantoic septum is (continuous with the
inner allantoic limb (appearing as the points b, c, d, in a cross-section
like Diag. 10), will always lie on the albumen-sac. The line AA
represented in Figs. 51-56 is, in fact, this line. The i-elation of the
inner edge of the inter:dlant(nc septum (DKG) in Figs. 17 and LS
to the cave-like diverticulum of the albumen-sac will make this evi-
dent. These changes are accompanied by gradual enlargement of the
amniotic cavity and gradual decrease of the yolk. Tiie stage in
which these conditions are observed is given in Figs. '22-'2b, of which
the first two re])resent surface views of the foeral membranes frcnn
two opposite sides; and the last two give theii* outlines. The letters
as far as possible refer to the same parts as in Figs. 14-lN. Tlie
bloodvessels first become visible in passing from beneath the amnion

AXD THE FOETAL MEMBRANES, IN THE CHICK. ßfj^

(Figs. 23 and 25) : the allantoic vein and the Jeit allantoic artery on
one side and the right allantoic artei-v on the other (Z, Fig. 25). The
allantoic vein passes towards the right along the inner limh (Fig. 28)
to the point marked F on Fig. 25, then passes from the inner to the
outer limh along the line between F (Fig. 23) and E (Fig. 24)*
(like the line FE, Fig. 15) and emerges on the outer limb at the
point E (Figs. 22 & 24). The full line EG corresponds to the
line EM in Fig. 15 and the fold FEOMF to the fold FECMDF in
Fig. 15. The right allantoic ;u"tei-y stîu-ting from the point Z (Fig.
25) runs towards the left, but soon divides mainly into two branches
(Fig. 23), each of which together with a third smaller one produces
a fold in the allantois thus making three minor septa. No septum
formed by the terminal portion of the left allantoic artery is present in
tliis case. All the septa above referred to come together at the point
marked 0 (Fig. 24). Fig. 2() represents the allantoic septa which have
thus come together, as seen from tlie vegetable pole. Full lines show
the tracings of these septa on the outer limb of the allantois. In
addition to the |)i-imary interallantoic septum (E'O) there are tliree
minor sej)ta. formed by three branches of the right allantoic artery
which run along the free edges ZX, ZY, and MN. Notice also how
the albumen-sac, now pit(;her-sliaped, has decreased in size.

l)y the sixteenth day yolk, as well as albumen, is much reduced in
volume and the princi])al part of the e^xg is now the amni(^n occupied
by the embryo proper (Figs. 27-30). Fig. 31 represents the septa (only
three in all in this case) come together at the point F, as seen from the
vegetable pole. In this figure we see that the right allantoic artery is

* The reiider will understand that in tracing a bloodvessel or other structure from the
surface rejj resented in Figs. 23 and 25 to that given in Figs. 22 and 24, the upper edge of the
former two figures should 1)P applied to the lower edge of the latter two figures and also that as
eggs have a considerable thickness, a structure leaving one surface may not l)e found at exactly
the corresponding point of the other.

3ß4 HIROTA ; ON THE SERO-AMNTOTIC CONNECTION,

divided into two brnnrhes at the point Z (cf. Figs. 14 and 23), and
that the two branches ascend directly, along the free edges ZY and ZX,
townrds the outer limb. At this stage the albumen-sac has assumed a
peculiar elongated form, and the ' omhilic omhilicaV of Uuval, by which
the albumen-sac is united to the yolk-sac, has l)ecome conspicuous and
easily observable. The sac is supplied Avith numerous blood vessels on
the yolk side, and albumen is absorbed through that singular placental
structure discovered by Duval in the linnet. The position of the
umbilicus may vary in different individuals (Figs. 29 and 34), but
the papillatcd placental structure of the albumen sac is always con-
fined to the vicinity of the umbilicus. It is to be added that from
this staçfe the blood vessels of the outer allantoic limb undero-o certain
changes in their distribution. One Ijasal branch of the allantoic
vein, which runs with the right allantoic artery, becomes conspicuous
by anastomosing with other branches of the same vein (Figs. 27, 2(S,
32, and 33).

On the eighteenth day, the albumen-sac appears as a slender tube
which opens into the amniotic cavity thnjugh the perforated connec-
tion (Figs. 32-35). Fig. 3(3 represents the septa come together at the
])oint P as seen from the vegetable pole. [n this case the spot
where the riofht allantoic artery divides is concealed beneath the
yolk-sac, and the two branches ascend along the free edges ZX
and ZZ'Y respectively towards the outer allantoic limb. In this
example the primary interallantoic septum has iKjt l)een folded along
an axis in the allantoic cavity (cf. Diag. 1), lî), and moreover is not
influenced by the left allantoic artery.

At length, the albumen becomes entirely absorbed and the vitel-
line membrane which had previously been imbedded in albumen
is alone left in the former albumen-sac, often partly ])rojected into
the amniotic cavity through openings in the sero-amniotic connec-

A.VD THE POETAL MEMBRANES. IN THE CHICK. 3^5

tion. The yoJk-sac becomes then enclosed by the abdominal wall and
the entire s])ac(' of the egg becomes filled by the amnion which is
occupied by a fuJly orown embryo.

Contrai-y t(j the observations of some writers, the Jeft aJhmtoir
artery is found to [)ersist. th(ju;^-h faintly, till the last day (Figs. 27,
2S, o2, and o3), while the allantoic vein and the right allantoic artery
are of great functional impoi-tance throughout the allantoic life. The
inner allantoic limb is always sup|)lied by much fewer blood vessels
than the outer limb.

Finally, the sero-amniotic connection is quite without relaticju
to the emergence of the embryo, which pushes out its beak into the
air chamber through the point of the membrane far distant from tlie
connection.

10. Comparison betiveen llw Sero-Amniodc Connection
of the Chick and lliat of Chriniia.

'I'lie sero-amiiiotic connection of the chick, as des('ril)ed above, and
that of Cliclcjnia, as studied by Mitsukuri in Clrnnni/s und Trionyx, mav
be compared as follows : —

Tlie essential points in irliich fJiei/ atjrce. — a). In the chick, there
is found at the p(Jsterior edge of the amniotic fold a constant deltoid
area free from the mesoblast. The same sti-ucture is observed in
Chelonia in comparatively later stages of the progress of the fold
backwards.

/)). As long as the amniotic fold grows p(3steriorlv the sei'o-
amniotic connection or the remnant of the epiblastic deltoid ai-en is
equally prolonged posteriorly until it becomes a long string.

c) The sero-amniotic connection is more or less widened on
both sides of the epiblastic bridge by the entrance of the netted meso-
blastic tissue.

3ßß HIROTA; ON THE SERO-AMNIOTIC CONNECTION,

d). The epiblaatic layers of the extra-embryonic parts are stratifi-
ed, at least near the connection.

e). The connection between the serous envelope and the amnion
persists durino- egof-life and no direct continuity is ever made between
the extra-embryonic coelomic cavities of the two sides along the length
of the connection.

f). The growth of the allantois is greatly influenced in later
stages by the presence of the sero-amniotic connection.

Thi' essential points in which they disaijree. — a). In the chick the
amniotic fold arises in comparatively later stages than in Chelonia, and
the extra-embryonic coelomic cavity is from the first extended backwards
in the amniotic fold almost as far as the epiblast has progressed, while
in Chelonia the mesoblastic cavities insinuate themselves secondarily
from the two sides in the latero-median direction. It is owing to this
difference tliat in the amniotic fold of the chick there is found no
conspicuous area free from the mesoblast, and that there is no meso-
blastic septum in Chelonia before the epiblastic sero-anuiiotic connec-
tion appears.

h). In Chelonia the epiblastic delta of the amniotic fold is
a.lwavs larger than that of the chick, but in the latter it is enormously
thickened dorso-Aentrally.

c). In Chelonia the anuiicjtic fold passes over the tail end before
the lattei- is yet differentiated and there is therefore formed no proj»er
tail fold. In the chick, the head and tail folds fuse together at the
]e\ el of the rudimentary right hind limb and therefore there is pro-
duced no trace of the posterior tube, which forms so (conspicuous a
feature of the chelonian amnion.

d). The epiblastic bridge of the connection seems to be persistent
in Chelonia, while in the chick it is replaced by mesoblast and becomes
extended on both sides.

AND THE FOETAL MEMBRANES, TN THE OHTOK-. '^^J

e). Thei'e. tîikes plficc no ])eri"or:ilioii of the .scro-amniotic con-
nectioM in Clielonia.

f). The sero-amniotic coiiiiectioii is quite witlioiit relation to
the «^uieroeiice of the yoniio- in the chick.

g). Tliere is produced no alhumen-sac in (-helouia and therefore
the sero-amniotic connection is not enclosed within the foetal mem-
branes. Consequently the allantois of Chelonia is very dilterentlv
affected by the sero-amniotic connection.

77, Methods for the Preparation of Spechiiens.

From the second week of incubation it becomes ahiiost hoj)eles.s \o
try to take out the contents of an e^^^ entire in the fresh condition,
and the whole eg'g- con tents wliould tlierel()i'e be hardened, in the fol-
lowing manner.

When ail incubated ^gg is looked at by transmitted light it should,
to be a promising one, be semi -opaque after the first week and entirely
so after the first ten days. Having obtained such an egg it siiould
be patiently tapped with a glass rod or something of the sort, until
the greater part of the shell is broken into small pieces, the shell then
taken off, piece by piece, leaving tlie underlying shell-membranes
intact. From this point the proeeduiv will be somewhat different
according as the egg has been incubated more than two weeks <m- less.

a). lllieit tjie e<fif luis been in inciihation less than tiro weehs.
Care should l)e taken not to injure large blood vessels of the allantois,
which is closely attached to the inner shell-menil)]'ane especially near
the large pole: whenever they are injured by accident, the blood
may be cooled by bhjwing at the point, when coagulation soon sets in.
The shell is taken off in this way over an area about one inch square,
and an opening made into the air chamber, the position of which is
easily found out by the sound on tapping.

•>(;(s; HIROTA : ON TTIR SERO-AMXTOTTr rTjNXErTTOX,

The eii'g i« then \mt for a few minutes in l\leineiil)erg'« picro-
sulphiiric acid in :i cup of moderate size, to partly dissolve and
soften the cMlcareous shell. The shell membranes then become
easily separable from the allantois, mid the whole contents can he
entirely freed from the Jion-cel hilar envelopes. At this stage, the
contents being yet soft, a large piece of the shell, disunited from the
contents, shcMild be left to suj)port them in their relative positions
until they have become hardened in Kleinenberg's fluid. Xext day,
the hardening fluid is replaced by ;ilcohol, by taking it away in small
portions by a ])ipette from the bottom of the cup, then gently pouring
on alcohol (70 ^/o) so that an alcoholic layer rests on it and re])eating
this ])i-ocess until it has been entirely replaced by alcohol. \)y this
method Kleinenberg's fluid is easily and economically rej)laced by
alcohol without disturbing the contents.

h). Whe?! the egg has been in incuhation mmr tJuni tiro irtuis. There
is liere no dangei* of injuring the lilood vessels, for the calcareous
shell is now brittle and easily separable from the outer shell -membrane
which is white, dry. and leathery. Besides, the contents are now
firm and consequently the shell can be taken oif very extensively,
and even frimi the entii'e sui-face. Howe\er. the serous envelope
being intimately attached to tlie inner shell -membrane, it is almost
impossible to separate them in the fresh state witln^ut bleeding.
But if we put the whole in Kleineijberg's fluid for ten to thirty
minutes, there is no longer anv danger. When, however, we wish to
examine the cellular structure of the serous en velo])e, the inner shell-
membrane should be left in position, for the safety of the underlying
layers. When tlui eontents are thus entirelv cleared from the non-
cellulai- envel()|)es, the specimen is put aside in Kleinenberg's fluid for
about half;i day. In this case, the contents, being firm the harden-
ing fluid (^iin be easilv repinced by alcohol after it has done its \voi"k.

AND TFIR FOETAL MEMI'.P.ANKS, TN M'HE CHJC'K.

0(3 H

Kin;illy, in l)()tli eases, Ça and /^) the vicinity of the connection
should i)e freed from yolk, albumen, and the amniotic finid, l)efore
the egg is inany lioars in alcohol, as otliei'wise these substances
oong'ulate aiid ean not then be removed. The outlines of the foetal
meinl)ranes should, however, be examined or sketched before tlie last
operation.

;]70 HTUOTA: OX TTIE SERO-A MXIO'I'TC < 'ONXKCTT* )X,

Explanations of Abbreviations and Colours
used in Figures.

By tlio >i\de of every ti^'iivo. tlio ago of tlie ombrvo, and the scale of eiilavgcinent
ave given. XitincrdJ.s in smull ii//>i: on some of tlie surface views sliow tJie levels of
the sections, represented in fignres wliicli ave niarl^ed with the coi'i'esponding nuniher.
Abbreviations used in the plates are : —

Alb for Albumen,

All „ Allantois,

All. Vein „ Allantoic Vein,

Aiun „ Amnion,

Coel.' „ Extra-Embryonic Coelomic Cavity,

Inall. Sep ,, Literal Ian toic Septum,

L. All. Art , „ Left Allantoic Artery,

Proam „ Proamnion,

11. AIL Art ,, Plight Allantoic Artery,

S. A. C ,, Sero-Amuiotic Comiection,

Ser. Env ,, Serous Envelope,

Hcil represents in the surface views (Figs. 1-5G) arteries — those of the inner
allantoic limb being formed of dotted lines — and the purely mesoblastic part of the
sero-amniotic connection. In the figures of sections (Figs. 57-94) red represents the
protoplasma of mesoblastic cells:

Blue represents veins — those of the inner allantoic limb being in dotted lines.
Blue also represents albumen in Figs. 51-5(i, und the liypoblast in the tigures of
sections (Figs. 57-94).

Black represents in the surface views (Figs. 1-5G) the epiblastie bridge of the
sero-amniotic connection, and in tlie figures of sections (Figs. 57-94) the nuclei
of cells.

Grei/ represents the interallantoic septa (Figs. 15-20). In the figures of sections
it represents the epiblast.

Yelloir represents yolk in Figs. 17-18.

(rreen represents tiie outer surface of the outer allantoic limb.

Plate XV.

(For explanation of abbreviations and colours see p. 370.)

Figs. 1-5 : — Dorsal' views of embryos in different stages from 48 to 78 lionrs. The
sero-amniotic connection in —

Fig. 1 is represented apart in Fig. 38,
)> ^ 11 ■>■> It "")

11 " )) 11 11 ^^j

4 41

11 ^ 11 11 )) ^'-i

„ 5 „ „ „ 42.

Figs. 6-9 : — Dorsal views of embryos in different stages from 104 to 168 hours. The
embryo proper and the splanchnic mesoblast are left out. The serous envelope
is also for the sake of simplicity not represented. The sero-amniotic con-
nection in —

Fig. 7 is represented apart in Fig. 43,
„ 8 „ „ 1, 45,

„ 9 „ „ „ 46.

Figs. 10-11 :— Tlie first is the surface view of an egg taken out from its non-cellular
enveIo[)es, and the second of the vicinity of the sero-amniotic connection in the
same egg. The sero-amniotic connection has a peculiar fish-like shape.
The anterior i)art of the sero-amniotic connection is in this specimen
exceedingly elongated dorso-ventrally and being bent down on the plane of
the paper appears t)'iangular, looking like the tail fin of a fish. The connec-
tion is thus twisted at the point A. The sero-amniotic connection is repre-
sented apart in Fig. 47.

Figs. 12-13 : — The first is a surface view of an egg taken out from its non-cellular
envelopes, and the second oï the vicinity of the sero-amniotic connection of
the same. The amniotic cavity, the cave-like diverticulum and the three
allantoic septa are represented l)y dotted lines. In the second the area
F' E' M F' represents a part of the interallantoic septum and the space
CBAD the cave-like diverticulum beneath the overlapping allantoic lobe.
The sero-amniotic connection is represented isolated in Fig. 48.

Fkss. 14-16 : — Fig. 14 is a surface view of an egg taken out from its non-cellular
envelopes; Fig. 15 an outline of the ibetal meml)ranes ; and Fig. 16 a surface

view of the vicinity of their sero-amniotic connection. The allantoic septa of
Fig. 15 shonld he examined along witli the hlood vessels represented in Fig.
14 : — a fall description of them is given in tlie text. The areas of the amnio-
tic cavity and of tlie albumen-sac are represented by dotted lines, but a part of
the yolk-sac is concealed beneath the albumen-sac. In Fig. 16, the albumen-
sac is advanced as a cave-like diverticulum, GPKL, towards the amnion,
which it meets along the line KP. the sero-amniotic connection. The
area UE"E'CHCtDF'E" is a part of the iuterallantoic septum, Avhich is
folded along the line DP7'. The sero-amniotic connection is represented
isolated in Fig. 49.

Figs. 17-18 : — The first is a diagrammatic representation of the allantoic septa alone,
and the second of all the foetal membranes of the egg represented in Figs. 14
and 15. In both, the outer allantoic limb is removed to a great extent. In
Fig. 17 the folded area FEDGrHGE is a part of the iuterallantoic septun]. The
line ED is the axis along which the iuterallantoic septum is folded in two. The
area ABC is the minor septum formed by the left allantoic artery. The septum
formed by the right allantoic artery is not represented. The deep black
areas represent duplicated parts of the septa. The positions of the albumen-
sac, the yolk-sac, the umbilical stalk, and the amniotic cavity are shown by
artificial incisions. In Fig. 18 the areas of the anniiotic cavity and albumen-
sac are given in addition, but a part of tlie yolk-sac is concealed beneath
the cave-like diverticulum of the albumen-sac.

Figs. 19-21 : — Fig. 19 is a surface view of an egg, 264 hours old, taken out from
its non-cellular envelopes. Fig. 20 represents the outlines of the foetal
membranes ; Fig. 20 bis the minor septa formed by the right allantoic artery
as seen from the surface ; and Fig. 21 the surface view of the vicinity of the
sero-amniotic connection. In examining the allantoic septa of Fig. 20 re-
ference should be made to the blood vessels represented iu Fig. 19. The
areas of the amniotic cavity and alimmen-sac are represented by dotted lines,
but a part of the yolk-sac is concealed beneath the cave-like diverticulum of
the albinnen-sac. In Fig. 21 the areas KDEF'K and KGHE are part of the
folded iuterallantoic septum, and the areas ADK and XZÖY (Fig. 20) are the
minor septa formed by the left and right allantoic artery respectively. The
sero-anmiotic connection is represented isolated in Fig. 50.

Jour. Sc. Coll. Vol. VI. PI. XV.

Plate XVI.

(For explauatiou nf ;il)lirt'viiitious aud colours see ]■. ;i7())

i'lGs- 22-36 : — These are surface views of tJirec eggs aged respectively 812, 385,
and 432 hours. They are arranged mostly in three horizontal rows, each
row being devoted to o]ie egg. The first two coloured figures of a row are
the surface views of opposite faces of the same egg, taken out from its non-
cellular envelopes, and the two last figures of a row are the outlines of the
foetal membranes of the corresponding faces. In following a blood vessel or
any other structure from one face to the other in these rows, it should be
remembered that the lower edge of the first and third figures of a row
should be applied to the upper edge of tlie second and fourth figures.
It should also be noticed that as an egg has a considerable thickness, a
structure leaving one face m;i,y not be found entering at exactly tlie corres-
ponding point of the other. Each of Figs. 26, 81, and 36 represents the
allantoic septa, come together at the vegetable pole of the respective egg,
as seen from that pole. P\-ill lines are tracings of the septa on the outer
limb of the allantois. In Figs. 27-28 and Figs. 32-33, the ijranciies of
the allantoic vein are anastomosed. In the example given in Figs. 32-36
the interahantoic septum is not folded on itself. In these examples of later
stages the interallantoic septum is not intluenced by tlie left allantoic artery.
The sero-amniotic eon])ection of the specimen represented in —
Fig. 22-2(> is ri'prescïjited ;i|);ii-t in Fig. 51,
„ '^7-Hl „ „ „ 54,

,. o2-3<i „ „ „ 56.

Fig. 87 ; — iJorsai view of tlie \icinity of the sero-amniotic coimection of an embryo
in the same stage as that represented in Fig. 7. Tlie dotted lines show the
extension of the lateral fold formed within the amniotic cavity (see Diag. 5).
The points A, B, &G correspond to those marked with the same letters in the
section (Fig. 92) of that level. The blood vessels in the inner allantoic limb
are for the sake of simplicity not represented.

Figs. 38-42: — Dorsal views of the sero-amniotic connection represented in Figs. 1.
2, 3, 4, and 5 respectively. The partly obliterated mesoblastic septuin is
represented by a broken line. All are magnitied to the same scale ( x 10).

Figs. 43-50 : — ^The ruesoblastic sero-amiiiotic connection at the corresponding stages
projected on an imaginary straight line, showing its relative width in different
individnals and in different parts of the same individnal. In breadth all
are stretched as much as possible, but in length, each represents the shortest
distance between the two extremities of the connection. The ends, where
the connection is incomplete, are represented by transverse interruptions.
Tile black lines in the middle represent the epiblastic sero-amniotic con-
nection, their interruptions showing breaks in it. All are magnified to the
same extent ( X 10). The connection represented in —
Fig. 43 is that of Fig. 7,
,, 44 is from a specimen, 144 hours old.
,, 45 is that of Fig. 8,
„ iö „ „ y,

„ 47 „ „ lü,

„ 48 „ „ 11,

49 14

„ 50 ., „ 19.

Figs. 51-56 : — Eepreseutations of the perforated states of the sero-amniotic coiniec-
tion, as seen from the amniotic cavity. The albumen-sac approaches the
amnion as a sort of tube or cave. Albumen is coloured blue. The line AA is
the line along which the interallantoic septum is continuous with the imier
allantoic limb. The circular line (Amn.) represents a part of the amnion.
Differently magnified. The specimen represented in —
Fig. 51 is that represented in Figs. 22-26.
,, 52 is from an egg fourteen days old.
,, 53 „ „ fifteen ,, ,, .

,, 54 is that represented in Figs. 27-31.
,, 55 is irom an egg seventeen days old.
,, 56 is that represented in Figs, 32-36.

Jour. So. Coll. Vol. VI. PI. XVI.

Lilt' et Imp i Kcshiba.

PLATE XVII.

Plaie XVII.

(For explanation <it' aMireviations ami colours sec p. 370)

Ail tlio l'if>nr('s arc tv.uisvovse sections of tlie embryos cut throivi>-li tlio levels
)iiark((l with riniiiei-als in small ty[)(! in the correspoiidiuü; surface views.
Fi(is. '>1-{J0 ai'c taken IVom the specimen represented in I'ig. 1. I'lic iiiosohlastic

septum in Fig. öS is niagnitied in Fig. öd.

Fig. (il is taken frona the specimen represented in Fig. 2,

Fios. ()2-05 are „ „ ., ,. 8,

. <i<i-ti7 .. „ „ „ ., 4,

„ 68-Ü9 „ „ „ „ „ 5,

„ 70-72 ., ., „ „ „ (5.

Fu;. 71. An epihlastic cell iji tlie connection is here in division.

l''ic,.s. 73-74 ;ire taki'ii horn the .specinaen represented in l''ig. 7,

n 7:î-7S ., „ „ „ ,. „ 8.

Fio. 7<S. An epihlastic cell is here in division.

I'kt. 79 is tak'en h-oiu the specimen represented in Fig. 11.

The dorso-ventrally elongated sero-amniotic connection is pi-essed down to a
horizontal position.
Figs. 80-82 are taken from the specimen represented in Fig. 16,
. 83-84 „ „ „ „ „ „ 21.

The comiection of Fig. 83 is magnified in Fig. 84.
Figs. 8.')-87 are taken from the specimen of the twelfth day showing ]ici-f(H-ations

in the sero-amniotic connection.
Figs. 88-89: Eepresent a stage nearly ecjual to that shown in Fig. .'j. The first is
anterior to the second by yV min. At the level given in Fig. 8H tlic anmion is
just closed. In these two figures nuclei are represented by white spots.
Fig. 90 Section Just in hoiu of the ailantois of au embryo, which is of the sanae
stage as that represented in Fig. 5. The point A is the lat(n-al angle of the
amniotic fold.
Figs. 92-93 : Sections of the specimen represented in Fig. 37, through the levels
marked witli the corresponding number on the latter. The points A, B, & C
inFig. 92 con-espond to the points marked witii the same letters in the
surface view.

Jour. Sc. Coll. Vol. Ml. PI. XVII.

On a New Human Tape-worm
(Bothriocephalus sp.).

by
!. Ijima, Ph. D.

lîigaknshi; Eigakiihakushi ; Prof, of Zoology, Sei. Coll., Imp. Univ., Tokyo.

and.

T. Kurimolo,

Igakushi ; Prof, in the Medical Department, oth Higher Middle School, Nagasaki.

With Plate XV TIL

The hiiiiKiii tape-worm to be noticed in tliis paper is, as will be
borne out in the se(|ue], a very large Bothn'occpliaivs species furnished
with double genital ducts and openings in each proglottis. So far as
we are aware, any similarly characterized species of tliat genus has not
hitherto been included in the human ]>arasitic fjuma, though a few such
have long been known to occur in seals and certain fishes. We hold it
highly prob;il)le that tliis human Botlirioccpliahis is a near relative
of those already described from seals,' and possibly identical with one
of them. The supposition that the patient who discharged it, had
acquired his from a source similar to that which furnishes seals with
theirs, seems to be not improbable from the fact of his continual
residence near the sea-shore.

The specific determination of this tape-worm, whether species
nomrum or not, we have preferred to leave to the discrimination of

I I!i)t]ir. riiriiil)i/ix K'raMH', Hotlir. faaciatiin Krabbe and Bothr. tetrajyteriis y. Siehold, irom.
sevei'al species of I'lioca.

372

I. I.IIMA A XI) T. KÜRIMOTO

those inveritiu-ntor« more Ihvoiinibl}' situated th:in we are with respect
to liter:itiire :iri(l s])ecimens to compüre with.

We owe our material to the com-tesy of Mr. Soiclnro Xakamura,
who, in 1(SÎ)2, while acting as physician to the hos])ital attached to the
Takashima Coal Mines, near Nagasaki , ol)tained the worm from one of
his patients.

I'articnlars concerning this p'ltient, partly furnished us Ijy Mr.
Nakamura and partly obtained by one of us from the patient's family,
run to the following effect:

Taniaji Murazato, male, born 1865, at Taira-mura (a village on
the Ariake-Sea, near the town of Shimabara) in the Province Hizen.
In boyliood healthy but never muscular. Remained in his native
village until 1879, when he went t(j Nagasaki. Here attacked by
cholera but recovered. Up to 1892 resided at several places in the
ijeighlxjurhood of Nagasaki • and other sea-side localities witliin
the Province Hizen, outside of which he seems to have ne\er
travelled. Calling : emanuensis, post-office clerk, school-teacher,
&c. 1891 settled at the Takashima Coal Mines, where he had been
engaged in Ijook-keeping business until his death by accident in
November of the following year. Some hve years ])revious to this
period, lie liegan to suffer occasional dizziness and colic. Medical help
h;id not mucli effect, beyond palliating the latter. Gradual anaemia
su]>er\ened. During October 1892, a piece of tape- worm about one
foot lono- was disdiaro'ed. About this time violent colic is said to
have returned. He was then taken into the hospital before mentioned
and submitted to medical treatment by Mr. Nakamura, whose notes
taken at the time are as follows :

" Initient aged 28 years. Medium bodily constitution. Badly
nourished, weary. Symptoms of cyanosis on face. Liable to fdl
into insensibilitv while sitting or otlierwise occu])ied. l^ilse weak

ON A NEW HUMAN TAPE-WORM (BOTHRIOCEPHALUS SP.). 373

îind frequent, niimberinsf 120. Palpitation -somewhat accelerated.
Temperature 36. S° C. Tongue with yellowish covering. Appetite
ordinary, sometimes vigorous. Gastric region swollen out and
frequently giving spasmodic pain, radiating towards the back and
ceasing gradually or suddenly, followed by a feeling of pressure on
intestines. This feeling either remains at one place or shifts its posi-
tion. Tlie [ittack occurs after taking food but also at other times.
Pressing the gastric region from outside has soothing effect on the
pain. Sometimes ])ain also in the i)elvic region. Diarrhd'a, oi"
costiveness for many days.

"From above symptoms, the presence of AiicijlDslounnu duDtloiale
was suspected. Microscopictd examination of the frees liowever
unexpectedly revealed a number of eggs, resembling very much those
QÏ Vistoumm riHficrl both in size and appetirance. Irrespective of wliat
parasite these eggs might belong to, a dose of cxlr. filic. mas. was
tried and the result was the discharge of a tape-worm measuring
10 iiicters in lengtli and, at the broadest portion, 25 milHmeters
in breadtli. The broad hind end had its extreme tip shrunk, nmch
macerated and easily detachable. Of the other end, a ]iortion as thin
as 1.5 millimeter was found but no head could be discovered. From
the following day, all the complaints the man had sutfercd fr»jm for
so many years entirely dis;ip[)eared."

But the man did not live long to enjoy this relief, for during the
following month a collision with a coal truck broke his back and killed
him. Post mortem examination was not allowed to Mr. Xakamura
notwithstanding his appeal.

Judging from Mr. Nakamura's statements, the tape-worm, a
Botlirioceplialm as already mentioned, must have been an extra-
ordinarily long and broad one, gradnaJly tapei-ing anteriorlv into an
almost filamentous colliun.

374 I- IJIMA AND T. KURIMOTO.

We did not see the entire specimen, Init sam])le pieces from four
different regions of the bod}^ were kindly placed at our disposal.
They were preserved in strong spirit that liardened tliem into stiffness
but kept the tissues in excellent condition.

Sample No. 1 is a piece from near tlie anterior end of the original
specimen. It measures 3 mm. in breadth and 0.5 mm. in thickness
at the middle. No trace of reproductive organs is visible in tliese
seo'ments.

o

iSam[)le No. 2 is a piece somewhere from the anterior quarter of
the (original specimen. At this region the 1)ody already presents con-
sidernljJe dimensions, being 18 mm. broad and about 1.5 mm. thick.
Tlie reproductive organs are partly developed ; l)ut of this, later on.

Sample No. o is from the middle ]3ortion, 14-16 mm. broad and
about 1.5 mm. thick. The reproductive organs are fully developed
and the uterus is already partially filled with eggs.

Sample No. 4 consists of two pieces from the posterior p(jrti<3n,
one of them cut off' -10 cm. from the hind end. Breadth varies
from 10 mm. to 15 mm ; thickness measures 1.5 mm. or some-
what more. The varying breadth is certalnl\- due to different states
of contraction and accordingly, where the brcadtli is less, the proglottis
is comparatively longer. The uteri in this section are nuich distended
and tilled u]j with eggs.

Mr. Nakamura's measurement of maximum lireadth, rjamely :25
mm., was no doubt taken whem the worm was (piite fresh. This
accounts for the fact that nowhere in the alcoholised and contracted
saiu{)les before us is that great Ijreadth attained. Tlie foremost
portion of the original specimen, stated by Nakamura to \\ii\e been only
1.5 mm. broad, must have belonsfed to a section more anteriorlv
situated than our sample No. 1.

One of the very striking features of our Bothriocejihabts species

ON A NEAV HUMAN TAPE-WORM (BOTHKIOCEPHALUS SP.)- 375

1*8 the extreme shortness aiid coiisequeiit narrowness in antero-posterior
direction, of the ja-oi-lottides. They almost [)resent the appearance of
closely set transverse AvrinkJes to the naked eye (ride figs. 1 and 2,
PL XVIII).' In the middle and hind regions, the length of the pro-
glottides averaged only 0.15 nnn., and their breadth 14-16 mm.
Even where most distended (l)readth 10 mm.), their length did not
exceed 0.(S mm. The average length given above was calculated by
counting /ro;;i tlw outside the number of [»roglottides within a measured
space aloiuj tlw median line. It is important to note here that
wliat appeared externally to be two distinct and C(nisecutive pro-
glottides as indicated by the usual boundaries, very often proved to be
one internally, i. e. with respect to certain genital arrangements.
For instance, a piece oO mm. long from sample No. I, showed but 57
uteri in a series, while the number of proglottides as coLuited in the
way mentioned amounted to about 6H. Besides the superficial super-
numerarv boundaries, both dorsal and ventral, that extend throuofhout
the entire breadth of the body, others which disappear at the middle of
it after runnino- for a greater ov less distance from the marg-in, are of
(juite frequent occurrence; so that, counting the proglottides by the
marginal serration wc^ald give a still greater numl)er than when
connted along the median region. Thus, in the j)iece above referred
to, the number of segments as counted ne^r the margin amounted to
Do or thereabout. The proglottis that has sucli an incomplete
supernumerary boundary on the one side generally shows the same also
on the other. Sometimes such a false proglottidal boiuidary in its course
joins a neighbouring true one or loses itself on the general surface to
appear again after a short interruption. In rare instances, apparently
two incomplete filse boundaries in succession were interposed between
two true ones in the marginal region. The antero-posterior lengtlis
of consecutive segments, separated by false proglottidal boundaries

376

I. IJIMA AND T. KURIMOTO.

and belonging to one internal proglottis, are quite variable bat when
taken together may be said as approximately equalling or somewhat
surpassing the length of those proglottides that show no trace of false
demarcations whatever (Figs. 4, 5, and 0). The features of marginal
serration as also the manner of indentation of the body-surface
essentially agree in all proglottidal boundaries, lx)th true and fnlse,
excluding all possibility that the latter might be some mechanical or
accidental production. We are inclined to view the phenomenon in
the light that the present species of Cestodes has a tendency to
produce superficially more proglottides than it does internally, con-
trary to the well-known case of Ligalidae in which the external
strobilation remains more or less obsolete. In other words, under the
crowded state of proglottides in our Bothriocephakis species, one
proglottis seems to remain but partially developed, i. e. only super-
ficially marked, in order to give necessary space for the full de\elop-
ment of certain genital parts (especially uterus, cirrus, and ovary) in
its immediate neighbour. The widely distributed testes and yolk-
glands develope themselves as well in the abortive as in the other
proglottis and are apparently related to the genital ducts and openings
of the latter as if they were its own.

What further seemed to us t(j be of interest with respect to the
strobilation of the present species, is the presence, in our sample No. 1,
of indications that certain prcjglottides are undergoing re|)eated
division. In this anterior region as many as o8 proglottides were
counted within a space of 10 mm., giving to each proglottis an
average length of 0.26 mm. (l)y o mm. in breadth). The actual
lengths were tolerably uniform, excej)t only that the latest formed
proglottides were only half or less than half as long as the
others. Division of a proglottis int») two takes place, not at its
middle, but invariably at its anterior portion; consequently, of the two

ox A NEW HUMAN TAPE-WORM (BOTTTKTOCEPHALUS SP). '^JJ

iiew ])r()gl()[ri(le.s, flie anterioi'ly sitiuited is ;il\v:iyy the shorter until the
norinnl len<:;th is attained by growth. Examining with a hand-lens of
Jow magnifying ])ower, onr attention was at once called to the focts
tliat the bonndai'ies of ])roglottides were not all alike in their shar])-
ness and de])ths of marginal indentation and that they succeeded one
anotlier with a certain degree of regularity. The differences are evidently
duo to the oldness or lateness of their formation, usually four or
live consecutive proglottides together f(.)rmed a grou]) or what might
conveniently be called a primary segment, terminated anteriorly and
posteriorly by much better defined proglottidal boundaries. In
other words, every fourth or fifth boundary was genei'ally the most
pronounced and presumably the oldest formed. Where four proglot-
tides made up such a ])riniary segment, the boundary between its 2nd
and ord proglottides was usually the next well-deiined, whereas that
between the 1st and :^nd and also that between the ord and -fth were
com])aratively somewhat less sharply ])ronounced. We m.ight inter-
pret this so, that such a primary segment is composed of two
secondary segments, each of which again consists of two tertiary
segments or proglottides. Often the front ])roglottis of a ])rimary
segment had more or less distinctly divided into two, at
apparently a quite late period, in which case that primary segment
seemed to consist of five, instead of four proglottides. Sometimes the
next tertiary proglottis also showed signs of a similar t|uaternary
division. Thus then, within a given primary segment, the
division of ])roglottides takes place successively backward beginning
from the foremost proglottis. This corresponds to the fact already
mentioned that in an individual proglottis division occurs at
its anterier jiortion. It is easy to conceive that by a continued
process of such division, an individual proglottis would in course
of time come to rank as a secondary and this again as a jirimary

;-J7,S I- IJTMA AND T, KURIMOTO.

.segmerjt. lUit this process seems not to take place unilornily
throughout, as indic:ited by the fact that primary segments, with and
without proglottides undergoing (juaternary division, showed no
reofularity in their order of succession, and also bv the fact that some-
times, between two more or less typical primary segments, there were
interposed two or more proglottides, which showed no ti-ace of division
and were separated from one another by boundaries as marked, and
hence presumably as old, as those that 1)ounded any primary segment.
At all events, the generally accepted idea that in a tape-worm the
more posteriorly situated proglottis is always the older, seems not to
hold tiaie in the present species of Bothriocephaliis. The repeated
serial di\ision here described naturally reminds one of a somewhat
similar process in MicnMomwn and certain annelids, but we abstain
here from entering into comparisons. We regret that Fig. 11, Plate
XVIII, which should represent a portion of sample No. 1, has failed
to ilhistrate exactly the ordinal distinctions of proglottidal boundaries
plainly visible on the real object.

How far backwards in the entire tape- worm the subdivision of
proglottides is repeated, coidd not be ascertained further than that it
probably ceases with the beginning of tlie development of genital ducts,
somewhere between the two portions represented liy onr samples Ko. 1
and No. "2. In the latter, in which the formation of sfenital ducts is
almost completed, abortive proglottides before described are plentifully
met with. One might regard these as indications of divisions taking
place here, though certainly only external, were it not for the
fact that similar abortive proglottides are also found in sam])le
No. 3 or No. 4 in about the same jiroportion. Hence, we nither
consider all those present in our samj^les No. 2-4, as representing the
proglottides that were formed about the time when the genit:d ducts
were beginning to develo])e but too late to develope their own.

ox A NEW HUMAN TAPE-WORM (BOTHUTOPEPHALUS SP). ;^79

Moreover, the body of tlie present Bothriocephnlns species is
longitiulinalh' trnvei'scd ])y severni, more or less deep, furrows on l)oth
its ventrîd and dorsnl surfaces. These are few and insignificant on
sample No. 1. l)ut numerous on all other samples, in Avhidi the most
conspicuous are the two on either surface, that run almost iminter-
ruptedlv and piu'allel to each other along the double series (^f main
genital ducts, dividing the tape-worm hody into a middle and two
lateral longitudinal zones (see figs.). They are slightly nearer to each
(3ther than to either body-margin. They may attain the depth of
about 74 the thickness of the body and must plainly be constantly
present in fresh specimens. The same can hardly be asserted of
all other hnigitudinal furrows seen on the middle and lateral zones
above mentioned, which -are, as seen in alcoholic samples, of
quite variable depths and sharpness, often interrupted or losing
themselves in their course, and by no means definite in their number.
However, some 8-5 in the middle and some 5-7 in the lateral zone are
the usual numbers to be met with.

As already indicated, there are, to each (true) proglottis, two
sets of genital openings, situated riglit and left and communicating
externall v at the l)ottom of the tAvo most conspicuous longitudinal
furnjws of the ventral surface (J>, h, figs. 5 and ()). Each set consists,
in antero-posterior succession, of a cirrus Çcir. 0., figs. 10 and 1:^), a
vaginal {vag. 0.) and an uterine (vt. 0.) o])ening, lying close to one
another. On account of their secluded position Avithin the longitu-
dinal furrow, they are usually not recognizable from the outside, but a
pit-like depression of the latter, associated with a short cross- furrow or
two, sufficiently marks their position and at the same time serves as
the index to distinguish the ventral from the dorsal surface. In many
proglottides of sample No. 4, the cirrus is externally visible as a minute
rounded protrusion, evidently the result of its ])artial evagination (fig. G).

p),SO T. TJTMA AND T. KUPJMOTO.

As might l)t' iiiieiTed from the ahove mention of o'enital openinp-s,
tlie arrangements of sexual organs in the present species are typically
bothriocephaline. To hegin with the male organs :

These develope earlier than the female sexual organs, as in other
species of Cestodes. The testes, which present the usual features, are
present from sample No. 2 downwards, [n the sample just mentioned
they are not yet fully mature and are separated I'rom one another by
somewhat wider spaces than in sample No. 3 or 4, in either of which
the production of spermatozoa is actively going on. They may nttain
a diameter of 0.07 mm. Generally arranged in a single layer, they
occupy the usual position in the " Mittelschicht " (Ji, fig. 8). The area
of their horizontal distribution is divided into three parts by the
regions taken up by the double sets of main genital ducts. In cross-
sections passing midway between the anterior and posterior limits of
proglottides, we have counted 30-40 testicular vesicles in each of the
three parts.

The cirrus (t'?V.) is a round or oval-sha])ed body, essentially
agreeing in its fine structure with the same organ of other Botlirioce-
pliahis species. It lies w^ith the axis of its tortuous lumen slightly
inclined from above downwards in an antero-posterior direction (fig. 12).
At its superior end, the cirrus is directly continuous with tlie muscular
wall of a, sjiherical cesiciila seniinaUs. Tlie latter presents an appearance
as though it were a posteriorly bent, knob-like, terminal portion of
the cirrus itself. Its cavity, as also the lumen of the cirrus, is narrow
and empty in sjimple No. 2, but filled up with and much distended
by spermatozoa in samples No. 3 and No. 4. In these the seminal
vesicle measures about 0.12 mm, and the approximately spherical
cirrus sibout 0.25 mm. in diameter. In sample No. 2 both are
mucii smtiliei'. the cirrus being liere decidedly oval-sha])ed as seen
in fig. 12. (In this figure, the lettering cir. referring to the second

ON A NEW HUMAN TAPE- WORM (BOTHRIOCEPHALUS SP.). 381

cirrus from left, points to the resiciila >ieminalis^ which is better seen on
the cirrus next to the right.)

The vas deferens (ed., üg. 10) is seen in sample I^o. '2 as a thin
cellular string with no recognizable lumen and in samples No. o and
No. 4 as a thni-walled, much convoluted tube filled with spermatozoa.
After startinîT from the vesieida seniinalis it turns towards the median
line, i.e., to the rio-ht if it belonofs to the left set of ofenital ducts, and vice
eersa. It then pursues its irregular course for a variable distance but
always stopping sh(.)rt before the end of the first uterine loo]) of the
corresponding side is reached. How it c(jnnnunicates witli the testes
could not be observed. —

Of the female sexmd organs, the eilellariiun (dis., fig. ,S) is yet
very scantily and weakly developed iji sample No. 2, but fully
developed in samples No. o and No. 4. In these it consists of very
numerous lobules about 0.025 mm. broad and about 0.045 mm. long,
with their long axis directed perpendicularly to the body -surface.
Arranged in a single layer, they occupy the usual position outside of
the layer of strongly developed longitudinal nuiscles and are nuich
more numerous on the ventral than on the dorsal side. On the
former they are distributed almost uniformly throughout except at
regions occupied by the two sets of main genital ducts (fig. 7). At
the body-margin they i)ass in a continuous layer mU) those <jf the
dorsal side, where, tracing them in the median direction, they l)ecome
scarcer and irregularly distributed, and finally disappear at a greater or
less distance fn^m the position of the uteri. In the middle regic^n of
the dorsal side, many proglottides showed none of them, while in others
they were develo[)ed also in this part but in irregular patches com-
paratively few in munber.

The ovary is [)resent to each set of genital ducts. It is a hori-
zontally and transversely stretched cellular sti'ing that divides on

3<S2

I. IJIMA AND T. KURIMOTO.

either side into an elongated bundle of anastomosing branches (oc, fig.
10). At its middle, whence the oviduct arises, it measures only
0.01 mm. or less in thickness while the lateral, divided portion may
be nearly 0.08 mm. broad. In functional activity, it may measure
1 nnu. from the origin of the oviduct to either of its extremities. Its
position in the proglottis is on the ventral side of the il/'i/,^c/.S6'//R'/?^,
along the posterior proglottidal border.

The exact course of the female ducts could only be conveniently
studied by combining sections (^f sample JSTo. 2, where they were yet
without contents except some masses of yolk-like granules that occurred
here and there in the uterus. The oviduct (ovd., fig. 10), after its
origin from the ovary, at first runs backwards, soon to take an
irreo-ular ascendins; course, which is at the same time directed out-
wards, i. e., away from the median line of the tape-worm. During this
course, the oviduct is j(3ined first l^y the vagina that approaches it
from the median side and then, after a short interval, by the yolk-
duct that comes u[) from below. In tracing the lumen of the oviduct
from tlie ovary, it appears to be directly continuous into that of the
vagina (as shown l)y the unshaded passage in fig. 10), rather than
into the remaining portion of the oviduct leading towards the junc-
tion of the yolk-duct.

The vagina (I'dfJ.) i« a fine duct, that makes a few windings
beneath the uterus but crosses the ovary on its dorsal side. Ptjsteriorly
and close to the junction with the oviduct, the vaginal tube swells uj)
into an oblong vesicle, which we consider to be the receptaculuui seminis
(d)., fig. 10). Anteriorly it descends almost perpendicularly along the
posterior border of the cirrus and opens at the vaginal pore (auj. o.,
fig. 12) just behind the cirrus opening.

The yolk-duct (dtg., fig. 10) is a very thin tube that after descend-
ing: ÎI short distance from the oviduct soon becomes untraceable.

ox A NEW HUMAN TAPE- WORM (BOTHRIOCEPHALUS SP.). 3|^3

Although yolk-granules were often found in its lumen, we could never
make out its connection with the vitellarium. Nor could we ever
recognize the shell -glands. Nevertheless there can be no doubt that
the present species essentially agrees in these as in so uuiny other
points with BotJu-ioccjiltdliis lalus.

The uterus {ut., figs. 10 and 12), as seen in sample No. 2, occupies
for the most part the dorsal I'egion of the Mittehcliiclit. Its anterior
portion descends ventrally, almost in a straight line to <3pen at the
uterine pore situated at about the middle of the length of the [)r()-
glottis (^ut. 0., fig. 12). The rest of the uterus, notwithstnnding some
irregularity in its course, describes in general two loops on each side.
The distance between the ends of two (opposite loops measures only
about h-alf a millimeter. No eggs are yet found and the uterine lumen
is for the greater part obsolete.

In sam[)le No. 3, the production of eggs is considerably advanced.
They fill the uterus and distend the latter into a wide tulje, but not
by far to such a great extent as in sample No. 4. The latter, when
viewed by holding it against the light, shows the uteri of successive
proglottides as blackish spots, about 1 mm. or slightly more in
breadth and ai-ranged in two longitudinal series. In each series
some may lie somewhat to the rio-ht or left of the ü'eneral line. When
compressed and clarified, they are certainly very distinctly visible.
In such preparations they present varial)le forms but the most general
and the least disturbed condition is that figui-ed in tig. 7. Each
uterus appears as consisting of two pairs of loloes, corresponding t<) the
Coin- loops of the uterine tube in .•sample No. 2. The anterior pair is
the shorter of the two and clasps the posterior side oi' the cirrus. The
numerous iiggs contained in the uterus, or more properly their shells,
give colour to the uterine loljes which varies from the colourless
transparency of the portion nearest the oviduct into the dark brown of

384 i- IJIMA AND T. KUEIMOTO.

the portion containing older eggs.

An esfof, with brown «hell and taken fn^m the uterus near its
external opening, is represented in fig. 9. In alcohol the shell is
collapsed but soon returns to its proper form by leaving it a while in
water. It is rather thick. The general form is oval, 0.063 mm. long
and 0.048-0.05 mm. broad. The diameter of the operculum measures
about 0.02 mm. The contents are oil-globules and a morula-like mass
of finely granular spheres, evidently cleavage-spheres. —

With regard to other points in the structure of the present Botli-
rioceplialus species, we have made but casual ol)servations. The
main longitudinal nerve runs outside the series of uteri, on each side
of the body (///., fig. 8). It lies nearer to the uteri than to the body-
margin of the same side. A short distance inside of the longitudinal
nerve, the main trunk of the excretory vessels finds its course on both
sides. Sections of excretory vessels were sometimes also n.et with at
other places. The muscular system is essentially the same as in
Bollirioccplhilus laliLs. Finally, we ought to mention that nowhere
in our samj)les have we found a single calcareous body in the mesen-
chyma.

UX A NEW UUMAX TAPE-WORM (BOTHRIOCEPHALUS SP.).

885

Explanation of Plate XVIII.

a, dorsal groove along the series of main

genital ducts.

b, ventral groove along the same.
fir, cirrus.

(•/;•, 0., cirrus opening.

dtH-, yolk-duct.

(Iti<., yolk-gland or vitellarium.

//., testes.

Im., longitudinal muscular layer.

In., longitudinal nerve.

/»—/»., median-line of the specimen.

07'., ovary.

ord., oviduct.

,9h., receptaculum seminis.

tm., transverse muscular layer.

lit., uterus.

lit. 0., uterine opening.

var/., vagina.

vap. 0., vaginal opening.

ihL, vas deferens.

Fig. 1.

Fig.

2

Fig.

3

Fig,

4

Fig.

5

Fig.

G

Fig. 7.

Fig. 8.
Fig. 9.
Fig. 10.

Fig. 11.

Fig. 1'2.

A piece from sample No. 4, 40 cm. from the liind end of the original speci-
men. Dorsal view. Nat. size.
Ventral view of the same. Nat. size.

Cross-section of the same. Nat. size. The two hlack spots represent the
uteri filled with eggs.

Dorsal view of a portion of the same. Magnified 10 times.
Ventral view of a portion of the same. Magnified 10 times.
Ventral view of another portion of the same, with cirri partially protruded.
Magn. 10 times. By an oversight this figure has been placed in a posi-
tion the reverse of that intended.

Portion of a preparation from sample No. 4, seen from the ventral side.
Magn. 20 times. The piece was somewhat compressed between two
glass-plates, coloured and clarified. The numerous dot-like bodies on
either side of the uteri represent yolk-gland lobules of the ventral side.
Portion of a cross-section of sample No. 4. Magn. 20 times.
An egg taken from the uterus. Magn. 440 times.

Half-diagrammatic representation of a left set of main genital ducts, as
seen from the ventral side; made out by combining sections of sample
No. 2. Magn. 150 times.

A portion of sample No. 1. Magn. 10 times. This figure is a failure in
so far as it does not properly show the ordinal distinctions of proglottidal
boundaries.
Portion of longitudinal section passing through tlie cirri, from sample No. 2.

Jour. Sc. Coll. Vol. VI. PI. XVIII.

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