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Digitized by the Internet Archive
in 2009 with funding from
Ontario Council of University Libraries
http://www. archive.org/details/journalofanatomyO08anatuoft
THE
JOURNAL
OF
ANATOMY AND PHYSIOLOGY
CONDUCTED BY
G. M. HUMPHRY, M.D. F.R.S.
PROFESSOR OF ANATOMY IN THE UNIVERSITY OF CAMBRIDGE,
HONORARY FELLOW OF DOWNING COLLEGE;
AND
WM. TURNER, M.B.
PROFESSOR OF ANATOMY IN THE UNIVERSITY OF EDINBURGH,
2 ie
VOLUME VIII.
(SECOND SERIES, VOL. VII.)
MACMILLAN AND CoO.
Cambridge, London and seo Work.
1874.
I ry >
Cambridge:
PRINTED BY C. J. CLAY, M.A.
AT THE UNIVERSITY PRESS.
‘————
CONTENTS.
FIRST PART. NOVEMBER, 1873.
Dr Ganasin, The Causes of the Secondary Waves in the Pulse (Addi-
tional Note, p. 112). P -
Mr A. Liversipcr, The Rite nemo ‘of the Eee
Dr Turn, The Structure of Tactile Corpuscles
Dr Marzin, The Structure of the Olfactory Mucous Reenitan
Dr M. Fosrsr, The Effects of a rise of Temperature on Reflex Actions
in the Frog : : - - : : : :
Mr A. H. Garrop, The fiw hen regulates the frequency of the
Pulse ‘ :
Mr W. Kitcuen se Morpliolasieal! Bae ents of the Skutl
Mr A, G. Dew-SuirH, Double Nerve Stimulation A
“5 7” The Presence of an Insoluble sugar- A aii
Substance in Penicillium :
Dr M. Watson, Contributions to the Anatomy of iio Taian: mienhant
Part Il. The Head :
Dr Brouyron, The apparent production of a new “effect 7 ‘fie: join
action of Drugs within the Animal Organism. : -
Mr A. Davrpson, An Account of the Madagascar Ordeal Pelion :
Dr SrrurHers, The Subdivision of the Scaphoid Carpal Bone
- 35 Rudimentary Finger Muscles in a Whale .
Dr Hotuxis, Tissue Metabolisn
Mr C.J. F. Youur, The Mechanism of ppenae me Goats ‘he Musto
chian Tube :
Proressor Turner, Note on a sapidental Skull aks a Nagyhal:
Dr Dwicut, Abnormal Ischio-Trochanteric Ligament :
Proressorn Humpury, Depressions in the Parietal Bones of an Oring
and in Man—Supernumerary Molars in Orang
ProFessor Turner, The Relations of the Cerebrum to the ee Sur-
face of the Skull and Head
Mr J. A. Russzewt, Two cases of Beraiatent: Gonmditontion ert
the Umbilical and Portal Veins : 5 - - :
Mr 8. H. West, A Peculiar Digastric irae A Variety of the Oc-
cipito-Hyoid :
Dr Ferrier, Experimental deeanaise in ‘Cerebral Phy Sidley and
Pathology . :
Dr Curnow, Two ineteneo of Irregular Ophthalmic ia Middle Me-
ningeal Arteries : ; : : : 2 : ;
Notices of Books .
Report on the Progress of henane by Deneadaon TURNER .
Report on Physiology, by Dr Sriruine
Report on Pharmacology, by Dr Fraser
lv CONTENTS.
SECOND PART. MAY, 1874.
PAGE
Prorressor Bryz, Some effects of Alcohol on Warm-blooded animals . 233
Dr Buaxe, The Action of Inorganic Substances when introduced directly
into the Blood : , 243
Proressor Cienanp, Double- potted Monetenes aa the ‘Develagmeds A
the Tongue. . 250
Dr Reyuer, The Carnes. a Beant Womiheares of the es -, », 261
Jas. Reocu, M.B., The Acidity of Gastric Juice . ‘ 274
PRorEssor ieee Additional Observations on the heeiges of the
Greenland Shark (Lemargus Borealis) . : 285
Proressor Savory, The Use of the Ligamentum Teres of oo He
joint F : 291
Prorrssor TURNER, Further Be neantes of eens in the Arran
of the Nerves of the Human Body : f . “397
Dr Ravcurre, The Synthesis of Motion, Vital fl Physical : . 800
Mr Lesniz Ocinyiz and Mr Cuartes W. Carucart, Dissection of a
Lamb with Fissure of the Sternum and Transposition of the Origin
of the Right Subclavian Artery P 321
Proressor Crum Brown, The Sense of Rotation and FN Renters ana
Physiology of the Semicircular Canals of the Internal Ear . - | Sat
Dr Brunton, Effect of Warmth in preventing Death from Chloral . 332
Mr F. Cuaupneys, The Septum Atriorum of the Frog and the
Rabbit. : 340
Mr J. C. Ewart, Ee yciereeree Note on an Epithelial ec eaeee in
front of the Retina and on the External Surface of the Capsule of
the Lens ; : ‘ . 853
Dr O«txr, Note of an Tnkoreatiae Cae of Mialtgers nticn “ 358
Proressor Turner, An Illustration of the Relations of thes Gare
volutions of the Human Cerebrum to the Outer Surface of the
Skull : 3 : : ; » aan
Prorresson Turner, The ‘Piaventeten of the Sloths : : ; . 862
ProrEessor Curnow, Notes on some Muscular Irregularities : 377 ©
Mr G. J. Mauconm Suite, Notes of a Dissection of an Excised
Elbow : . 880
Mr J. A. Russext, Note on an Unnsialiy ee Renal Caines . 882
Mr Epwarp Beniamy, Singular Malformation of Wrist and Hand . 3883
Notices of Books . : : . 884
Report on the Progress of Annem ic Sea Caos : . 386
Report on oo by Dr Sriguine ; j 2 : : Pees 3 15)
5 6.00) 0): a = : : : é : F ; , ; >, 432
EE
Hournal of Anatomp and Phpstology.
ON THE CAUSES OF THE SECONDARY WAVES
SEEN IN THE SPHYGMOGRAPHIC TRACING OF
THE PULSE. By A. L. GAvAsin, M.A., M.D., Fellow
of Trinity College, Cambridge.
Ir is a fortunate circumstance for the application of the
Sphygmograph, as a means of clinical research, that a know-
ledge of the cause of the several secondary waves of the pulse
is not necessary for their practical interpretation. This may
be learnt empirically by watching the association of the dif-
ferent forms of pulse-curve with the known conditions of the
patients from whom they are obtained. Thus we find that
authors who differ totally from each other as to the causation
of any particular form of pulse, and even as to the state of
circulation which it implies, yet are quite in agreement as to
its clinical import, and the prognosis to be derived from it.
But it requires much experience of tracings to be able to draw
these inferences, and those who have not this are apt to inter-
pret what they see from theory, and thereby easily fall into
error. For instance, it has often been supposed that a high
sharp primary summit, followed by a sudden fall, is a sign of
aortic regurgitation, but this may occur just as much in the
pulse of simple excitement. Hence, from the practical point
of view, the study of causes is useful, and it is essential for
the arriving at general physiological conclusions, such as the
determining what is the true state of the circulation in fever,
or in Bright’s disease, or what is the effect on the vascular
system of various drugs.
It needs but little study of the literature of the subject
to discover that scarcely any two authorities agree together
VOL. VIII. 1
2 DR GALABIN.
as to the cause of the several waves, and of their variation.
The question is one which must be settled by having regard
both to the principles of mechanics, and to the results of
observation. And it is impossible to avoid suspecting that
some of those who have treated the subject experimentally,
would have been assisted in the interpretation of their results,
if they had possessed some theoretical knowledge of Hydro-
dynamics.
I have adopted an experimental mode of investigation by
the use of a combination of bifurcating elastic tubes to repre-
sent the arterial system. I have not attempted to imitate the
individual arteries of the body, for although such an apparatus -
looks well in a lecture-room, it does not any the more re-
semble the actual complexity of the human circulation. To
these tubes I have adapted, in some cases, the heart of a
sheep, in others, an artificial heart of india-rubber, and the
contraction of the heart has been imitated by manual com-
pression. It will be seen that counterparts have thus been
obtained of the most important types of pulse found in the
human body. Tracings 1 to 4 were procured with the real
heart, those from 5 to 9 with the artificial. In the case of the
real heart, I have found it more convenient to use the right
side, for the thick walls of the left ventricle form an obstacle
to manual compression. The aortic, or pulmonary valves act
efficiently after death, but it is not so with the mitral or
tricuspid. It follows from this, that the real heart will only
work against a low pressure, and the highest which can be
obtained with it (fig. 3) corresponds to nearly the lowest ever
found in the living body, while the tracings at lower pressures,
such as fig. 1, represent a state of things which never occurs in
arteries. Tracings from the real heart, at the higher pressure,
resemble closely those obtained at a similar pressure from the
artifical heart, thus showing that the action of the latter is
sufficiently like that of the real heart for the purpose of these
experiments (compare tracings 2 and 5). It has been con-
sidered by Dr Burdon Sanderson impossible to make the con-
traction of the hand sudden enough to imitate that of the
living heart, but I have not found this difficulty at all insu-
perable.
SECONDARY WAVES IN PULSE, 3
I had, in the first place, made use of a single elastic tube,
corresponding to the schema constructed by Dr Burdon San-
derson. In this case I found the tracings different at different
parts of the tube, and the variation was such as to show the
presence of retrograde waves due to reflection from the end of
the tube. This reflection took place as much from an open as
from a constricted orifice. In my final combination, I made
the several tubes of very unequal lengths, and in using com-
pression in order to vary the tension, I applied it to each of the
smallest tubes, not at one, but at several points, so that no
reflected waves might be called forth of a kind which could
not occur in the body. Applying then the same test of taking
tracings at different distances, their resemblance to each other
showed, as it does in the case of the arteries of the body, that
all the waves were direct and not retrograde. The effect of
friction in the eapillary circulation was thus imitated by that of
compression at several points of each small tube, and if it may
be said that reflection might still possibly take place in the
schema from the latter, so it has been held by many that it
occurs in the body from the former. I shall refer to the several
tracings obtained under different conditions as the points arise
which they illustrate.
It was shown so long ago as in 1833 by Weber, that the
motion of the pulse is that of a wave. By this term is meant
the transmission with a definite velocity, not of matter, but of
a state of motion and of pressure. It follows from the theory
of waves, that on the large wave other waves may be super-
posed, which run each their own course almost exactly as if they
existed alone, and which may be added together to form com-
pound waves. ;
Among those who have not given special attention to the
Sphygmograph, almost exclusive regard has been paid to one of
the secondary waves of the pulse, namely, the dicrotic wave,
and no doubt this is, to a great extent, justified by its import-
ance in the theory of the pulse, its constant occurrence, and its
great development in fever. But there is another wave pre-
ceding it, which sphygmographers find to be of quite as great
clinical significance, and which varies as widely, for, while often
absent, it may attain considerable dimensions. Being but
1—2
7 DR GALABIN,
slightly marked in a healthy pulse (fig. 10), and often disap-
pearing in that of a student (fig. 19), it forms a stronger feature
in the pulse of an athlete, and shews itself also in that of ladies
under the influence of excitement. Under another form again,
with advancing years, it acquires a magnitude which is of
ominous significance, but gains perhaps its greatest develop-
ment in cases of chronic Bright’s disease (fig. 12). I believe
this wave would be better known if it had a convenient name.
“ First secondary wave” is too Jong, so is “systolic pressure-
wave,” and, moreover, that asserts a theory. The only short
name applied to it is that of “tidal-wave,” used by Mr Maho-
med in his papers in the Medical Times. Although this also
is intended to imply a theory, which I believe to be erroneous,
yet, as it does not assert it so directly, I shall adopt it for its
convenience. I propose, therefore, to discuss first the tidal,
and afterwards the dicrotic wave, and in connection with the
former it is necessary to consider the primary upstroke.
I will notice first existing theories. Dr Burdon Sanderson,
writing in 1866, says that the contraction of the heart pro-
duces two waves, one of accelerated movement, and one of
increased tension; that these, starting together, become sepa-
rated in the distant arteries, because the former travels with a
velocity of about 90 feet per second, and so is practically in-
stantaneous. He compares the former to the communication
of a blow along a line of ivory balls, but afterwards, accepting a
correction on the ground that water is not elastic, admits that,
but for the effect of the elastic receptacle, the resemblance
would be rather to balls of clay. Now in the popular sense of
the word elastic, that of compressible, water is almost inelastie,
but in the more exact sense, in which elasticity is measured by
the perfection of recoil to any force impressed, water is very
elastic, as is shown by the fact that sound is conveyed much
better in water than even in air. So far it is more like ivory
than clay, but in truth the behaviour of neither has the
slightest analogy to that of a fluid. It is the first principle
of the mechanics of fluids, that at every point of a fluid,
whether at rest or in motion, pressure is the same in all direc-
tions. It follows from this that there can be no wave of
forward pressure (and without that there can be no accelera-
-
SECONDARY WAVES IN PULSE. 9)
tion), which is not at the same time a wave of lateral pressure,
expanding the walls of the tube.
The view of Dr Burdon Sanderson is therefore mechanically
impossible: it is also contrary to observation. Jn the first
place, the distinction of the two waves should, according to him,
be most when tension is lowest. But, on the contrary, the tidal
wave is most separated in pulses of high pressure, like that of
Bright’s disease, altogether absent in the febrile pulse of low
tension, and precisely the same relation was found in my expe-
riments with tubes. Secondly, the tidal wave should be farther
apart from the primary wave in the more distant arteries; but
a comparison of tracings 15, 17, and 18 from the brachial, radial,
and dorsalis pedis, shows that this is not the case. And thirdly,
it is not the fact that the wave producing the first upstroke is
practically instantaneous. In the case of elastic tubing the in-
terval is very perceptible, if a length of 6 or 8 feet be taken.
The velocity varies altogether according to the diameter of the
tube, its material, and tension, but in one case I found it to be
less than 20 feet per second. The same thing has been shown
as regards the body, in this Journal, by Mr Garrod, who by his
Cardio-Sphygmograph measured the interval between the heart’s
contraction and the first upstroke of the pulse tracing. Weber,
again, reckons the average velocity of the pulse wave to be
about 28 feet per second.
Dr Anstie, writing in conjunction with Dr Burdon Sander-
son, yet differs somewhat, at least in expression, for he speaks
of the tidal wave as an expansive wave, which is a movement
in the arterial wall, and so slower in propagation. This would
seem to be the same as the theory adopted by Volkman to ac-
count, not for the tidal, but for the dicrotic wave, that one wave
is transmitted in the fluid, another in the arterial wall.
Dr Balthazar Foster, as represented by the article in the
last edition of Dr Aitken’s Medicine, holds that the primary
wave is a vibration of the blood column, travelling instanta-
neously, and that the next is a wave of distension by blood. It
is obvious that a wave of vibration is quite a different thing
from a wave of forward motion, yet Dr Burdon Sanderson would
seem to have combined this view with his own, for he says in
1871, “The bursting open of the aortic valves produces a vibra-
tv) DR GALABIN.
tory movement of the blood transmitted instantaneously (that
is, in about =, second).” Now one vibratory wave would indeed
be transmitted, namely, that of sound, with a velocity due to
the compressibility of water, not of 90, but of about 5000 feet
per second, but this produces no motion in the lever. Tracing
22 was obtained by giving blows to a rigid part of the tubing.
It shows that even a coarse and violent vibration produces
‘hardly any upstroke. Dr Burdon Sanderson has published
some tracings of waves produced by percussion, but in that case
the blows were applied to the elastic tube, which would of
course give rise, not only to vibration, but to waves of forward
motion and expansion. Most of the other objections to the
views of Dr Burdon Sanderson will apply also to those of
Dr Foster.
Yet another theory of the tidal wave is maintained by Mr
Garrod. Discarding the notion of an instantaneous wave due
to the closure of the mitral valve, or the first impulse of the
heart, he yet holds the tidal wave to be an instantaneous wave
due to the closure of the aortic valves. I think this theory will
commend itself to no one who has watched the variation of the
tidal wave in many tracings; but, if any refutation be necessary,
I would refer to tracing 8, showing its prolongation by a pro-
tracted contraction of the heart.
The view of Mr Mahomed, as far as I understand it, is the
same as that of Dr Foster, except that he does not consider the
first wave to be instantaneous.
Proceeding next to my own explanation, I have to remark
first, that, since the sphygmograph is possessed of inertia, and
is itself subject to the laws of motion, its construction must
necessarily have some influence, however small, on the tracing
produced. In sphygmographs having a secondary spring to
depress the long lever, the tidal wave is often broken into two
waves (fig. 21), and if this spring has a short time of vibration,
a jagged line may appear. In the instrument I have used, of
the form devised by Mr Mahomed, this spring is, with advan-
tage, omitted, and no such waves are then ever seen. Thus
from many tracings published, an experienced person may draw
inferences, not only about the patient, but about the form of
sphygmograph used. In order, therefore, to determine how
SECONDARY WAVES IN PULSE. i.
much is due to the instrument, since inertia cannot be got rid
of, I have adopted, what is called by Mill, the “ Method of Con-
comitant Variations,” altering the moment of inertia of the
lever about its axis of motion by a small sliding weight. The
results are tracings 15 and 16. It will be seen that the rela-
tion of the primary and tidal waves is altogether altered, while
the position of the dicrotic wave remains unchanged, although
its amplitude is diminished. The method of measuring the
- position of these waves is first to draw a horizontal line of
reference, then to place the tracing again on the moving plate
of the sphygmograph, and draw curved lines with the writing
point of the lever. It is obvious that if the tidal wave were a
wave passing in the artery, its relative position could not be
altered by weighting the lever of the sphygmograph; or, at any
rate, if any effect at all were so produced, it could only be an
apparent retardation, and not an acceleration of the wave. The
separation of the primary and tidal waves is therefore due to
an oscillation in the sphygmograph, caused by the inertia of
the instrument, and the relation of the tracing to the true
pulse wave is something like what I have drawn in fig. 23. In
some cases the lever may be separated slightly from the knife-
edge on which it rests, but generally the oscillation takes place
in the instrument as a whole, and it may be followed by others
in a descending series. Thus if the lever be moved, not by a
knife edge, but by a rack and pinion adjustment, the tidal
wave still occurs. Such an arrangement probably makes the
tracing resemble the true pulse wave a little more closely, but
for clinical purposes it is not so good.
I may mention that in the pulse shewn in fig. 15, in which
the tidal and dicrotic waves are both so marked, one secondary
beat could be felt by the finger, but only one. Such pulses
do indeed give at first to the finger the impression of several
secondary waves on account of their thrilling quality. But the
sphygmograph gives no record of a thrill, as is shown by its
application to the heart in the case of mitral stenosis. If the
stenosis be moderate, the prolonged auricular contraction is
shown in the tracing; but if it be so close that a thrill is con-
tinued throughout the whole period of rest, no record of it any
longer appears.
8 DR GALABIN.
My general conclusion is confirmed by the application of the
little instrument which has been called a sphygmoscope, whereby
the motion of the pulse is displayed by the variation of a jet of
gas. By this means the dicrotic wave may readily be seen, but
not a single tidal wave. Its place, however, is supplied by a
slight quivering motion, which is due to the vibration of the
elastic diaphragm, upon which the pressure of the pulse is
received. ‘This vibration varies with the size and tension of the
diaphragm, and it might be possible so to adjust these that a
wave should appear like the tidal wave of the sphygmographic
tracing.
Another argument may be drawn from the fact that the
development of the tidal wave varies in some degree according
to the pressure which is applied to the artery. Thus in the
pulses in which, at ordinary pressures, no tidal wave can be
seen, it may sometimes be made to appear by using an excess-
ively low pressure.
The explanation which I have applied to the whole tidal
wave is adopted by Dr Burdon Sanderson, to account for the
first of the two waves into which it is broken by the use of the
secondary spring. But this must be entirely due to that spring,
sinee it never occurs in its absence.
The view of M. Marey is, up to a certain point, the same
which I have taken, for he says that the first pointed summit is
due to the acquired velocity of the long lever. But he regards
the tidal wave in some of its forms as an instantaneous wave
due to the closure of the aortic valves: the dicrotic wave he
believes to have no connection with those valves. We have
seen that Mr Garrod has adopted the same theory as far as
concerns the tidal wave.
Of all the diagrams in the work of M. Marey, one of the
most interesting is that in which three simultaneous tracings
are shown, of which the first represents the pressure within the
right auricle, the second the pressure within the right ventricle,
while the third is the tracing of the apex beat, obtained by
means of an ampulla inserted within the thoracic wall. In all
of these the primary summit is followed by two or more
secondary eminences, much resembling the small waves which
may be seen in the place of the tidal wave in the pulse tracing
SECONDARY WAVES IN PULSE. 9
as drawn by M. Marey’s sphygmograph (Vid. fig. 21). I believe
that their origin is similar, and that they illustrate the mode in
which eminences may be produced in consequence of the
inertia of the instrument. By M. Marey himself, however, they
are attributed, in the case of the first two tracings, to an
oscillation in the tension of the auriculo-ventricular valves,
occurring after their closure, and causing a corresponding rise
and fall of pressure within the two cavities. Now it is evident
that if the oscillation of the valves causes a rise of pressure in
the ventricle, it must, at the same moment, cause a fall of
pressure in the auricle, and conversely. Hence the elevations
in the tracing of the one cavity, if due to this cause, ought to
be synchronous with the depressions in the tracing of the other.
But on referring to the diagram of M. Marey, it will be found
that the elevations in the auricular curve correspond, not to
depressions, but to elevations in the ventricular curve, and
therefore the explanation given by him would seem to be
inapplicable.
While I thus believe that waves occur in the tracing, which
have no separate existence in the pulse, I am yet of opinion that
the instrument is more clinically useful than if it followed the
artery more closely, for I think that slight differences in the
form of the pulse wave, and in the suddenness of its commence-
ment, are thus translated into a form much more manifest to the
eye. The constancy of the form of the pulse tracing in the
same person under similar circumstances, proves that it contains
no casual oscillations, but that its form has, at least, a fixed and
definite relation to that of the true pulse wave. My experi-
ments with elastic tubes showed that the tidal wave could not
be produced unless the pressure exceeded a certain point, and
also the length of elastic tube were limited, or rigid tubing sub-
stituted for a part of it (fig. 7), and it could then be prolonged
to a great extent by increasing the length of the heart’s con-
traction (fig. 8). Its development thus indicates three things,
high tension, diminished elasticity, and long laborious action
of the heart. This conclusion agrees entirely with clinical
experience of the kinds of pulse in which it is most manifested.
Passing on next to the dicrotic wave, it may be thought
almost superfluous to consider its cause, since the common view,
10 ; DR GALABIN,
ascribing it to a recoil produced by the closure of the aortic
valves, appears so probable and intelligible. Yet the case is
not so simple as it seems. Thus we find that Dr Burdon San-
derson, who, in 1866, appeared to adopt the common view,
says in 1871, “The dicrotic wave has nothing whatever to do
either with the closure of the aortic valves, or the cessation of
the heart’s contraction.’ His present theory is a little difficult
of comprehension. He says, “In the largest arteries the expan-
sion is ebbing, while in the smallest it is still culminating: so
that for an instant the pressure is greater in the latter than in
the former. The restoration of the equilibrium must take place
by increase of pressure towards the heart, and diminution to-
wards the periphery. This restoration of equilibrium consti-
tutes the second beat.” In another placehe says that owing to
the cessation of the heart’s contraction the capillary arteries
become relaxed, the capillary circulation retarded, and the aorta
simultaneously distended in consequence of the increased resist-
ance in front: and that this distension is in its turn propagated
towards the periphery. These two accounts seem to me different,
nor can I clearly gather whether or not Dr Burdon Sanderson
considers the dicrotic wave to be retrograde. But such a trans-
mission of influence from the periphery to the centre could only
take place as a retrograde wave. To determine therefore
whether this occurs I have compared two tracings from the same
dicrotic pulse, one from the femoral artery just below Poupart’s
ligament, the other from the posterior tibial below the ankle.
These are 19 and 20. ‘The corresponding position of the dicrotic
wave in the two shows that it is not retrograde but direct, and
there is no retrograde wave at all present, for such a wave would
be close to the primary wave in the posterior tibial, and farther
from it in the femoral. The same tracings refute the theory
that the dicrotic wave is due to reflection at the bifurcation of
the aorta, for then it would be absent below that point. While
I have thus opposed most of the views of Dr Burdon Sanderson,
I should be the last to undervalue the service which he has done
for the practical application of the sphygmograph, for I agree
with his clinical inferences as completely as I differ from his
mechanical explanations.
Perhaps the oldest theory of the dicrotic wave is that of
SECONDARY WAVES IN PULSE. 11
Dr Barlow, who observed it before the invention of the sphyg-
mograph, and held it to be a reflection from the periphery. He
believed dicrotism to indicate an obstructed capillary circulation,
and therefore high tension, and to imply a stage of irritation
and contraction, which, in disease, preceded the stage of fever
and relaxation. The sphygmograph, however, has shown that
great dicrotism belongs especially to the state of fever itself, and
is found neither in the preceding stage, nor in that of exhaustion
which follows. M. Marey and Dr Carpenter likewise consider
the dicrotic wave to be a reflection from the periphery. Now
this reflection is exactly what occurs in a single elastic tube, but
not in the body, as already shown by the comparison of tracings
19 and 20. Duchek believes the dicrotic wave to be an oscilla-
tion, not in the aorta, but in the peripheral arteries. Vivenot
regards it as an oscillation, but does not explain how it arises.
The view of Volkman has been already mentioned.
Now, looking at tracing 1, from the elastic tube, and 14
from the radial pulse, it would seem that a dicrotic wave, equal
in magnitude to the primary, could hardly be due to the simple
closing of a valve. Experiments with tubes show that the
dicrotic wave is only the first of a series, and when pressure is
very low the aortic valves, being in glass tubes, can be seen to
open and close a second time, after their first closure. In that
case therefore the closing of the valves is not so much the cause
as the effect of the secondary waves. I have tried the effect, in
the case of the real heart, of dividing one of the semilunar
valves, in that of the artificial heart of removing them altogether.
The results are shown in fig. 4 and fig. 9. In both the dicrotic
wave is less than before the alteration, but still very consider-
able. This fact of the occurrence of the dicrotic wave without
any aortic valves has also been noted by Duchek, by Vivenot,
and by Dr Burdon Sanderson. It agrees entirely with experi-
ence of tracings in the case of aortic regurgitation, for although
the dicrotic wave is diminished when regurgitation is free, it is
yet never absent, and even in a splashing pulse often retains
considerable size. The use of the sphygmograph is thus dis-
appointing for the diagnosis of aortic disease, although, the fact
of regurgitation once known, it is of service in determining its
extent.
12 DR GALABIN.
My own view is, that, as the tidal wave is due to the inertia
of the sphygmograph, so a wave occurs which is due to the
inertia of the arterial walls. If it were not for this inertia,
their distension would always be such as to be in equilibrium
with the pressure of the fluid within at every moment. As it
is, it takes a little time to reach this point of equilibrium, then
by acquired velocity is carried a little beyond it, and so again
passes within it as it recoils, and thus makes a series of oscilla-
tions about the equilibrium point. Thus there occur oscilla-
tions of expansion and contraction of the largest arteries, due
to the effect of the inertia of the arterial walls on their lateral
motion, but modified also by the inertia of the fluid. The first
of these, the only one which commonly occurs, forms a part of
the dicrotic wave. Be it especially observed that I attribute
the oscillation to the inertia of the arterial wall and not to its
elasticity, although that elasticity is of course necessary for
this, as for every other part of the motion of the pulse wave,
and its degree affects the period and extent of the oscillation.
Nor, again, would any oscillation occur from the inertia of the
fluid alone, so far as that affects its forward or backward motion;
but such an oscillation being once set up, it is more ample the
greater the momentum of the fluid, because the motion of the
tube and of the contained fluid can only take place as a whole.
It is thus that dicrotism is increased, as shown by M. Marey,
if a denser fluid, as mercury, be taken instead of water. There
is however another way also in which the inertia of the fluid
may come into play, and that is by its effect on that slight
lateral motion of the particles which must take place in conse-
quence of the expansion of the tube. The effect of the-ac-
quired lateral velocity of such particles would be to expand the
tube a little beyond the point which it would otherwise have
reached, and by that means set up an oscillation. The effect of
such acquired lateral velocity is generally disregarded in mathe-
matical investigations of similar waves, as being too minute to
have any appreciable effect. It would be difficult to ascertain
whether in this case the part it plays in the general result
ought to be taken into account. The aortic valves would pro-
duce a wave of their own without any oscillatory wave, and
they also reinforce that wave by reflection. The second oscil-
SECONDARY WAVES IN PULSE. 13
latory wave I have found in but few pulses, of which one is
shown in tracing 14. That pulse was in the highest degree
compressible, and was taken only a few hours before death.
The condition was just the opposite of that of the common
tricrotic pulse (vid. figs. 11 and 15), in which the second wave
is the tidal wave.
I think an argument for my view may be drawn from the
possibility of the occurrence of a monocrotic pulse. According
to the common theory of the dicrotic wave, this would imply
that the aortic valves never close at all, in which case we can
hardly suppose that the circulation could continue; but upon
the other view, it only means that the rate of the pulse is
equal to the rate of oscillation in the aorta. The dicrotic wave
seen in a tracing may thus be made up of three waves super-
posed, the recoil from the aortic valves, the first oscillatory
wave of the large arteries, and the second (or sometimes the
first) oscillatory wave of the sphygmograph.
An opinion has been expressed by one of the ablest mathe-
maticians of the day, especially in relation to physical pro-
blems, I mean Professor Maxwell, that no mathematical solution
could be usefully applied to the theory of the pulse, and that
for two reasons,—because the blood is not truly fluid, and
because the motions of the arteries would be so much affected
by their external attachments. These difficulties however
would not apply to experimental elastic tubes, and since the
principal forms of pulse can be imitated in them, I am led to
the conclusion that blood is really fluid while contained in the
arteries, and that their external connections are too loose greatly
to modify their motion.
The introduction of the inertia of the arterial wall makes
the question very complex for mathematical treatment. Disre-
garding that, a differential equation may be obtained of a fourm
similar to that occurring in other kinds of wave motion. Its solu-
tion gives a velocity for the wave, which involves the materi. 1
an| diameter of the tube, and the pressure of the fluid, beirg
greater when pressure is greater. This last result agrees with
observation as to the retardation of the pulse, and is a much
likelier explanation of that phenomenon than to suppose that
what is felt by the finger is in some cases not the primary
14 DR GALABIN.
upstroke, but the tidal wave, which is only a convexity in the
descending curve. I think it will be found that the retardation
is most in dicrotic pulses, where tension is low, and therefore
the wave velocity less, but in which the tidal wave is entirely
absent. Calculation gives no indication of the existence of any
other wave travelling with a different velocity, except the wave
of sound, whose velocity is due to the compressibility of water,
and is nearly 5000 feet per second. ;
As a rule my experiments showed the dicrotic wave to be
increased by diminution of pressure. This is the general view,
and has been denied only by Mr Mahomed, on experimental
grounds. But his schema differed from the arterial system by
the introduction of a spherical elastic bag, which could hardly
fail to introduce a set of oscillations of its own. It need hardly
be said that it would be mechanically most unlikely, as well as
for the sake of inference most unfortunate, that the same dicro-
tism should at different times employ opposite conditions.
There is however one limitation to be made. If tension be
increased by compressing a tube at a single point, dicrotism is
often not diminished but increased, because the oscillatory wave
is then kept in and reflected. This explains why dicrotism is
increased by placing a tourniquet upon the abdominal aorta.
Theoretically both components of the dicrotic wave should
be increased as pressure is lowered. The recoil from the aortic
valves is not indeed greater, and could never produce such
a dicrotic wave as that in fig. 1 and fig. 14, but it becomes more
marked because preceded by a greater reflux, and consequent
fall of pressure, when the valves close slowly. Oscillatory waves
again are always more ample when tension is low.
In my experiments I have found that besides the variation
of pressure, one other condition increases dicrotism, provided
pressure be also low, namely to make the action of the heart
short and sudden. In that case, however, the tracing has a
rather different aspect, for sharper points aré seen in the
curve. This agrees with observations on the human pulse, for
in that the rounding off of points is well known to be of bad
prognosis. Ifthe action of the heart be jerky, but at the same
time pressure not low, the result is the common tricrotic pulse,
in which the second wave is the tidal wave, and the first sum-
SECONDARY WAVES IN PULSE. 15
mit high and sharp. This state of things occurs in the body in
the case of muscular exertion, mental excitement, from the
smoking of tobacco, and, with a greater proportionate develop-
ment of the tidal wave, in acute nephritis.
As an application of the foregoing principles, certain conclu-
sions may be drawn as to the state of the circulation in fever.
The degree of dicrotism together with the rounding off of sharp
points indicates that arterial pressure is very low, and at the
same time the action of the heart short and sudden, but that
the former of these conditions preponderates. This would be
explained by supposing a paralysis to occur of the nerves which
cause contraction of the arterial walls. The rapidity of the
heart would then be in part the direct consequence of low
arterial pressure, according to the relation demonstrated by
M. Marey, but its short and sudden action appears to indicate a
disturbance of its own innervation in addition. he increased
rate of circulation would however depend less on the action of
the heart, than upon the arterial relaxation, which is therefore
the most important element in the state of the vascular system
existing in fever.
Experimental tracings from schema of elastic tubing, combined
with sheep’s heart.
1. Lowest pressure. The dicrotic wave is about as high as
the primary, and is followed by other oscillatory waves.
16 DR GALABIN,
2. Low pressure. The dicrotic wave is still large, and the
tracing resembles the hyperdicrotic pulse of fever.
3. Higher pressure. This tracing is less dicrotic, and resembles
the pulse of slight fever, or of feeble health.
4. One semilunar valve divided. The dicrotic wave is less
than before the alteration (compare figs. 1 and 2), but still con-
siderable. It is followed by a second oscillatory wave.
Tracings from the same schema, combined with artificial heart.
5. Low pressure. This tracing resembles 2, obtained with the
sheep’s heart, and 13 the pulse of fever, but the rate being slower,
a trace is seen of the second oscillatory wave.
SECONDARY WAVES IN PULSE. 17
6. High pressure. This tracing resembles one of the forms of
healthy pulse.
7. High pressure: rigid tubing. The tidal wave here for the
first time makes its appearance, and the dicrotic wave becomes less
in proportion. This tracing resembles the pulse of atheroma.
8. High pressure: rigid tubing: long contraction. The tidal
wave is here much prolonged.
9. Aortic valves entirely removed : pressure low. The dicrotic
wave is still considerable, the tidal wave absent.
VOL. VIII, 2
18 DR GALABIN.
Pressure 23 ounces.
10. Healthy pulse. The tidal wave is seen preceding the
dicrotic wave, but only slightly marked.
Pressure 3} ounces.
11. Atheroma without kidney disease. The pressure is some-
what greater than normal, the tidal wave large, and the dicrotic
wave small in proportion.
Tressure 5 ounces.
12, Atheroma with granular kidney. The pressure and height
of upstroke are both increased, and the tidal wave is very large.
SECONDARY WAVES IN PULSE. 19
Pressure 1 ounce.
13. Dicrotic pulse of fever. Tidal wave absent; dicrotic wave
large: pressure low.
Pressure 3 ounce,
14. The pulse of a case of Bronchitis with chronic Alcoholism
taken a few hours before death. The pressure is excessively low :
the tidal wave absent: the dicrotic wave almost equal to the
primary, and followed by a second oscillatory wave, which is hardly
ever seen in the human pulse.
Pressure 5 ounces,
15. G. L. aged 18. Granular kidney. Tracing of radial
pulse. The pressure is very high, the tidal wave large and dis-
tinctly separated: after it is seen the dicrotic wave, and after the
dicrotic wave a third secondary wave, which is the third oscillatory
wave of the sphygmograph, the second such wave being superposed on
the dicrotic wave.
2—2
20 DR GALABIN.
Pressure 5 ounces.
=) 2¥6. Ga: Radial pulse. Lever of sphygmograph weighted.
The tidal wave is thus brought nearer to the primary wave, while
the dicrotic wave retains about the same relative position.
Pressure 4 ounces.
17. G. L. Tracing from brachial artery.
Pressure 4 ounces.
18. G. L. Tracing from dorsalis pedis. The tidal wave is
not more widely separated from the primary wave than it is in
the brachial or radial arteries, but, if anything, rather the reverse.
SECONDARY. WAVES-IN PULSE. 21
Pressure 3 ounces.
19. Tracing of dicrotic pulse from the femoral artery just below
Poupart’s ligament.
Pressure 3 ounces.
20. Tracing from the posterior tibial artery below the aukle
of the same person. The dicrotic wave is not nearer in propor-
tion to the primary wave, as it would be if it were a reflected or
retrograde wave.
21. Tracing copied from M. Marey, shewing how the tidal
wave is broken into two small waves by the use of a secondary
spring in the sphygmograph.
22. Tracing shewing the effect of percussion on the exterior
of a rigid part of the experimental tubing.
22 DR GALABIN. SECONDARY WAVES IN PULSE.
23. Diagram to illustrate the relation of the sphygmographic
tracing to the true pulse-wave. The thick line is intended to
represent the true pulse-wave, the thin line the sphygmographic
tracing, the dotted line the tracing drawn by a sphygmograph having
a secondary spring to keep down the lever.
a. Tidal or “first secondary” wave.
6. Dicrotic or “principal secondary” wave.
c.d. Two small waves into which the tidal wave may be
broken by the action of the secondary spring, as shewn in fig. 21.
ON THE AMYLOLYTIC FERMENT OF THE PANCREAS.
By ARCHIBALD LIVERSIDGE, F.C.S., late Scholar of Christ's
College, Cambridge.
(From the Physiological Laboratory in the Unversity of
Cambridge.)
THE (so-called) ferment of the pancreas was chosen as the
object of the following study rather than that of the saliva,
simply because it could be obtained much more readily in
sufficiently large quantities.
1. Mode of Preparation.
The method of V. Wittich (Pfliiger’s Archiv, 11. p. 193) was
selected as affording a means of obtaining the ferment in as
pure a form as possible in quantities sufficiently large.
Fresh pig’s pancreas having been freed from fat and
finely minced, the pulpy minced mass was placed in a flask and
covered with spirit; at first ordinary strong methylated spirit
was employed, but afterwards, so as to render the proteids more
completely insoluble, absolute alcohol wasused. After standing
for a few days the spirit was removed by straining through fine
muslin—the residue was next treated with pure strong glycerine
for 24 or more hours (sometimes the glycerine was left on the
pancreas for a month even). The glycerine extract was sepa-
rated by squeezing the mass through muslin, and then clarified
by allowing it to slowly filter through thick flannel bags (after
many long and patient trials of other methods, with filters of
various kinds and with the use of Bunsen’s aspirator, this plan
was finally adopted, for it was the only one which gave a clear
and transparent glycerine extract). The residue was again
similarly treated with three or four additions of fresh glycerine
until an appreciable quantity of the ferment was no longer
extracted. The filtered clear glycerine extract thus ob-
tained was then treated with many times its bulk of strong
spirit in large glass cylinders; after standing for some time
the clear supernatant fluid was syphoned off from the white
flocculent precipitate. This precipitate, when the alcohol had
24 MR LIVERSIDGE.
been removed by careful filtration and drying at a low tempera-
ture, was partly soluble in distilled water. The filtered aqueous
solution thus obtained was extremely active in converting starch
into sugar. But concentrated solutions (i.e. solutions obtained
by treating large quantities of the precipitate with small quan-
tities of distilled water) always gave most unmistakable proteid
reactions (xanthoproteic reaction and colouration with Millon’s
reagent). I never succeeded in getting this first glycerine
extract free from proteids.
To further purify the ferment thus prepared, the above
precipitate was taken, washed with strong clean spirit, and after
partial drying, by the spontaneous evaporation of the spirit, was
again treated with glycerine, and this second glycerine extract
was in turn precipitated by spirit.
This second precipitate was far less in bulk than the first.
Its aqueous solution had about the same activity in converting
starch into sugar as that of the first extract, while, though the
precipitate itself still gave some evidences of proteids being
present, the solution gave hardly any appreciable reactions
of them.
2. The Composition of the Ferment.
The ferment prepared as above by making two successive
glycerine extracts was dried at 100°C. On boiling with KHo
a small quantity of ammonia was evolved.
Ash. Heated on platinum foil the ferment intumesced and
evolved copiously a gas which burnt with a clear luminous flame,
and emitted a slight nitrogenous odour. A bulky coke was left,
but burnt off very readily and left a white very fusible ash.
I was prevented by my departure for Australia from com-
pleting, as I had intended to do, the analysis of the ferment thus
prepared, but Mr Frank Clowes was kind enough to make for
me the following determinations of the carbon, nitrogen, and
ash. Two determinations of each were made, ‘05 gramme being
used in each trial.
I. Il. Mean.
Carbon ‘017611 ‘017314 ‘017462
Nitrogen 005521 ‘005499 005510
Ash ‘1147 grm. gave ‘0174 ash.
THE AMYLOLYTIC FERMENT OF THE PANCREAS. 25
Taking the mean we have in percentage,
Carbon 34925, Nitrogen 11-020, Ash’ 157070)
A specimen of the ferment which had been prepared by a
single glycerine extraction oniy, gave in ‘05 gramme,
Carbon “022963, Nitrogen ‘006884,
or in percentage,
Carbon 45°926, Nitrogen 13:768.
A quantity of pepsin prepared in an exactly similar manner
by glycerine extraction from pig’s stomach gave, also in -05
gramme in two trials,
if Il. Mean.
Carbon ‘019199 0205927 | ‘0198958
Nitrogen ‘004995 0051115 0050532
Ash ‘1177 grm. gave ‘0194 of ash,
or in percentages,
Carbon 39°792, Nitrogen 10°106, Ash 16°48.
I do not think that any great importance can be attached to
these analyses. The material was certainly, in spite of my
care, mixed with some amount of foreign substance (filter frag-
ments, &c.); and, independently of this, one would be very rash
to infer that the substance obtained, even after a second treat-
ment with glycerine, was ferment and nothing else. The large
quantity of ash (principally potassic phosphate apparently) will
alone serve to indicate this. Still it is not uninteresting to
observe that the pepsin and pancreatic ferment have a very
similar constitution, widely different from that of ordinary
proteids. My own preliminary determinations had indicated a
still lower percentage of nitrogen and of ash.
3. Action of an aqueous solution of the ferment upon rodide
of starch.
Iodide of starch is almost immediately decolourized on
adding an aqueous solution of pancreatin, and the same quan-
tity of ferment will decompose successive quantities of iodide.
This decomposition takes place in the presence of free starch ;
26 MR LIVERSIDGE.
for on adding more iodine to the decolourized solution it again
acquires its blue colour. On adding potassium nitrite (KNO,)
and a drop of sulphuric acid the iodine is set free and the blue
restored. Hydroxyl’ also does this. But in neither case is the
restored blue quite so intense as the original. Chlorine water
does not answer well for this reaction. After standing for about
30 minutes the restored blue disappeared again, shewing that
the body, active in splitting up the iodide of starch, was still
unchanged.
There is then in the purified ferment solution a something
which splits up iodide of starch and brings the iodine into a
combined condition. V. Wittich is inclined to attribute impor-
tance to this reaction as throwing light on the mode of action of
the ferment. But I always obtained the same reaction with
boiled ferment which had completely lost its power of transform-
ing starch into sugar. Blood serum and other proteids, as is
well known, also decompose the starch iodide, and I find that
the action is quite similar to that of the ferment, being restored
by potassium nitrite in presence of an acid, &c.
4. Action of pancreatic ferment on other substances.
The modus operandi of an amylolytic ferment being the
adding on of a molecule of water, it seemed possible that the
ferment might be induced to bring about not only the particular
transformation of starch into glucose, but also other transforma-
tions which consist in the addition of a molecule of water. The
results were entirely negative, but are perhaps worth recording.
Salicine was tried first because the statement that saliva
and other amylolytic ferments will convert salicine into saliginin
and glucose is often found in text-books.
From a very large number of experiments it was found that
an aqueous solution of pancreatic ferment did not decompose
salicine into glucose and saliginin. A temperature of 40°C.
was used, and the time allowed was from 24 to 48 hours. The
pancreatic solution was always previously proved to be good by
allowing it to act upon starch paste; while, on the other hand,
if a solution of salicine was left for a longer period, even in a
1 The com!. 103 solution.
THE AMYLOLYTIC FERMENT OF THE PANCREAS. 27
stoppered bottle, more or less saliginin and glucose were pro-
duced from its so-called spontaneous decomposition.
Large quantities of the ferment were tried, but with no
‘result. Hence it seems probable that this spontaneous split-
ting up of salicine has been referred to the action of amylolytic
‘ferments.
It was thought, however, possible that the action which did
not take place in a mixture of the ferment and salicin alone,
might be brought about if the ferment were first of all thrown,
so to speak, into an active condition by the presence of starch ;
that the action of the ferment on the starch might be carried
on to the salicin. To a quantity of starch in active change
under the influence of pancreatic ferment, salicin was added,
but no saliginin was found at the end of 48 hours. On the
third day a little was found, but by that time the whole mix-
ture had become decomposed, and was covered with penicilium.
The same negative results were met with in attempting to
produce change in urea, tartaric, and oxalic acids, both alone
and in the presence of starch in active change.
The effect of the ferment on indigo blue was also tried with
like negative results.
The purified solution of pancreatic ferment, like all impure
amylolytic and other ferments, decomposed hydric peroxide with
evolution of oxygen; and the action was wholly absent when
the solution of the ferment had been previously well boiled.
5. Upon the regeneration of the ferment in the previously
exhausted pancreas.
Over some minced pancreas which had been twice extracted
with glycerine a stream of water was passed for 12 hours in a
tall glass. The washings still contained sufficient ferment to
decompose starch paste; this was ascertained by allowing the
washed residue to stand in a small portion of the washing
water for 30 minutes and then trying its effect upon starch
paste. But on continuing to wash for a longer period the whole
of the ferment appeared to be removed, for the residue ceased
to render a small quantity of water active. This washed and
inactive residue was now transferred to a filter and allowed to
remain on it exposed to the air for six hours: when on again
298 ; LES MR LIVERSIDGE.
treating it with a small quantity of distilled water it was found
to render the water very active. One and the same portion of
pancreas was in this manner deprived of its ferment four succes-
sive times, and each time the ferment reappeared. This form
of experiment was repeated on four different portions of
pancreas.
As a crucial experiment two quantities of pancreas were
placed in April, 1871, in 40z. flasks and the flasks filled up
with glycerine; after standing some time the glycerine was
filtered off (i.e. squeezed through muslin) and fresh added; even
after the seventh application of glycerine the pancreas still
yielded ferment. Altogether, these two quantities of pancreas
were treated with eleven successive doses of glycerine, and were
not rendered free from ferment until June, 1872, i.e. after
standing in comparatively large quantities of pure strong gly-
cerine for fourteen months.
Glycerine extract from flask No. 1, after standing on the
pancreas for eight days from the time of last filtermg and
washing (for it should be recorded that not only was the gly-
cerine removed in each case by squeezing through muslin,
but also the residue was well washed under the tap for some
time, so as effectually to remove every trace of glycerine), was,
on June 3, 1872, at last found to be inactive. Yet this same
residue, after standing on the muslin filter for six hours,
readily gave not only an active aqueous extract, but also yielded
an active glycerine solution.
On June 3rd the glycerine extract from flask No. 2 was
slightly active, but the residue after one more washing became
inactive, ie. did not yield any ferment to glycerine, and yet
after six hours’ exposure to the air it readily did so. As the
washed pancreas decomposes it evolves an odour of rotten cheese,
and the aqueous extract is then very active upon starch paste.
These experiments seem to shew pretty conclusively that the
ferment is regenerated in the exhausted pancreas.
The assertion of Bernard and others, that all decomposing
proteids are amylolytic has been disproved by Dr M. Foster
(see this Journal, 1, 1. 107); and we may fairly infer that there
is in the pancreas some substance (or substances), not in itself
active as a ferment, which in the processes of decomposition
THE AMYLOLYTIC FERMENT OF THE PANCREAS. 29
becomes converted into the ferment. It is worthy of note, that
though the solid washed tissue of the pancreas thus readily gave
rise to ferment, the aqueous solution of the ferment itself as
obtained by the glycerine method, when once its amylolytic
powers had been destroyed by boiling, never regained them
afterwards, at any stage of the decomposition which subse-
quently took place in it.
It was my intention to have attempted the isolation of this
antecedent (or antecedents) of the ferment, but my departure
for Australia put an end to my studies in this direction.
SyDNEY, Nov. 1872.
ON THE STRUCTURE OF THE TACTILE CORPUS-
CLES. By Grorce Tun, M.D. (Plates I, IL)
THE structure of the tactile corpuscles of Wagner is a subject
on which anatomists are not agreed. It is unanimously ad-
mitted that one or more medullated nerve-fibres have a course
through the corium to the corpuscle with which they come in
contact, that the corpuscle itself presents a well-defined contour,
that on its surface are to be observed a number of rounded or
oval bodies—the so-called transverse elements (quer-elemente)
of German authors—which by their more or less parallel ar-
rangement and number contrast with the cellular elements in
the contiguous tissue, and finally that, when subjected to the
influence of the agents commonly employed to prepare the
skin for microscopic examination, the substance of the corpuscle
seems to be composed of waving fasciculi of fibrille. The points
on which observers are not agreed are the fate of the nerve
after it comes in contact with the corpuscle, the nature of the
transverse elements, and of the substance of which the corpuscle
is composed.
Dr Allen Dalzell presented to the University of Edinburgh in
1853 an “Inaugural Dissertation on the General Integuments of
animals and their appendages’,” in which he records the results of
his examination into the nature of the touch-corpuscles as follows :—
“In very thin sections of integument, and when the knife had
evidently been carried in the same perpendicular as the nerves, a
power of 400 linear with acromatic light from a Ross condenser
never shewed the division (either within or without the corpuscle)
of the double contour nerve into pale filaments; an appearance often
observed by Wagner, twice confirmed by Gerlach, but never de-
tected by either Nuhn or Kélliker.” And again, “In touch-bodies
with a hyaline centre no nervous structures had been seen in section,
but repeatedly in those in which the so-called ‘cortical appearance’
was continuous throughout the touch-body, the appearance of two or
more circular apertures had been detected which might be the cut
ends of tubes.”
1 This Thesis is a manuscript volume in the Library of the University. It
has never been printed.
tr etna
DR THIN. STRUCTURE OF THE TACTILE CORPUSCLES. 31
Tomsa (Wien. Med. Wochenschrift, 1865) experimented by
boiling the skin in a mixture of alcohol and hydrochloric acid. The
theory of this procedure is that it leaves intact the axis-cylinder,
whilst the surrounding elements are dissolved. He states as the
result of his researches, that the nerve-fibre gradually loses its
medulla as it approaches the corpuscle, and making a number of
spiral windings before entering it, “splits up into a varying number
of branches whose section presents a polygonal contour.” The
corpuscle itself is, he holds, formed by cellular elements more or
less transversely disposed. “These flattened cells” are through
their very short prolongations continuous with each other and with
the fibrille of the axis-cylinders which enter the pedicle of the
corpuscle.” To other corpuscles he assigns a different composition.
“Other tactile corpuscles in the palmar surface of the hand are
composed of a coil of thickened axis-cylinder, an aggregation of
nerve-material, in which are embedded nuclei which are either
transversely arranged at the periphery, or are irregularly distributed
over the surface. The prolongations of the cells are here in the
background, and the corpuscle consists entirely of a compact mass of
nerve-substance formed by the confluence of the cell-protoplasma of
the nerve.” He states emphatically that the cellular elements of
the corpuscle do not belong to connective tissue elements.
Kolliker’ describes the corpuscle as composed of a cortical layer,
a nucleated capsule, and an interior of homogeneous clear connecting
substance with fine granular elements. He cites the opinion of
Krause that the transverse elements are nerves, and maintains, in
opposition to him, that they are transversely disposed nuclei (querste-
hende Kerne). In regard to their nature, he states that “ they perhaps
all belong to cells similarly arranged, to which the value of connective
tissue corpuscles might be assigned.” But he also adds—* that
there are often also in addition transversely disposed nerve-fibres
in considerable number is certain, but these are not the chief
cause of the transverse linear appearances.” In regard to the nerve
he believes that one to two, or even three to four, nerve-fibres run
along with or surround the corpuscle, and then “apparently ” enter
it, “ending in the superficial parts of the inner substance in pale
terminal fibres.” He further states, ‘but it would seem that the
nerves end in the superficial parts of the corpuscle and never pass
through its centre.”
Biesiadecki (in Stricker’s Handbook) refers briefly to these
poiuts as follows: ‘The transverse lines have been differently
explained as connective tissue, elastic, and nerve-fibres, the trans-
verse nuclei sometimes as connective tissue cells, sometimes as the
nuclei of the membrane of Schwann. According to some, after the
disappearance of the medulla the sheath of the nerve enters a
depression in the corpuscle, in which it has a free ending, analogous
to the corpuscles of Krause in other situations.
“Successful gold preparations decide some of these doubtful
1 Handbuch, 5th Edition.
ae DR THIN.
points, as they shew the nerve-fibres coloured dark violet, whilst
the remaining tissue appears of a pale reddish hue. The ,margin
of the corpuscle is indicated by a faintly marked contour in which
oblong nuclei lie. In fine sections 4—6 violet-coloured nerve-
fibres can be seen which are sometimes obliquely, sometimes longi-
tudinally arranged, and which are accompanied by more faintly
coloured small nuclei.
“But these fine sections teach nothing concerning the course
of the nerve in the interior of the corpuscle; they give no explana-
tion as to whether the fibres divide and as to how they end.”
Virchow (Cellular pathologie, 1871, p. 284), after enumerating
the various opinions held regarding the relations of the nerve to
the corpuscle, and expressing his conviction that Meissner is in
error in considering the corpuscle to be composed of nervous
matter, remarks, ‘“‘ It seems to me to be a matter of doubt whether
the nerve ends in the interior of the corpuscle or forms a loop in its
circumference.”
My investigations, most of which were conducted in the
spring of this year, in Professor Stricker’s Laboratory for Ex-
perimental Pathology in Vienna, have yielded me the results
which I have now to describe ’.
The methods I employed consisted chiefly in treating freshly
amputated skin by Chloride of Gold, Osmic Acid, and Carmine
and Acetic Acid.
Osmic acid has the great advantage of not shrinking the
tissues, and, when employed successfully, of giving remarkably
distinct and perfect specimens. Tuctile corpuscles consist of two
classes, Single and Compound. The greater number, inclusive
of all the larger ones, belong to the latter category.
A vertical section through the meridian of a corpuscle which
has been properly treated by Osmic acid, shews either a simple
homogeneous, more or less, rounded body, enclosed in a capsule,
or two or more such simple capsulated bodies arranged in a
row, parallel to the vertical axis of the papilla, and enclosed in
a common oblong capsule. The former I propose to designate
as Single, the latter as Compound Corpuscles. Compound
Corpuscles may again be conveniently divided into twms and
triplets, according as they are composed of two or three single
corpuscles, and for convenience im description, each of these
individuals may be termed a member of the Corpuscle. Exam-
1 Many of these results I submitted to the Academy of Sciences in Vienna,—
(Sitz.-Bericht. May, 1873.)
STRUCTURE OF THE TACTILE CORPUSCLES. 33
ples of Singles will be found in Plate L., Figures 3, 6, and 10,
of Twins in Figures 7 and 8, of Triplets in Figures 9 and
11. The space which separates the members of a compound
corpuscle varies in breadth (Figures 7, 8, 9 and 11). The
compound corpuscle is thus a conglomeration of Singles, and
must be regarded not as one, but as consisting of several
organs.
The well-marked unmistakable separation between the
members of compound corpuscles I have been able to demon-
strate only in osmic acid preparations, and then only when
the section happened to be through the meridian of the cor-
puscle. If the section has laid bare the capsule of a compound
corpuscle, the line of demarcation between its members can
only, even in osmic acid preparations, be inferred from the
depressions in the capsule (Figure 3). In gold preparations
the depressions in the contour corresponding to the bounda-
ries of the members are sometimes visible as in Figure 1, and
sometimes not as in Figure 2, probably according as more or
less shrinking has been produced in the preparation of the
tissue for examination. In whatever way the skin is prepared
the bulging of the members of compound corpuscles is often
seen (Fig. 12).
On account of the winding course of the nerve it is in the
majority of sections only seen in detached portions. Occa-
sionally no nerve is seen, which happens when the section has
hit the external portion of the corpuscle. When the prepa-
ration is successful, the nerve-fibre is seen with perfect dis-
tinctness within the substance of the corpuscle, the extent
to which it is visible depending on how far the direction
of the section has been parallel to the course of the nerve.
The result of the examination of a very large number of
specimens has shewn me that each single corpuscle, and each
member of a compound corpuscle, is penetrated by one, and
never by more than one, nerve-fibre.
I have never seen a nerve leave the substance of the cor-
puscle after having penetrated it.
Although a perpendicular course in the corium is the rule
for the nerve-fibres that terminate in the tactile corpuscles
it is not invariable. In triplets especially it is not rare to find
VOL. VIII. 3
34 DR THIN.
that the nerve of the upper member approaches it by winding
transversely from the summit of another papilla.
That the nerve penetrates the substance of the corpuscle
is demonstrable both in gold and in osmic acid preparations.
Before piercing the capsule it sometimes describes a curve, or
a complete spiral, or it may suddenly twist round itself and
form a loop, or it may simply enter in a straight line.
If the corpuscle is single, the nerve, as soon as it has tra-
versed the capsule, at once penetrates into its interior (Fig. 10).
If it is compound the nerve after it has penetrated the capsule
either immediately enters the member opposite its point of
entry, or takes an upward course within the common capsule
towards the member to which it is destined, mostly parallel to
the vertical axis of the papilla, but it may in its course cross
the corpuscle once or even twice transversely, and generally in
the grooves between the component members (Fig. 11). In its
course towards the member in which it terminates it follows no
fixed rule, sometimes running parallel to the long axis of the
corpuscle inside, or outside the common capsule, and sometimes,
as has been indicated above, approaching it in a line parallel to
the transverse axis of the papilla and on a level with its point
of entry to the capsule substance.
In gold preparations it can be distinctly seen that in com-
pound corpuscles a varying number of nerves enter at different
altitudes, and in osmic acid preparations it is seen that the
points of entry correspond to the relative position of the mem-
bers of the corpuscle.
When the nerve has entered the substance of the corpuscle
it penetrates to a certain depth either in a straight line or in
a curve, and then bends round and describes part of a circle.
The largest curve I have seen has not exceeded an are of 270°.
In this terminal course the nerve retains its medulla, and be-
tween the medulla and the corpuscle substance there is a clear
space visible in the osmic acid preparations (well seen in Fig.
10) which exactly corresponds to the position of the sheath of
Schwann.
I have not once seen the nerve-fibre divide either external
or internal to the capsule, and the number of preparations in
which I have obtained a clear view of the nerve before and
STRUCTURE OF THE TACTILE CORPUSCLES. 35
after its entry through the capsule is so considerable that I
do not hesitate to deny that the alleged division of the medul-
lated nerve in the tactile corpuscle exists’.
The inference from these facts 1s that each single corpuscle
and each member of a compound corpuscle represents the termi-
nation of a single medullated nerve-fibre.
The transverse elements, seen with varying distinctness
according to the manner in which the skin is prepared for ex-
amination, are especially prominent in gold and in acetic acid
preparations. They are best demonstrated by making a thin
section from fresh skin, colouring it in carmine, and then
allowing it to macerate in concentrated acetic acid for about
48 hours. Fig. 12 is drawn from a section so prepared.
By teasing gold and carmine and acetic acid preparations,
I succeeded in isolating fragments of corpuscles, and was able
to demonstrate the accuracy of the conviction I had acquired
from examining macerated specimens, that the transverse ele-
ments are the nuclei of oblong cells which anastomose with
each other by means of prolongations of elastic tissue fibres.
The distinction between the cell and the nucleus is not often
accurately defined, but I have seen it repeatedly and unmis-
takably both in gold and osmic acid preparations. ‘The cells
colour dark.in gold, which, in conjunction with their elongated
form, gives them an appearance which it is sometimes difficult ©
to distinguish from that of nerve-fibres. They are very abun-
dant in every part of the corpuscle, which may be described as
being in great part formed by a dense network of cells and
elastic tissue. The cells and elastic fibres seem to be as nu-
merous in the deeper as in the superficial parts, every change
of focus in a suitably prepared section bringing a new layer
into view.
1 Appearances are often seen both in the corium and within the capsule that
might easily be mistaken for the division of a nerve-fibre. For example, in
Figure 2 three nerve-fibres are seen ascending towards a corpuscle, and in one
part of their course one of the fibres lies so directly under another that only
two fibres are then visible. If in such a case the section did not happen to
include the deeper portion of the course of the fibres the illusory image might
be presented of two nerves in the upper part of the corium apparently becom-
ing three before entering the corpuscle.
In Figure 5 a second nerve is seen emerging at the side of a corpuscle almost
immediately under a first that is seen ascending through the corium, and so
producing an appearance that might be mistaken for a division of the first
nerve.
3—2
36 DR THIN.
In a successful attempt to colour the skin with silver, I
found the position of the corpuscles indicated by oval masses
of beautifully distinct granulated nuclei, which changed by
the slightest touch of the fine adjustment.
The capsule of the corpuscle is formed by a circular layer of
elastic tissue formed by the anastomosing continuations of cells.
The network of elastic tissue in the corpuscle and the cells
connected with it communicate in no way with the medullated
nerve-fibres.
The connecting matter of the corpuscle clears up with acetic
acid, and is coloured by gold and osmic acid the same shade as
the connective tissue of the surrounding corium.
The division of the papille of the skin into vascular and
nervous, I have not found borne out by wy investigations.
Virchow (Cellular-pathologie, Berlin, 1871, p. 285) remarks’,
“Tn the narrower papillae there is always a simple or dividing
vascular loop but no nerve. This observation is in so far important
as through it we have acquired the knowledge of a new nerve-less
part. In the other kind of papille, on the contrary, very frequently
no vessels but nerves and the peculiar structures designated as
touch-corpuscles are found.”
Again (Op. cit. p. 285) he remarks, ‘Setting aside the ana-
tomical and physiological question, the instance of the skin papillae
has a great value in explaining pathological appearances, because we
find here in parts in themselves perfectly alike two conditions that
are in absolute contrast”: one set of papille that have no nerves and
are rich in vessels, and another set that have no vessels and are only
provided with nerves.”
Isidor Neumann, in his text-book of Skin-Diseases (Second
Edition), in the Chapter on the Anatomy of the Skin, represents
the general opinion of German anatomists when he remarks that
“Compound papillae only, and these only rarely, contain both a
touch-corpuscle and a vascular loop; with this exception papilke
with corpuscles have no vessels.”
Dr Dalzell had already in 1853, in the Thesis from which
I have quoted, expressed himself in the following clear and decided
terms —“ Wagner’s division of the papille into purely nervous
+ In den schmalen findet man constant eine einfache, zuweilen eine
verastelte Gefiiss-schlinge aber keinen Nerven. Es ist diese Beobachtung
insofern wichtig, als wir durch sie zur Kenntniss eines neuen nervenlosen
Theiles gekommen sind. In der anderen Art von Papillen findet man dagegen’
sehr haufig gar keiue Gefiisse, sondern Nerven und jene eigenthiimlichen.
Bildungen, welehe man als Tastkérper bezeichnet hat.
2 Emerseits nervenlose und gefissreiche, andererseits gefisslose nur mit
Nerven versehene Papillen.
STRUCTURE OF THE TACTILE CORPUSCLES. 37
and vascular has no anatomical foundation either in man or the
lower animals, and even those papillae in which his corpuscula tacti
are placed very frequently contain one or more vessels.”
T have satisfied myself that when skin from a finger that
has been amputated whilst the capillary vessels are full of blood
is treated with osmic acid, nerves, touch-corpuscles, and rows of
blood corpuscles arranged in loops indicating the position of
the capillary vessels, are distinctly seen in the same papille.
(Plate IL, Figs. 13 and 14.) It is capable of easy demonstra-
tion that in a majority of so-called nerve-papille a thin
vertical section contains at the same time a section of the
corpuscle and one or more capillary loops. In the smaller
number in which nerve or touch-corpuscle is seen and no
capillary loop, it is quite possible that in the wide papilla
the vessel may not have been included in the section. What
is certain is, that at least the majority of papillee which con-
tain touch-corpuscles contain vessels also, and that the dis-
tinction of papillae into nervous and vascular has no foundation
in fact. Nerve-fibres are found in papille which have vessels
and no touch-corpuscles, and in a proportion that seems to bear
a relation to the number of fibres found approaching the rete
from the corium in a given area.
EXPLANATION OF PLATES.
Plate I. Figures 1 and 2 are compound corpuscles prepared by
treatment with Chloride of Gold. The nerves are only visible to
their point of entry.
Figure 3 is part of a compound papilla prepared by Osmic Acid,
showing two corpuscles with part of the course of the nerve in their
interior, and with part of a capillary loop indicated by blood cor-
puscles.
Figures 4 and 5 are compound corpuscles prepared by Chloride
of Gold, showing the course of the nerves in the interior of the
corpuscles,
Figure 6 is a single corpuscle prepared by Osmic Acid.
Figure 7 is a twin corpuscle prepared by Osmic Acid. Two
nerve-fibres are seen on one side of the corpuscle and a capillary
38 DRTHIN. STRUCTURE OF THE TACTILE CORPUSCLES.
vessel filled with blood corpuscles on the other. The nerve of the
upper member is seen entering the corpuscle substance.
Figure 8 is a twin corpuscle prepared by Osmic Acid.
Figure 9 is a triplet corpuscle prepared by Osmic Acid, with
sections of two nerve-fibres.
Figure 10 is a single corpuscle prepared by Osmic Acid, with its
nerve lying in its substance.
Figure 11 is a compound corpuscle prepared by Osmic Acid.
The nerve to the upper member is seen crossing in the grooves
between the members under the common capsule before entering.
Figure 12 is a compound corpusele which has been coloured with
Carmine and macerated in Acetic Acid, showing the nuclei of the
cells and the continuity of the latter with elastic fibres. (The relative
position of the figure in the plate is inverted.)
Plate II. Figures 13 and 14, from Osmie Acid preparations,
illustrate the co-existence of touch-corpuscles and vessels in the
same papillae. The figures represent vertical sections through the
skin. c. ¢. capillary blood-vessels ; n. nerve-fibre, t. touch-corpusele.
ns = ud
ey
NOTES ON THE STRUCTURE OF THE OLFACTORY
MUCOUS MEMBRANE. By H. Newer. Marty, D. Sce.,
M.B. Lond., Scholar of Christ's College, Cambridge. (PI. III.)
(From the Physiological Laboratory im the University of
Cambridge.)
I. THe EPITHELIAL CELLS.
SincE Max Schultze first described two forms of epithelial cell
as occurring in the olfactory region of the nasal mucous mem-
brane, and divided them into the two classes of epithelial or
supporting cells, and olfactory or special sensory cells, most sub-
sequent observers have confirmed him on every essential point.
Recently, however, Exner’ has denied the distinctness of the
two forms of cell described by Max Schultze, and maintained
that they gradually shade off into one another, numerous trans-
itional forms being found between them. The following notes
are based on observations which I have not been able as yet
to carry on to a systematic conclusion, but which, so far as
they go, lead me to confirm decidedly the anatomical distinct-
ness of the two forms of cells described by Max Schultze,
whilst, at the same time, they verify in very many points the
accuracy of Exner’s descriptions. The animals with which I
have principally worked are the newt and the dog, but I have
also employed the frog, rabbit, guinea-pig, and rat.
Olfactory Epithelium of the Newt.
In this it is quite easy to make out two distinct kinds of
cell, and I have never succeeded in discovering any intermediate
forms. In the newt, indeed, the two forms of cell are more
widely different than in any other of the animals mentioned
above ; it affords therefore an excellent subject on which to begin
the microscopical study of this tissue.
If the olfactory mucous membrane of this animal be treated
with Miiller’s fluid, and then teased carefully out in water,
1 Sitz. der K. Acad. der Wissensch. Wien. I. Abth. Jinner-Heft, 1870, u. III.
Abth, Janner-Heft, 1872.
40 DR MARTIN.
besides isolated cells floating about the field, there will be found,
here and there, groups of cells, each group consisting of one of
the so-called “epithelial” cells, surrounded by a number of
“olfactory” cells (Fig. 1). This appearance is so frequent that I
believe the whole epithelium in its natural state to be made up
of groups of this kind; so that its surface might be divided into
a great number of small areas, each of which would consist of a
central “epithelial” cell, with a number of “olfactory” cells
grouped round it. This arrangement is still more easily demon-
strable in preparations made by hardening the membrane in
alcohol, and then teasing it out in glycerine. The groups of
cells thus obtained form much more coherent masses, the “ ol-
factory” cells appearing imbedded in a uniting granular sub-
stance, which is apparently dissolved away by Miiller’s fluid
(Fig. 2).
When the epithelial cells (Fig. 3) are obtained from Miiller’s
fluid preparations, each possesses a large oval granular nucleus,
which sometimes contains a few large particlés looking like oil
globules. Around this nucleus les a homogeneous structureless
layer, with well-defined inner and outer margins; from one end
of this layer proceed, sometimes one, but usually several “ cen-
tral processes,” which are also homogeneous and structureless.
These processes often branch several times, presenting here and
there little angular enlargements which, but for their angularity,
might be called varicosities, and appear like prolongations of
the layer round the nucleus, no line of demarcation being ob-
servable between them and it. The “peripheral process,”
arising from the opposite end of the cells, is, on the contrary,
always single, and distinctly marked off from the layer sur-
rounding the nucleus. It is much stouter than the central pro-
cess, finely granular, and often obscurely longitudinally striated,
By staining these cells with iodine after treatment with
Miiller’s fluid, I have frequently been able to observe in the
peripheral process a central stained part, surrounded by a deli-
cate unstained loosely fitting outer membrane, as if the middle
part had shrunk away from a cell-wall (Fig. 4). I have never
succeeded in finding cilia on the peripheral process of the epi-
thelial cells, as described by Exner; but I have several times
met with the appearance shewn in Fig. 5, the central mass
STRUCTURE OF THE OLFACTORY MUCOUS MEMBRANE, 41
being produced at one side of its free end into a tolerably thick
prolongation.
The nucleus of these cells varies considerably in appearance
with the reagent employed in preparing the membrane. In
gold-chloride preparations it is granular, as in Miiller’s fluid;
but in chromic acid it is hardly granular at all; and, judging
from analogy with the dog, it probably appears non-granular in
osmic acid preparations also.
In some few cases (Fig. 4) it has seemed as if the central
processes were replaced by a delicate crumpled membrane ; but
I am inclined to believe that this is an optical illusion. Ba-
buchin* has, however, described these cells as having such a
membrane at their deep end, and says that he has stained it.
The olfactory cells in Miiller’s fluid preparations exhibit a
spherical nucleus very distinct from the oval one possessed by
the “epithelial” cells. This nucleus has a peculiar appearance
—it is not granular in the ordinary sense of the term—but
looks as if some highly refracting body were wrinkled or broken
into fragments so as to refract the light unequally and get a
sort of pseudo-granular appearance (Fig. 1). It is surrounded
by a hyaline transparent layer, exactly like that round the
nucleus of the “epithelial” cells; and from this proceed in
many cases a peripheral and a central process, both of which
are hyaline and transparent. The peripheral one is rather the
thicker of the two. The central one exhibits varicose enlarge-
ments, but does not divide, like the central processes of the
epithelial cells.
Many cells are always to be found about the field resem-
bling in most points those just described, but differing in having
no process, or only one. They have usually been considered as
mutilated olfactory cells; but the most careful examination so
frequently fails in detecting any indication of a broken-short
process, that I am inclined to believe that some of them may
be normal, and lie naturally imbedded in the network formed
by the deep processes of the “epithelial” cells. In some of
them the nucleus is double (Fig. 6). Cells resembling the olfae-
tory cells, but having their processes unusual in position or
number, are also occasionally found (Fig. 7).
1 Stricker, Handb. d. Lehre v. d. Geweben, Cap. xxxv. p. 969.
42 DR MARTIN.
Olfactory Epithelium of the Frog.
In this animal the “epithelial” and “olfactory” cells are
not so readily distinguished as in the newt. One chief cause
of this is, that the nuclei of both are oval, instead of that of
“olfactory” cells being round as in the newt. The peripheral
processes of the “epithelial” cells also are not relatively so
thick, compared with the corresponding processes of the olfac-
tory cells, as in the newt; and the central processes are fewer
(usually only one), and less branched. In these three points,
then, the two forms of cell approximate to one another; but
still, so far as I have seen, they can always be distinguished by
the smaller amount of protoplasm around the nucleus of the
olfactory cells, by the smaller diameter and less granular cha-
racter of their peripheral, and by the greater fineness of their
central processes.
In some cases I have seen the deeper end of the central
process of an “epithelial” cell ending in an enlargement, from
which again several small processes arise. This may possibly
be an indication of a less developed stage of that peculiar ar-
rangement of the deep ends of the corresponding processes
which is seen in the dog. (See Exner’s first paper, Taf. 1.
Fig. 12.)
Olfactory Epithelium of the Dog.
In this animal also the two forms of cells have always
appeared to me to be quite distinct; although both differ
somewhat from the corresponding cells of the newt.
In the “epithelial” cells the nucleus is smaller and rounder
than in the newt; and in osmic acid preparations it is often
very indistinct. The central process is comparatively thick,
and has little angular prominences here and there upon it. It
is always single, and never branches until towards its deeper
end. There it swells out into a large knot, exactly as described
by Exner in the rabbit; and from this a number of short thick
processes arise, whose general direction is in the plane of the
mucous membrane, and which appear to join those of neigh-
bouring cells, thus forming close to the sub-epithelial tissue a
sort of irregular network with small meshes and thick inter-
STRUCTURE OF THE OLFACTORY MUCOUS MEMBRANE. 43
secting trabecule. Just above its terminal enlargement each
process contains a number of granules, which stain very deeply
with osmic acid, forming a black patch; and the black patches
of neighbouring cells being apposed, form, in vertical sections
of the mucous membrane, a black line reaching all along it
near its base. This is well shewn in Exner’s second paper,
Taf. . Fig. 4. Similar stained granules are usually found in
small numbers in the terminal enlargement, and here and
there in the central process.
The peripheral process of these cells is essentially like the
corresponding process in the newt.
The “olfactory cells” are proportionately less numerous
than in the newt; and their central processes are finer. Their
nuclei are rather smaller than those of the epithelial cells, and
are oval, but rather pointed, at their poles. I have never
succeeded in tracing out the deep process of one of these cells
to its end, and cannot offer any opinion, as to whether it ends,
as Exner has described it in the human infant and some other
cases, in the network formed by the deep processes of the epi-
thelial cells. The peripheral processes of these cells are very
fine, but otherwise not peculiar.
The conclusion to which I have been led is, that the two
forms of cell met with in the olfactory region are anatomically
quite distinct, as described by Max Schultze, and do not
shade off into one another. I think the contrary opinion at
which Exner arrived is due in great part to his having at first
and chiefly worked with the frog, where the olfactory and epi-
thelial cells certainly do approximate to one another in several
points; although, even in the case of that animal, his own
figures do not seem to me to support the view of the existence
of such a transitional series of cells as he describes. With all
his other statements, so far as I have followed them, I entirely
agree.
In my descriptions I have adhered to the names epithelial
and olfactory as those by which these cells are best known, but
I am very doubtful whether they possess any such difference of
function as is thus implied. Under the best circumstances
anatomical structure alone affords a very uncertain basis from
which to deduce physiological function; and both these forms
4c DR MARTIN. THE OLFACTORY MUCOUS MEMBRANE.
of cell differ so much from those of any ordinary form of epi-
thelium, that there appears to me no reason for ascribing special
sensory functions to one more than to the other, and I am
inclined to regard them both as concerned directly in the sense
of smell. If both end, as Exner has described, in the same’
deep network, with which fibres of the olfactory nerve are con-
tinuous, it would seem to settle decisively the similarity or
intimate relationship of their functions.
DESCRIPTION OF THE FIGURES.
Fig. 1. Group of cells from olfactory mucous membrane of
newt. Miiller’s fluid.
Fig. 2. Cells from same hardened in alcohol.
Fig. 3. Epithelial” cell from olfactory region of newt. Miil-
ler’s fluid.
Fig. 4. Ditto Ditto. Treated with iodine solution.
Fig. 5. Ditto. Ditto.
Fig. 6. “Olfactory” cells with only one apparent process from
newt. Miiller’s fluid.
Fig. 7. Unusual forms of cell from newt. Miiller’s fluid.
Fig. 8. “Epithelial cells” from olfactory region of dog. Osmic
acid preparation.
Fig. 9. Ditto Ditto Miiller’s fluid. °
ON THE EFFECTS OF A GRADUAL RISE OF TEMPE-
RATURE ON REFLEX ACTIONS IN THE FROG.
By M. Foster, M.A., M.D., F.R.S., Prelector in Phy-
siology in Trinity College, Cambridge.
(From the Physiological Laboratory in the University of
Cambridge.)
Gotz’ observed that if a brainless frog be placed in a vessel of
water, and the temperature of the water be very gradually
raised to 40°C., no movements (beyond a few flickering spasms)
take place; the frog becomes at last perfectly rigid and dies
without any attempt at escape.
An uninjured frog, on the other hand, becomes violent in
its attempts to get away as soon as the temperature rises to 30°,
or thereabouts.
These observations I have verified repeatedly. They are
justly urged by Goltz as a very striking instance of the differ-
ence between the conditions of a frog with and without a
brain; but they present a new difficulty :—why the brainless
frog is not excited to reflex action by the stimulus of the hot
water. This difficulty is increased by the following facts.
Obs. 1. If a frog, from which the brain has been removed,
be suspended by the jaw with the legs hanging freely down, and
the toes dipping into a vessel of water, on gradually heating the
water the toes are withdrawn by reflex action as soon as the
temperature of the water reaches a little over 30°. The result
does not essentially depend on the rapidity of the rise. How-
ever slowly the water be heated, the feet are always withdrawn
at a temperature of 35°, or earlier. Rapid heating may possibly
lower the degree at which the feet are-withdrawn ; but to this I
have not paid particular attention. Whether heated slowly or
rapidly the feet are withdrawn at about 35°C. or at a lower
temperature.—Obs. 2. If the whole body thus suspended be
similarly immersed and heated, no movements (or only the very
slightest spasms of the muscles of the legs) take place; and on
1 Functicnen der Nerven-Centren des Frosches.
46 DR FOSTER.
still further raising the temperature the body becomes rigid
(rigor caloris).—Obs. 3. If both legs be immersed up to the anus,
and similarly treated, they also become rigid without move-
ments either of the legs or of any part of the body, save only a
few spasms.—Obs. 4. If one leg only be immersed and simi-
larly treated, it also becomes rigid without movements, or with
only slight movements.— Obs. 5. If both legs (or one leg) be
immersed up to the knee, they are sometimes withdrawn ; but
sometimes no movements take place, and the portion immersed
becomes rigid. The results in this case are not so constant as
when either more or less of the body is immersed.— Obs. 6. If
the feet only be immersed, they are invariably withdrawn at
35° C., or under.— Obs. 7. If a frog be suspended over a vessel
divided by partition, with water at unequal levels on the two
sides, so that one leg is wholly immersed and the foot only of the
other leg, and the vessel be surrounded with water, the tempe-
rature of which is gradually raised, neither the leg nor the foot
will be withdrawn, if care be taken that the water on both sides
of the partition be equally and uniformly raised in temperature.
If in this last observation the water on both sides be reduced
to the same level, both feet are withdrawn. This result shews
that warm air and vapour have not the same effect as warm
water, and that the absence of movements is not due to the
unavoidable contact of the thighs of the animal with the top
of the partition giving some support to the legs, and thus
diminishing the tendency to the withdrawal of the feet.
The above observations shew that when the toes (alone im-
mersed in water) begin to be affected by the high temperature,
say 30°C., the stimulus of the hot water causes a reflex action
which results in the withdrawal of the foot. When the whole
leg or body is immersed, the same stimulus is still at work,
but no reflex action occurs. What is the reason that reflex
action is absent?
The following explanation is perhaps the first to offer itself.
The warmth applied to the leg diminishes the irritability of the
nerves or of the muscles, or of both; and thus the impulses
generated by the warm water in the sensory terminations of the
nerves of the foot are not carried up to the cord owing to the
diminished irritability of the sciatic trunk, or, being so carried,
a
rrr
EFFECTS OF TEMPERATURE ON REFLEX ACTIONS IN THE FROG. 47
the reflex process taking place in the cord cannot manifest itself
on account of the diminished irritability of the muscles or motor
nerves.
But this view is clearly untenable. It requires that the
nerves and muscles, covered and protected by the skin, should
be affected before the sensory terminations in the skin itself.
Moreover, no appreciable difference in the irritability of the
nerves, trunks or muscles of a leg thus exposed to 35° C., could
be detected. And it is directly contradicted by Obs. 7, where
the immersion of one leg prevents movements in the other.
Two other views then suggest themselves.——(1) The blood
returning from the legs being warmer than the normal, raises
the temperature of the spinal cord above the normal; this
reduces the irritability of the cord, and hence reflex actions set
going by a feeble stimulus, which in a normal cord would mani-
fest themselves, are here absent—(2) From the stimulation of
the whole leg as compared with that of the foot, a multitude
of impulses, arising from all parts of the skin exposed to the
warm water, reach the spinal cord. These produce such an
effect upon the cord that the simpler reflex action resulting
from the stimulation of the toes alone is prevented.
At first sight it seems easy to separate these two different
agencies. For if the reflex excitability of the cord be lowered by
the heated blood, it will be lowered all over, and reflex actions
will be lessened not only in the immersed legs but in the trunk
and fore-legs above the water. Practically, however, in the case
of the frog it is extremely difficult to.estimate excitability quan-
titatively by means either of mechanical or of chemical stimuli,
in any other part of the frog’s body than the legs, when only
slight variations have to be accurately determined. With the
legs one can estimate by the dilute sulphuric acid method (che-
mical stimulus) with tolerable accuracy slight variations in exci-
tability; though even here mechanical or electric stimuli are
unsatisfactory. But one cannot easily apply the acid method to
other parts than the feet, and hence the difficulty. or it is
with slight variations that we have here to deal; the legs, which
remain perfectly quiet in warm water, are at once drawn up
when even slightly pinched or touched.
I attempted to eliminate the effects of rise of temperature
48 DR FOSTER, z
by using some other form of gradually increasing and uniformly
applied stimulus. .
I tried to do this by using dilute sulphuric acid, the
strength of which was steadily increased. The legs were sus-
pended in a small beaker of water, into which water from
a very large beaker was continually flowing at a very slow rate,
the surplus being removed by a syphon at the same rate. Into
the larger beaker dilute sulphuric acid was dropped, with con-
tinual stirring. The frog’s legs were thus brought in contact
in as uniform a manner as possible with dilute acid of gra-
dually augmenting strength. I invariably found that when the
acid reached a certain strength violent movements took place,
whether the foot only were immersed or the whole legs. This
result however is not conclusive, for the even slight movement
in the fluid of the small beaker might be considered as suffi-
cient to prevent uniform stimulation of the skin. For exactly
the same results were obtained when warm water was applied
in the same way as the acid. Instead of the legs remaining
quiet, as when the water is still (v.e. moved only by the cur--
rents of heating), they were withdrawn, with violent movements,
when a certain temperature was reached. I further attempted
to eliminate the effects of heated blood by ligaturing the legs
underneath the sciatic nerves so as to cut off all vascular con-
nection between the legs and the trunk, while leaving the
nervous connection intact, and immersing the legs of the animal
so disposed in gradually heated water. The diminished irrita-
bility however due to the lack of blood-supply and the expo-
sure of the plexus of nerves directly to the vapour and elevated
temperature, interfered with the course of events, and I could
get no satisfactory results.
Similar attempts were made to obtain analogous results
with cold instead of heat; but they resulted in failure. In
the first place it was found that (with winter frogs) immersion
of the feet directly in water at 0°C. produced no reflex action ;
a fortiori none was produced by the gradual cooling of the
water in which the feet were immersed. Only when the feet
became actually entangled in the forming ice which spread
from the sides of the vessel towards the centre, was any move-
ment visible. Here the stimulus was probably mechanical, due
EFFECTS OF TEMPERATURE ON REFLEX ACTIONS IN THE FROG. 49
to traction from the fixed foot, as the animal swung to and fro
in the fluid, from the vibrations of the room in which the obser-
vation was carried on.
An attempt was also made to use olive oil instead of water ;
but this failed too, partly from the difficulty of reducing suffi-
ciently and uniformly the temperature of the large body of oil
needed for immersion, and partly because the legs were fre-
quently withdrawn when immersed in the oil at the ordinary
temperature.
I took the trouble to make these observations because, I am
free to confess, I had first leaned to the idea that the chief
factor in the matter was the uniform stimulation of a large sur-
face. I called to mind the fact that when we dip one foot into
hot water we localize the sensation of heat as most intense in a
ring round the ankle marking the level to which the hot water
reaches. What we are really conscious of in this case is the
contrast between the condition of the surface of the skin in the
hot water, and that of the surface outside the water, and this
contrast we feel most intensely at the junction of the two sur-
faces. Normally we are not conscious of the condition of our
whole skin when not affected in any particular spot; and yet
we have as it were an unrecognized background of such a con-
sciousness with which we compare any local affection. Thus we
feel more acutely the temperature of a fluid when we plunge
our hand or foot only into it than when we immerse our whole
bodies in it. So also it is a matter of common experience that
tickling is most effective when the stimulus is applied to a very
small surface. The touch of the tip of a feather on the sole of
the foot at once produces reflex movements; but the contact of
a large pad of cotton wool applied with the same pressure as
the feather over a large surface of the sole, may be borne
without any uncomfortable sensations, though almost each point
of the same surface stimulated separately would at once cause
the sensation of tickling. The mutual effect of two neighbour-
ing sensations may also be shewn in the following way. Esti-
mate on any surface of skin by Weber’s method the distance at
which the sensations of two points merge into one. Then sur-
round the spot of skin so tested with a rim of metal, pressed
down with sufficient force to be distinctly felt, but not more.
VOL. VIII. 4
50 DR FOSTER.
Test again the power of localisation in the skin within the
rim; the distance at which the two points first appear at one
will be much increased. Remove the rim and test again. The
distance will be found to have returned to its former limit.
The simultaneous sensations of the rim have dulled (in some part
or other of the nervous mechanism) the sensations arising
within the rim. A similar explanation may be given of the
fact that it is much more difficult to call forth a reflex action
by applying a galvanic stimulus to the nerve-trunk of a frog
than by applying the same stimulus to a portion of skin to
which some of the fibres of that nerve-trunk are distributed.
In the one case a multitude of sensory impulses reaches the
cord, in the other a few only, yet movement is absent in the
former, though present in the latter. The immersion of the
whole leg or of the body in the observations described above
may be taken as analogous to the stimulation of the whole
nerve-trunk ; the immersion of the foot only as corresponding
to the stimulation of a portion of skin.
On the other hand, in all observations on the effect of a rise
of temperature on living animal tissues, the state of exhaustion
or depression which ultimately ensues is preceded by a stage of
exaltation in which the functions of the tissue are raised above
the normal. This is well shewn in the case of muscles, nerves,
and the heart. In none of the observations recorded above was
there any indication of such an initiative stage of increased
action. Had there been it would naturally have led to the
withdrawal of the feet in all cases. And the absence of this
presented a great difficulty to considering the results obtained ~
as being merely due to a depression of the powers of the spinal
cord by reason of the increased temperature. Some observations,
however, made in the Laboratory here by Mr T. O. Harding,
afforded a clue, by pointing out a distinction between simply
and directly raising the temperature of an organ or a tissue,
and indirectly heating it by supplying it with blood heated
beyond the normal in some distant part of the economy. Thus
the heart of a frog, either empty or filled with serum, when
heated beats with a more frequent rhythm and, at first, with
greater force. But the same heart when indirectly heated by
the immersion of the legs of the frog in hot water (the heart
EFFECTS OF TEMPERATURE ON REFEX ACTIONS IN THE FROG, 51
remaining in the body and the brain and spinal cord being
destroyed) is lowered at once both in the force and frequency
of its beat, by reason of the heated blood with which it is
supplied. This result leads us to expect that in the same way
the spinal cord, if heated by being supplied with blood heated
beyond the normal, would be depressed without any preceding
stage of exaltation, and thus reflex actions which otherwise
would have occurred be prevented.
The observation (Obs. 7) where the heating one leg prevents
reflex action in the other, seems to point distinctly to such an
explanation. But the following observation shews still more
clearly that, whether or no the stimulation of a large surface
may assist in producing the effects described, the main cause is
the heating of the spinal cord.
Obs. 8. A brainless frog was so placed in a vessel with a
hole in the bottom, that the body and forearms could be ex-
posed to the action of water the temperature of which was
gradually raised, while the whole of both legs from the hips
downwards hung freely from the vessel, and were not subject to
the action of the heated water. Though in an unusual posi-
tion the frog remained quiet in the absence of stimulation, and
executed reflex movements when stimulated so long as the
water in the vessel above remained at the ordinary tempera-
ture ; thus when the toes were made to dip in water gradually
warmed, the legs were drawn up after a while as usual. When
however the temperature of water in the vessel above, and that
in which the toes were dipped below, were both raised part
passu, no movements at all took place, and ultimately, as the
temperature continued to rise, the body above and the toes
below became rigid from rigor caloris (the legs and thighs
remaining supple), without any save the slightest spasm.
Tested by the dilute sulphuric acid method (which was here
practicable) the reflex excitability of the spinal cord diminished
as the temperature in the vessel above rose, without any signs
of an initiative stage of exaltation.
The depressing éffect of a rise of temperature (especially up
to or beyond 35°C.) on the energies of the spinal cord of frogs
is well shewn in the case of tetanus. Two frogs A and B were
each poisoned with a similar small dose of strychnia. To A
4—2
52 DR FOSTER.
nothing further was done. B, as soon as the spasms manifested
themselves, was immersed in water at 37°C.; the tetanic con-
tractions almost immediately disappeared; and the animal
when taken out was perfectly flaccid, neither tetanus nor ordi-
nary reflex action being excited by stimulation. After a short
while the tetanus returned, and was again removed by a second
immersion. ‘This was repeated three times with a like result,
the frog A all the while remaining im a state of complete
rigidity, and ultimately dying long before B.
We may conclude then that the absence of reflex action in
Goltz’s experiment, and the other modifications of it, are due
primarily and chiefly to the depressing influence of heated
blood carried from the skin to the spinal cord. But this depress-
ing influence comes into play by virtue of the gradual character
of the stimulation. Dipping a frog either wholly or partially
into water of 27°C. or above, at once produces violent move-
ments. When the temperature however is raised gradually the
effect on the sensory organs of the skin is much less, and a
higher temperature has to be reached before a sensory impulse
is generated strong enough to give rise to a reflex action. But
by the time that higher temperature is reached the spinal cord
has already begun to flag and needs a still stronger impulse,
and therefore a still higher temperature in the water acting
on the skin; when that still higher temperature is reached the
energies of the spinal cord have sunk still lower, and so on
stage by stage, until the frog is boiled without having made
a sign.
The absence of reflex action with a gradual rise of tempera-
ture is still further insured by the stimulus being uniformly
applied, @.e. by the water being kept as still as possible, and
assisted, we may add, by the exposure to the stimulus of a large
amount of sensory surface at the same time. Both these cir-
cumstances tend to put off the reflex action till a higher tem-
perature is reached, and thus assist in preventing it altogether.
It would be interesting to inquire how far the distinction I
have suggested above between directly heating an organ and
indirectly heating it by supplying it with blood heated in the
extremities is general; and if so, to what changes in the blood
the effects are due. They would probably be best shewn in the
EFFECTS OF TEMPERATURE ON REFLEX ACTIONS IN THE FROG. 53
frog, whose sensitiveness to elevated temperatures is unfor-
tunately only too well known to physiologists working in the
summer, ‘The effects of heating the blood of mammals has
already been shewn to be peculiar (see Fick, Pfliiger’s Archiv. v.
p. 38).
There remains the question—why does the frog in posses-
sion of a brain not behave in the same manner? Why are not
his sensations and cerebral processes dulled in the same way by
the heating of the blood? The answer simply is, that a less
intense sensory impulse is needed to call forth a movement of
volition, that is, a movement carried out by the encephalon,
than an ordinary reflex action, that is, a movement carried out
by the spinal cord alone. The water as it is being warmed
suggests a movement to the intelligent frog long before it is
able to call forth an unintelligent reflex action. The very first
movement of the frog, the removal of any part of his body out
of the water, increases the effect of the stimulus; for the return
of the limb to the water already warm gives rise to a stronger
stimulus than contact with water raised to the same tempera-
ture while the limb is still in it; and thus one movement leads
to another, and the frog speedily becomes violent. It is nearly
the same with the brainless frog, when a movement has for
some reason or other been started; only in the observations we
have been dealing with this initial movement is wanting.
ON THE LAW WHICH REGULATES THE FREQUENCY
OF THE PULSE. By A. H. Garrop, B.A. Cantab.
(Continued from Vol. vu. p. 219.)
In a paper on Cardiograph tracings’ from the human chest
wall, in the 5th Vol. of this Jowrnal, I have endeavoured to
substantiate a law respecting the elements of the heart’s beat,
which may be thus enunciated :—
The heart’s beat consists of two parts, which for any given
pulse-rate do not vary in their ratio to one another; but the
length of the first part varies inversely as the square root of
the rapidity of the pulse.
A second series of measurements of the cardio-arterial inter-
vals, published in the Proc. of the Royal Society, London, have
further verified the law just stated, and in the rest of this paper
it will be assumed as proved. No theory respecting the cir-
culation throws light on its significance; but the one which
it has been my endeavour to demonstrate above gives a very
satisfactory explanation of it, which will now be considered in
detail.
First, the heart’s beat consists of two parts, which for any
given pulse-rate do not vary in their ratio to one another. It
having been proved previously that the pulse-rate does not
depend on the blood-pressure, and, as shewn now, the length
of the first part of the heart’s beat not varying when the pulse-
length is constant, it is evident that the length of the first
part of the pulse-beat does not depend on the blood-pressure in
any way.
Again, the first part of the pulse-beat is compound, for it is
the interval between the commencement of the cardiac or ven-
1 Since writing the paper referred to, a further comparison of tracings has
shewn me that in the slow pulses taken while lying, I mistook the primary sys-
tolic rise for the auricular, and was so led to the conclusion that the length of
the cardiac intervals depended in some measure on the position of the body.
This is incorrect, as subsequent measurements shew me, and the length of the
first part does not vary with the position of the body; the proper equation for
finding the cardiac first part under all circumstances being ry =204/z.
LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 55
tricular systole and the closure of the semilunar valves; there-
fore it may be divided into the systole and the valve-closure
interval.
Physiologists have laid very little stress on this valve-
closure interval, it generally being considered as instantaneous.
But in the study of cardiograph tracings it is to be remem-
bered that the distances between events occurring within one-
fiftieth of a second of one another can be appreciated without
much difficulty, and there is every a priorz reason for believing
that this interval has a longer duration than that. In my
paper on the Cardiograph trace, reasons have been given for
the belief that in quick pulses the commencement and the end
of this valve-closure interval are indicated by separate and
distinct changes of direction in the curve, and its length as
obtained by measuring from these points agrees entirely with
that required from arguments to be mentioned further on. It
may be called the diaspasis, that is, the period during which
the heart is being opened out by the regurgitation of blood
from the arteries.
The length of the combined wen: and diaspasis not de-
pending at all on the pressure, and it being constant for any
pulse-rate, it is infinitely probable that the systole and dia-
spasis separately are independent of the pressure; and this is
extremely interesting, as it gives a further insight into the
mechanism of the heart. For, in order that the duration of the
diaspasis should not vary with different blood-pressures, it is
evident that with higher pressures there must be greater ob-
struction to the heartward flow of blood, otherwise the valves
would then close more quickly. And this is exactly what
would be expected from the combination of Mr Bryan’s obser-
vations concerning the shape of the heart, and Briicke’s theory
of the active diastole of the ventricles’, According to the
latter author the cardiac muscular tissue has no inherent power
of opening out the ventricles, but remains inactive after systole,
during diaspasis in fact, until the regurgitation from the aorta
1 Mr Bryan’s paper is in the Lancet, Feb. 8th, 1834.
Briicke’s theory appeared in Sitzungsberichte der Wiener Akad. der Wiss.
1854, Vol. x1v. p. 345. See also a paper on the same subject by myself in this
Journal. May, 1869.
BO MR GARROD.
-has closed the aortic valves and so uncovered the orifices of the
coronary arteries, immediately upon which, the resulting sud-
den turgescence of the heart’s walls makes them open up.
-Mr Bryan has shewn that during systole the whole heart alters
its position as a result of its change in shape during contraction,
and recovers it during diastole ; therefore the greater the force
of contraction the more will it alter its shape, and the more
difficult will it be for it to resume the original one, which has
to be done partly by the regurgitating arterial blood; but
the greater the blood-pressure, the greater will be the facility
for overcoming this greater work, which two, as must be the
case, vary together. This argument explains how the diaspasis
need not vary in length with different blood-pressures.
Next, with regard to the systole. As the first part of the
heart’s beat varies as the square root of the length of the beat,
and as the diaspasis, a part of that first part, does not vary
with the blood-pressure, upon which, and the rapidity of ten-
sion fall (which must be a comparatively insignificant force), it
can alone depend, it is necessary that the other component of
that interval must vary more than as the square root of the
pulse-length. And to find how much more quickly, it is
necessary to obtain the actual length of diaspasis in a par-
ticular case, from which a close approximation can be arrived
at as to its duration at all rapidities. Careful measurements
of a cardiograph trace, beating 102 in a minute, give the ratio
of the systole to the whole beat as 1 to 3:1915, and that of
the first part to the whole beat as 1 to 2°0, which leaves the
ratio of the diaspasis to the beat as ‘187 to 1, or the diaspasis
length as (00183 of a minute. <A very similar length of dia-
spasis is found from quicker pulses.
The great interest attaching to these figures is that, when
with the diaspasis equal to ‘00183 of a minute, the ratio of
the systole to the diastole is enquired into, it is found that
there is a very simple relation between them, and that after
subtracting this diaspasis of nearly constant length, there re-
mains the systolic varying as the square root of the diastolic
period, and with no other diastolic length is so simple a ratio
obtainable, which is all-important, because it will be seen that
the systole must depend directly on the previous diastole.
LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 57
Next, considering the systole itself; the fact above demon-
strated, that its length does not depend on the blood-pressure
is extremely important, and can only be explained by assuming
that when the pressure rises, the circulation through the coro-
nary vessels increases to a sufficient extent to enable the heart
to get through the extra work it has to perform without alter-
ing the duration of its action, or in more precise terms, the
nutrition of the walls of the heart must vary directly as the
blood-pressure in the aorta.
But the systolic length varies as the square root of the
diastolic if the argument which is developed above is correct,
in other words, the longer the time of nutrition of the heart, the
longer is the systole. This at first sight seems an anomaly,
but the theory that the pulse-rate depends on the fall in tension
only, presents a most complete explanation, and so throws great
light on cardiac action in general.
Consider the heart as a pump working against a certain
pressure, and filling an elastic reservoir with a certain resist- —
ance to the outflow of its contents. Varying the pressure has
been shewn to have no effect on the lengths of the different
parts of the pulsation for the reasons given above; and it has
next to be considered how it is that varying the resistance
changes the lengths of the elements of the revolution. This
pump, directly its muscular fibres begin to contract, exerts its
full pressure, for there is nothing to prevent it doing so. But
during the previous diastole it was supplied by blood at a cer-
tain definite pressure and for a definite time, both of which
factors limit the force of the systole. Consequently the ven-
tricles produce directly their full systolic pressure, and main-
tain that pressure until they are empty. But it is evident that
the time necessary for emptying them of a definite amount of
blood under these conditions must depend on the rapidity of
the flow from the capillaries, for when the flow is halved the
systolic time must be doubled, if no other force come into play ;
in other words, the length of cardiac systole is a function of
the arterial resistance ; and the pulse-rate has also been shewn
to be a function of the same, upon the fall of tension theory.
It has been proved that the systole varies as the square
root of the diastole, not directly with it, as might be supposed.
58 MR GARROD.
This clearly shews that the time of diastole influences the
length of the systole and shortens it, in other words, strengthens
the heart, according to the law which may be stated thus, the
nutrition of the heart varies as the square root of the time
during which the coronary circulation is maintained.
It will strike some as peculiar that no mention has yet been
made of the influence of the nervous system on the heart. But
it appears to me that the facts which have been brought for-
ward have not called for any special reference to it. May not
the law it has been my endeavour to prove, be but an expres-
sion of that action in the healthy body? For it must depend
on a somewhat complicated mechanism, as is shewn by the fact
that it is almost impossible to contrive a self-acting engine
which would pulsate in accordance with its requirements.
As is well known, the effect on the kymographion trace,
of slightly stimulating the pneumogastric nerves, is greatly to
amplify the oscillations, and at the same time to lower the
mean pressure ; while cutting them produces the reverse effects.
The larger oscillations of the hemadynamometer column in the
former case, shew that the proportionate tension-fall and the
time of pulsation are both greatly increased, and from previous
considerations it is evident that these are necessarily associated,
when as now, no influence is being exerted on the peripheral
vessels’.
Further, these amplified oscillations must be attended with
an abnormal enlargement of the ventricular cavities during
diastole, for the time intervening between the beats being
increased, the amount of blood which flows through all segments
of the circulation between any two pulsations, must be also
more considerable. Having arrived so far, it is extremely in-
teresting to observe how an augmentation in the degree of
cardiac dilatation during diastole, as a cause, will include and
correlate all the peculiarities which are observed when the pneu-
mogastric nerve is thus operated on; and it is not unreasonable
therefore to suppose that this is the direct effect of its action.
As the quantity of blood contained in the heart at the end
1 In rabbits the normal fall of tension as judged by the hemadynamometer
trace is about ,th of the whole, while when the pneumogastric is stimulated it
may increase to ;',th or more.
tei
ee ee
LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 59
of diastole has been shewn to depend on the circulation through
the coronary vessels, it is evident that the explanation of any
variations in the capacity of the ventricles must be referred to
changes in the cardiac walls themselves. Just as the degree
of rigidity of an india-rubber tube through which a current of
water is flowing, can be made to vary by changing the diameter
of the orifice from which the fluid is allowed to escape, so the
turgescence of the ventricular walls, or what is the same thing,
the amount of active diastole of the heart, can be altered, by
varying the diameter of the small arteries of the coronary sys-
tem, their contraction producing a greater, and their dilatation,
by facilitating the flow of blood through the capillaries, a less
degree of diastolic enlargement of the ventricular cavities.
From the above argument, therefore, the amplified range
of pressure and time depending on change in heart-capacity,
and the change in capacity being caused by modification in the
calibre of the smaller coronary arteries, it 1s almost a logical
necessity that the function of the pneumogastric nerve is to
regulate the degree of tonicity of those vessels, and Dr Brown-
Séquard, from entirely different facts, has also published it as
his belief that the pneumogastrics contain fibres which contract
the small coronary vessels’.
It will be noticed that throughout this paper it has been
assumed that the systole never recommences until the ventri-
cular cavities are completely filled, that is, until a pressure
equilibrium has been arrived at in the interior of the heart.
Perhaps it is the absence of pressure which admits of the heart
recontracting, but this is a doubtful point, and until more is
known as to the mechanism of muscular action in general, it is
probable that the question as to the reason why the heart re-
commences to beat at a particular moment will remain unset-
tled. Sir J. Paget’, when he pointed out the relation of
rhythmic nutrition to rhythmic action of nerves and muscles,
laid the foundation for a scientific treatment of the subject,
and the law which it has been my endeavour to substantiate, is
only a precise method of expressing that relation.
1 See Principles of Human Physiology. By Dr. Carpenter, 1869, p. 219,
foot-note.
3 Croonian Lecture. Royal Society, 1857.
60 MR GARROD.
The following summary of the main features in the circula-
tion, as they appear to me, may assist in explaining some of the
previous arguments.
The circulation of the blood is maintained by the repeated
contraction of the heart. Each cardiac revolution is divided
into three parts, the systole, the diaspasis, and the diastole.
The following laws hold with regard to the length of these
intervals.
I. The systole together with the diaspasis, or in other
words, the first cardiac interval varies as the square root of the
whole revolution.
II. The systole varies as the square root of the diastole.
III. The diaspasis varies very slightly with different pulse-
rates.
The amount of work that the heart has to perform in main-
taining the circulation depends on two sets of changes which
may occur in the system; 1. Variations in the blood-pressure.
2. Variations in the resistance to the outflow of that fluid from
the arteries.
As the capacity of the arteries including the ventricles,
varies directly as the blood-pressure, and as the flow of blood
from the capillaries does the same, the frequency of the heart’s
beats is dependent on the resistance to the capillary outflow,
and not at all on the blood-pressure; in other words, the
heart always recommences to beat when the blood-pressure
in the systemic arteries has fallen a certain invariable pro-
portion.
Variations in blood-pressure result from: 1. Absorption
into, and excretion from, the vascular system, of fluids. 2.
Changes in the capacity of the arterial system, which occur
on the contraction or relaxation of the muscular arteries. 3.
Changes in the amount of available blood, which result from
the hemastatic dilatation of some of the yielding vessels on
altering the position of the body. As changes in the first of
these cannot be very sudden, and those in the latter are never
very considerable, the mean blood-pressure in health varies but
little during short intervals.
Variations in peripheral resistance result from: 1. Different
degrees of tonicity or patency of the muscular arteries. 2. Dif-
LAW WHICH REGULATES THE FREQUENCY OF THE PULSE. 61
ferent resistances in the venous system. The former may occur
independently in one or other system of vessels, as the cuta-
neous or the alimentary; also mechanically from pressure on a
part of the body. The latter are insignificant in health.
The heart depends for its power of doing work on chemical
properties in the blood it pumps into the systemic vessels, and
as the blood reaches it direct from those vessels, the cardiac
intramural circulation varies with the changes in the former ;
and the length of the systole varying only as the square root
of the time of diastole, the degree of cardiac nutrition varies
directly as the systemic blood-pressure, and as the square root
of the diastolic time. The coronary arteries supplying the
whole heart, the work done by the right ventricle is governed
by that done in the left; thus the supply of blood in the left
auricle is always rendered sufficient for the requirements of the
systemic circulation ; though, as there is no reason for believing
that the resistance in the pulmonary vessels varies with that
of the systemic, there must be some peculiarities in the former
circulation (which may explain the variations in the ratio of
the number of pulse-beats to respirations, In some cases).
The auricular contraction is a very small force, and its
function is most probably to close the tricuspid and mitral
valve.
The heart commencing its systole as a whole, it is highly
probable that the impulse for action is given by a force which
affects both ventricles; such is found in the coronary circulation
and the active diastole produced by means of it.
In conclusion I have to present my best thanks to Dr
Michael Foster, Professor Sanderson, and Professor Pritchard
of the Royal Veterinary College, for the opportunities they have
afforded me in trying the experiments above detailed; without
their assistance it would have been impossible for me to have
put the law of the relation of blood-pressure to pulse frequency
on any satisfactory basis.
MORPHOLOGICAL ELEMENTS OF THE SKULL.
By W. KircHEen Parker, F.R.S. (Pl. IV.)
THE skull, or cranio-facial skeleton, would be easily compre-
hended as a morphological structure, were it no more complex
than the skeleton of the trunk.
In that part of the vertebrated animal we have merely a
repetition of essentially similar segments, each, typically, form-
ing a ring above and a hoop below. Over these hoops at two
points at the most there are the paired fins or limbs, with
their imbedded and free portion—limb-girdles and limbs proper
—and in fishes the unpaired upper and lower fin-rays, with
their imbedded cartilaginous or osseo-cartilaginous roots.
But the essential, primary parts of the axial skeleton have a
totally different “habit” in the region of the head, to what
they have along the spine; the notochord, truly, is there, and
the substance which invests it (“investing mass”), but this
never shews the least tendency to break up into “somatomes,”
and “metasomatomes,” as in the body. Moreover, although the
cranium contains a continuation of the common neural axis,
yet in this region, instead of a simple nervous tube, grey within,
and white without, in the head the nervous mass always’ swells
into three principal vesicles, which are grey without, and white
within.
Also let it be noted that it is only the medulla oblongata,
an evident continuation of the spinal chord, and the cerebellum
which are enclosed in the occipital ring, or that part of the
cranium which is constructed out of the fore end of the
notochord, and its undivided investing mass.
The box, then, in which the cranial vesicles and their often
immense and very complex outgrowths are enclosed, is a com-
pound structure. Before it is treated of it will be necessary to
describe and to classify the other skeletal parts of the head.
1] purposely omit the Lancelet (Amphioxus), from all consideration, at
present; I begin with the Suctorial Fishes (‘‘ Myxinoidei,” ‘‘ Marsipobranchii”),
We get no proper series that can be considered through and through until we
come to the Lamprey and its relations,
MR PARKER. MORPHOLOGICAL ELEMENTS OF THE SKULL. 63
I believe that the only truly cartilaginous parts of the
Amphioxus or Lancelet are the “palpi” or “cirrhi:” these proto-
skeletal parts re-appear in the higher fishes, and in some of the
air-breathing types.
Then besides these we have in the suctorial fishes a sub-
cutaneous system of cartilages, immensely developed in the
lips (“labial cartilages”), and in the pharynx; forming in the
Lamprey the beautiful branchial basket.
In these fishes, besides the outer cartilages, which I propose
to call “ extra-visceral,” there is a cartilaginous skull, and also
three pairs of what are called “visceral arches” by embryolo-
gists. One pair of these bars is in front of the mouth, and
two pairs behind; there is also the rudiment of a second pre-
oral pair; in embryological works, the first post-oral, or mandi-
bular arch, is called the first visceral arch.
These “Marsipobranchs” also possess “sense-capsules,” which
more or less modify the form of the cranium, the last pair,
or auditory sacs, being compacted into the posterior part of
the cranial side-wall.
In the next group of fishes, the Sharks and Rays, or
“ Elasmobranchii,” the cartilage, which is soft in the “Suctorii,”
becomes calcified externally, and sometimes within; but the
ossifications that occur in the cutaneous system with its aponeu-
rotic enfoldings and its various strata, looser or more dense,
do not become modified correlatively to the cartilaginous skele-
ton. This first shews itself as a new morphological specializa-
tion in the “Ganoid” fishes.
The elements that go to form the skeletal parts of the
head of a vertebrate, some height up in the scale, are as
follows :
Curhi.
Extra-visceral cartilages.
Sub-cutaneous bony plates.
Visceral cartilages (arches).
Sense-capsules (mostly cartilaginous).
Cranium, proper; composed of
Me QAeors
1. Notochord and undivided part of “investing
mass,”
64 MR PARKER.
2. General membranous coat, which is chondrified
below and behind by the “investing mass;”
below and above by the arch formed from
that mass (occipital arch); and anteriorly by
an independent growth of cartilage.
3. Bony plates developed in subcutaneous tissue
outside the chondrified external layer of the
membranous cranium, such as the frontals and
parietals.
But the eye-balls are the only pair of sense-capsules that
grow free of the cranial walls—they indeed modify the shape of
the box, being nested in its sides. The other sense-organs
are most intimately built up into the very structure of the
cranium.
This is not all, for certain segments that appear in the
adult cranium in the higher types, notably in the mammalia,
are formed by ossification of compound cartilaginous tracts,
formed by early fusion of the first pre-oral (actual first)
visceral arches.
It is also to be remarked that the more the brain develops
—as we ascend among the types—the less does it owe to the
primordial cartilaginous thickening of the outer stratum of its
membranous investment. The inner stratum (“dura mater”)
always retains a fibrous character.
So also as to the visceral arches: these are huge in fishes,
especially when they are only partially calcified, as in Sharks
and Rays; but as we ascend they become more and more
liable to histological metamorphosis; becoming arrested, and
often being replaced, functionally, by subcutaneous bones, or by
these together, with “extra-visceral” cartilages.
An instance of this is at hand in the Pig—which undoubt-
edly represents a large group of mammals—where the primary
mandibular bar is arrested above, absorbed below, and gives
place functionally to the combination of a labial cartilage with
a subcutaneous ossification.
Both in the nasal sacs and in the organs of hearing the
true cartilagimous investment of the sense-organ itself forms
metamorphic combinations with contiguous visceral arches.
MORPHOLOGICAL ELEMENTS OF THE SKULL. 65
- With regard to the viseeral arches if we examine a series
from the Chondrosteous Ganoids (Sturgeon, &c.) upwards, we
shall find a very large amount of variation in the metamor-
phosis of their substance. elow, a thin ectosteal plate may
invest a permanent core of unchanged hyaline cartilage; but
above, as in the hot-blooded classes, the arches may _ ossify,
whilst as yet their substance is a soft granular mass of “ indif-
ferent tissue.”
Not only so, the arch, originally visible as a solidish grani-
cular tract, may chondrify here and there; large tracts being
arrested, and becoming lost in the general fibrous stroma
amongst which it was developing.
Segmentation of the “visceral arches.”
The morphological observer notes especially in the skull
how it is compacted together by that which each element
supplies to it,—for we have just seen that this important box is
formed of very diverse parts and structures.
But whilst all tends to solidity in the cranium, in the face,
in many instances, the parts are specialized in relation to the
most varied movements: hence the large amount of segmenta-
tion seen in the visceral bars.
The first pair of visceral bars, the “trabecule cranii” of
Rathke, are least mobile, and therefore least segmented; the
most mobile are the parts that lie around the mouth and throat.
Before describing the very uniform manner in which, on
the whole, the visceral arches break up, it may be well to
remind the reader that they must never be confounded with
the costal arches that spring from the bodies of the vertebre.
These oro-faucial bars are developed about the mouth and
throat, at first as thickenings, whilst the face of the embryo is
growing into a ridge-and-furrew structure: the ridges develop
a pith, which is the visceral arch, and the furrows are the first
sign of that dehiscence which takes place to form the “visceral
clefts.”
These visceral clefts are the doors and windows through
which the vertebrate creature keeps up its necessary com-
merce with the surrounding world for food, and drink, and
~
VOL. VIII. D
66 MR PARKER.
air. As a rule the clefts are in distinct pairs, but the pair
which form the mouth run into each other at the ventral line.
The metamorphic changes of the visceral arches rise, as it
were, one above another in the scale of life, the higher type
having “nothing over,” no more change than its habits demand;
and that which undergoes but little modification has “no
lack ;” for though its parts suffer less change, yet they answer
to the necessities of the creature as perfectly as in the nobler
forms.
We have seen that the true visceral arches are few in num-
ber in the “Suctorial Fishes;” but amongst the “Placoids” or
Sharks and Rays there are, normally, five pairs of “ branchial
arches” behind the hyoid or arch of the tongue. In the embryo
of the shark (e.g. Scyllium canicula) both the mandibular and
hyoidean arches carry caducous (external) gills, and the hyoid
carries a half-gill in the Placoids, generally, and in most of the
“ Ganoids;” yet in the Teleostean or Osseous Fishes the gills
are confined to the arches behind the hyoid. The last arch
is abranchiate, and in Osseous Fishes forms the “nether mill-
stone’? to the grinding apparatus, the upper stones of which
are formed by the apical segment of the gill-arches in front of
it: the papille on that, the “ pharyngo-branchial” segment, be-
coming developed into teeth, and not into gill-plates. That the
arches in front of the mouth are the equivalents of those be-
hind is shewn by the development of the mouth itself, and also
by the presence of a pre-oral cleft, the passage which in us
conveys the tears into the nose by the lachrymal duct.
Normally, in Fishes above the Suctorii there are seven post-
oral arches: in the Anourous Amphibia only six. In the Shark
and its congeners I find no pharyngo-palatine or second pre-oral
arch ; and for the time of the greater part of the tadpole-life
of the frog this arch is suppressed, or nearly so, and it is never a
distinct bar. I am satisfied that it is also suppressed perma-
nently in the Tailed Amphibia, and that it is only a rudiment,
not free, in the Suctorii. These things are not, however, all
quite certain at present.
As a rule the segmentation of the visceral arches takes
place some time anterior to ossification, that is in the Gill-
bearing, or Ichthyopsidan types: in the higher forms, in some
2 RY Or: Rie ee m8 i go
{
:
MORPHOLOGICAL ELEMENTS OF THE SKULL. 67
of the arches, at least mostly those in front of the mouth, the
osseous deposits determine the segmentation, and now and
then in Birds a single bony part breaks up into two sepa-
rate bones.
In the “ Branchiata” segmentation is generally greatest in
the hyoidean and branchial arches. In the latter it may be
said to be typical; but the hyoid arch of a Shark is arrested as
to its subdivisions. That arch in the Teleostei runs into parts
beyond what is typical.
Furthermore, whilst in the Branchiata the whole seven of the
post-oral arches become converted into hyaline cartilage, at least,
if not bone, in the Abranchiata only the first three of these
are developed, the last of the three only partially.
The form of each arch is sigmoid, much like the italic
letter f, they generally turn both apex and distal end inwards
and backwards; but sometimes the direction of the apex is for-
wards, at others simply inwards. The arches about the mouth
have a very regular habit of grafting themselves upon other
parts, parts having a diverse morphological origin ; for instance,
the trabecule thus seize upon the apices of the “investing
mass,” the pterygopalatine seeks the fore edge of the mandi-
bular pier, and that pier, the apex of the first post-oral arch,
whilst the hyoid grows to and grafts itself upon the auditory cap-
sule. As a rule, the proper branchial arches turn their apices
over the pharynx; and in the Shark, especially, it may be seen
that by stretching or contracting that large tube, these cartilages
that enring it may be turned inwards and forwards from their
normal backward direction.
Passing suddenly from one type to another in my slow
researches—a year on an average to a type is not an unusual
period, so that I may say ten years have I consumed over as
many skulls—I became conscious that the same essential thing
was still before my eyes. Not troubling myself with the motto,
“Nihil per saltum,” but passing at once from the Pig to the
Shark, I became conscious of a nearness in nature of these two
most diverse subjects, that was not a little startling.
Filling-in this space, and looking this way and that for
light, it has been gradually revealed to me that there is a
marvellous uniformity in number, in the segments of a visceral
5—2
68 MR PARKER.
arch, wherever found. At least the same number will turn up
so frequently that there can be no doubt that there is some
law to be discovered causing this uniformity. Not to lay too
much stress upon this subject, it yet appears to me to promise as
much fruit in morphology as the number of members in the
whorls of a flower in various natural orders—three, four, or five,
as the case may be. Running the eye over the branchial
arches of those three great groups of Fishes called “ Elasmo-
branchii,’ “Ganoidei,” and “ Teleostei,” we shall find that
Professor Owen’s elegant nomenclature for these parts is appli-
cable throughout, namely, from above downwards the segments
are known as “ pharyngo-” “epi-” ‘‘cerato-” “hypo-” and “basi-
branchial”—the latter element uniting the right and left
moieties of the arch (fig. 9).
The same prefixes may be made applicable to the segments
and regions of all the visceral arches; and they may serve for
the generic as well as specific names of these arches.
Thus for the whole genus, “visceral arch,’ we have seg-
ments or regions that may be called, throughout the series of
arches and in all the types of the vertebrata, “ pharyngo-,”
“ epi-” “cerato-” “hypo-” and “ basi-visceral.” These prefixes
can as well also be made to fit on to the fore-front of such
specific terms as “trabecular,” “palatal,” “mandibular,” and
“hyoidean,” as well as “ branchial.”
To apply this, the main segments in the much-divided
hyoid arch of a mammal are as follows: namely, at the apex
the “os orbiculare” next below the “incus;” then the “stylo-
hyal,” below this the “cerato-hyal,” and, uniting the two sides,
the “basi-hyal” (fig. 9). These all exactly correspond with
the typical segments of those well-developed visceral arches,
the “branchials” of the three largest orders of Fishes—the
eroups in which these visceral arches have their fullest develop-
ment, and are most typical, morphologically. It may be ob-
jected that there is an additional piece in the hyoid arch of a
mammal that I have not mentioned, namely the little “inter-
hyal” segment which gets attached to the neck of the “stapes,”
or auditory plug.
There is this additional piece, and in the hyoid arch of the
osseous Fish it is also present, but it is a secondary morpho-
) 6
MORPHOLOGICAL ELEMENTS OF THE SKULL. 69
logical element, and is of the same value in morphology
as a “meniscus” between two segments. The hyoid arch
of the osseous Fish also goes beyond the typical visceral arch by
having two centres of ossification for each of three of its seg-
ments, namely for the so-called hyo-mandibular, the main bar,
and the distal piece or “ hypo-hyal.”
This tendency to further segmentation by the superaddition
of fresh osseous centres, is conjoined in these fishes with the
absence of the apical piece, which is so well developed in the
branchial arches as the “ pharyngo-branchial” segment.
For a long while the meaning of the hyoid arch of the Frog,
after metamorphosis, was hidden from me, for, besides the chief
segments of the osseous Fish, the Frog, and its congeners, the
other “ Anoura,” have an additional segment above the hyoman-
dibular (= 7ncus); which I saw long ago must be the counter-
part of our little “os orbiculare,” wedged as it is between the
incus and stapes. This part, therefore, seemed to be a new
thing, a specialization not to be accounted for by any normal
segmentation of visceral arches: it is, however, merely a “ pha-
ryngo-hyal” segment, and is perfectly normal.
The manner in which the “cornu-minor” of the mamma-
lian “os hyoides,’—the hypohyal segment—turns backwards to
articulate with the basal piece, is also quite normal (fig. 9) ;
in like manner the hypo-trabecular region of the first arch is
always hooked backwards, and the two diverging hooks are the
“trabecular horns” (fig. 10). The manner in which the right and
left moieties of these arches join each other, and the mode of
junction of a bar with one in front of, or behind it, is very
interesting,
The junction of the right and left parts of the same arch
at the mid-line below may be called the visceral “ commissure,”
the yoking on of arch to arch a visceral “conjugation,” and
such projections or parts as are developed may be termed con-
jugational processes. The “basi-pterygoid processes” of the
trabeculee, and the “orbital processes” of the palatals, are con-
jugational processes: the “basi-visceral” elements are speciali-
zations of “ visceral commissures.”
Now the azygous, or commissural pieces, have been long
known in the post-oral arches, as in the basi-hyal, and basi-
70 MR PARKER.
branchials, but basal elements have not hitherto been described
in the pre-oral arches. In various papers on the bird’s skull
I have made mention of a “ pre-nasal rostrum,” a cartilage
that projects from the fore-end of the nasal septum, and is
the model on which the very prognathous pree-maxillaries of
the bird are formed. In the paper on the fowl’s skull (Phil.
Trans. 1869, Plates 81—87), the reader will find the genesis of
this cartilage, and it is one of great interest: it is, 1 now see,
the “basi-trabecular” element (figs. 5—8). JI am now familiar
with this part in rays, sharks, serpents, tortoises, birds, and
mammals: the degree of modification and development this
bar undergoes determines very largely the form of the face.
In my first plate of the fowl’s skull (Plate 81, fig. 3, beginning
of 2d stage) this cartilage (p. .) is a rounded bud, looking
downwards and backwards. This is similar to what I have
lately described in the embryo pig, 7} lines in length; in this
latter animal it is arrested at this stage, and the pre-maxil-
laries are formed beneath it. In the turtle, Chelone midas, it
straightens down, and the pre-maxillaries are formed in front
of it. In birds (as in sharks and rays) it straightens still fur-
ther (Op. cit., Plate 81, fig. 4, Plate 82, figs. 1 and 2, and Plate
83, figs. 1, 2, 4, 5, p. m.), and projecting in front of the face
forms a most fit model for the pra-maxillary parostoses (figs.
6—8). The least change in the direction of these two bones,
whilst still growing upon the transitory cartilage, makes all the
difference between the beak of a crow, a hawk, a hornbill, or in-
deed any type of this beaked class. In the snake (fig. 3), as in
the mammal, it is arrested, and keeps the backward position it
has in common with the “hypo-trabecular horns.” It was a
satisfaction to find a morphological “ pigeon-hole” for the pre-
nasal cartilage, and at the same time a basal element for the
newly-discovered trabecular visceral bars’; but my labours have
1 JT had long satisfied myself that the old term inter-maxillary was better
than Professor Owen’s new name premaxillary, and I had stated this to Pro-
fessor Huxley, for I saw that the base of the nasal septum was with its “‘pre-
nasal rostrum” the axis or primordial part of the foremost pre-oral arch.
But it was Professor Huxley who shewed that the whole of each trabecular bar
belonged to this part; it was suggested to his mind by the manner in which
the nasal diverticulum of the Lamprey passes between the ethmo-vomerine
cartilage (cornua trabeculz) and the skull-bone. But more than thirty years ago
Johann Miller (in the Chapter on Generation in his Physiology of Man),shewed
MORPHOLOGICAL ELEMENTS OF THE SKULL. 71
been further cheered by the unveiling of a like part in the very
variable pterygo-palatine bar.
T have already stated that the second pre-oral arch is ossi-
fied in the “ Abranchiata” whilst quite soft. This is true on
the whole; but I had long ago noticed that in the higher birds
certain parts of it were slow to ossify, and were developed into
hyaline cartilage first. It then turned up that in the wood-
peckers (“Picidze”) the contiguous edges of the palatines be-
came cartilaginous, and that this “commissure” was afterwards
formed into a dagger-shaped bone—a “basi-palatine.” The
same thing takes place in Caprimulgus europeus; it has two
bones in this part, and so also has Podargus; and I expect to
find it in many others’.
The “hypo-palatine” region is developed in the Salmon
into a large knob of cartilage in front of the trabecular conjuga-
tion.
In no creature have I yet seen a distinct “ pharyngo-pala-
tine” element in a separate state; but in most of the Lacertians
the “‘epi-palatine” is more slowly ossified than the rest of the
arch, which bifurcates behind, the upper becoming a sub-carti-
laginous rod; this rod, ossifying, becomes the so-called “colu-
mella”—my “epi-pterygoid.” This element also exists in the
Chelonians, and becomes flattened and wedged-in between the
descending parietal wall and the floor-shaped “pterygoid.” In
birds this part is shewn as a “process,” and is a very elegant
hook in Passerine birds, and in such forms as Scythrops, Podar-
gus, Megalema. In mammals also it is a notable part, most
elegant in Tragulus javanicus, but known long ago in man as
the “hamular process of the internal pterygoid plate.”
These instances of parts shewn to be classifiable must suf-
fice at present; they can easily be referred to by the student,
and I flatter myself that they will commend themselves to his
judgment.
An attempt to harmonize the “labials” and outer branchial
cartilages as “ extra-viscerals,” will, perhaps, be made at further
leisure; but it may not be thought out of place if I remark
that the inter-maxillary apparatus was developed independently of the palato-
maxillary. j
2 A supplementary paper to the one just quoted on the Fowl is ready for pub-
lication, in which these structures will be illustrated and described.
"2 MR PARKER.
upon the relation of the cephalic skeleton to that of the
trunk.
In the present state of our knowledge it is better for us to
consider the morphology of the skull as quite distinct from that
of the body.
' Firstly. The formation of somatomes in that part of the
“mesoblast,” which lies on each side of the notochord, takes
place very early, and is never seen in the cephalic region.
Secondly. The pituitary body, growing downwards and
backwards from the first cerebral vesicle, beneath the second,
effectually stops the notochord in its forward growth: the “ in-
verting mass”’ ends in a pair of blunt, or even squared extremi-
ties, opposite the end of the notochord.
Thirdly. The three well-known basal pieces of the skull,
which beguiled the older observers, who saw, or thought they
saw, in them three more vertebral centrums, are very diverse,
morphologically, and only the last contains any of the noto-
chord.
Fourthly. Ribs are direct down-growths from the sides of
each vertebral “centrum:” nothing of the kind is developed
in the head, where the basi-occipital represents several poten-
tial centrums.
Fifthly. Any arches in the head and embracing the throat-
region, which should correspond to ribs, would enclose the
heart: the visceral arches would lie inside such hoops, and the
heart Jie outside (below) the visceral arches.
Sixthly. The “extra-visceral” bars, outside the branchial
arches of the shark, which lie close under the skin, like ribs,
have no similarity in their origin to those arches of the trunk-
vertebree.
EXPLANATION OF PLATE.
Magnified figures of certain visceral arches of Vertebrata.
1. Side view of third branchial arch of adult dog-fish (sey/lium
canicula), p. or. pharyngo-branchial ; e. br, epi-branchial ; c. br, cerato-
branchial ; h. br. hypo-branchial ; b. by. basi-branchial ; br. 7, branchial
rays; ew. br, extra-branchial.
MORPHOLOGICAL ELEMENTS OF THE SKULL, 73
2. Under view of snout of the same; tr. trabecule; c. tr, cornua-
trabecule ; b. t. basi-trabecular rostrum; J. labials.
3. Under view of snout of embryo snake (natrix torquata) ;
letters as above.
4. Side view of the same.
5. Side view of snout of passerine bird (“ Avis Aigithognatha”).
lst Stage.
6. Ditto 3 $5 Ditto 2nd Stage.
12: Ditto a 3 Ditto 3rd Stage.
8. The same as last; under view.
9. Hyoid arch of embryo pig (sus scrofa); 0. 0. os-orbiculare
(pharyngo-hyal) ; 7. incus (epi-hyal) ; st. h. stylo-hyal (cerato-hyal) ;
¢. m. cornu minor (hypo-hyal) ; 6. h. basi-hyal; 2. h. inter-hyal.
10. Side view of snout of embryo pig: lettering as in figs. 3—8.
ON DOUBLE NERVE STIMULATION. By A. G. Dew-
Smith, B.A., Zrinity College, Cambridge. Pl. V.
(From the Physiological Laboratory in the University of
Cambridge.)
THE problem, the solution of which I have attempted, and
which, as far as I know, has not hitherto engaged the attention
of any observer, is as follows :
If two pairs of electrodes be placed on a nerve, one pair
being nearer the muscle than the other, what happens when a
stimulus is sent through both pairs of electrodes at the same time?
Taking the contraction of the muscle supplied by the nerve as
a measure of what is going on in the nerve, is the contraction
which results from a stimulus thrown through either pair of
electrodes singly at all influenced, and if so in what mamner, by
the simultaneous stimulation of the nerve at the other pair of
electrodes ?
The first observations were naturally made with a single
induction shock and with the opening or breaking shock. It
seemed desirable also to begin at least with an equal, and that
a weak stimulus, or one below the maximum, at the two pairs of
electrodes. But here four different cases presented themselves,
according to the direction of the current; these cases would
-need to be distinguished, as the results might be different in
the different cases. Calling the electrodes farther away from
the muscle A, and those nearer the muscle B, we have:
Case I. where the current in A is descending, in B descending,
ee ease 3 ES Sk B ascending,
acre 6 ee ee A is ascending, in Bdescending,
ENE at eee >: ny oes B ascending.
In Case I. and LV. the currents have the same direction, in
II. and III. opposite directions.
The majority of my observations were made on Case I.; the
other cases were examined afterwards for the purpose of ascer-
taining any differences from that case. I first attempted to use
a divided circuit, interposing both the A and B electrodes in
’ 2S" 2eei@eere } 6.1 i el
DOUBLE NERVE STIMULATION. rae
the circuit of the secondary coil. The electrodes used were plati-
num wires fixed in paraffin blocks ; the distance between the anode
and kathode of each pair (about 3 mm.) was thus maintained per-
fectly uniform. An observation was conducted as follows. The
sciatic nerve of a frog was placed on the B electrodes alone, the
upper part of the nerve resting on the clean paraffin block. On
the electrodes A was placed a portion of the other sciatic nerve
of the same frog, carefully selected, so as to correspond as exactly
as possible to that portion of the nerve which would le on the
electrodes A when both A and B had to be stimulated at the
same time. The gastrocnemius muscle belonging to the nerve
was attached to a simple lever writing on a smoked surface,
the excursions of the muscle being magnified several-fold,
in order to be able to use only a very slight stimulus, and
yet measure successfully the variations which occurred. It
was hoped that in this way the resistance at the electrodes A
might be maintained the same, both when B only was stimu-
lated, and when A and B were stimulated together, and that
consequently the amount of stimulus reaching B would be
the same, whether the nerve were stimulated at the same
time at A or not. Similarly, when it was desired to stimulate
A alone, a piece of the nerve of the other leg, corresponding
to the piece which would lie on B when the nerve was placed
on both electrodes, was carefully laid on B.
After a few observations, however, it was soon evident that
this method was impracticable. However carefully done, the
shifting of the nerve on A caused marked variations in the
resistance at the electrodes A, and consequently variations in
the amount of stimulus sent into B. ~
An attempt so to increase the total amount of resistance in
the circuit that variations at A might be neglected, introduced
other elements of confusion, and I was consequently led to
make use of two separate and independent induction machines,
with independent primary circuits, the secondary coil of each
machine being connected, of course, with its own electrodes
alone. By means of two pairs of mercury cups, the two primary
circuits could be made or broken, singly or together, at plea-
sure; or by means of a key either secondary circuit could be
short circuited off from the electrodes. By cither method A
76 MR DEW-SMITH.
and B might be stimulated together, or either of them apart.
The two machines were placed at right angles to, and at some
distance from, each other; and by placing a separate nerve
and muscle on each pair of electrodes, I satisfied myself that
the two machines had no influence on each other.
In this way I made a large number of experiments, without
however arriving at a result which I could consider thoroughly
satisfactory. For instance, when the contraction resulting from
the stimulation of A was about equal to that resulting from the
stimulation of B, the contraction given by A and B together
was in the majority of cases equal to that of B alone. A small
number of exceptions, however, always occurred, in which the
contraction of A + B was greater than B. Upon consideration
it seemed probable that the cause of these exceptions was to
be sought for in the want of exact simultaneity in the breaking
of the primary circuits of A and B, and consequently in the
stimulus arriving at the one ‘pair of electrodes a small fraction
of a second sooner or later than at the other. That this cause
was the true one became evident when, instead of employing an
ordinary rotating cylinder for registering the contractions, I
began to make use of the pendulum. myographion.
In as much as the instrument in the Laboratory was fitted
with a single contact-breaker only, I at first so arranged matters
that the pendulum in its swing broke an independent current,
which, by means of a steel slip and an electro-magnet, was
keeping two platinum points dipping in two cups of mercury.
-At the breaking of this current the steel was released from
-the electro-magnet, the two platinum points raised from the
mercury cups, and, consequently, the circuits of which they
respectively formed part broken. By raising (or lowering) one
or other of the platinum points, the moment at which contact
was broken in the one cup was thrown in front or behind that
in the other.
This method, however, proved on trial inexact, and I
consequently replaced the original single contact of the myo-
graphion by a double contact-breaker, similar to the one
employed by Helmholtz, so arranged that the one contact-
breaker could be shifted in front of or behind the other, and
thus the catch of the pendulum would break the two contacts
DOUBLE NERVE STIMULATION. Vi i
at an interval of time, varying according to the distance
through which one had been moved.
My results then became for the first time satisfactorily
uniform. When contact is broken in both the A and B circuits
simultaneously there is no increase of attraction; the resulting
curve is the same as if the nerve had been stimulated at B
only. Thus, in Fig. 1, which reproduces a photograph taken
from the myographion plate, 1 represents the muscle-curve
resulting from the stimulation of A alone, with a single open-
ing induction shock derived from a single Daniell with the
secondary coil of a Du Bois Reymond’s machine at 25. 2 is in
like manner the muscle-curve resulting from the stimulation of
B alone, also with an opening induction shock, derived from a
single Daniell with the secondary coil of a second Du Bois
Reymond’s machine at 30. The mark indicates the moment
at which the primary currents were broken, being exactly the
same in both A and B. In both cases the direction of the
induced current was descending. It will be observed that,
allowance being made for the difference in the base lines,
the curves are very nearly of equal height. The curve 3 repre-
sents the muscle-curve resulting from the simultaneous stimu-
lation of A and B, « as before marking the breaking of both
primary currents. Allowance being made for differences in
the base line, the curve 3 is the almost exact duplicate of the
curve 2. In other words, when the breaking of both primary
currents is simultaneous, the contraction produced by the
stimulation of B is in no way affected by the stimulation of A,
Curve 4 represents the muscle-curve resulting from the
stimulation of A and Bb, when the primary circuit of A is broken
as before at z, and the primary circuit of B broken at y, the
contact breaker of B having been shifted 5mm. in advance of
its previous position. The tuning-fork-curve below the muscle-
curves marks 180 double vibrations per second, the distance
between x and y measured by this gives 5%, or =4, second, as
the interval of time elapsing between the stimulation of B and
that of A. The muscle-curve is, as will be observed, immensely
increased, and at the same time altered in form, so that its
maximum point is deferred, being very nearly identical in time
with those of the previous curves.
78 MR DEW-SMITH.
Fig. 2 represents a series of curves obtained by increasing
the interval between A and B, B being in all cases thrown in
advance of A. Thus the curve 1 is the muscle-curve resulting
from the simultaneous stimulation of A and B with the
secondary coils in each case at 30 and a single Daniell’s to each,
x as before marking the moment of breaking the primary cur-
rents. Curve 2 is the curve resulting when B is thrown
2°5 mm. (= ;5,sec.) in advance of A; the contraction is most
distinctly greater than in 1. In curve 3 the interval between
A and Bis 5 mm. or 4, second; there is still an increase of the
curve. In curve 4 the interval is 10mm. or j4 second, the
contraction produced by B has arrived at its maximum be-
fore the contraction produced by A has begun, a division into
two curves is already apparent, and the total contraction has
reached its maximum. In 5 (15 mm.= , second) the analysis
of the curve has proceeded further, while the total contraction
=
has diminished; and in curve 7 (25 mm.=-,second) we have
simply two curves, the earlier one being that resulting from the
contraction of B, the later one being due to A, and all but
coinciding with the original curve of A and B.
In all the above cases B was made earlier than A, but
exactly similar results are obtained when dA is made earlier
than B.
With simultaneous stimulation at two different points of
the nerve the contraction resulting is the same as when the
near point only is stimulated; but when a small interval of
time is allowed to elapse between the two stimulations, the con-
traction increases, rising to a maximum as the interval is
lengthened, and afterwards dividing gradually into two indepen-
dent curves.
What are the inferences which may be drawn from these
results? At first sight they might seem to be mere variations
of Helmholtz’s well-known researches on the summation of
contractions. But there is a most important difference. Helm-
holtz has shewn that when a maximum induction shock (i.e. an
induction shock giving a maximum contraction) is made to
follow. (fallmg upon the nerve at the same point) a preceding
similar shock, no increase of contraction is observed when the
interval between the two shocks is so small that by the time
CO LRN PN. ee IR a
EEO
ee eee ee eee eee
DOUBLE NERVE STIMULATION. 79
the muscle has begun to contract in obedience to the second
shock the contraction due to the first shock has not appreciably
begun. This interval he puts at about =1,sec. But this is
true of maximum shocks only. Whether applied simultane-
ously, or with an interval of time between them of any suft-
ciently small magnitude, two minimum (or at least not maxi-
mum) shocks produce a contraction greater than that due to
either of the shocks acting singly. In the above experiments
the facts illustrated by Fig. 2 have to do with the summation
of contractions in the muscle; and would come out exactly the
same if a maximum stimulus were employed, the interval
between curves 1 and 2 being greater than is necessary to obtain
summation of the contractions. The same may also be said of
the curve 4 in Fig. 1; the interval there also being considerable.
But in such a case as that illustrated by curve 3, Fig. 1, we
have to deal with something different. Here the stimulus of
both A and & being below the maximum, we ought to have an
increase of the contraction on double stimulation; whereas the
curve is as nearly as possible of the same form.
Similar considerations shew that the absence of increase of
contraction in simultaneous contraction cannot be due to any
action taking place at the electrodes of B. For example, we
might suppose that the stimulation of 6 diminishes for a definite
small fraction of time the conductivity or irritability of the
nerve between the electrodes B in proportion as it excites it to
action, that consequently from the nervous impulse originating
in A there is taken away when it reaches B just so much as is
required to give the contraction due to the excitation of B, and
that therefore A and B being equal, the resulting contraction
is the same as when B only is stimulated.
This is in itself exceedingly unlikely when we remember that
a maximum stimulus is not in question. Granted that B is ex-
hausted (if we may use the phrase) in proportion to the stimulus,
there remains, when the stimulus used is below the maximum,
a surplus of irritability at B, of which the impulse at A can avail
itself, and so produce an increase of the contraction just in the
same way as increasing the stimulus at B itself would increase
the contraction. Besides, this loss of conductivity or excitability,
or this exhaustion, whatever it be called, is coincident with
80 MR DEW-SMITH.’
the stimulation, or at least can last only an exceedingly short
time afterwards, for the stimulus repeated at B after as short
an interval as possible gives rise to an impulse which by sum-
mation on the muscle increases the contraction. Hence if the
results arrived at were in any way due to such a cause acting
at B, they ought to be influenced by the distance between the
two pairs of electrodes A and B. If when A and Bare near
together no obvious increase of contraction makes its appear-
ance on simultaneous stimulation, yet it ought to become more
and more distinct the farther apart A and Bare placed. This
however is not the case; the absence of increase of contraction
in simultaneous double stimulation helds good at whatever dis-
tance apart A and B are placed.
The only explanation which I can offer is that we have here
to deal with a block of nervous impulses. I avoid the word
interference, as having already been appropriated by the phy-
sicists in a more limited sense. When A or # is stimulated, a
nervous impulse is originated and travels in two directions:
downwards towards the muscle and upwards towards the central
end of the nerve. The evidences of this double direction of
impulses are, Ist, the fact that the negative variation does travel
equally in both directions; 2nd, the experiments of Phil-
lipeau, Vulpian, and others, on the mixing of sensory and motor
nerves.
When A and B are stimulated simultaneously the up-
ward impulse from B meets the downward impulse from A,
and the two block each other; i.e. the two impulses moving
in opposite directions mutually antagonize each other, so that
neither is carried on in its own direction beyond the point
where they meet. Consequently, the impulse from A not reach-
ing the muscle at all, the contraction in the muscle is caused by
the impulse from B only. And naturally the effect is the same
whatever be the distance between A and B.
When however one of the two, for example, Bis stimulated a
certain fraction of a second earlier than A, the upward impulse
of B has passed beyond the electrodes A on its way to the cen-
tral end of the nerve before A is itself stimulated. Hence the
downward impulse from A meeting with no opposition reaches
the muscle where the contraction it gives rise to is summed up
oa ee ee
DOUBLE NERVE STIMULATION. sl
with the contraction due to the stimulation of B, and an
increase of the curve is the result.
In the same way, when 4 is stimulated before B, the down-
ward impulse from A passes B before B is stimulated, and
hence meets with no opposition.
In both cases it is obvious that the result will be affected by
the distance between A and B. When A and Bare near each
other the time necessary for the upward impulse starting from
B (when B is stimulated first) to have passed by A is less than
when A and B are far apart. The difference however, with the
extremest range possible on the sciatic nerve of a frog, is a very
small fraction of a second; and the contact breaker which I
used was not sufficiently delicate to appreciate this fraction in
a satisfactorily accurate manner. I am obliged therefore to
postpone the completion of these observations till a new con-
tact-breaker has been constructed for me. I may add that this
method evidently offers an opportunity of indirectly measuring
the velocity of a nervous impulse.
Until these further observations have been carried out, I
cannot claim to have established satisfactorily the theory of a
block. Meanwhile I may notice an objection which readily
presents itself. If the sciatic nerve of a frog consisted of a
single fibre, a block might be intelligible enough. But the
nerve is composed of a multitude of fibres, each of which we
have reason to think may act independently of the others; and
one would imagine that while the upward impulses from B
were passing along certain fibres, the downward impulses from
A might pass independently along other fibres, without the one
in any way interfering with the other. And one might further
suppose, that in any position of a nerve on the electrodes some
fibres are excited by a shock passing through the electrodes and
others not. But this appears not to be the case. As far as we
know, when a nerve is placed between a pair of electrodes 3 mm.
apart, all the fibres in that part of the nerve are excited by
a stimulus passing through the electrodes. This is shewn by
the contraction of all the muscles supplied by that nerve, and
not of any particular set or sets of muscles.
I have used the phrase “block” rather than interference,
because the latter has a special reference to wave-movements.
VOL. VIII. 6
82 MR DEW-SMITH.
The phenomena which I have described are rather those of the
mutual neutralisation of two opposing forces. They may be
interpreted on either a ‘molecular mechanical’ theory, or a
‘chemical’ theory of nervous action. One may imagine, by
way of an exceedingly rough illustration, two movements start-
ing from each pair of electrodes and clashing midway, or indeed
the block may be felt at the very beginning of the movement
and no actual movement affected at all, as when the two
extreme members of a series of balls are struck at the same
moment. One may equally imagine two chemical transforma-
tions stopped at midpoint by a common want of material for
the time being, each having exhausted half that of the other,
just as the explosion of a train of gunpowder stops in the middle
when lighted at both ends. I simply use the phrase, without
committing myself to any special interpretation of it.
All the above observations refer to what I have called
Case I., where the momentary induced current had a descending
direction in both A and B.
I have also made observations on the other cases. Before
making use of the pendulum myographion, I was inclined to
believe that the direction of the currents (whether opposing or
otherwise) affected the magnitude of the contraction. But
careful observations with the pendulum have convinced me that
such is not the case, that the direction of the current makes
no difference whatever to the general result, that with simulta-
neous double stimulation there is no increase of contraction.
This is shewn by the figures 3, 4, 5, which illustrate Case IL,
IIL, IV. respectively.
I have also made observations using the make and break
of a continuous current in place of a single induction shock,
and likewise on tetanus, but they are as yet incomplete, and
_ I must defer their publication for the present.
NOTE ON THE PRESENCE OF AN INSOLUBLE
SUGAR-FORMING SUBSTANCE IN PENICILLIUM.
By A. G. Dew-Smitu, B.A., Trinity College, Cambridge.
PENICILLIUM, grown in a solution of inorganic salts with am-
monium tartrate and cane sugar, very rapidly manufactures
SUGAR-FORMING SURSTANCE IN PENICILLIUM. 83
considerable quantities of both protoplasm and cellulose, but no
starch. It seemed to be a matter of interest to ascertain whe-
ther any body at all analogous to starch or glycogen could be
detected in the mould when grown in large quantities. For, on
the one hand, the association of glycogen with living functional
protoplasm is at least common, and may possibly be universal
(the quantity of glycogen associated with any given quantity of
protoplasm varying within wide limits). On the other hand,
we have no direct evidence that penicillium growing in saccha-
rine solutions obtains its cellulose by the immediate trans-
formation of the sugar; on the contrary, analogy would rather
point to the conclusion that the cellulose is a product of the
changes going on in the protoplasm which itself feeds on the
sugar. If such a view be correct, one would naturally expect
to find in the plant small quantities of carbohydrates analogous
to starch, either as stages in the transformation into cellulose
or as by-products.
With these views I grew large quantities of penicillium,
making use of Mayer's (Untersuch. iiber d. alkohol. Gahrung)
normal solution, Potassic Phosphate ‘1 grm., Calcic Phosphate
‘01 grm., Magnesic Sulphate ‘1 grm., Ammonium Tartrate
15 grm., Sugar, 20 cc of a 158 aqueous solution, Distilled Water
1000 ce.
The spores were sown in this fluid in shallow saucers. In
a few days a thick crust of the mould appeared. This was
removed before the gonidia had formed; in this way the
mycelium was roughly separated from the fluid in which it
was growing.
The fluid after filtration and evaporation to a small bulk
was treated with alcohol as long as any precipitate occurred.
The precipitate was then washed with alcohol until free from
sugar. The watery decoction of the precipitate thus freed from
sugar gave no evidence of the presence of glycogen, or of sugar
after being acted upon by ferment. The ferment used was
pancreatic ferment purified by means of glycerine after Von
Wittich’s method.
The mycelium was thoroughly washed with alcohol until
free from sugar. A watery decoction of this washed mycelium
was filtered until a clear light yellow solution was obtained.
6—2
84 MR DEW-SMITH. SUGAR-FORMING SUBSTANCE IN PENICILLIUM.
This decoction gave no evidence of glycogen. Evaporated down
to a small bulk, it was treated with alcohol as long as any pre-
cipitate occurred. The precipitate washed with spirit and
redissolved in water, was also free from glycogen and from any
body giving rise to the reactions for sugar after the action of
ferment.
The portions of mycelium left after aqueous decoction, and
therefore insoluble in water, were treated with alcoholic potash,
and afterwards thoroughly washed free from alkali. A quantity
of this purified mycelium was boiled in water, and the aqueous
filtrate again examined for glycogen or for sugar-forming sub-
stances. But none were found.
Up to this point my results were entirely negative. A
quantity however of this purified mycelium which was insolu-
ble in boiling water, gave none of the reactions for starch
and was not coloured by iodine, was suspended simply in
water, a small quantity of ferment perfectly free from sugar
added, and the whole left at the temperature of 40°. In a short
time the fluid contained considerable quantities of sugar, re-
ducing the cupric solution with great readiness.
There is present then in the mycelium of penicillium a
substance which resembles glycogen in not being acted upon
by alcoholic potash, and in becoming converted into sugar
under the influence of amylolytic ferments, but which differs
from glycogen in being insoluble in water, and gives neither
the starch nor the glycogen reaction with iodine. In its in-
solubility it resembles cellulose, but differs from that body in
its behaviour towards ferment. An insoluble glycogen has
been described as being present in the liver in company
with the ordinary glycogen, but it is more than probable the
insolubility in this case was due to the fact that the other
solid substances protected the soluble glycogen from solution.
In the case of the penicillium mycelium such an explanation
is invalid. The mycelium was reduced to a fine powder and
so thoroughly boiled that everything soluble in water must
have passed into solution.
I attempted to isolate this substance in various ways, but
entirely failed.
CONTRIBUTIONS TO THE ANATOMY OF THE IN-
DIAN ELEPHANT (ELEPHAS INDICUS). Parr III.
THE HEAD*. By M. Watson, M.D., Demonstrator of
Anatomy in the University of Edinburgh.
I HAVE no intention in this communication of writing a full
description of the anatomy of the head of the Indian Elephant,
but merely to direct attention to certain structures which have
either been overlooked by previous anatomists, or have been
described as possessing arrangements differing from those which
I have met with in the course of my dissection,
TEMPORAL VENOUS RETE MIRABILE.
In Froriep’s Notizen, October, 1832, p. 39, the following
passage occurs :—Otto “showed a drawing of a peculiar arterial
network, which he found in the neighbourhood of the peculiar
excretory gland of the head of the elephant, situated between
the eye and the ear, and seemingly occupying the entire side of
the head, and remarked that similar arterial retia and anasto-
moses were to be found in several tardigrades, as well as in the
extremities of many plantigrades.” The same author, in his
Erléuterungs-Tafeln (Heft 6), gives a figure of this rete, but in
the explanation appended to the plate does not state whether
the rete is venous or arterial. As, however, the figure is
‘coloured red, we must conclude that he believed it to be an
arterial network, as in all the other figures blue is the colour
used to distinguish the veins when present. Farther, Mayer
in his paper (Nova Acta XxXIL) refers to this description; and
although he had himself dissected the animal, does not disagree
with it. In my own dissection I found, by means of injection,
that this rete was not arterial but venous. It is formed by
numerous anastomosing veins, which occupy the whole of the
temporal fossa, lying superficial to the temporal muscle, but on
1 PartI. On the Thoracic Viscera, appeared in this Journal, November, 1871 ;
Part II. On the Urinary and Generative Organs, in November, 1872. This part
was read as a communication at the Bradford meeting of the British Associa-
tion, Sept. 22, 1873.
86 DR WATSON.
a plane deeper than that of the temporal arteries, or that of the
peculiar temporal gland. These veins are small in size round
the margins of the temporal fossa, and are formed by the junc-
tion of numerous rootlets derived from the skin and superficial
structures. Increasing in size as they are traced down to the
temporal fossa, and communicating freely with one another to
form the rete, they finally converge toward the root of the zy-
goma to form three or four main trunks, which uniting together
give rise to the temporal vein. By means of several branches a
communication is opened up with the facial vein in front, but
this latter trunk takes no part in the formation of the rete,
which moreover receives several branches from the substance of
the temporal gland. It is worthy of remark that there are no
valves present in the veins forming this rete, as the entire net-
work can be freely injected from the trunk of the temporal
vein. This trunk finally unites with that of the internal maxil-
lary vein, and both open into that of the zxternal jugular, the
external jugular vein being absent in the elephant.
The temporal artery, after crossing the zygoma, divides into
two main trunks, from both of which numerous branches are
given off to the surrounding parts, but they have no tendency
to form a rete, as described by the authors before referred to.
What may be the function of a venous rete in this situation it
is difficult to determine.
EYE.
In the next place I wish to draw attention to some inter-
esting points relating to the arrangements of parts within the
orbit of the elephant. First, as regards the muscles of the
orbit, Mayer (Nova Acta) states that there is a depressor of
the lower eyelid in this animal, in addition to the other muscles
usually contained within the orbit. So far as I can ascertain,
he is the only author who has, up to this time, observed the
muscle in question. It arises along with the recti and obliqui
from the bony canal posterior to the orbit, passing forward
beneath the globe of the eye in the same manner as the ele-
vator of the upper lid passes forward above the eye, and is
inserted into the cartilage of the lower eyelid. It evidently
ANATOMY OF THE INDIAN ELEPHANT, 87
depresses the lower eyelid. In addition to this muscle there is
farther to be observed a very extensive and well developed
pericsteal muscle, which has not hitherto been observed in this
animal. It corresponds exactly in position to the similar muscle
in the sheep and deer’. The orbit itself is completed poste-
riorly and inferiorly by periosteum, and it is in relation with
the orbital surface of this periosteum that the muscle referred
to is situated. The fibres composing it, which are of the in-
voluntary or non-striated description, form a large sheet cover-
ing nearly the whole of the periosteum, and run from without
obliquely forward and inward. The function of this muscle,
although generally stated to be that of a protractor of the eye-
ball, is, I think, difficult to determine with precision, Camper,
Harrison, and Mayer, refer to two small but distinct muscles,
which pass to be inserted into the cartilage of the third eyelid.
Of these, one arises from the lower, and the other from the
upper eyelid, and both pass inwards to be inserted into the
cartilage just referred to. On careful dissection I have not
found them to be distinct muscles, but formed by certain of
the palpebral fibres of the orbicularis palpebrarum, which pass
inward to be inserted into the third eyelid. According to
Mayer, one of these museles acts by drawing the third eyelid
outwards across the globe of the eye, whilst the other retracts
it toward the inner canthus. That this is the action of these
muscles seems to me extremely doubtful, as both being formed
of prolongations of the orbicularis, it is difficult to imagine that
two parts of the same muscle supplied by one nerve (the 7th)
should have actions so opposed to one another. So far as a
study of the anatomical arrangement of the parts would enable
me to decide, I am inclined to think that both those muscles
- will tend by their contraction to draw the third eyelid outwards
across the eye, but by what agency this lid again regains its
position is more difficult to determine. It would be interesting
to learn from those who may have opportunities of watching the
elephant during life, whether the third eyelid is drawn across
the eye when the upper and lower eyelids are separated, or
1 See Professor Turner’s papers on the Periosteal Muscle of the Orbit in Man,
the Sheep and Deer, in Proc. Roy. Physical Soc. of Edinburgh, Dec. 19, 1861, and
Natural History Review, Jan. 1862.
88 DR WATSON.
only when these are closed. If the latter supposition be cor-
rect, it will establish the views I have advanced regarding the
action of these little museles.
Regarding now the lachrymal apparatus of the elephant, we
find that various statements have been made by different au-
thors. Camper and Harrison, on the one side, maintain that
no portion of a lachrymal apparatus is present in the elephant,
while, on the other hand, Mayer (the most recent writer on the
subject) says, “The puncta lachrymalia are small, the lachrymal
duct single and very narrow, the laechrymal gland of tolerable
size. Its excretory duct is as large as a knitting needle, and
opens on the external angle of the eyelids;” and he adds, “It
is striking that Camper should neither have discovered this
gland, its excretory duct, nor the lachrymal canal.” Perrault
also mentions the presence of lachrymal glands in the elephant.
My own observations agree with those of the authors who have
not discovered any portion of a lachrymal apparatus, although
each separate element was carefully looked for. It is difficult
to explain the statements of those authors who maintain the
existence of such an apparatus, more especially when it is borne
in mind that the ethmoid bone in the elephant is quite imper-
forate, and consequently affords no way of escape for the lachry-
mal secretion. True, a Harderian gland, similar to that which
exists in connection with the third eyelid in birds, is to be
found in the elephant. It does not, however, occupy the usual
position of the lachrymal gland at the outer angle of the orbit,
but rests between the inner wall of that cavity and the internal
rectus muscle. Its excretory duct, moreover, opens on the sur-
face of the third eyelid, and not in the usual position of the
ducts of the lachrymal gland. That this gland to some extent
fulfils the function of the lachrymal gland is rendered probable
by the statements of African travellers, one of whom (Cumming)
describes an elephant, after suffering from the effect of several
balls, as weeping profusely. The mode, however, in which the
secretion of this gland is got rid of, under ordinary circum-
stances, is difficult to determine in the absence of all trace of
an excretory apparatus.
ANATOMY OF THE INDIAN ELEPHANT. 89
THROAT.
I shall now direct attention briefly to certam peculiarities
in the formation of the throat of the elephant, which up to this
time have escaped notice, and which seem to be of importance,
inasmuch as they afford an explanation of some of the creature’s
functions which have not hitherto been explained. In my first
communication I had occasion, when remarking on certain pecu-
liarities in the thoracic viscera of the elephant, to refer to the
following statement made by Sir Emerson Tennent in his work
on Ceylon, to prove that the animal possesses the power of
withdrawing water stored within the cavities of the stomach by
means of the trunk inserted into the mouth. Sir E. Tennent
says, with reference to the Indian elephant, “I have elsewhere
described the occurrence to which I was myself a witness of
elephants inserting their probosces into their mouths and with-
drawing gallons of water, which could only have been contained
in the receptacle figured by Camper and Home;” and he (Ten-
nent) farther quotes from the author of the Ayeen Akberry as
follows: “An elephant frequently with his trunk takes water
from his stomach and sprinkles himself with it, and it is not in
the least offensive.”
That the same thing is true as regards the African elephant
has been observed by Cumming, who, in his travels in South
Africa, when speaking of these animals, says: “They seemed
heated by the pace at which they had retreated, and were now
refreshing themselves with large volumes of water which Nature
enables them to discharge from their capacious stomachs and
shower back upon their bodies with their extraordinary trunks.”
This regurgitation of water from the stomach I showed in my
first paper to depend not on any peculiarity of structure in the
elephant as compared with that of other animals, but that it
was a function similar to the physiological regurgitation of food
in the ruminant, and performed by means of the combined
actions of the diaphragm and other abdominal muscles. It now
remains to show in what the peculiarity of construction of the
throat of the elephant consists to enable the trunk when placed
in the mouth to withdraw the water regurgitated from the sto-
mach. For it is evident that were the throat of this animal
90 DR WATSON.
similar to that of other mammals, this could not be accom-
plished, as the insertion of a body, such as the trunk, so far into
the pharynx as to enable the constrictor muscles of that organ
to grasp it, would at once give rise to a paroxysm of coughing,
or were the trunk merely inserted into the mouth, it would be
requisite that this cavity be kept constantly filled with water at
the same time that the lips closely encircled the inserted trunk.
The formation of the mouth of the elephant, however, is such
as to prevent the trunk ever being grasped by the lips so as
effectually to stop the entrance of air into the cavity, and thus
at once, if I may so express it, the pump-action of the trunk is
completely paralysed. We find therefore that it is to some
Fig L
Explanation to Figure 1. A, superior aperture of pharynx. B, root of
tongue. C, soft palate with larynx projecting through the centre. D, pharynx.
ANATOMY OF THE INDIAN ELEPHANT. 91
modification of the throat that we must look for an explanation
of the function in question, and this we find to be as follows :—
The superior aperture of the pharynx, Fig. 1 A, is extremely
narrow, so much so as to admit, with difficulty, of the passage
of the closed fist. Immediately posterior to this narrow aperture
the pharynx dilates into a pouch of large size, Fig. 2 E, capable
of containing a considerable quantity of fluid. This pouch is pro-
longed forward beneath the root of the tongue, and is bounded
in the following manner. The floor extends from the epiglottis
as far forward as the root of the tongue, being formed from be-
hind forward by the thyroid cartilage, thyro-hyoid membrane,
and hyoid bone. Its lateral walls are completed by the sides of
the pharynx (that is, by the superior constrictor muscles, Fig. 2 F),
in addition to the stylo- G, and hyo-glossi H, muscles. The root
of the tongue forms the anterior boundary, whilst the posterior
wall is completed by depression of the soft palate, or when the
latter is elevated the pouch then communicates freely with the
cesophagus. In connection with this pouch is to be observed the
very peculiar form of the hyoid bone, which being deeply concave
on its upper surface forms as it were the greater part of the floor
of this pouch. Between the pouch and the concavity of the
hyoid bone, moreover, there is placed a large quantity of loose
and distensible connective tissue, which permits of the expan-
Fig. 2.
Explanation to Figure 2. E,E pharyngeal pouch. F, superior constrictor.
G, stylo-glossus. HH, hyo-glossus. K, small muscle which diminishes the depth
of the pharyngeal pouch. L, genio-glossus muscle.
92 DR WATSON.
sion of the pouch. The size of the latter is, moreover, liable to
alteration by the actions of several muscles. These are more
especially the hyo-glossi muscles, and two little additional
muscles, Fig. 2 K, the homologies of which I have not yet
been able to determine, which, springing from the middle line of
the hyoid bone in front of the pouch, pass up, one on either
side of the middle line, and blend with the other muscles form-
ing the root of the tongue. By the action of these muscles the
pouch may be diminished in depth ; but in consequence of the
narrow interval existing between the hyoid cornua, the length
of the pouch from before backwards cannot be altered, as the
thyroid-cartilage is thereby prevented from being approximated
to the hyoid bone. I have now to complete the anatomical
description of this pharyngeal pouch by a reference to the
formation of the soft palate. This, Fig. 1 C, which is of very
large size, forms almost a complete muscular diaphragm, through
the central aperture of whieh projects the superior extremity
of the larynx, which thus in some respects approximates to
the arrangement of the corresponding parts in certain cetacea
as described by Dr James Murie’. With reference to the mus-
cles entering into its formation, we find that the palato-glossus
is entirely absent, its place being supplied by a wide and ex-
tremely distensible fold of mucous‘membrane. The palato-
pharyngeus, on the other hand, is of large size, and forms, in
fact, the principal feature in the soft palate. There is neither
a levator nor a tensor palati present. Such being a brief de-
scription of the anatomical arrangements met with im connec-
tion with this pharyngeal pouch, a few words may now be said
on their physiological bearing. An elephant can, as the quo-
tations sufficiently prove, withdraw water from his stomach in
two ways; first, it may be regurgitated directly into the nasal
passages by the action of the diaphragm and abdominal mus-
cles, the soft palate being at the same time depressed so as to
prevent the entrance of water into the mouth. Having, in this
manner, filled the large nasal passages, communicating with
the trunk, the water contained in them is then forced through
_ the trunk by means of a powerful expiration ; or, in the second
place, the water may be withdrawn from the cavity of the
1 Trans. Zoolog. Soc. Lond. Vol. v1.
ANATOMY OF THE INDIAN ELEPHANT. 93
mouth by means of the trunk inserted into it. Now, in this
case, it is manifestly impossible that the water can be contained
within the cavity of the mouth itself, as | have already shown
that the lips in the elephant are so formed as effectually to
prevent this. The water regurgitated is, however, by means
of the elevation of the soft palate, forced into the pharyn-
geal pouch. The superior aperture of this pouch being much
narrower than the diameter of the pouch itself, and being com-
pletely surrounded by the muscular fibres of the stylo-glossus
on each side and the root of the tongue in front, which is pro-
longed backwards so as to form a free sharp margin, we have
thus as it were a narrow aperture surrounded by a sphincter
muscle, into which the trunk being inserted and grasped above
its dilated extremity by the sphincter arrangement just referred
to, air is thus effectually excluded, and the nasal passages
being then exhausted by the act of inspiration, water is lodged
within these passages to be used as the animal thinks fit, either
by throwing over his body or again returning it into his mouth,
as observed on one occasion by Cumming, who says: “'Through-
out the chase this elephant repeatedly cooled his person with
large quantities of water, which he ejected from his trunk, over
his back and sides; and just as the pangs of death came over
him, he kept pouring water into his bloody mouth until he died.”
The increase in the size of this pouch is accomplished mainly
by the depressor muscles of the hyoid bone, which, in conse-
quence of the tongue being fixed and restrained in its move-
ments by the muscles attaching it to the lower jaw, depress the
hyoid bone and thyroid cartilage, which are both freely move-
able, at the same time that the tongue itself is almost fixed, and
in this manner the depth of the pouch is materially increased.
Such is the explanation of a function which, so far as I am
aware, has not up to the present time been satisfactorily ex-
plained, and it will be of much interest to examine the corre-
sponding region in the African elephant, to ascertain if arrange-
ments similar to those I have described in the Indian species
are to be found in that animal. If similarity of function im-
plies similarity of structure, then I have little doubt that such
will prove to be the case. The modifications in the throat of
the elephant are not without interest from two points of view,
94 DR WATSON. ANATOMY OF THE INDIAN ELEPHANT.
In the first place, these modifications are such as show that,
in respect of this portion of its structure as in several others,
the elephant closely resembles certain forms of the cetacea, and
are thus of importance inasmuch as they furnish one more item
of evidence in favour of a relationship which has been long
suspected to exist between those two groups, which when taken
by themselves, although they seem sufficiently widely separated,
are, nevertheless, connected by a number of intermediate forms:
and, secondly, they are not without interest inasmuch as they
afford food for reflection as to the origin of these modifications
when compared with the corresponding parts of other mammals.
Did they arise gradually in accordance with the law which
tends ever to bring the organism into harmony with external
conditions, and so to adapt the functions of such an organism
as finally to give rise to that chain of circumstances which is
formulated in the expression, “survival of the fittest”? Or are
we to believe, with the teleologists of the previous century, that
these modifications were occasioned by the direct intervention
of a great First Cause ever attempting to remedy imperfections
which he had at first created? Which of those alternatives is
to be accepted must be left to the private judgment of each
individual enquirer.
oN THE APPARENT PRODUCTION ‘OF A NEW
EFFECT BY THE JOINT ACTION OF DRUGS
WITHIN THE ANIMAL ORGANISM. By T. LAupER
Brunton, M.D., D. Sc. Edin., Lecturer on Materia
Medica and Therapeutics at St Bartholomew's Hospital.
THE admirable researches of CrumBrown and Fraser have
demonstrated that the physiological action of several alkaloids
may be completely altered by their union with such bodies
as iodide of methyl. The compounds thus produced some-
times act on organs which do not appear to be affected by
either of the components separately, the ends of the motor
nerves for example being paralyzed by iodide-of-methyl-strych-
nia, though neither iodide of methyl alone, nor strychnia alone,
seems to have much influence over them. Although chemical
action outside the body alters in this way the action of alka-
loids, Iam not aware that any instance has been noticed in
which a similar modification appears to be produced by the
joint action of two drugs after their introduction into the
animal organism. I have lately observed an example of this
sort in the case of strychnia and nitrite of amyl. The experi-
ments which I made on this subject were performed in several
ways, but I will only describe the two most important. In the
first series of experiments a solution of strychnia was injected
into the dorsal lymph sac of a frog, and as soon as tetanus
came on the animal was put into a vessel filled with the
vapour of nitrite of amyl. A second healthy frog was also
introduced along with it for the purpose of comparison. They
were left in the vessel till both were motionless, when they
were removed and the sciatic nerves of both were exposed.
On irritating these nerves by the application of a Faradic
current, vigorous contractions occurred in the limbs of the frog
poisoned by nitrite of amyl alone, but those of the animal
poisoned by strychnia and nitrite of amyl together remained
motionless. The skin was then removed from the legs of both
and the muscles irritated by the application of the current
96 DR BRUNTON. JOINT ACTION OF DRUGS.
directly to them. In many instances, those of the frog poi-
soned by the nitrite and the strychnia together contracted
nearly as strongly and readily as those poisoned by the nitrite
alone. It is therefore evident that their failure to contract
when the nerves were stimulated must have been due to
paralysis of the nerves themselves, just as it 1s In poisoning
by woorara. The second series of experiments was made by
ligaturing the artery supplying one leg of a frog before injecting
strychnia into the lymph sac. The poison was thus carried
by the blood to every part of the body except the leg whose
artery had been tied. The animal was then placed in a vessel
filled with the vapour of nitrite of amyl as before, and after
motion had ceased the sciatic nerves were exposed and irri-
tated. It was then found that the muscles of the ligatured
leg which had been exposed to the nitrite of amyl, but pre-
served from the strychnia, contracted vigorously when the
corresponding sciatic was irritated, while those of the other
leg did not respond at all. When the skin was stripped off,
however, and the muscles irritated directly, in many instances
no great difference could be noted between their irritability.
The muscles of frogs which had been poisoned either with
strychnia and nitrite of amyl, or with nitrite of amyl alone,
passed more quickly than usual into a state of rigor mortis;
and I therefore regard nitrite of amyl as a muscular poison.
It is not improbable that the apparent paralysis of the motor
nerves may be partly due to diminution of the irritability of
the muscle itself, but the results of direct stimulation show that
this is not sufficient to explain it entirely, and we must there-
fore believe that the nerves themselves are also paralyzed.
Besides nitrite of amyl, I have tried the nitrites of sodium,
ethyl, butyl and capryl; but my researches on these are not yet
completed. They seem however to show that the nitrites are
muscular poisons, but their actions differ according to the bases
which they contain.
AN ACCOUNT, HISTORICAL AND PHYSIOLOGICAL,
OF THE MADAGASCAR ORDEAL POISON, THE
TANGHINIA VENENIFERA. By ANDREW Davipson,
F.RC.P.E, Medical Missionary and Physician to the
Court of Madagascar.
ORDEALS of various kinds have been devised in certain stages
of civilization as a means of testing the guilt or innocence of
suspected persons. Records have come down to us of the
widespread existence of this usage in remote antiquity, and
in more recent times ordeals by fire, water, and wager of
battle were prescribed by law and sanctioned by religion
throughout the whole of Europe. Ordeal by pozson is, how-
ever, peculiar to Africa, although philology renders it probable
that the same practice may have prevailed among the progeni-
tors of our own race in prehistoric times.
It is to be observed that these ordeals are chiefly employed
for the detection of witchcraft, by which African Jurists under-
stand the use of poisonous drugs for evil purposes. It is in fact
equivalent to the dapuaxeia of the Greeks; and as the terms
gappaxos and veneficus were applied by the ancients to signify
alike a physician, a sorcerer, and a poisoner, so in many of the
African languages the same peculiarity obtains. This arises
from the fact that amoxg these and other primitive races the
physiological effects of drugs, whether poisonous or medicinal,
are ascribed to some magical power, either inherent in the sub-
stance itself, or imparted to it by sorcery. Medicines are thus
employed as charms both for causing and curing disease. With
such superstitious notions of the properties of poisons, it was
only natural that they should ascribe the differences in the
results observed to follow their administration to a sort of
discriminative faculty or intelligence possessed by the sub-
stance, and thus have come to employ poisons in the detection
of occult crimes, such as witchcraft.
Although we know that the custom of ordeal by poison
prevails over a great part of the continent of Africa, we are
-_
MOT aVAULIs (
98 MR DAVIDSON.
as yet unfortunately ignorant, in most instances, of the poisons
employed by the different tribes’; and, with the exception of
the Calabar bean, none of them have been subjected to a
satisfactory examination. This is to be regretted from a medi-
cal as well as a scientific point of view, as remedial agents of
high value will probably be found among these powerful ordeal
poisons.
As the advance of civilization has now abolished the use of
one of the most celebrated of these ordeals—the ‘ Tangéna’ of
Madagascar—it seems desirable to put on record the mode of
administering it and its effects on man, while such information
may still be obtained from those who were acquainted with its
employment, and had witnessed or experienced its effects,
Shortly before my arrival in the island, in 1862, the Tangéna
ordeal was abolished, but as it happens that an officer, now
attached to the hospital, was formerly from hereditary office an
administrator of the poison, I have in this way had the oppor-
tunity of obtaining trustworthy information upon this subject.
Ffstorical—There is no certain evidence when or how the
Tangéna first came to be used as an ordeal in Madagascar. We
know that some such method of trial has long been practised in
the Island. The testimony of Flacourt, who visited Madagascar
in the middle of the 17th century, is conclusive upon this
point; but if his statements are to be considered as strictly
accurate, some other poison must at that time have been used
in the district visited by him’. Ordeals of other kinds, such as
that by plunging the hand into boiling water, were at one time
practised in some parts of the country; and there seems reason
to believe that the Tangéna was not generally or frequently
employed until the beginning of the present century.
1 Dr Livingstone observes that this custom ‘‘is common among all the
Nezro nations north of the Zambesi.” The natives of that part of Africa
employ a plant called the ‘Goho’ which is possessed of purgative and emetic
properties.
The poisonous juice of the Erythrophlaeum Guineense is employed for the
same purpose on the coast of Guinea, and the Physostigma venenosum by
the natives of Calabar. In the inland regions near the equator, according to
Du Chaillu, the natives use as an ordeal the root of a plant or tree called
‘‘Mboundou” conjectured by Prof. Torry of New York to be a species of
strychnos,
2 He says that the Malagasy administer for this purpose ‘‘ Manrechetsi,
qui est de quelque sorte d’herbe ou de racine qui est poison et fait mourir celui
quien mange.” Histoire de la grande tle de Madagascar,
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 99
It was seldom had recourse to in ordinary judicial cases in
which more rational modes of trial were followed, but was
reserved for the detection of those guilty of infamous crimes,
for the discovery of whom ordinary evidence either could not
be obtained or would not suffice. Such crimes were treason
and witchcraft, and indeed the latter comprehended the
former, and for the detection of these it was administered
either by order or permission of the sovereign, and in presence
of officers appointed by him. An experiment was, however,
frequently enough made, in corpore vile, in the instance of
individuals suspected of minor offences, or in order to decide
which of two or more persons was guilty of a crime known or
believed to have been committed by one or other of them.
In the former case, a dog having been selected as a substi-
tute for the suspected party, the Tangéna was given to the
animal in the same way as when administered to a human
being. When again it was given with a view to decide be-
tween two or more accused persons, then dogs of similar size
and condition were selected, and the party whose representative
first succumbed to the poison was treated as guilty.
Mode of administration.—The ordinary mode of administra-
tion was as follows:—Two Tangéna almonds, or nuts as they are
often styled, were taken, and the half of each rubbed down
with water. This custom of taking the half of two different
almonds was adopted in order to increase the probabilities
that the poison administered should be only of the average
strength.
The suspected party now ate a little rice and was afterwards
made to swallow three small pieces of fowl’s skin, and this was
followed by the Tangéna emulsion. After a few minutes, vary-
ing however according to the result desired by the adminis-
trator, tepid water was given in considerable quantities, and
violent, long-continued vomiting usually ensued. If the three
pieces of skin were discharged the suspicion of guilt was dis-
missed, as a rule, and the friends of the unfortunate were then
left to do their best for his recovery. Not unfrequently, how-
1 I say ‘‘as a rule” for some other omens of an unfavourable kind, which
I do not require to detail here, occasionally affected the result.
7—2
£00 , ; MR DAVIDSON.
ever, the poison operated more as a purgative than as an emetic,
and then it often happened that with or without the stigma of
crime (according as the pieces of skin were retained or rejected)
the case terminated fatally. It can easily be understood that
state policy readily attained its crooked ends by the administra-
tion of the Tangéna. It was observed that those who might
be called “the opposition members of the government” seldom
recovered from the ordeal. So far as can be ascertained, it
proved fatal in as many as one in ten cases when given with
no hostile intention. As it was often administered to whole
villages at once, it will be understood that the numbers
destroyed by this poison were immense.
The points especially affecting the result seem to have
been:—(a) The colour of the kernel; the very red ones are said,
and probably with truth, to be more poisonous than the less
ripe ones, which are whiter in colour. (6) The amount admin-
istered was in every case enough to prove fatal, if not speedily
rejected. From what I have learned from the natives, as
well as from the results of my own experiments, I have no
doubt that the weight of one almond is amply sufficient to
poison an adult, if not got rid of by vomiting. (¢) If admin-
istered on an almost empty stomach, it was more dangerous
than when a larger quantity of rice had been previously taken.
(d) A great deal depended upon the seasonable and reasonable
administration of diluents. Experience enabled the expert to
judge the time when to give drink, and the amount required to
effect his object, whether that might be the death or recovery of
the victim.
Symptoms.—As the result of a careful examination of seve-
ral who have been themselves subjected to this ordeal, and of
many who have witnessed its effects on others, I conclude that
the symptoms produced by it when given in poisonous doses, in
the manner just described, are as follows:—A peculiar numb
tingling sensation is felt in the mouth and fauces, due to its
topical action. Several of those who have undergone the
ordeal have assured me that they experienced a similar feeling
more or less over the whole body, but especially in the hands.
This point is important, for my experiments on warm-blooded
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 101
animals have not indicated any noticeable disturbance of sensa-
tion. Sickness ensues with vomiting, intense, distressing and
repeated—first of the contents of the stomach—then of bile
and mucus. The vomiting is attended by a feeling of great
debility and anxiety. If the greater part of the poison has
been thus ejected, the patient recovers perfectly within a short
time. Where more of the poison has got into the circulation
the sufferer is said to feel giddy. The Malagasy, however, use
their word for vertigo in a loose sense. J am therefore inclined
to think that partial paralysis of motion with unsteady gait
may be the condition indicated. The patient, under the influ-
ence of the Tangéna, staggers if he attempts to walk, is unable
to support his own weight, and falls down helpless and para-
lysed. Although the mind is usually clear, yet delirium occa-
sionally occurs. Along with these nervous and cerebral symp-
toms, purging and urination appear and are more or less urgent.
The faecal discharges do not contain blood or mucus. Nothing
abnormal has been observed by the natives in the appearance
of the urine. The patient in cases tending to a fatal issue
becomes unable to rise. In other instances, according to the
testimony of observers, he lies as if asleep, and when roused
answers like a drowsy man, then lapses back into his former
condition. In other cases the patient remains conscious to the
last, without either stupor or delirium. Death is preceded by
spasmodic movements of the fingers and toes. Purging is a
bad symptom and worse the more urgent it is. Almost none
recover when the stage of stupor has been reached.
The natives know of no antidote for this poison, but they
think that the application of cold and draughts of lemon-juice
are of service.
I may remark upon the condition of sleepiness described
above as of pretty frequent occurrence in the advanced stage of
poisoning by this substance, that I do not believe that my
informants were able to distinguish between narcotism and a
state of prostration. I have however given their statements
literally.
Upon this point I may further observe that my experiments
on the lower animals do not seem to countenance the opinion
of some, that there is any narcotic property in the substance.
102 MR DAVIDSON.
There is only one exception to this statement of the result of
my experiments. In two instances in which I administered the
Tangéna to fowls, they appeared to be overcome by sleep.
No post-mortem examination has been made of those who
have died by Tangena.
Botanical note.—The Tangéna, or Tanghin (J'anghinia Venent-
Jera; Cerbera Tanghin, Hooker), is a large tree of the natural order
Apocynaceae. The poisonous part of the plant is the fruit, which is
a drupe, almost the size of an apple. The colour of the fruit is
a greenish-yellow; the external pulp which surrounds the kernel
is soft, somewhat grey in colour, destitute of smell, and possessed
of a slightly bitter, disagreeable taste. The kernel is hard, ligneous,
and brown, and elliptical in shape. Within this is the almond,
which is divided into two cotyledons, of the consistence of a newly-
plucked bean, varying in colour from a white to a brownish-red;
and weighing from forty to seventy grains. For a minute botanical
description of the tree and its fruit the reader may consult Hooker’s
Botanical Miscellany, 11. 290. The Tangéna grows abundantly in
the forests on the east coast of Madagascar, it is rare in the central
provinces, and towards the south of the island.
Chemistry.--Two crystalline principles are said to have been
obtained from the Tangena: the one, the bitter principle Tanghinin ;
the other, the poisonous principle which has been named Vanghicine,
and is described as transparent plates obtained by ether, insoluble
in water, bitter and poisonous. I have no access however to any
account of the process followe:l in the separation of these; and I shall
state in a few words what little I know upon this point.
(a) ‘The kernel contains a large quantity of an inert, bland oil,
and if rubbed up with water it forms a white emulsion.
(b) Its active principle is insoluble, or at least nearly so, in
water, readily soluble in alcohol, zther, and chloroform, as is proved
by the activity of the extracts made by means of these solveuts.
I have obtained by means of chloroform impure crystals, in the
form of long, flattish needles, arranging themselves under the micro-
scope as if branching out at acute angles from a centre.
(c) By treating a carefully prepared alcoholic extract with water
a white precipitate is obtained.
In my experiments, I have used the simple emulsion, and extracts
made with ether or alcohol, and in a few instances the impure
crystals mentioned above’.
1 T have given the above as it was written more than three years ago. As
my object was not to investigate its chemical properties, but its physiological
action, I had neglected to note down the steps of the process by which I
obtained the crystals alluded to; and I cannot now add anything from memory.
The recent work by Chatin (Recherches pour servir &@ Vhistoire botanique,
chemique et physiologique du Tanguin de Madagascar. Par Joannes Chatin,
Paris, 1873) upon this poison would render it prubable that the active principle
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 103
Physiological Action.—Results of experiments on the lower
Animals.
The Tangeéna proves fatal by absorption however introduced,
whether into the alimentary canal, the serous membranes, or
into the cellular tissue. It acts less actively if swallowed,
because partly got rid of by vomiting. When a concentrated
solution of the poison is applied to the frog’s foot it is slowly
absorbed and causes death. Ligature of the blood-vessels pre-
vents or delays its action.
(A). Topical Effects.
1. Its topical action on the mucous membranes is to alter
and diminish the sensibility in the part. The same results
follow its application to the skin.
2. Its local action on muscular tissue is no less evident.
Applied to the exposed heart of the frog it produces immediate
paralysis. It acts with equal rapidity on the excised heart
which is still pulsating. The electric contractility of the poi-
soned muscle is diminished or destroyed. Applied for about
thirty minutes to the leg of a frog, the part is paralysed, and if
pricked or pinched, is insensible or nearly so; all the while the
animal is lively, and the blood continues to circulate naturally
in the web of the affected foot. After the lapse of about an
hour, the other leg becomes paralysed, then the upper ex-
tremities, and finally death occurs in an hour and a half, or
two hours.
3. When applied to the exposed sciatic nerve, paralysis
followed by death has also been the result; but I am unwilling
to deduce any conclusion from this, as, from the difficulty of
is not a neutral body, but an alkaloid which he obtained thus :—Having first got
rid of a considerable part of the oil by pressure, he made an etherial extract,
which was treated by warm alcohol, and on evaporation left a residue which
he thus describes: ‘‘la liqueur évaporée dans la vide laissa un résidu assez
considérable, brundtre, légérement amer et comme granuleux en certains
points; facilement fusible, ce produit, chauffé au contact de l’air, se comportait
comme un corps gras. Le produit ainsi obtenu était toxique; je le traitai
alors par de l’acide acétique étendu et j’obtins, par l’évyaporation des liqueurs,
une petite quantité de poudre blanchatre, agsez soluble dans leau, beaucoup
plus soluble dans l’alcool, Elle fut en conséquence traitée par ce dissolvant et,
par l’éyaporation dans la vide, elle donna de petits crestaux, d’un blanc vitreux
et appartenant au systéme diclinorhombique” (p. 30). This subject will still
require further investigation.
1040: MR DAVIDSON.
localizing the poison, I am not quite satisfied that it did not
come into contact with the exposed muscles.
4. When applied to the conjunctiva it does not affect the
pupil.
(B). Action through the Circulation. (On warm-blooded
Animals.)
1. However introduced it produces violent vomiting and
purging in all animals capable of these actions. As an emetic,
Tangéna is even more violent, and operates sooner when intro-
duced directly into the circulation than when swallowed.
2. It produces marked paralysis of motion, apparently more
intense at first in the lower extremities, and last of all affecting
tbe muscles of the trunk and neck’. |
3. When it acts through the circulation, sensation seems
slightly if at all paralysed. Pinching the tail of a lemur makes
it suddenly exhibit signs of pain, long after it is incapable of
motion. Other facts seem to prove that sensation is altered,
not abolished.
4, The animal remains conscious and observant to the
last; excepting in fowls, as already mentioned, I have never
seen narcotism induced.
5. The action of the heart is first somewhat increased (at
least in some cases), then it becomes weak, irregular and slower,
and finally stops a short time before the respiration. The heart's
action is arrested in systole. The ventricles usually contain a
little blood but are never distended. The blood is sometimes
slightly coagulated, but more commonly fluid. The auricles and
venee cavee are engorged.
6. In one case I noticed peristaltic movements of the right
auricle, continuing for half an hour, or longer, after death,
excited by mechanical or chemical stimuli; and in another
case, this peristaltic movement was observed to occur spon-
taneously, twice a minute, producing a wave-like motion in the
ven cave.
1 It must be remarked, however, that in these experiments the poison was
injected into the cellular tissue over the loins. This fact should be kept in view
as it suggests an explanation of this observation.
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 105°
7. Idio-muscular twitchings of the pectoral muscles. are
commonly observed if the body be examined soon after death.
8. After death by Tangéna, I found the muscles to con-
tract on applying strong magneto-electric stimulation, but. less
perfectly and persistently than natural; but after the direct
application of the poison to a muscle, its irritability by elec-
tricity is much more distinctly diminished, or even destroyed.
9. The liver is always much congested, and notwithstand-
ing the urgent vomiting, the gall-bladder is sometimes full.
There are usually no signs of irritation, or injection of the
mucous membrane of the stomach or bowels. There is no con-
gestion of the brain or cord. The lungs are always exsanguine
and collapsed.
(C). Action on Frogs.
1. However introduced, Tangéna tetanizes the heart, with
varying degrees of rapidity, according to the size of the frog,
the amount of poison used, and the mode of administration.
The pulsations usually first increase in number, then become
less frequent, while from the first they tend to be irregular—
afterwards this irregularity becomes more marked. After some
time, tetanic contractions of distinct portions of the substance
of the ventricle take place, the tetanized portions remaining
pale; then complete tetanus of the organ ensues, preventing all
further entrance of blood. The auricles continue to contract
after the ventricle has ceased. The ventricle sometimes stops
for a little and then begins to act again. After death the
auricles are distended and the ventricle is contracted and pale.
As already stated, the heart’s action is immediately arrested by
the topical application of the poison to its substance. In this
case it is paralyzed not tetanized.
2. The heart ceases to beat before respiration and reflex
action are abolished. The decapitated frog, whose heart has
been arrested by Tangéna, will draw up its legs on being
irritated, unless indeed the paralysis of the extremities has been
very complete before the experiment has been tried.
3. Destruction of the medulla, or decapitation, does not
prevent, although it perhaps somewhat delays, the action of
Tangéna on the heart.
106 MR DAVIDSON.
4. The pulsations of the posterior lymphatic hearts become
rapid, irregular and excessively weak, and seem to cease at the
same time that the heart’s action is arrested.
5. The extremities are always more or less paralyzed.
After a pretty large dose, the frog remains with its posterior
extremities extended, and it requires strong stimulation to
excite any movement.
6. Reflex action is probably diminished, not immediately,
but soon after the administration of the Tangéna. This I ascer-
tained by suspending two frogs, of the same size, by the lower
jaw. I gave one of them a small dose of Tangena (a grain of
the semi-liquid spirituous extract), and then by the application
of similar mechanical and chemical stimulants, observed the
readiness with which in either case reflex movements fol-
lowed. ,
7. The muscles can be made to contract by the magneto-
electric current after the paralyzed limb has been amputated,
but less actively and for a shorter time than in a healthy one.
If a ligature be applied to the sciatic artery and vein, the limb
below the ligature will be protected from the poison and will
contract actively, if the magneto-electric current be applied ;
while the poisoned limb remains motionless, or nearly so,
whether the stimulus be applied through the nerve, or to the
muscle.
Experiments on warm-blooded Animals.
Exp. 1. On a Lemur.—At 7.58 am. Ten grains of a liquid
extract of Tanyhinia venenifera, mixed with one drachm of water,
were injected into the cellular tissue in the lumbar region of a full-
grown lemur. 8 a.m. The extremities are weak—scarcely able to
walk. 8.7. Vomits frothy mucus. 8.15. Continues to vomit, and
when it attempts to walk, its movements are slow and uncertain,
it is unwilling to move; the heart’s pulsations are reduced in fre-
quency, pupils normal. 8.17. Lying on its belly with its legs
stretched out and flaccid, sensation perfect. 8.24. Limbs paralysed;
pinching the tail makes it look round, and it makes vain attempts to
change its position, pulsations of the heart less frequent and weaker,
It is perfectly conscious, for although unable to move its limbs, it
follows with its eyes the movements of any one approaching it. 8.30.
Heart irregular, two pulsations and a pause; pupils a little con-
tracted (7), sensation unaffected. The heart is now becoming slower
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 107
and slower, the respiration panting. 8.35. The heart only beats
occasionally, sensation seems perfect, it is able to move its head a
little, but the rest of the body is paralyzed. 8.40. Respiration
panting, and forty per minute. 8.45. Paralysis complete, sensation
unaffected, the pulsations of the heart very infrequent, after a con-
siderable interval a few hurried beats succeed each other. Slight
shivering movements in the tail, fingers and toes (spasmodic). 8.47.
The heart has ceased to beat, the pupils are dilated, the lower jaw has
fallen, a few gasping respirations. 8.47. Slight spasmodic move-
ments in the feet. 8.48. Died.
9.4. Opened the body and observed spontaneous twitchings of
the pectoral muscles, which also were excitable by mechanical irrita-
tion. Auricles and great veins very much congested; a little semi-
coagulated blood in right ventricle; a drachm of fluid blood in left
ventricle, which coagulated after removal. For half an hour after
death the application of dilute sulphuric acid or mechanical irritation
produced contractions, especially of the auricles, which from being
dark red became temporarily pale and bloodless; pinching the
substance of the ventricles made them contract, the contraction of
the latter did not extend to the auricles; lungs collapsed and pale;
liver congested, gall-bladder full, vena porta congested, stomach and
alimentary canal healthy.
Exp. II. On a Cat (nearly full-grown).— After applying a
spirituous extract of Zanghinia v. to the conjunctiva in order to
observe its action on the pupil, with a negative result, and having
ascertained the pulsations of the heart to number nearly 120 per
minute, at 12.15 p.m. we injected three grains of the same extract,
mixed with one drachm of water, into the cellular tissue. 12.20.
The animal is able to walk, but its legs seem weak. 12.31. Begins
to vomit. 12.37. The action of the heart weak, pulsations about
110 per minute, the vomiting urgent, bowels moved. 12.43. Vomiting
still continues, great debility, the pulsations of the heart very much
reduced in number and strength, the respirations fewer and panting,
pupil normal. 12.50. The animal is now scarcely able to move
about, it lays its head down upon the ground, and after a little
changes it into a new position, without moving its body. How far
this condition results from paralysis, and how far from pure debility,
it is difficult to say. 12.52. Pulsations of the heart 20 per minute,
and weak, 12.54. Clonic spasms of extremities, with trembling mo-
tion of the skin of the back, expulsion of urine and feces, a sigh, the
pupils dilated, the action of the heart has ceased, a few gasps, and the
animal died at 12.57.
Examination immediately. Lungs pale and collapsed, the coro-
nary veins of the heart full, the substance of the heart congested, all
the chambers contain blood, the auricles however are engorged, the
left one full of bright red and watery blood, the vene cave full,
The liver, vena porta and its branches, are very much congested, the
whole intestinal canal abnormally pale. A mild magneto-electric cur-
108 MR DAVIDSON.
rent produces contractions of the muscles to which it is applied, the
contractions are less powerful when the current is transmitted through
the nerve, slight contractions cau be produced by connecting the two
limbs, the heart does not respond to the magneto-electric stimulus. I
have observed paralysis to be much less distinctly marked in the cat
than in the lemur.
Exp. III. On a species of Civet about the size of a small cat.—
The poison mixed with a little water was injected into the cellular
tissue at 9.24 am. At 9.25, makes violent efforts to vomit. 9.28.
Restless, and vomiting. 9.37. Posterior extremities are paralyzed,
sensation seems unaffected. 9.40. A slight shivering motion all over
its skin; extremities paralyzed, but it is still able to move its head a
little. 9.42. It is now quite unable to move any part of its body.
Died at 9.45.
Examination immediately. Great congestion of liver, kidney,
portal vein, and substance of the heart. Right auricle engorged.
The right auricle was seen to make about two slight contractions
every minute, producing a wave in the blood, filling the vena
cava descendens. These contractions were spontaneous; but after
they ceased they could be excited again by mechanical irritation of
the muscular substance for about half an hour after the organ was
exposed.
Haperiments on Frogs.
The experiments made on frogs were very numerous, and _
every experiment repeated frequently by my assistant, Andri-
analy. ‘l'wo species of frog were used, the one very consider-
ably smaller than the common English frog, Rana Temporaria ;
the other about one-third larger. I shall select a few experi-
ments from my note-book to illustrate some of the points
referred to in the preceding observations.
Exp. IV. To ascertain the effect of the local application of the
poison.—A little of the commen extract was applied to the right leg
of a frog at 9.20 am. At 9.58, the right leg is observed to be pa-
ralyzed, pinching it does not produce any sign of sensation, the left
limb normal. 10.30. The left limb is now paralyzed. 10.40. The
paralysis has extended to the whole body, and is gradually becoming
more marked. The animal died about 11.15.
The two following experiments illustrate the action of Tan-
ghinia on the heart.
Exp. V. At 9 A™M., exposed the heart of a large frog, the pul-
sations were about 52 per minute. 9.5. Injected about one grain of
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. 109
the extract of Zanghinia ven. into the peritoneum. 9.8. Pulsations
52. 9.15. They became reduced to 39. At 9.20, the tetanized
ventricle was seen to contract imperfectly, and at 9.21 the heart
and respiration ceased almost simultaneously; but although the heart
has ceased to beat, the animal made a few leaps. On looking at the
posterior lymphatic hearts, we find they have ceased to beat. 9.28.
The animal still continues to withdraw its legs if they are drawn
out.
Exp. VI. Having exposed the heart of a large frog, and found
the pulsations 42 per minute, at 7.15 we administered one grain of
the extract of Tanghinia v. by the mouth. The action of the poison
on the heart was as follows :—
7.20. The pulsations were 34.
7.25. - F 29.
7.30. 3 be 28.
7.40. 5 as 26, and irregular.
7.50 ss - 24. The ventricle contracts very im-
perfectly.
8. . rs 22. Slght vermicular motions of the
ventricle.
8.10. The ventricle has ceased to beat, the auricles however
make 12 pulsations per minute. 8.20. The auricles have stopped ;
but respiration has not quite ceased, and the animal is still able to
move. The head was now removed, leaving the lower jaw connected
with the body. Reflex movements could be induced by pinching, or
by the application of electricity. The ventricle was contracted, and
the auricles dilated.
Exp. VII. We selected two large frogs of about the same size,
and having exposed their hearts, we administered two grains of the
extract of Tanghinia v. by the mouth to one of them; and in order
to test the action of the poison on the reflex functions of the cord,
suspended them by the lower jaw, and observed the length of time
which elapsed between the application of dilute sulphuric acid and
the appearance of the corresponding reflex movements. We found the
reflex movements to be less energetic, and slower in appearing in the
poisoned frog ; and after about an hour, scarcely any irritation could
induce reflex movements ; yet after death we found the muscles to
contract, when magneto-electricity was applied, although less actively,
and for a shorter time than the muscles of the non-poisoned frog. We
observed in this same experiment paralysis of the lower jaw, which
appeared almost immediately after the acministration of the poison,
and was probably owing to its topical action on the muscles and
nerves. The pupils became much dilated within ten minutes after the
poison was given, and continued at least double the size of the pupils
of the non-poisoned frog. The posterior lymphatic hearts were also
observed in this experiment, inasmuch as they receive their nervous
supply from the cord. In the non-poisoned frog, the posterior lym-
110 MR DAVIDSON,
phatic hearts continued to beat throughout the experiment at from
48 to 50 per minute, and the pulsations were distinct, strong, and
- regular. In the poisoned frog, they very soon rose in number: ten
minutes after the administration of the poison they had risen to 63 ;
after half an hour they were 77, and weak, and they gradually be-
came weaker and more irregular, and after the lapse of an hour were
quite imperceptible.
There is very great difficulty in deciding to what extent the
conductivity of the motor nerves is affected, inasmuch as the
poison affects the contractility of the muscles, and this compli-
cates the matter very considerably. We constantly observe
voluntary motion remarkably affected before the reflex functions
of the cord are much diminished, but this loss of voluntary mo-
tion may of course be due to other causes than the mere loss of
motor conductivity. Although I do not consider it proved, I
think it probable that the helplessness and dragging of the
limbs are partly owing to poisoning of the muscular tissue, and
partly to paralysis of the motor nerves. It seems impossible to
doubt that the irritability of the nerves and contractility of the
muscles are lessened, and they are thus less able to respond to
the influence of volition, while, on the other hand, this manifes-
tation of reflex movements in the decapitated animal poisoned
by Tanghinia proves that the muscles nevertheless retain enough
of irritability, and the nerves enough of conductivity, to make it
impossible to ascribe the paralysis entirely to these two causes.
Admitting then these conditions as existing, I am inclined to
think that the anterior columns of the spinal cord are also
implicated, and that the paralysis, as I have already said, is
partly due to this cause.
Exp. VIII. We decapitated a small frog in order to test the
length of time that reflex movements may be excited by mechanical
and chemical stimuli. 9.11. The frog was beheaded. 9.16. It
draws up its hind legs if disturbed. 9.18. Makes attempts at leap-
ing. 9.22. Pinching the foot produces movements in all the extre-
mities. 9.31. Applied some dilute acid to the skin, and strong reflex
movements followed. 9.40. Reflex movements cannot be any longer
induced. In this experiment reflex movements persisted for nearly
half an hour. A frog of similar size had its heart exposed, and a
grain of Tanghinia injected beneath the skin of left leg. The pulsa-
tions of the heart rose from 80 to 100, and in two minutes stopped.
The posterior lymphatic hearts stopped at the same time as the blood.
heart. We now cut off its head. Touching the eye induced pro-
AN ACCOUNT OF THE MADAGASCAR ORDEAL POISON. I1]
tective motions of the eyelids. Pinching the feet did not bring on
any reflex motion, but acetic acid applied to the fore-legs did. Sia
minutes after the heart ceased to beat, all reflex action ceased.
In repeated experiments in which the amputated limbs of
frogs have been immersed in a mixture of Tanghinia the con-
tractility of the muscles has been lessened in a very few minutes,
and after a somewhat longer time abolished.
Exp. IX. We selected a large active frog, and tied the vessels of
the left posterior extremity, and then poisoned it with extract of
Tanghinia given by the mouth. The muscles of the limb so protected
were found to contract much more energetically than those of the
other, and that whether the stimulus was applied through the sciatic
nerve or to the muscular substance itself.
Several experiments were made to ascertain whether de-
struction of the medulla would affect the action of the poison
on the heart. In one case in which we decapitated a frog, and
then injected one grain of the extract into the peritoneum, the
heart continued to contract for twelve minutes after the injection
of the poison, and dilute sulphuric acid induced reflex move-
ments for some time after the heart was arrested. We may
conclude that Tanghinia does not arrest the heart through any
action on or through the vagus.
General Conclusions.
(a) The Tangena must be classed among the cardiac poisons.
It uniformly causes death by arresting the action of the heart.
(b) It does not act on the heart through the vagus nerve.
When applied to the exposed heart its rapidity of action is re-
markable. The fact that it arrests the pulsations of the ex-
cised heart of the frog is conclusive proof that its influence,
when topically applied, is direct, either on the muscular sub-
stance, or the muscular substance and cardiac ganglia.
(c) There is sufficient reason to believe that the Tangéna
acts on the spinal cord, producing paralysis and diminishing
reflex action.
(dq) Voluntary motion is abolished, and the irritability of
the motor nerves lessened by the poison, When it acts through
112 . MR DAVIDSON. MADAGASCAR ORDEAL POISON. -
the circulation in mammalia, sensation is not remarkably af-
fected ; muscular contractility is very much diminished. More
exact knowledge of the degree and order in which these various
functions are affected, can only be obtained by carefully per-
formed experiments made in Europe, where the more delicate
electrical instruments can be had.
(e) It is exceedingly fatal to man, in doses of thirty grains
of the kernel, if not promptly ejected.
(f) It causes a numb, tingling sensation in the part with
which it comes into contact, and also throughout the body.
(g) It is powerfwly emetic and purgative, produces great
nausea and debility, paralysis of motion, occasionally delirium,
narcotism, and perhaps vertigo.
(hk) It may be inferred to cause death in man, as in all
other animals, by tetanizing the heart.
NOTE by Dr Gatasin, in continuation of his Paper (p. 22).
Yet another has been added to the already numerous theories of
the dicrotic wave by Mr Mahomed, in a paper lately published in the
Medical Times. It is the more deserving of notice since it is to the
able researches of its author that we owe the most recent contribu-
tions to our knowledge as to the clinical use of the sphygmograph.
Mr Mahomed then finds a sufficient cause for the dicrotic wave in
the mere fact that the coat of the aorta is elastic, and considers that it
originates from the contraction of this elastic coat during diastole. It
appears to me that this theory involves a misconception of the nature
of elasticity. If a surface is said to be elastic, nothing more is
meant than that it is extensible in such a way that the degree of its
extension has a definite relation to the forces extending it. Its con-
traction is not an active proceeding, but is merely the effect of the
diminution of tension. Thus the contraction of the aorta is the
consequence of the diminished pressure within, and therefore the
mere fact that it contracts cannot at the same time be a cause of an
increase of that pressure originating a second wave.
CASE OF SUBDIVISION OF THE SCAPHOID CAR-
PAL BONE. By Jonny Srrutuers, M.D., Professor of
Anatomy in the University of Aberdeen.
THIS condition was found in the right hand of a male subject,
aged 68. My attention was called to it just after the igaments
of the first carpal row had been divided. The scaphoid is
represented by two bones of nearly equal size. The lower or
radial division carries about a fourth part, in length, of the
articular surface for the radius, less than half of the articular
hollow for the os magnum, and the whole of the surface for the
trapezium and trapezoid bones. The direction of the articula-
tion between the two bones is nearly at right angles to the
general axis of the scaphoid. The surfaces are nearly flat from
above downwards (i.e. in the direction between the surfaces for
the radius and the magnum) allowing free gliding motion of the
two bones on each other in that direction, but in the opposite
direction motion is impeded by a little undulation of the sur-
faces. This articulation between the two bones occupies the
entire thickness of each, and is continuous across the whole
breadth with the synovial cavities of the surfaces for the radius
and magnum.
So far the case looked like one of natural variation, but
I was led to doubt this by the following considerations. The
articular surfaces between the two bones, though polished and
synovial looking, are hard, the knife turning up no articular
cartilage till it reaches the edge of the other articular surfaces ;
and there are irregular depressions on parts of the polished
surface, more distinctly seen when examined with a magni-
fying power. The trapezoid and os magnum, and the second
and third metacarpals, where they meet at the carpo-metacarpal
articulation, shew some bony excrescence on the dorsal aspect ;
and the shafts of the second and third metacarpals, especially
that of the second, are altered above their middle, being
elevated on the dorsal surface where they begin to be left
uncovered by the dorsal interossei muscles, and having an
VOL. VIII. 8
114 PROFESSOR STRUTHERS.
unusually projecting border on the palmar aspect. These two
metacarpals, and the carpus when they join it, appear to have
suffered some injury and to have undergone subsequent change.
The scaphoid bone is naturally so placed that it would receive
the shock of an injury transmitted from these two metacarpals
through the moving trapezoid and magnum, and would be most
likely to give way about the middle where it presents a kind of
constriction or neck. My inference is that this has happened
here, and that this case is not one of natural variation, but one
of fracture of the scaphoid followed by the formation of a false
joint. It might, however, readily have been mistaken for a
case of variation, especially if the carpus had been already
macerated, and the attention of observers may be called to
the importance of looking narrowly to see whether some of
these cases of additional carpal or tarsal ossicle have not had
their origin in fracture. It should be added, that in this sub-
ject the left hand and both feet were normal and healthy.
ACCOUNT OF RUDIMENTARY FINGER MUSCLES
FOUND IN A TOOTHED WHALE (HYPEROODON
BIDENS). By Joun Srrutuers, M.D., Professor of
Anatomy in the University of Aberdeen.
It has been supposed that muscles passing from the forearm to
the hand do not exist in any of the Delphinoid Cetacea. In a
foot-note on page 115 of my paper “On some points in the
Anatomy of a Great Fin-Whale” (Balzenoptera musculus), pub-
lished in this Journal for November 1871, I noticed briefly that
I had found that such muscles are present in Hyperoodon
bidens, a specimen of which had just been stranded on our coast.
Having since dissected them more fully, and examined them
also in the other paddle, I am now able to give an account of
their arrangement. In order to save repetition, I shall take
the muscles in the great Finner as a standard for comparison,
and shall note the differences between them and the correspond-
RUDIMENTARY FINGER MUSCLES IN A TOOTHED WHALE. 115
ing muscles in Hyperoodon; and beg to refer to my detailed
account, and to the drawings given, of these muscles in the
paper above referred to.
These finger muscles, so far from being absent, are nearly
all better developed in this toothed whale than in the great
Finner.
(a) Internal, or palmar, aspect. 1. Flexor carpi ulnaris.
The differences in this muscle are that implied in the different
form of the cartilaginous olecranon, and that the tendon is flat
and not placed so far from the shaft of the ulna. The cartila-
ginous olecranon is long, tapering, and curved backwards, and
the origin of the muscle is continued to the point, running
along the concave edge and some way on the inner surface of
the cartilage. The upper convex edge of the cartilage gives
insertion to the greater head of the triceps, though not back to
the point. A muscular expansion from the trunk is inserted
into the cartilage at its point and for some way along the inner
aspect of the insertion of the triceps. The cartilage is very
flexible inwards and outwards, from its flatness, and will be
raised a little by the action of the triceps and depressed a, little
by the flexor carpi ulnaris. The insertion, as in the Finner, is
entirely into the pisiform cartilage. This cartilage moves
freely laterally, but has very little longitudinal mobility. It
articulates distally with the fifth metacarpal bone as well as
with the epiphysis of the ulna and the neighbouring carpal
bone. The presence of this muscle cannot be satisfactorily
accounted for on the theory of action upon its insertion or upon
its origin.
2. Flexor digitorum ulnaris. The chief differences in this
muscle are, that it is considerably more developed than in the
Finner, that it lies more upon the ulna without being sunk into
the interosseous hollow, and that the tendon retains its thick-
ness instead of expanding before it divides. The fleshy fibres
of origin reach farther forwards on the humerus, as far as to
touch and even to pass a little under cover of the teres major ;
and further across the ulna so as to reach the flexor carpi
ulnaris, the two bellies being in contact by their edges, the
nerve disappearing between them, The belly lies upon the
8—2
116 PROFESSOR STRUTHERS.
proximal half of the ulna, the tendon obliquely across the
distal half. The breaking up is just as it is getting upon the
first carpal row, and is into three tendons. The ulnar tendon,
after a course of an inch, divides for digits Iv. and v.; the
radial tendon, after nearly an inch, is joined by the tendon of
the radial flexor.
3. Flexor digitorum radialis. This muscle, though less
developed in Hyperoodon compared with the last muscle, is as
well developed in proportion to the limb as it is in the Finner.
Its fibres arise not only from the radius and interosseous tissue,
but from the ulna fully as much as from the radius. It is sunk
in the interosseous space. The tendon throws itself entirely
into the radial tendon of the flexor ulnaris.
The tendons of these two muscles are disposed thus. The
radial tendon, one-third of which is formed by the entire
tendon of the radial flexor, is fully twice the size of the tendon
of any other digit. It goes almost entirely to digit 11, a few
fibres going off to blend with the terminal ligament which
connects the end of digit I. to the edge of the second bone of
digit 11. Its earliest insertion is into the metacarpo-phalangeal
cartilage and broadly into the first phalanx, these insertions
maintaining the previous obliquity of the tendon. Digit IL,
though but little longer than digit I11.,is the most robust of the
digits. The middle tendon goes straight on to digit m1. The
radial subdivision of the ulnar tendon goes on to digit Iv., the
ulnar subdivision, the larger of the two, goes very obliquely
towards digit v., and is broadly inserted into the first phalanx,
which is mainly cartilaginous. The obliquity is maintained by
this earlier part of the insertion.
(b) Hxtensor, or dorsal, aspect. In the Finner there is
simply one common extensor, giving a tendon to each of the
four digits. Here there is at least one other extensor, and,
proportionately to the limb, a considerably greater bulk of
muscle. There is the general division of the mass into a radial
and an ulnar extensor, the latter somewhat complex. The
fleshy fibres of both arise as high as the ligament of the elbow.
The fleshy bellies occupy the proximal two-thirds of the fore-
arm. That of the radial flexor reaches farthest back, is larger
RUDIMENTARY FINGER MUSCLES IN A TOOTHED WHALE. 117
by a third than the other, arises by its distal portion from the
ulna as well as from the radius and interosseous tissue, and is
mostly sunk into the interosseous hollow. The ulnar belly
arises from the ulna and interosseous tissue, and both bellies
have fibres from an intermuscular septum between them, and
from a strong fascia over them.
The tendon of the radial extensor divides opposite the first
carpal row into two, one straight on to digit 11, the other, the
larger, to digit 1. The latter gives from its radial side a slip as
if for digit L, but it is inserted into the distal part of the first
joint of digit 11, very nearly reaching the terminal ligament of
digit I.
The ulnar extensor divides above the middle of the forearm
into two fleshy bundles, from which tendons soon proceed, the
most ulnar tendon beginning first. So far, these might be
looked on as representing separate muscles, if the tendons bore
out that view. The tendon of the radial portion passes to
digit Iv., adhering to the carpus on its way. The tendon of
the ulnar portion after giving off a slip, one-third of its bulk,
which joins the tendon of digit Iv., opposite the second carpal
row, passes on to digit v., and is inserted into the distal end of
the metacarpal bone and into the phalanx beyond.
These details apply to both sides, but in the right paddle
there is an intermediate fleshy slip, leaving the ulnar side of
the radial flexor, soon ending in a tendon which divides at the
carpus into two (adhering to the back of the carpus on its way),
one joining the tendon of digit 111., the other joming that of
digit Iv. The radial extensor thus, on this side, sends a tendon
to digit Iv. as well as to 1. and 1. A farther want of sym-
metry is seen in the presence, also on this side, of a small ten-
dinous slip from the radial portion of the ulnar flexor to join
the slip which the ulnar portion of that muscle gives to the
tendon of digit Iv.
Viewing these extensor muscles homologically, it might be
held that the extensors of the carpus, at least on the ulnar
side, are represented by the adhesions to the carpus and to the
fifth metacarpal bone ; but it may be sufficiently comprehensive
to regard them as representing, the radial, the internal common
extensor of the quadruped, the extensor communis digitorum
118 PROFESSOR STRUTHERS.
of man; and the ulnar, as representing the external common
extensor of the quadruped, the extensor minimi digiti of man.
The difference on the two sides illustrates the tendency to va-
riation in rudimentary structures.
The extensors on the whole appear to be somewhat less
powerful than the flexors, but the difference between the flexor
and extensor powers is not so great as in the Finner. In ac-
cordance with the greater development of these muscles in
Hyperoodon, the finger-joints are more moveable than in the
Finner, the cavity reaching completely across between the car-
tilages, but the surfaces are quite flat. The same remark ap-
plies to the carpo-metacarpal joints and to those of the carpal
cartilages, but to a less extent, the mobility at the joints in-
creasing distally. To the merely passive resistance which a
purely ligamentous condition would afford, these muscles will
add some activity; but, though somewhat stronger than in the
Finner, they are small and feeble relatively to the size and
condition of the parts on which they act; in striking contrast
with their fully developed neighbours, the muscles of the
shoulder.
Rudimentary Brachial Muscles in Hyperoodon. In both
paddles alike there is a short thick muscular and fibrous mass
lying, like a cushion, along the lower border of the distal half
of the humerus. It begins on the humerus just beyond the
deltoid elevation, and terminates just beyond the elbow on the
lower edge of the radius. It is about as thick as a fore-finger,
flat next the humerus, rounded on the opposite aspect. It is
largely mixed with fibrous tissue. There is some slight lateral
mobility at the elbow, but none in the direction in which this
muscle would act. Along the opposite border of the humerus
lies a rudiment of the external short head of the triceps ex-
tensor cubiti muscle, passing from the border of the humerus
to the early part of the olecranon. It is about as thick as a
finger, 1} inch in length, and very largely composed of fibrous
tissue. The nerve and artery are seen to pass between it and
the humerus, from the flexor aspect, and some tissue situated
below this passage may be considered a remnant of the third
head of the triceps. Under the microscope the reddish tissue
aimee
RUDIMENTARY FINGER MUSCLES IN A TOOTHED WHALE. 119
contained in these rudimentary brachial muscles is seen to be
composed of well-striped good-sized muscular fibre.
The scapular head of the triceps is reinforced by a bundle
from the teres, forming the distal part of the insertion into the
olecranon. The limb having been removed from the trunk, it
is now impossible to decide how far the latissimus dorsi may
form part of what I have termed the teres major.
Size, age, sex, &c. This Hyperoodon was stranded, alive, at
Fraserburgh, on the Aberdeenshire coast, on August 17, 1871.
Length, along the curve of the back, 20 feet 9 inches; along the
side, in contact, 19 feet 9 in.; in a straight line, 19 feet 3 in.
Measurements of the paddle, in inches—length, from tip to
axilla, 15; froni tip to shoulder joint, 25, of which the humerus
has 7}, the forearm 6}, the carpus 2, and the longest digit 9.
Breadth, at axilla 64, at broadest part 7. Girth, at axilla 15, at
broadest. part 15$. Compared with the great Finner (B. mus-
culus), the paddle is shorter in Hyperoodon in proportion to the
size of the animal; and comparing the segments of the respec-
tive paddles, Hyperoodon has, in proportion, a longer humerus,
a considerably shorter forearm, and fingers of very nearly the
same length. It was believed, when lying on the beach, to be
a male, but circumstances prevented me from seeing the viscera
afterwards. The maxillary crests are about five inches apart at
the middle, and about three inches thick. The epiphyses of
the bodies of the moveable vertebre are all separate. The rudi-
mentary teeth, one on each side, near the front of the lower
jaw, are seen, in the preparation which I have preserved, to be
buried half an inch below the surface of the dense gum.
TISSUE METABOLISM, OR THE ARTIFICIAL INDUC-
TION OF STRUCTURAL CHANGES IN LIVING
ORGANISMS. By W. Arysiie Hotuts, M.D., Cantab.,
Part III, Annulata continued.
In two previous papers I have given the results of some
experiments made by me upon actinie and lumbrici; and I
have there shewn that the tissues of such creatures are readily
acted upon by caustic irritants (such as blistering fluid, strong
acetic acid, &c.), and that a definite sequence of phenomena is
thereby induced, which, in many respects, is similar for these
two classes of animals. I shall in the following pages give a
short résumé of some further experiments undertaken with a
view to extend our knowledge of ‘tissue metabolism’ to other —
branches of the animal kingdom than those mentioned above.
The first observations were made by me upon the medicinal
leech (H. medicinalis), and are detailed below.
I. The action of certain irritants wpon leeches.
Blistering fluid and acetic acid appear to act similarly
when they are applied to the integuments of leeches. Either
of these vesicants induces, immediately after its application,
great swelling of the tissues, and a greater or less loss of the
power of contraction at the injured part according to the
extent of the injury. These phenomena are quickly followed
by a copious secretion of mucus of a bluish white colour, and
by the infiltration of the tissues with blood-plasma. In the
course of an hour or less ecchymoses with intense congestion of
the superficial blood-vessels are observable. The swelling of
the parts with the vascular congestion usually lasts for twenty-
four hours, subsequently contraction of the injured tissues
takes place with puckering and the permanent loss of their
contractility. During the second stage of symptoms detailed
above, and before their fina] contraction, the tissues become
softer than natural and are more easily torn. The mucous
fluid exuded during the second stage consists of a viscid
DR HOLLIS. TISSUE METABOLISM. TZ
hyaline plasma with numerous corpuscular, granular and
pigmentary elements intermingled with it. The first of these
bodies are about the size of human leucocytes, of oval shape
(occasionally almost fibrillar); and they possibly possess
amceboid movements. In two experiments, wherein blistering
fluid was used, a distinct bulla was produced at the point of
vesication; in one case this blister was filled with a colourless
serum, in the second with a fluid of a reddish colour, and
containing the corpuscular elements above described.
If we compare the above phenomena with those observable
under like conditions in the structures of the actiniz and
lumbrici, we shall find a considerable similarity in them all, a
resemblance which leads me to conclude that the processes in
each instance are almost identical. I shall not at this stage
recapitulate the various phenomena noticeable in these different
animals, as they have been elsewhere detailed, and I shall have
occasion to refer to them again at a future period.
II. The application of the actual cautery to a leech.
When a part of the dorsal surface of a leech is placed
momentarily in contact with the end of a glass rod heated to
redness, the burn instantly becomes of a bluish white colour and
its surface is depressed. About ten minutes after the injury is
made, the surrounding parts begin to swell, and subsequently
mucus to a small extent is exuded from them. Within twenty-
four hours the burnt tissues are covered with a shreddy bluish
white débris, easily detached, and leaving the surface much
depressed. The surrounding tissues at the same time become
puckered and drawn towards the seat of injury. No bulla
appears to be formed in this case; nor is there apparently any
great exudation of serous fluid. It is necessary to notice these
particulars, as I shall hereafter refer again to them.
ARTHROPODA AND MOLLUSGCA.
My experiments upon several species of these two sub-
kingdoms have been attended with no marked results. I have
on various occasions tried the effect of irritants and the actual
cautery upon members of the class arachnida; but, owing
to the thickness of their chitinous integument, the ordinary
122 DR HOLLIS.
irritants appeared to have little effect upon them, whilst more
- powerful corrosives or the actual cautery destroyed life without
inducing any of the phenomena noticed in the other classes
of animals. Like results have followed my attempts to induce
metabolic processes in various insects. The larve of the
lepidoptera (upon which most of my experiments were per-
formed), although manifesting by their movements that blister-
ing fluid, nitric acid and other irritants were local sources of
uneasiness to them when applied to their chitinous investment,
failed entirely to manifest any of those palpable changes of
structure noticed by me in the annulata, although they
sometimes rapidly died after the operation.
Slugs, snails and other gasteropods, upon which I have
experimented as in the preceding cases, appear to a great extent
to be protected from such possible sources of injury by the
rapidity with which a large quantity of mucus is evolved from
the affected surface. This flow of mucus carries away with it
the local source of irritation. When the actual cautery was
applied for a moment to the foot of a snail a like evolution of
mucus took place at the point of contact, and a whitish eschar
was visible; but on the following day this local change had
been removed with a large quantity of mucus, and was entirely
separated from the animal, whilst the surface of the mollusc
appeared in all respects natural.
VERTEBRATA. AMPHIBIA.
In dealing with the subject of ‘tissue metabolism’ among
the higher animals, it will be necessary to encroach somewhat
upon the labours of others in a similar direction, notably on
those of Ryneck, Stricker and Cohnheim, in their researches
upon the process of inflammation. I trust, however, that the
importance of the subject, and a certain novelty in the method
of performing the experiments, may plead an excuse for pur-
suing a path already trodden by such able physiologists.
The experiments I shall now detail were performed upon
the common smooth newt (Lissotriton punctatus).
I. The application of certain irritants to the abscised tail of
a newt. When the tail of a newt is freshly removed from the
4
TISSUE METABOLISM. 123
body it will under favourable conditions continue to give signs
of life by occasional movements for eight-and-forty hours and
upwards. It therefore appeared to me to be a favourable
object upon which to pursue my experiments on ‘tissue meta-
bolism.’ With this view I applied blistering fluid to the recently
abscised tail from a decapitated newt, and set it aside in a small
quantity of water for a few minutes. The part vesicated
was then of a bluish white colour and covered with a copious
exudation of mucus. In two hours time the portion of the
tail untouched by the Liquor Epispasticus contracted readily
on the application of a slight stimulus, while the rest of it
remained uncontracted. Twenty-four hours subsequently several
bullze were observable on the injured portion, and these con-
tained a fluid and a minutely corpuscular plasma. The cutis
was easily detached from the subjacent structures at the seat of
injury, and these were found to be swollen and infiltrated
with a watery plasma containing numerous granules and a few
leucocytes. I have since repeated this experiment with similar
results. If all movement has ceased in the tail before the
application of the blistering fluid, beyond the whitening of
the integument, no further phenomenon takes place. This I
think clearly shews that the exudation of plasma and the other
concurrent symptoms are due to the active vital properties of
the tissues. Now the structures in a newt’s tail after its
removal from the body appear to me to approach closely in
their constitution to the whole body of those invertebrate
annelids we have lately been considering, with this difference in
favour of the greater heterogeneity of the latter, namely, that
they possess a definite circulatory fluid driven through their
tissues by appropriate pulsatile organs, whilst the detached
caudal extremity of the newt has no such provision for its
general tissue nutriment. In such cases the various phenomena
observed after irritation must, in the first place, I think, be
considered entirely due to the local action of the irritant on the
individual living elements of a part; for it is difficult to sup-
pose that either the nerves or blood-vessels, owing to the
absence of the blood, can exert any more than a very slight
influence over the tissue-changes that take place. If at the
same time that an irritant is applied to the abscised tail of a
124 DR HOLLIS.
newt a similar injury is induced on the tail of a healthy newt,
the various phenomena detailed above will correspond in each
case—although in the latter the secondary effects on the
blood-circulation will be evidenced in the well-known phe-
nomena accompanying its stasis.
Il. The effects of a burn on the abscised tail of a newt.
When the end of a red-hot glass rod is momentarily applied
to the integuments of a recently abscised tail the burn becomes
depressed and whitish, while the adjacent tissues are rapidly
elevated and covered with a bluish white débris—as a rule no
bullae are formed in such a case—and this corresponds with
what I have observed to take place in leeches. In twenty-four
hours after the injury the burnt surface is covered with a ragged
epithelial and granular débris and the surface is still much
depressed ; the surrounding tissues have usually resumed their
natural shape, and no further phenomena are observable. This
cessation of further changes in the part is probably due to the
greater shock of the cautery to the tissues themselves, and the
more rapid death of the tail generally than is the case when
the blistering fluid is used.
Ill. The application of blistering fluid to the excised heart
of a newt. Upon touching the ventricular apex of a heart still
pulsating, and freshly removed from the body of a newt, with
about ;4,th of a drop of Liquor Epispasticus, contained in a
capillary tube, there was an immediate pallor and loss of con-
tractility in the part, and subsequently a slight swelling. The
loss of contractile power was very manifest in this case, as the
seat of injury was sharply defined by the change in colour. The
remainder of the organ continued its usually rhythmical pulsa-
tions, whilst the pallid portion was uncontracted after the
application of the vesicant.
General Remarks. If we exclude the arthropoda, and in
some cases the molluscs, from these observations for reasons
elsewhere stated, these experiments tend to shew that there is
a general correspondence throughout the animal kingdom in the
sequence of phenomena observable after the application of an
irritant to a living surface. These may be briefly summarised as
below :
TISSUE METABOLISM. 125
1, Swelling and loss of contractility. 2. Exudation of fluid
(muciform or serous). 3. Elevation of the external integument
with the production of a bulla, containing corpuscular elements
of various shapes. (This phenomenon is only observable where
there is a distinct epiderm, and occasionally not then.) 4. In-
filtration of the subjacent tissues with fluid plasma and their
subsequent disintegration, when the cause of irritation is pro-
longed. 5. Subsequent contraction of the subtegumentary
tissues and permanent loss of their contractility, should the
injury extend to them.
What then do these phenomena imply?) Dr Ryneck* has
shewn—by first removing the blood from the vessels in the web
of a frog and “then subjecting their internal surfaces for a few
moments to an agent, which, by virtue of its chemical action,
might be expected to modify or destroy its vitality ; and finally,
after replacing the injurious liquid by milk or defibrinated
blood, observing the effects of local irritation”—that no stasis
is produced in webs which were thus treated, while the pheno-
mena of stasis occurred in vessels after local irritation, even
when their natural contents were replaced by milk. Such
“results seem to make it perfectly clear that the local changes
which lead to the production of stasis must have their seat
either in the walls of the vessels or in the tissues which im-
mediately surround them.”
Upwards of twenty years ago H. Weber® shewed that even
after ligature of the thigh of a frog the blood gathers from all
sides to the irritated part of the web, when ammonia is applied
to it; and quite recently Cohnheim*® has shewn that the
opinions of Hering and Schlarewsky, who ascribed “the phe-
nomenon of extravasation to the slow filtration of a colloid
substance through the walls of the vessels,” is incorrect, by
placing an animal in the required conditions for physical
transudation, when none of the phenomena which he (Cohn-
heim) has described elsewhere will take place. “ Not a single
1 Zur Kenntniss d. Stase des Blutes in d. Gefassen entziindeter Theile, in
Rollett’s Unters. a. d. Instit. f. Phys. u. Hist. in Graz. 1870, p. 103, noticed in
Holmes’ System of Surgery, 2 ed. vy. p. 757. (The process of inflammation by
Sanderson.)
2 Miiller’s Arch., 1852, p. 361. (Holmes, Op. Cit. v. p. 763.)
3 See article in Brit. Med. Journ. Sept. 27, 1873, on Cohnheim’s paper, Neue
Untersuch. ti, die Entziindung, Berlin, 1873.
- 126 DR HOLLIS. TISSUE METABOLISM.
red or white corpuscle passes through the walls of the vessels,
in spite of the walls of the veins being lined with white cor-
puscles, and of the increased blood-pressure in the capillaries.”
In summing up the results of his observation, Cohnheim be-
lieves that the essence of inflammation consists in some local
change in the walls of the vessels of the affected part, by which -
the extravasation of the corpuscular elements of the blood can
take place. What this change is, he is however unable to say,
as he has not succeeded in finding any structural alteration.
The results of my own observations lead me to believe
that the effect of irritants of all kinds on living structures is to
diminish or destroy their vitality, and with this their tonicity;
in the case of the epiderm, the destruction of tissue leads to its
removal from the body as a dead substance, and its subsequent
renewal by the subjacent structures without modification of its
original form. When, however, the deeper structures are in-
volved in the destructive process, they are never replaced,
except as lowly organised cicatricial tissue. This tissue never
inherits the voluntary contractile power of the muscle it fre-
quently represents, nor does it actually equal in volume the
original structures, for a permanent depression always marks
the place of injury.
It seems that we must look upon the “ essence of inflamma-
tion” to be a weakening or loss of vitality in the tissue-web of
a part, and that the changes which take place in the capillaries,
their dilatation and the permeability of their coats, are due to
this loss of tonicity in their walls, and herein their parietal
elements only share with the muscles and the nerves (if any)
the same general change in their conditions, which produces on
the one hand loss of contractile power, on the other loss of
sensation.
If we enquire the prime cause of these functional changes,
I think it will hereafter be found to be due to a molecular
change in the tissue-elements induced by the modified con-
ditions of their environment, whether such modifications be
brought about by mechanical, chemical, or purely physical
methods; but to this question we have at present no means of
replying satisfactorily.
ON THE MECHANISM OF OPENING AND CLOSING
THE EUSTACHIAN TUBE. By C. J. F. Yuts, BA.,
Scholar of St John’s College, Cambridge, and Fellow elect
of Magdalen College, Oxford.
(From the Physiological Laboratory in the University of
Cambridge.)
Dr Toynbee was the first, I believe, to state the proposition
that the Eustachian tube is normally closed, and that it is
opened only during the action of swallowing. Up to his time
the reverse was believed to be the case. The chief experiment
which he adduced in support of his view was the old one of
swallowing with the mouth and nostrils closed; this, as is
easily felt, produces a distressed feeling in the ears, which he
described as a “fulness or distension.” He proceeds to say,
“this sensation arises from the air which is slightly compressed
in the fauces, passing into and distending the tympanic cavities;
upon removing the hand from the nose it will be observed that
the feeling of pressure in the ears does not disappear, but
remains until the act of deglutition is again performed while the
nose is not closed.” This observation is very pertinent and
valuable. Its value, moreover, is not lessened by the mistake
into which Toynbee fell, as pointed out by Dr Jago, of sup-
posing that in the experiment air was forced znto the tympanic
cavity, while it is in reality sucked out. The fact remains the
same, namely, that to cause any flow of air out of the tube
considerable foree has to be used; and also that when an
adequate effort is made, the action takes place suddenly, as
if an obstacle to its flow had been burst open, and not gradually
overcome, as would be the case if the tube were patent. Now
the tympanic membrane is very sensitive to distension, as
anybody may convince himself by blowing or sucking ever so
slightly, while the Eustachian tube is kept patent with a cathe-
ter adapted for the purpose. It cannot, therefore, be objected
to Toynbee’s experiment that the air is all the time passing
out through a patent tube, but that the unpleasant sensations
128 MR YULE.
are only felt when the pressure on the tympanic membrane
reaches a certain maximum, and that the attainment of this
maximum is mistaken for the opening of the tube. Toynbee
also describes the inverse experiment, where the pressure on
the outside of the tympanic membrane is greater than within
it, as during a descent in a diving bell. In both these expe-
riments the inequality of pressure is removed by swallowing,
owing to the fact that the muscles which open the Eustachian
tube are also implicated in swallowing; and, moreover, that
after the effort of swallowing the closure of the tube takes
place without effort. Ifa slight modification of the first expe-
riment be adopted, and air be blown into the tympanum instead
of sucked out, considerable pressure can be resisted by the
Eustachian tube before the air finds a passage. The amount
of this resistance I have endeavoured to ascertain by various
methods—first, the injection of fluids; this was performed upon a
patient suffering from a large perforation of the tympanic mem-
brane, but possessing at the same time a healthy Eustachian
tube. The apparatus used was an ordinary ear-syringe with
T-piece and manometer with mercury in the bend, introduced
before the nozzle. The results obtained were nearly valueless.
A dilute solution of potassium carbonate passed freely at a
pressure of four inches of mercury, while a solution of alum so
acted upon the tube as to constrict it and prevent the passage
of all fluids. After injecting the potassium carbonate solu-
tion, blowing into the ear failed to force a passage, although the
pressure was raised as high as seemed to be compatible with
the safety of the fenestre. At the same time that I was
blowing, air passed into my own tympanum from the guttural
orifice; so that this experiment may shew either that there is
a valvular action in some part of the tube, allowing passage to
take place more freely from the throat to the tympanum than
in the reverse direction; or that there was some unsuspected
disease of the patient operated on. Of these two alternatives
I think the former is the more probable, because I find it cer-
tainly easier to distend than to exhaust the tympanum.
Finally, as a crucial experiment, to convince myself of the
closure of the tube, I had a catheter passed into my own
Eustachian tube, in order to compare the sensations felt in the
_— eS 7,
OPENING AND SHUTTING OF THE EUSTACHIAN TUBE. 129
normal ear with those in the catheterized one. This was
kindly done for me by Mr Hinton, to whom, for his courtesy
and patience, I desire to express my thanks. I then found, as
Dr Jago has suggested, that sounds produced in the larynx
were rendered very much louder, indeed, so much so, that on
humming a note just loud enough to be heard by a person
standing close to me, the noise produced appeared as powerful
as the loudest notes of my own voice; while by singing loud
the noise was painfully intense, and closely resembled the
sounds heard when a person puts his mouth near the external
meatus and sings. The catheter used was an ordinary Eusta-
chian tube gum one, with a hole cut in the outer side of the
knee. As it was impossible on account of this hole to percuss
the tympanum in the usual manner, several failures resulted,
the open end of the catheter becoming blocked by mucus, but
by passing a catgut bougie along the catheter, and for about
a quarter of an inch beyond its terminal orifice, this obstacle
was removed. My attention was first directed to the question
of the opening and closure of the Eustachian tube by a faculty
of which I had been long aware, that while singing I could, by
an effort of some faucial muscles, cause the sound to appear
very much louder, in a manner which I have compared to the
“swell” arrangement of an organ. The increase in apparent
loudness was very marked indeed. It naturally occurred to me
that this was due to the opening of the Eustachian tube, and
this idea was afterwards confirmed by the following experi-
ments :—First, when the tympanic cavity was blown full of air
and the membrane sensibly distended, the air which was
retained by the natural closure of the Eustachian tube, at once
escaped on the contraction of these muscles. Second, the effect
of passing a catheter of the form before described into the
Eustachian tube, was to produce a modification of hearing
exactly identical in every respect with that observed when the
contraction of these faucial muscles takes place.
That the Eustachian tube opens during the act of swallow-
ing has been long admitted, but owing to the extreme compli-
cation of the act, no observer has yet been able to ascertain
clearly which the efficient salpyngeal muscles are. By a careful
examination, however, of my own pharynx during the opening
VOL, VIIL, )
130 ' MR. YULE.
of the Eustachian tube, when not complicated by swallowing,
their action is easily understood. In this matter Dr Durham
of Guy’s Hospital has been good enough to assist me, and
I wish to express my obligation to him for his kindness and
material assistance during the laryngoscopic examinations ne-
cessary. It is noticed during the contraction for opening
the tube: First, that the velum palati does not change
either its position or shape, in fact, that it remains unmoyed ;
and further that it does not become tense, but hangs as soft
and flaccid to the touch as at ordinary times of rest. Secondly,
that the only parts which do move are the two posterior pillars
of the pharynx; and their motion is ample and decided and
altogether unmistakeable. They both move inwards simul-
taneously towards the middle line, moving from their old posi-
tion from one-half to three-fourths of an inch. This action is
not spasmodic, but perfectly steady, and can be sustained for
some considerable time at will, the pillars maintaining their
new position all the while.
Now I am quite satisfied and certain that during this
period the Eustachian tube is open. It will be noted that
from the flaccid condition of the velum, and also from the fact
of its position and form remaining unaltered, the tensor and
levator palati can have no participation in the opening of the
tube, and that the muscles most evidently concerned are the
palato-pharyngel.
Another point to which I wish to call attention is the sound
which accompanies the opening of the Eustachian tube. It is
a sharp crackling sound, which is referred to some part of the
tympanum, or perhaps to the membrane itself. This, I have no
doubt whatever, is caused by the separation of the walls of the
Eustachian tube; I have not succeeded in imitating it in the
dead human subject, but it can be very readily done in the
sheep. By taking hold of the fold of mucous membrane, which
half covers the lumen, and drawing it gradually inwards, the
tube is opened, and at the moment of separation of its walls a
sound is heard closely resembling that which I have described.
‘The sound can be heard by anybody at the commencement of
the act of swallowing, and forms the prelude to a series of
noises arising from the moving of mucus, &c. about the pos-
ee =
|
OPENING AND SHUTTING OF THE EUSTACHIAN TUBE. 131
terior nares and pharynx. I find it is most distinct when a
bolus of tolerably dry food is swallowed.
On examining the anatomy of the parts, the appearances
presented during the opening of the tube will be readily under-
stood. The cartilagmous continuation of the Eustachian tube,
or salpynx, presents at its upper part a massive lobe, to this is
attached the tendon of the salpyngo-pharyngeus. The direc-
tion of this muscle from above is downwards and a little out-
wards when the palato-pharyngeus is at rest, and its action in
this state would be to press the internal lobe of the salpynx
inwards towards the orifice; thus assisting by its tonicity the
elasticity of the cartilage in keeping the tube closed. The two
muscles upon which the opening of the Eustachian tube depends
are the salpyngo-pharyngeus and the palato-pharyngeus. ‘The
former muscle, as before explained, is attached above to the
lobe of the salpynx by a tendon, and below its fibres mix with
those of the palato-pharyngeus; it has a separate existence,
however, for a distance of about an inch and a half. The
palato-pharyngeus is divisible into two parts, a lower vertical
and an upper curved one, arching inwards towards its fellow on
the other side; when the muscle contracts it is evident. that
the curved part becomes straighter, and that the straight part
is drawn inwards; and as the salpyngo-pharyngeus arises from
the lower part its origin is also carried inwards. The effect of
this is to give the action of the salpyngo-pharyngeus a new
direction, such as to pull the lobe of the salpynx towards the
middle line and out of the orifice of the Eustachian tube, thus
opening the cavity of the tube. This is the rationale of the
approximation of the posterior pillars of the pharynx.
Of the other muscles around this part of the pharynx nearly
all have at various times been stated to have an opening or
closing effect on the Eustachian tube. The tensor-palati passes
backwards outwards and upwards past the inner edge of the
salpynx, but does not seem to be attached to the salpynx, or to
be attached in such a manner as to open the tube at all. Lower
down lies the levator-palati. This muscle may perhaps have
some little action in opening the tube when it contracts.
Slightly above its middle point it is crossed by the salpyngo-
eel
132 MR YULE. MECHANISM OF THE EUSTACHIAN TUBE.
pharyngeus; and it possibly may assist, when its diameter is
increased by contraction, in giving to the salpyngo-pharyngeus
its new direction; but that this action is very insignificant is
shown by the fact, that during the opening of the Eustachian
tube, the velum-palati remains flaccid.
In the sheep the salpynx has a different form from that in
man, the chief peculiarity being the absence of the bluff mass
of cartilage on its inner edge. The free edge is composed of a
thin flap or fold of connective tissue, covered by mucous mem-
brane. On examining it from the inside, by making a longi-
tudinal section of the head in the median plane, it will be seen
that the free edge of the salpynx projects very little—scarcely
at all above the surface of the mucous membrane. The orifice
of the tube is closed, most distinctly so, and is covered by a
layer of mucus. It is also to be noticed, that the free fold of
mucous membrane passes directly downward to the posterior
pillars of the fauces. Now the contraction of the palato-
pharyngei draws this fold of mucous membrane inward and
opens the tube. This action is easily imitated in the dead
subject.
For assistance in the dissections I am much indebted to
Dr Wilson of the Cambridge University school of Anatomy.
EXPLANATION OF PLATE.
Fig. 1. Cavity of the pharynx viewed from the right side in
Man. A. Hard palate. B. Velum Palati. C. Azygos Uvule.
D. Tensor Palati. E. Levator Palati. FF. Palato-pharyngeus.
G. Salpyngo-pharyngeus uniting below with the palato-pharyngeus.
K. Lateral cartilaginous lobe of the Salpynx. L. Tendinous inser-
tion of the salpyngo-pharyngeus.
Fig. 2. Anterior view of the pharynx during rest. A. Ante-
rior pillars of the Fauces. B. Posterior pillars of the Fauces.
Fig. 3. Anterior view of the pharynx during the opening of
the Eustachian Tube. Letters as in Fig. 2
Fig. 4. Anterior view of the Eustachian Tube in the Sheep
when closed.
Fig. 5. Ditto, when open.
_—
— c
——
tees
NOTE ON A BIDENTAL SKULL OF A NARWHAL.
By PRoFressoR TURNER.
Mr J. W. CLark of Cambridge, in his excellent notes* on skulls
of the Narwhal, in which two developed tusks have been found,
refers to a statement made by Dr R. Brown, in his account of
the Cetaceans of the Greenland Seas (P. Z S. 1868, p. 353),
who says, “ Among other double-horned skulls which have been
preserved, there is a fine specimen, presented by Captain Gra-
ville, in the Trinity House, Hull—one of the teeth is 3’ long,
and the other 4’.”. Mr Clark, being desirous of obtaining fuller
information respecting this specimen, wrote to the Curator of
the Hull Literary and Philosophical Society, who replied that
no skull of the Narwhal could be found in the Museum of the
Trinity House, and on enquiring of a friend of the Gravilles,
he could not hear that such a skull had been in their pos-
session.
As I had some recollection of hearing my former pupil, Mr
Charles Edward Smith, who acted as Surgeon to the whaling
ship “Diana” on her disastrous voyage, when Captain Graville and
others of her seamen died of cold and hunger, speak of a two-
horned skull of a Narwhal in the possession of Captain Graville,
I took the opportunity, as he was passing through Edinburgh in
the month of July last, of asking him to give me some account
of the specimen. Mr Smith very kindly complied with my
request, and furnished me with the following note :—
“ July, 1866. I went ashore to the Esquimaux settlement
at Button Point, on the north side of Pond’s Bay, to barter
knives and ammunition for Narwhals’ horns. The Esquimaux
brought out of a pool of water the skull of a Narwhal with the
two tusks projecting from the upper jaw, one was 7 ft. 4 in.
long, the other 7 ft. 1in., and the points were unbroken. The
tusks were not parallel, but diverged from each other, so that
a man could stand between them at their free ends. When
Captain Graville saw this skull he claimed it as his perquisite;
1 Proc, Zool. Soc, of London, Jan. 17, 1871.
134 PROF. TURNER. BIDENTAL SKULL OF A NARWHAL.
and in the month of September, about three months prior to
his death on board the ‘ Diana’ from exhaustion, he cut the two
tusks out of the skull. JI may add that I was much distressed
at witnessing the destruction of this specimen, and interposed.
in vain to save it.”
There can, I think, be little doubt that two of the Narwhal
tusks, which Mr Clark states were shown him by Mr Wareham,
the dealer in curiosities, who had bought them out of the
* Diana,” and who had been informed by the mate “ that two
of them were taken out of the same skull,” were the teeth ob-
tained from that particular cranium, the destruction of which
Mr Smith has given me an account of.
AN ABNORMAL ISCHIO-TROCHANTERIC LIGAMENT.
By Tuomas Dwicut, Junr., M.D., of Boston, United
States of America, Professor of Anatomy at the Medical
School of Maine. (Plate VIL, Fig. 1.)
THE anomaly was observed in February, 1873, on a male sub-
ject of large size and of remarkable muscular development.
In the left hip a smooth and glistening tendon (A in figure)
arose, above the tuberosity of the ischium, from the outer edge
of the cartilaginous surface over which the tendon of the obtu-
rator internus plays, and ran outward to be inserted into the
anterior part of the digital fossa. At its origin it was one-
eighth of an inch distant from the capsule, and it remained dis-
tinct from it for the first half of its course. After joining the
capsule the tendon expanded laterally, but without at all losing
its individuality. Below this there was another band (B in
figure), closely connected with the capsule, from which it re-
ceived in its lower border a broad band of circular fibres. B is,
no doubt, an uncommonly well-marked instance of Barkow’s
ischio-capsular ligament’; but the upper band (A), on account
1 Vide Henle’s Banderlehre.
DR DWIGHT. ABNORMAL ISCHIO-TROCHANTERIC LIGAMENT. 135
of its denser structure, its slighter connection with the capsule,
and its want of union with any of the circular fibres, cannot be
considered a similar hypertrophy, and admits, moreover, of a
more plausible explanation. As there can be little question
that in many positions of the leg the obturator internus plays
the part of a ligament restraining the great trochanter; as fur-
ther, in human anatomy at least, the gemelli are practically
parts of this muscle, their edges usually meeting under cover
of its tendon; and as the anomalous band arose close beneath
the gemelli, was inserted near the obturator internus, and must
have had a perfectly similar limiting function, it appears most
natural to consider it a repetition of that muscle. After the
drawing had been made, the joint was opened through the wall
of the pelvis for the examination of the round ligament, which
was found larger than usual.
The subject presented some other anomalies, all of which
were on the side of excessive development. The clavicles
were very strong and much curved; the sternal ends were pro-
longed backward as strong processes, and each bone had a true
synovial articulation with the first rib. The left semi-tendi-
nosus gave off a strong tensor fasciee suralis.
DEPRESSIONS IN THE PARIETAL BONES OF AN
ORANG AND IN MAN—SUPERNUMERARY MO-
LARS IN ORANG. By Prorressorn Humpury. (Plate
VII. Figs. 2 to 6.)
Fic. 2 represents the skull of an adult female Orang from
Borneo, lately presented by Mr W. Vores, of Caius College, to
the Anatomical Museum of the University of Cambridge, which
shews depressions on the exterior of the parietal bones similar
to those occasionally found in the human skull, and to which
I have directed attention in my Muman Skeleton, p. 242.
These depressions in the Orang are at the middle of the
parietal bones, placed, almost symmetrically, on the sides and at
a short distance from the sagittal suture. They look as if the
bones had been indented when in a soft state by the pressure of
two fingers; the deepest part, which is at the middle, is about
a line below the level of the surrounding bone. The bone
rises gradually to the circumference; and the whole surface is
smooth lke that of the remainder of the skull. The depres-
sion on the right side is irregularly oval, and is 14 inch in its
longest diameter, which is parallel with the sagittal suture, and
§ inch in its transverse diameter. That on the left side is
more circular and about an inch in diameter. A quarter of an
inch in front of this is another (a third) depression, more super-
ficial and smaller, measuring 8 inch by 3, with the longest
diameter antero-posterior, ‘There is no corresponding altera-
tion in the contour of the interior of the skull; and there is no
other peculiarity in the skull except the additional molar teeth,
presently to be mentioned.
Fig. 3 represents the calvarium of a human skull in the
Cambridge Museum, in which the parietal depressions are very
marked. They are ovoid, nearly symmetrical, slanting ob-
liquely from the posterior and inner part of the parietals, a
quarter of an inch from the sagittal suture and from its junction
with the lambdoidal on the two sides, outwards and forwards,
to about an inch from the coronal suture midway between the
sagittal and the spheno-parietal sutures. That on the right
PROFESSOR HUMPHRY. DEPRESSIONS IN PARIETAL BONES. 137
side measures 32 inches in the longest or antero-posterior direc-
tion and 2 inches in its greatest transverse diameter. That on
the left side is 2% inches long and 14 inch in its greatest width.
They are smooth, though finely pitted, and somewhat uneven
both in surface and contour. The edges are bevelled; and the
deepest parts, which are about the middle, extend almost through
the skull, leaving only a thin semi-transparent plate of the
inner table. They look as if a portion of the outer table and
diploe had been sliced off, leaving the inner table, which pre-
serves its proper level. At the middle of the sagittal suture is a
similar depression or indentation,14 inch by é Grins of course
both parietals. The interior of the skull presents. no alteration
in its contour corresponding with these external depressions,
Fig. 4 represents a transverse section through the parietal
depressions and through the median sagittal depression. It
does’ not indicate quite the thinnest part of the parietal bones.
It shews that the inner table retains its proper contour. The
bulging into the interior at the middle, which, it will be ob-
served, does not quite correspond with the sagittal depression,
has no relation to it, but is caused by the thickening of the
skull in the vicinity of the median suture.
I have met with similar depressions i in a few other instances
in the human skull. They exist in the skull from an elderly
woman in the museum of the College of Surgeons; and I
recently found them in the parietal bones of a woman, et. 73’,
who died suddenly of apoplexy, and in whom the skull, the
calvarium especially and the frontal bones in particular, are
thick and heavy from osseous deposit in the interior. The
internal surface of the frontal bones is remarkably uneven or
nodulated from this hard deposit, which would appear to have
taken place recently, during the latter years at least of the
patient’s life, and which was probably associated with, if not
dependent on, shrinking of the brain. The deposit has occur-
red upon the interior of the depressions as well as of other
parts of the parietals, but does not appear to have taken place
1 Dr Davis, in a note in the Crania Britannica, p. 6, mentions the skull of a
very aged Chinese, in which the central area of the parietal bones is thinned
and depressed, over an extent equal to four square inches, to about one-third of
an inch deep in the central part. See also Paget’s Lectures on Pathology, drd
Ed. by Turner, p. 101; and Rokitansky, Handbuch der Path. Anat. 11. 243.
138 PROFESSOR HUMPHRY.
upon the exterior of the skull. The specimen is in the Cam-
bridge Museum. The depressions correspond generally with
those in the specimen I have described; though the outline is
‘rather more irregular, and the inner edge of each is 14 inch
from the sagittal suture. There is also in this case a depres-
sion at the middle of the sagittal suture, though smaller and
shallower than in the specimen last described.
I can offer no explanation of the occurrence of these parietal
depressions or connect them with anything of embryological or
morphological interest, and I have not met with them in any
other animal besides this Orang and Man. They appear to be
due to an imperfection in the ossifying processes in consequence
of which this part of the skull is left thin, or, it may be, if the
specimen of the Peruvian, presently to be mentioned, is an in-
stance of the kind, perforated. They are not necessarily conge-
nital. It may be that the thinness is due to a want of proper
balance between the bone growth on the exterior of the skull
and the bone absorption on the interior, which accompany and
are caused by the enlargement of the brain, and which effect
the requisite enlargement of the cranial cavity after the sutures
are closed. But why the deficiency should especially mani-
fest itself at this part, on either side, I cannot tell. Their po-
sition is not at the parietal protuberances, where ossification
commences, or at the sagittal suture, where it terminates, or at
the parietal foramina, which are sometimes preternaturally
large. As we have seen, it is, in some instances, associated with
similar thinness in the course of the sagittal suture; and there
may be an approach to the same thing in other parts of the
calvarium, when it does not exist in this region of the parietal
bones. Thus in a skull in the Cambridge Museum there is a
wide shallow groove in the situation of the hinder part of the
sagittal suture, which is continued, on either side, in the course
of the lambdoidal suture; and in the skull of a South Austra-
lian, in the same museum, there is a slight depression in the
frontal bone, on either side of the median line, corresponding in
position and appearance with those I have described in the pa-
rietal bones, but much less marked. Indeed I should not have
noticed it had I not been close questioning the skulls in our
museum in reference to this point.
DEPRESSIONS IN PARIETAL BONES OF ORANG AND MAN. 139.
These parietal depressions commonly exist on the two sides?
and are symmetrical in position, and more or less so in form,
size, and depth. They do not present any indications of being
the result of disease. ‘They are certainly not the result of acci-
dent. They may be caused by absorption of the outer tables of
the skull. It is however as difficult to know why absorption
should attack this region as why deficiency of formation should
be manifested here. Pressure, interfering with formation of
the bone or inducing its absorption, would cause such depres-
sions. But whence the pressure? Clearly not from wens or
other cysts or morbid growths; and I am not acquainted with
any ordinary or extraordinary influence that would be likely to
produce the effect.
The depressions have a practical bearing, forasmuch as they
might lead to the supposition, after an accident, that there was
depression of the bone, or they might be supposed to indicate
that there had been loss of bone after accident or disease fol-
lowed by imperfect reparation. This mistake seems to have
been made in the case of the specimen in the museum of
St Thomas’ Hospital, referred to by me (loc. cit. p. 243). In the
skull of an ancient Peruvian, in the Cambridge Museum, there
is a slight depression in the usual position in the right parietal
bone; and in the corresponding situation of the left parietal
bone there is a large circular, uneven, slightly depressed space
in which at one part the depression is deeper; and at two other
parts in the same area the depression extends quite through
the skull, giving rise to two holes with sharp edges and bevelled
outer margins. The interior of skull presents, so far as I can
see through the foramen magnum, no alteration, except the
perforations. Now, in this case, it is extremely difficult to de-
cide whether or not the superficial and the deeper depressions
and foramina on the left side are congenital, as is probably the
depression in the corresponding situation on the opposite side,
or whether they are the result of accident or disease. Gall ap-
pears to have possessed “the skull of a full-grown man, whose
exterior lamella of the os bregmatis on both sides had been
1 Jn my treatise On the Human SkeletonI mention the case of an infant born
with a depression in the right parietal bone, which I thought to be of the same
nature es those I am describing; but the examination was made in the living
child, and I could not therefore form any decided opinion respecting it.
140 PROFESSOR HUMPHRY.
broken by Levret’s forceps, and had not been replaced in its
former situation. The whole form of the forceps is said to be
distinctly seen on the outside, but on the inside of the lamella
not the least impression is discernible, and consequently had
been restored to its due form through the action of the brain’.”
Doubtless, this was an instance of the depression in question ;
and one cannot be surprised at the supposition being enter-
tained, that the depression was caused by the pressure of the
forceps, for the appearance of the exterior (in the instances I
have, described) is suggestive of such a cause; though there is
an absence of any corresponding projection in the interior of
the skull.
In a paper in Virchow’s Archiv, vil. 338, Dr Maier describes
two specimens of this condition of the parietal bones, both from
aged females, and regards it as the result of senile osteoporosis,
the bone being yellowish, fatty, and porous, the porosity
being, partly, due to enlargement and fusion of the Haversian
canals. In neither of the specimens I have described is there
any appearance of osteoporosis. The bone in the region of the
depressions is as dense as it usually is, and as it is in other parts
of the calvaria. And sections taken from the human specimen,
represented in the figures, present, under the microscope, a
normal appearance with corpuscles and Haversian canals of
the usual size and appearance; and the bone structure shews
nothing abnormal’.
Another anomaly in this Orang’s skull is presented by the
presence of an additional, or sixth, molar tooth situated behind
the usual series on either side of the upper jaw and on the left
side of the lower jaw. These teeth, though smaller than the
other molars, are well formed, having each four cusps and the
normal complement of fangs, viz. one in front and one behind
1 Dr F. Gall’s System of the Functions of the Brain, extracted from C. A.
Bode’s account of Gall’s lectures, p. 62.
2 In Virchow’s Gesammelte Abhandlungen zur wissenschaftlichen Medicin,
S. 1000, is a paper iiber die Involutions-Krankheit (malum senile) der platten
Knochen, namentlich des Schiidels, in which specimens are described where the
parietal depressions were accompanied with more or less porosity of the
remainder of the roof of the skull and were associated, in one case, with thinning
of the blades of the scapule and of the middle of the iliac bones, and, in one,
with éhickening of the skull. The condition is regarded as the result of atrophy
commencing on the exterior.
DEPRESSIONS IN PARIETAL BONES OF ORANG AND MAN. 141
in the lower jaw, and two on the outer and one on the inner side
_in the upper jaw. On the right side of the lower jaw there is
no trace of an additional or sixth molar, though there is nearly
as much space for it as on the other side. Instances of such
additional molars in these animals have been seen by others;
though it has not happened to myself to meet with one
before. They are of interest not only in connection with the
fact of the presence of an extra molar (a third premolar) in
the monkeys of the new world, but also with the fact that in
some of the old world monkeys, as Macacus and Colobus, there
is a tendency to occupy the space which the elongated alveolus
of the jaw presents by the formation in the lower dental series
of an additional or fifth cusp at the hinder part of the last
molar, which is of large size.
ON THE RELATIONS OF THE CONVOLUTIONS OF
THE HUMAN CEREBRUM TO THE OUTER
SURFACE OF THE SKULL AND HEAD. By
PROFESSOR TURNER.
SoME years ago I planned in connection with an investiga-
tion into the arrangement of the convolutions of the human
cerebrum, on which I was then engaged, a series of observations
on the relations of the convolutions to the outer surface of the
skull and of the scalp. Various circumstances have, however,
combined to hinder me from carrying out this enquiry as fully
as I should have desired. I have thought, however, that it
might not be without interest to those engaged in the study of
the anatomy and physiology of the human brain to record some
of the leading facts, which I have up to this time ascertained,
as a preliminary to a more extended memoir which I hope in
due time to be ina position to publish.
In this communication, I shall direct attention to -the
general relations of the principal convolutions and sulci to the
surface as I have observed them in the heads of adult men,
and reserve to a subsequent memoir the relations of the corre-
sponding structures in the female, and the variations in position,
which to some extent occur in different individuals from
variations in the configuration of the brain and skull.
In conducting an investigation of this kind, it is in the first
instance necessary to have a clear conception of certain well-
defined landmarks, which can be seen or felt when the outer
surface of the skull and head are examined. The external
occipital protuberance, the parietal and frontal eminences,
and the external angular process of the frontal bone, are easily
recognised structures, the position of which can be determined
by manipulating the scalp, and still more readily on the
surface of the skull itself. The coronal and lambdoidal sutures
ean also be felt through the scalp in most heads, and on the
skull itself the position of the squamous, squamoso-sphenoid
and parieto-sphenoid sutures, and the curved line of the
temporal ridge can also without difficulty be determined.
PROF. TURNER. CONVOLUTIONS OF THE HUMAN CEREBRUM. 1438
‘With the aid of these structures, we may subdivide
and map out each lateral half of the surface of the skull or
scalp into ten well-defined regions or areas, and then localise
within these areas the convolutions on the outer face of the
hemisphere. The line of the coronal suture forms on each
side the posterior boundary of the pree-coronal or frontal region.
A vertical line drawn from the squamous suture upwards
through the parietal eminence to the sagittal suture lies
almost parallel to the coronal suture and subdivides the area of
the parietal bone into a post-coronal or antero-parietal, and
a pre-lambdoidal or post-parietal. The squamous part of the
occipital bone between the lambdoidal suture and the occipital
protuberance and superior curved line forms the post-lamb-
doidal or occipital region.
But these regions may be subdivided into smaller areas.
The temporal ridge starting from the external frontal process
curves backwards across the pre-coronal, post-coronal and pre-
lambdoidal regions to the lateral angle of the occipital bone,
and subdivides each of these regions into an upper and a lower
area. The lower areas may be termed the inferior frontal or
fronto-temporal, the lower antero-parietal, and the lower postero-
parietal. ‘lhe fronto-temporal area is bounded above by the
temporal ridge, behind by the coronal suture, below by the
fronto-sphenoidal suture. The lower antero-parietal region is
bounded above by the temporal ridge, in front by the coronal
suture, below by the squamous and sphenoido-parietal sutures,
and behind by the vertical parietal line already referred to as
subdividing the parietal region into an anterior and a posterior
area, The lower postero-parietal region is bounded above and
behind by the temporai ridge, in front by the above-named
vertical line, and below by the posterior part of the squamous
suture, and the parieto-mastoid suture. The upper area of
the pre-coronal region consists of the frontal bone above the
temporal ridge, and may be subdivided into a mid-frontal and
a superior frontal by an antero-posterior line drawn back-
wards parallel to the frontal suture from the superior orbital
border through the frontal eminence to the coronal suture.
The upper areas of the post-coronal and pree-lambdoidal regions
consist of the parietal bone above the temporal ridge, and are
144 PROFESSOR TURNER.
bounded below by that ridge, above by the sagittal suture, and
are separated from each other by the vertical parietal line
already referred to. They may conveniently be named the
upper antero-parietal, and upper postero-parietal areas. Eight
areas may thus be defined on each side of the head situated
beneath that portion of the scalp, which is so thin as to allow
their extent and boundaries to be ascertained on external
manipulation with a fair amount of precision.
That part of the side of the head, however, which les
below the line of the squamoso-parietal, sphenoido-parietal and
sphenoido-frontal sutures, is so covered over by the thick mass
of the temporal muscle that the determination of its limits on
the head itself is attended with more difficulty. On the skull,
however, it can be readily marked out, and may naturally be
subdivided by the lines of sutures into the squamoso-temporal
and the sphenoidal areas, which form the ninth and tenth areas
on the side of the head.
Before I proceed to state the position of the convolutions
within these different areas, it will be advisable to determine
the regions in which the great fissures are situated, which
subdivide the hemisphere into its five constituent lobes, frontal,
parietal, occipital, temporo-sphenoidal and the Insula.
The Sylvian fissure commences immediately behind the
posterior border of the lesser wing of the sphenoid bone, and in
its course backwards and upwards, is covered by the great wing
of the sphenoid where it articulates with the anterior inferior
angle of the parietal. It then passes obliquely under cover of
the anterior superior part of the squamous plate of the tem-
poral, and appears in the lower part of the antero-parietal
region, through which either it, or one of the small branches
into which it not unfrequently divides, may be continued into
the lower postero-parietal region.
The fissure of Rolando lies in the post-coronal region,
through which it passes from below obliquely upwards and
backwards, so that it traverses both its upper and lower
subdivisions. The distance of this fissure from the coronal
suture varies in different brains. I have seen its upper end as
much as 2 inches behind the top of the suture, its lower end
14 inch behind the inferior part of the suture, but I have
RELATIONS OF CONVOLUTIONS TO SKULL AND SCALP. 145
also seen the upper and lower ends not more than 14 and 13
inch posterior to this suture.
The parieto-occipital fissure lies in the upper area of the:
pre-lambdoidal region close to its sagittal border. Its exact
distance from the apex of the lambdoidal suture varies, partly
from variations in the brain itself, and partly from the not
unfrequent variations in the mode of ossification of the upper.
squamous part of the occipital bone. About 0°7 or 0°8 of an inch
will express its average distance from the apex of that suture.
The relations of the parietal eminence to the surface of the
hemisphere are apparently very constant. In the specimens I
have examined, the hollow corresponding to the cerebral surface
of this eminence was occupied by the supra-marginal convolu-
tion, which from this circumstance may appropriately be named.
the convolution of the parietal eminence.
As the fissure of Rolando, which I regard as the line of
demarcation between the frontal and parietal lobes, extends
obliquely upwards and backwards, in the post-corenal region, the
great central convolutions which bound it in front and behind,
and which we now name ascending frontal and ascending pari-
etal, necessarily occupy a considerable share of its two sub-
divisions. To define their positions more exactly, I shall now
state the contents of each subdivision.
In the lower antero-parietal area the lower third of the
ascending frontal and parietal gyri are found, and from the
anterior frontal arises, somewhat less than an inch behind the
lower end of the coronal suture, the inferior frontal gyrus.
Behind the ascending parietal, but separated from it by the
intra-parietal sulcus, lies a small part of the supra-marginal
gyrus or convolution of the parietal eminence. Below the
ascending gyri extends a short segment of the Sylvian fissure,
and quite at the lower and posterior boundary of the area a
little bit of the superior temporo-sphenoidal gyrus appears above
the squamous suture.
In the upper antero-parietal area the upper two-thirds of
the ascending frontal and parietal gyri, which extend close up to
the sagittal suture, are found. From the ascending frontal
arise the superior and middle frontal gyri; the former arises
1:2 or 1°3 of an inch behind the coronal suture, the latter about
VOL, VIII. 10
146 PROFESSOR TURNER.
an inch behind the same line. Immediately behind the upper
end of the ascending parietal gyrus, in the neighbourhood of the
sagittal suture, a little bit of the postero-parietal lobule may
or may not be seen; and below this a small portion of the con-
volution of the parietal eminence may appear.
The contents of the pra-lambdoidal region are as follows.
In the lower postero-parietal area lies the hinder part of the
convolution of the parietal eminence, behind which is the
angular gyrus, and below this the posterior ends of the three
convolutions of the temporo-sphenoidal lobe. In the upper
postero-parietal area the postero-parietal lobule occupies the
region close to the sagittal suture up to the parieto-occipital
fissure ; below it lies the upper portion of the angular gyrus, and
a part of the convolution of the parietal eminence; in the more
posterior part of the region the annectent gyri blend with the
three convolutions of the occipital lobe.
In the post-lambdoidal region, which is comparatively smal],
the three convolutions of the occipital lobe succeed each other
from above downwards.
The pre-coronal region is entirely occupied by the frontal
lobe. In the inferior or temporo-frontal area is situated a large
portion of the inferior frontal convolution, but a small bit of the
middle gyrus may be seen at its posterior superior angle. The
middle frontal area corresponds almost exactly to the middle
frontal gyrus, and the superior frontal area to the superior
gyrus. If the line of demarcation between the superior and
mid-frontal areas were prolonged backwards for about an inch
beyond the coronal suture into the upper antero-parietal area,
the extent and position of the superior and middle frontal
convolutions would be still more exactly defined. The frontal
eminence may lie opposite the sulcus between the superior and
middle frontal gyri, or if this sulcus, as is not unfrequent, is
bridged across by a small tertiary convolution, this may corre-
spond to the eminence.
The squamoso-temporal and the sphenoidal regions contain
the anterior two-thirds of the convolutions of the temporo-
sphenoidal lobe.
The lobes of the brain by no means precisely correspond to
the areas of the cranial bones, after which four of them are
F
{
RELATIONS OF CONVOLUTIONS TO SKULL AND SCALP. 147
named. The frontal lobe is not only covered over by the
frontal bone, but extends backwards for a considerable distance
under cover of the parietal bone. If we accept, as I have else-
where described', the fissure of Rolando as the posterior limit
of this lobe, then the larger part of the post-coronal region
corresponds with the frontal lobe, for not only does it contain
the origins of the superior, middle and inferior frontal gyri, but
also the ascending frontal convolution. The distance at which
the fissure of Rolando lies behind the coronal suture was pointed
out some years ago by M. Broca’, was referred to by myself in
my essay quoted above, and has subsequently been described
by Prof. Bischoff*, although the last-named writer, with Gratiolet
and some other anatomists, regards the convolution which I
have named ascending frontal, and not the fissure of Rolando, as
the posterior limit of the frontal lobe. But even if this mode
of looking at the ascending frontal gyrus were accepted, the
frontal lobe would still not be wholly localized under cover of
the frontal bone, for the superior, middle and inferior frontal
gyri all arise behind the line of the coronal suture.
The occipital lobe also is not limited to the region covered
by the squamous part of the occipital bone, but slightly over-
lapping the lambdoidal suture, extends forwards for a short
distance into the back part of the upper postero-parietal area,
and through the superior annectent gyrus reaches the parieto-
occipital fissure.
The superior temporo-sphenoidal gyrus, though for the most
part situated under cover of the squamous-temporal and great
wing of the sphenoid, yet ascends into both the lower antero-
and lower postero-parietal areas.
The area covered by the parietal bone, so far then from
being conterminous with the parietal lobe of the cerebrum, is
trenched on anteriorly, posteriorly and inferiorly by three of
the other lobes of the brain. The convolutions of the parietal
lobe are especially grouped around the parietal eminence, and
in the interval between that structure and the sagittal suture.
The Insula or central lobe does not come to the surface, but
1 Edinburgh Medical Journal, June, 1866, and separate publication ‘‘ The
Convolutions of the Human Cerebrum, topographically considered.”’
2 Sur le Siége de la Faculté du Langage articulé, Paris, 1861.
3 Die Grosshirnwindungen des Menschen. Munich, 1868,
10—2
148 PROF. TURNER. CONVOLUTIONS OF THE HUMAN CEREBRUM.
lies deep in the Sylvian fissure, and is concealed by the convolu-
tions which form the margin of that fissure anteriorly. It les
opposite the upper part of the great wing of the sphenoid and
its line of articulation with the antero-inferior angle of the
parietal and the squamous part of the temporal.
In conclusion I may again state that the above description is
not intended to be exhaustive, but merely to afford a general
conception of the chief relations of the convolutions to the
surface, neither does it take into consideration the variations
which may be occasioned by sex, race, or individual peculiarities.
It may however help the physiologist and pathologist to map
out on the surface of the living head, in a more precise manner
than they have hitherto been able to do, the position of the
convolutions, and in so far be of some use in the study of their
functions and diseases.
—————
TWO CASES OF PERSISTENT COMMUNICATION BE-
TWEEN THE UMBILICAL AND PORTAL VEINS IN
THE HUMAN SUBJECT. By J. A. Russett, M.A., M.B.,
Demonstrator of Anatomy, University of Edinburgh.
Mr Frank CuHampneys, in the sixth volume of this Journal, page
417, records a case of communication between the external iliac and
portal veins through the umbilical vein, and gives references to
previously recorded cases of communication between the portal vein
and various veins of the abdominal parietes.
In connection with this subject, the occurrence in the dissecting-
rooms of the University of Edinburgh last summer of two cases of
persistent communication between the umbilical and portal veins
where there was no obvious junction between the umbilical veins
and any veins of the abdominal wall, may be worthy of record. In
the first of those cases, that of a male, where there was a cirrhosed
liver considerably enlarged by fatty degeneration, the canal of the
umbilical vein, when traced towards the umbilicus, was found to end
within the abdominal wall in a cul de sac, just opposite the navel?
From this point to the suspensory ligament of the liver the vein was
larger than a goose-quill, and thin walled. Just before joining the
anterior edge of the falciform ligament it had a trilocular dilatation,
also thin walled, of the size of a walnut, continuous with its wall and
opening into it. An injection of melted lard failed to show any
communication with parietal veins. Immediately upon passing from
the dilatation and entering the suspensory ligament, which had much
fat between its layers, the coats became greatly thickened and the
vessel gradually enlarged, until, in the hepatic fissure for the umbili-
cal vein, where it became continuous with the anterior division of the
left branch of the portal vein, it was as big as the end of the fore-
finger. Branches of varying size entered the liver substance on both
sides of the fissure.
Proceeding from the dilatation and surrounding the vein were
three pairs of thin-walled, highly tortuous vessels, as large as crow-
quills at their origin. These vessels distributed branches to the fat
in the falciform ligament, and in the umbilical and part of the portal
fissures, and being diminished in size finally entered the liver sub-
stance in close contact with branches of the hepatic artery and
portal vein. The ductus venosus was quite impervious.
In the other case, also a male with a very slightly enlarged liver,
the connection between the portal and umbilical veins was exactly
similar to the above, but the pervious channel of the umbilical vein
did not reach the wall of the abdomen and only extended about
a couple of inches from the liver, where it was suddenly obliterated
within the falciform ligament, so that it formed, as it were, a diver-
ticulum on the portal system.
150 MR RUSSELL. UMBILICAL AND PORTAL VEINS.
In many of the cases to which Mr Champneys refers it is probable
that the patency of the umbilical vein was caused by its fusion with
Luschka’s Vena parumbilicalis, and that this channel was afterwards
enlarged by the obstruction due to liver disease. This could hardly
be the case in the subjects noted above, as the umbilical vein was
joined by no parietal vessels of any appreciable size, and such vessels
could not fail to have been distended by a force sufficient to enlarge
the umbilical vein, supposing the enlargement to have been due to
liver-obstruction.
In connection with the communication between the portal and
parietal veins, it may be interesting to note that Dr Monro’s prepara-
tion, referred to by Mr Champneys, is still preserved in the Ana-
tomical Museum of the University of Edinburgh (L 1 Old Catalogue,
and 346 New Catalogue). The liver which Monro described as scir-
rhous is, however, more probably cirrhosed.
ON A PECULIAR DIGASTRIC MUSCLE—A VARIETY OF
THE OCCIPITO-HYOID. By S. H. West, B.A., Junior
Student of -Christ’s Church, Oxford.
Tue muscle is of interest, as having been hitherto noticed, so far as
Iam aware, by only one anatomist, Mr Perrin, who has given a
description of it in this Journal, v. 251.
I have now seen it in an aged male subject, symmetrically placed
on both sides, digastric, slender, but of considerable length. On the
left side it arose by a tendon, which expanded and blended with the
fascia beneath the origin of the trapezius, below the superior curved
line of the occipital bone, and was crossed, just beyond its origin, by
the occipital nerve. From this origin it ran horizontally forwards
about half an inch below the superior curved line, having swollen into
a fleshy belly about one-fourth of an inch in diameter, as far as
the sterno-mastoid muscle, upon the external surface of which it
continued its course for about one-sixth of an inch. Here it
suddenly bent downwards at an angle of about 120° to a point
about one-fourth of an inch from the apex of the mastoid process,
and three-fourths of an inch from the anterior border of the sterno-
mastoid muscle, upon the external surface of which it lay throughout
the whole of this part of its course. Here it became tendinous.
Bending forwards to lie again parallel with its former course, 1.e.
horizontal, it became once more muscular, and was crossed by the
anterior jugular vein. It then turned directly inwards over the
anterior border of the sterno-mastoid muscle, lying behind the tendon
of the digastricus. Here it received a special artery, which arose
from the occipital just before the sterno-mastoid branch. Its fur-
2
MR WEST. PECULIAR DIGASTRIC MUSCLE. 151
ther course was downwards and inwards. It crossed the ascending
pharyngeal artery (the occipital artery in this subject rising very
high up), and passing immediately beneath the external and upon the
internal carotid, came into relation with the stylo-pharyngeus muscle,
lying parallel, but external to it. Continuing its course with this
muscle, it passed between the superior and middle constrictors of the
pharynx, and running under cover of the middle constrictor blended
inferiorly with the inferior constrictor on its posterior aspect. Some
of its fibres decussated with those of its fellow, and with those
of the inferior constrictor of the opposite side. It was contained in
all its superficial extent in a closely investing sheath of fascia,
which bound it down to the parts beneath.
The muscle of the right side pursued a similar course, and dif-
fered only in the smaller size of its muscular bellies.
I was unfortunately unable to make out its nerve supply. It
received its blood, as stated above, from the occipital artery by a
special branch. The stylo-hyoid muscle was present on both sides.
This muscle differs from those described by Mr Perrin, (i) in its
curved course, (ii) in not being attached to the hyoid bone, and (iii)
in receiving no accessory slips. Mr Perrin regards it as a differen-
tiation of the stylo-hyoid of birds; and Professor Humphry, who
refers to these peculiar abnormalities in his lectures (Grit. Med. Journ.
June 21st, 1873), considers it as ‘‘an appendage, a superficial append-
age, to the stylo-hyoid and digastric muscles,” and explains its
peculiar digastric condition as arising from causes similar to those
to which he supposes the two bellies of the digastricus to be due.
“The peculiarities of the digastric are in great measure due to the
continued blending of parts of two of these layers” (the layers into
which he states the primitive muscular sheet to be subsequently split),
“the posterior belly being derived from the deep layer, the anterior
belly from the superficial layer, and the intermediate tendon being
a remnant of one of the transverse septa of the primitive unstratified
muscle” (oc. cit.). For a more detailed account and for figures of
this muscle, I may refer to Mr Perrin’s paper quoted above.
EXPERIMENTAL RESEARCHES IN CEREBRAL PHY-
SIOLOGY AND PATHOLOGY*. By Davip Ferrizr,
M.D., &e. Professor of Forensic Medicine in King’s College,
London.
(Abstract by the Author.)
THE paper contains the chief results of a research commenced with
a view to test the accuracy of the views entertained by Dr Hughlings
Jackson on the pathology of Epilepsy and Chorea. As is well known,
Dr Jackson regards localised and unilateral epilepsies as dependent
on irritating or discharging lesions of the convolutions about the
corpus striatum.
In order to put this theory to the proof the author determined to
expose the brain in various animals, and apply irritation to the
surface. The method of irritation was suggested by the experiments
of Fritsch and Hitzig, who had shown that contractions of definite
groups of muscles could be caused in dogs by passing galvanic cur-
rents through certain portions of the anterior regions of the brain.
The progress of the research ultimately led to the endeavour to
establish the localisation of cerebral function, not merely as regards
motion, but also as regards sensation and the other faeulties of mind.
The paper in the West Riding Reports gives the result of experi-
ments on rabbits, cats, and dogs, and is confessedly only a preliminary
instalment of a more extended research.
Since this paper was written the author has been engaged in the
further prosecution of his inquiries, and at the British Association at
Bradford, on September 19th, he gave an account of his more recent
researches, at the same time entering more fully into the psychologi-
cal explanation of many of the phenomena which are partly described
in the published account. He has performed experiments on nume-
rous monkeys as well as other animals, but reserves a complete
account of his researches for the Royal Society.
The method of experimentation which the author has adopted is
to place the animal under chloroform, and gradually expose the
surface of the brain by successive trephinings and removal of the
skull by means of the bone forceps. In this way he has been able
to expose the whole hemisphere. After removal of the dura mater,
the points of blunted electrodes in connection with Du Bois Rey-
mond’s induction coil are applied to the surface of the brain without
injury to the cortical substance.
The first experiments recorded have special reference to the
production of epileptic convulsions; and the mode in which the
1 West Riding Lunatic Asylum Medical Reports, Vol. 111. 1873. London,
Smith, Elder and Co. 15, Waterloo Place.
PROF. FERRIER. RESEARCHES IN CEREBRAL PHYSIOLOGY, &c. 153
attacks begin, and the march of the convulsive spasms, are accu-
rately recorded.
’ It was found that in rabbits, cats, and dogs the application of
the electrodes for a few seconds induced almost immediately, on some
occasions after the lapse of a distinct interval, violent unilateral
epileptic convulsions. When the electrodes were applied, one at the
anterior, and the other at the posterior part of the hemisphere, the
convulsions were complete and violent in the whole of the opposite
side of the body. Asarule they commenced in the face, spread to
the neck and upper extremity, and then invaded the hind-leg and
tail. Dilatation of the pupil, clonic spasms of the jaws, foaming at
the mouth, and loss of consciousness were induced when the fits were
at their greatest intensity.
- Occasionally the spasmodic convulsions remained localised in one
or other limb, or in some one muscle or group of muscles, and fre-
quently, instead of a hasty epileptic attack, a series of choreic twitches
alone were manifested, without any affection of sensibility or con-
sciousness.
The march of the spasms is shown to be quite in accordance
with the clinical observations of Dr Hughlings Jackson in cases of
unilateral epilepsy in man. Peculiar variations in the mode in which
the attacks commenced, depending apparently on the position of the
electrodes on the surface of the brain, led the author to approximate
the electrodes and to apply very limited irritation, in order to
discover whether the convulsive spasms were not due to over violent
irritation of localised centres in the brain, whose special function
is to govern and direct the action of these muscles for definite
purposes, possibly such as might indicate volition and intelligence.
The results were such as to indicate, with a beautiful degree of
exactitude, the localisation in certain definite and easily defined regions
the cerebral centres for various apparently purposive combined
movements of the muscles of the limbs, as well as of the tail, the
facial muscles, and the muscles of the jaws and tongue. These are
all situated in the anterior parts of the brain in advance of the
Fissure of Sylvius, and the individual centres are marked off in the
various external convolutions, of which woodcuts are given. The
general plan is, that in the superior external convolution anterior and
posterior to the crucial sulcus the various movements of the paws, legs,
and tail are centralised, and it is shown that the differentiation of
these centres is to a great extent characteristic of the animal’s habits;
the centre from the fore-paw in cats being much more highly differ-
entiated than in dogs and rabbits.
The middle external convolution governs movements of the eye-
lids, face, and eyes, while the inferior and the Sylvian govern various
movements of the whiskers, angle of the mouth, depressors of the
lower jaw, and tongue.
In the convolutions posterior to the Fissure of Sylvius certain
movements are described as resulting from irritation, viz. of the ears,
eyes, &c. In the paper as yet published no attempt is made to
explain the signification of these; but the author, from his later
154 PROFESSOR FERRIER.
experiments, indicated at the Meeting of the British Association that
he had been able to obtain indications of the situation in these regions
of the centres of special sense, sight, hearing, and smell. These
results and conclusions are however not as yet detailed fully. The
author indicates, in a note in the paper published in the West Riding
Reports, that he had at that time explored the brain of the monkey,
and satisfactorily localised the regions and the homologues of the
centres already discovered in the brain of the cat, rabbit, and dog.
One of the more important conclusions drawn from the experi-
ments is, that the region which governs the movements of the mouth
and tongue in cats and dogs is the homologue of what is known as
Broca’s convolution in man, viz. the posterior part of the inferior
frontal.
This, it may be stated, is further borne out by experiments on
monkeys.
The pathology of Aphasia is thus rendered comparatively simple.
The memory of words is situated in that part of the brain which
governs the movements of articulation. It is shown, however, by
the experiments, that the brain is symmetrical, and that the corre-
sponding part of each hemisphere produces exactly the same effects
on opposite sides of the body. Generally the action is unilateral
and crossed, but as regards the mouth the action is almost bilateral,
and hence disease of one or other side alone does not cause paralysis
of the articulating muscles, because the other side is able to govern
as before. The occurrence of loss of speech with lesion of the left
side is attributed to the fact that most people are left-brained, and
that therefore a lesion of the left side causes such an interference
- with the voluntary recalling of words that the person is speechless,
not because memory of words is utterly lost, as this exists in the
undamaged side, but because he is unable to lay hold of the word
which he wishes to express. With the education of the other side
however the individual recovers the power of speech. During the
interval of recovery of speech only automatic expressions or inter-
jections are uttered, which are evoked by a sort of reflex action, and
unconnected with volition. Among other points discussed is the
hypothesis advanced by Dr Broadbent, that associated movements of
the body are bilaterally coordinated in each hemisphere. The author
thinks the experiments which he gives indicate not an anatomical
but a physiological coordination through the media of the lower
ganglia.
The results of experiments in the corpora striata, optic thalami,
corpora quadrigemina, and cerebellum, are also detailed.
The corpora striata are shown to be motor in function, and to
govern all the muscles of the opposite side; representing all the
muscles directed by the hemispheres, and having a physiological sub-
ordination to these as higher centres.
The optic thalami are shown to have no motor function, and the
author attributes the interference with motion, which is sometimes
described in connection with disease of these ganglia, to affection of
the motor strands with which they are in close relation.
RESEARCHES IN CEREBRAL PHYSIOLOGY AND PATHOLOGY. 155
The corpora quadrigemina have a special relation to the eyes and
also to the extensor muscles.
Trritation of the nates causes great dilatation of the pupils. The
action is crossed, but powerful irritation easily acts on both sides of
the body. Trismus and opisthotonus are induced when these ganglia
are powerfully stimulated.
The cerebellum is shown to have a function which has never been
allotted to it, viz. to be a coordinating centre for the muscles of the
eyeballs. The author has only given the results of his experiments
on the cerebellum of rabbits, but he has since extended and confirmed
them in cats, dogs, and monkeys.
The various lobules of the rabbit’s cerebellum are shown to have
the power of directing the eyes in certain definite directions.
These cerebellar oculo-motorial centres are brought into relation
with the cerebellum as a coordinating centre for the muscles con-
cerned in the maintenance of the equilibrium, and these functions
are indicated as mutually depending on each other.
A more complete exposition of the facts of experiment, and an
account of the results obtained from a further investigation of the
brains of the various classes of the vertebrata, is in process of pub-
lication.
TWO INSTANCES OF IRREGULAR OPHTHALMIC AND
MIDDLE MENINGEAL ARTERIES. By Joun Curnow,
M.D. Lond., Professor of Anatomy in King’s College, London.
I. In the summer session of 1872 I dissected an interesting
irregularity affecting the right middle meningeal artery. The foramen
spinosum was much smaller than usual, and through it passed a very
slender artery from the internal maxillary, which after giving off a
few small twigs to the Gasserian ganglion, entered the hiatus Fallopii.
The foramen ovale transmitted the inferior maxillary nerve only. A
large artery arose from the trunk of the ophthalmic soon after it had
entered the orbit, and passing out through the sphenoidal fissure
divided into two branches which were distributed as the normal
branches of the middle meningeal, except that they ran backwards
above the inferior border of the parietal bone, which was consequently
smooth, and without its usual deep grooves. In addition, a large
artery was given off from the internal carotid in the cranium, and
coursed backwards over the basilar processes of the sphenoid and
occipital bones almost to the foramen magnum. This recurrent branch
was also present on the left side, but the left middle meningeal and
ophthalmic arteries were quite regular,
156 PROF. CURNOW. INSTANCES OF IRREGULAR ARTERIES.-
A less complete instance than this is depicted by Barkow’ (Tab.
xvu. figs. 1 and 2), in which the anterior portion of the middle
meningeal comes from the ophthalmic through the sphenoidal fissure
into the cranium; but the posterior branch is given off from the
internal maxillary and enters through the foramen spinosum.
II. During the past winter session I met with an arrangement
almost the exact reverse of the preceding.
Besides its ordinary branches, the left middle meningeal gave off
a large artery, which entered the orbit through the sphenoidal fissure,
and from which all the regular branches of the ophthalmic arose, with
the single exception of the arteria centralis retine. It ended ina
long dorsal artery on the nose, running downwards as far as the tip,
and supplying angular and lateral nasal offsets. The ophthalmic from
the internal carotid was a very small twig; it passed through the
optic foramen, gave off the arteria centralis retine, and terminated by
joiming the posterior ethmoidal branch of the irregular artery from
the middle meningeal. The trunk of the left facial was smaller than
usual and ended as the superior coronary, from which proceeded the
septal branch as usual. On the right side, the ophthalmic, middle
meningeal, and facial arteries were quite regular.
Anastomotic twigs between the middle Meningeal artery and the
lachrymal branch of the ophthalmic are always met with; and
Cruveilhier mentions the occasional origin of the lachrymal from the
middle meningeal. Barkow figures an example of this irregularity,
and I have also seen it.
These anomalies are obviously due to the suppression of the main
trunk, and compensatory enlargement of the anastomosing branch,
similar to the mode of formation of the abnormal obturator from the
epigastric, the dorsal artery of the foot from the anterior peroneal, &c.
I have failed to find any reference to a deviation from the regular
origin so extensive in this direction as those which I have recorded
above. The only irregular origin of the ophthalmic described. by
Quain (Pl. x11. fig. 8) is of quite a different type. In this the place
of an absent internal carotid artery is supplied by two branches of
the internal maxillary, which enter the cranium through the foramen
rotundum and foramen ovale respectively, and from their junction the
ophthalmic is given off.
1 Comparative Morphologie des Menschen und der menschendhnlichen Thiere.
5ter Theil.
_ Note sy Epitors.—Blandin (Anatomie Topographique, p. 147)
mentions an accessory ophthalmic artery arising from the middle
meningeal, which sometimes also gives origin to the lachrymal artery.
C. Krause states (Handbuch der mensch. Anat. p. 892) that he once
saw the ophthalmic artery arise from the middle meningeal and pass
through the sphenoidal fissure. Tiedemann and Dubrueil have also
recorded similar cases.
REVIEWS AND NOTICES OF BOOKS.
The Anatomist’s Vade Mecum. By Erasmus Witson, F.R.S. Edited
by G. Bucnanan, M.D. 9th edition, London, 1873.
THE new edition of this well-known Student's Manual has been pre-
pared by Dr George Buchanan with the assistance of Mr Henry E.
Clark. To bring it up to the present state of knowledge, many addi-
tions have been made both to the text and wood-cuts,
Handbuch der systematischen Anatomie des Menschen. Von Dr J.
Hente. 3rd vol. 2nd division, 2nd part. Brunswick, 1873.
Proressor HENLE has now completed his great work on Human
Anatomy, by the publication of the descriptive anatomy of the cere-
bral and spinal nerves and the sympathetic system. Like all the
previous parts, it is very fully illustrated by original drawings from
dissections. Not only are the more usual arrangements of the nerves
described, but reference is made to the principal variations which
have been recorded by other anatomists.. The work constitutes the
most complete treatise on human anatomy in any language. It has
lost somewhat in unity by its mode of publication, as do all scientific
treatises published in parts, with long intervals between the appear-
ance of the different parts; for the earlier volumes are necessarily
not so well posted up to the present state of the subject as the one
just finished. So great, however, has been the demand for the earlier
numbers, that the one on the bones has reached a third edition, and
second editions of those on the ligaments, muscles and viscera have
been issued. We may congratulate Professor Henle on having lived
to complete so enduring a memorial of his learning, industry, and
power of description.
The Comparative Anatomy of the Domesticated Animals. By Prof.
CHAvvEAU. ‘Translated and edited by GEORGE NEEL, Vet.
Surg. Royal Engineers. :
Tue reviewer regrets that in the notice of this work in the last num-
ber of the Journal, in consequence of an oversight, it was intimated
that there is an omission of the mention of the sources from which
the drawings, which are not original, are derived. This, however, is
fully done in a table of illustrations which immediately follows the
table of contents.
158 REVIEWS AND NOTICES OF BOOKS.
The Harveian Oration, 1873. By Grorce Roueston, M.D., F.R.S.,
Linacre Professor of Anatomy and Physiology, and Fellow of
Merton College in the University of Oxford. London, 1873.
In this very able and characteristic oration, full of humour and
quotation, and full likewise of evidence of assiduity and acuteness of
perception, Professor Rolleston first expounds certain advances re-
cently made in the anatomy and physiology of the circulatory organs,
and gives an account of the ‘moderator band’ found by him in the
heart of a cassowary, pointing out the homologous band to be not
unfrequently seen in the human heart. Secondly, he gives “the
as yet unrecorded history of one of the many attempts to rob Harvey
of his rightful rank in the noble army of discoverers, which were
made in the latter half of the seventeenth century.” It is the claims
which have been put forward on bebalf of Walter Warner, the editor
in 1631 of WHarriott's Algebra, to the discovery of the circulation
of the blood. Having by a circuitous route come upon Walter
Warner’s original MS., the Professor gives some quotations from
it which point out “the real merits of the claimant before us,” and
show that Harvey might have truly said
‘“We were the first that ever burst
Into that silent sea.”
The oration concludes with a glowing eulogium of the scientific
character, mental culture and style of Harvey, based upon a perusal
and re-perusal of his works.
The Convolutions of the Human Brain. By Dr ALEXANDER EcKER.
Translated by Joun C. Gatton, M.A. London, 1873.
In our Half-yearly Report on the Progress of Anatomy, rv. p. 157,
we directed attention to Prof. Ecker’s Monograph on tho Cerebral
Convolutions, published in that year, and we briefly criticized some
of the statements contained in it. A translation of this Monograph
by Mr Galton has now appeared, and will be found useful to those
members of the medical profession, happily an increasing number,
who are studying the convolutions of the brain, with the object of
obtaining a more exact knowledge of cerebral physiology and patho-
logy. Mr Galton has done good service in compiling a more com-
plete bibliography than the author had himself appended to his
Monograph.
REPORT ON THE PROGRESS OF ANATOMY.
By Proressor TuRNER’.
OssEous System.—Wenzel Gruber has prepared a Monograph on
the Os Zycomaticum BiparTITUM in men and mammals (pamphlet,
Vienna, 1873). He gives in the first Instance an account of the
cases observed by other anatomists, and then proceeds to describe
ten cases of subdivision of the malar bone in the human subject,
which have come under his own observation, nine of which were
in male crania, and one in a female. In five specimens the sub-
division was on both sides, in two on the right, in three on the left.
The malar bone was subdivided into a superior or orbital, and an
inferior or temporo-maxillary part; the latter of which was always
smaller than the former, and sometimes so small as to be little more
than the inferior border of the bone. In all the cases the zygomatic
process of the temporal articulated with both the orbital and temporo-
maxillary portions. Gruber then describes an example of bipartite
malar bone in Phascolomys wombatus, and the arrangement of the
malar in various mammals.——Gruber also describes VARIATIONS IN
THE ForRAMEN MENTALE (Keichert u. du Bois Reymond’s Archiv, 1872,
738), and to the same Archiv, 1873, p. 195, ¢.s., he contributes a
number of other VARIATIONS IN THE ANATOMY OF THE CRANIAL
Bones, viz. a special ossification at the junction of the malar and
superior maxillary bones; an unusually deep depression of the
incisive fossa of the upper jaw; an adult male skull in which the
pre-maxillary element of the left upper jaw was marked off by a
fissure which extended from the posterior internal end of the palatine
foramen to the posterior internal angle of the lacuna incisiva; a
skull showing accessory processes of the upper jaw prolonged back-
wards to make good the posterior border of the hard palate, which,
on account of imperfect development of the hard palate, would
otherwise have been deficient; two additional ossifications in the
hard palate of a young male skull. On p. 208, a supplementary
memoir to that on the os zygomaticum bipartituwm, already referred
to.——In the same Archiv, p. 649, H. V. Jhering gives an account
of the DEVELOPMENT OF THE FRONTAL Bone. He states that this
bone possesses, in addition to the two chief centres of ossification,
Six accessory pieces; the two least important of which are for the
nasal spine and for the inner wall of the orbit in the region of the
fossa trochlearis, the two more important on the external lateral
angles of the bone. ‘The last are for a time independent bones, but
as a rule are blended with the frontal at the time of birth.
J. Balandin discusses the question of the CURVATURES OF THE
Human Spine (Virchow’s Archiv, vit. 481). After an historical
1 To assist in making the Report complete Professor Turner will be glad
ie receive separate copies of original memoirs and other contributions to Ana-
omy.
160 PROFESSOR TURNER.
introduction, he details his own observations and experiments, and
concludes as follows: The thoracic curve has been observed by him
at the 2nd month of embryo-life. The first indication of consolida-
tion appears in the 4th month. He believes that it is conditioned
in the first instance by the laying down of the skeleton, and that it
is perfected by the pressure of the growing and expanding thoracic
viscera. The cervical curve commences in the 3rd month of extra-
uterine life, and is consolidated in the 4th to the 5th month;
that is at the time when the infant sitting in the arms of its nurse
raises its head and removes the chin from the thorax. The lumbar
curve appears at the end of the first, or the beginning of the 2nd
year. It is not usually consolidated until growth is completed.
It takes place, therefore, at the time when the child assumes the
erect position by the extension of the lower limbs on the trunk.
Both the cervical and lumbar curves are occasioned by the action of
muscles on the spine.
Muscutar System. — P. Lesshaft investigates (Reichert u. du
Bois Reymond’s Archiv, 1873, No. 1) the arrangement of some of
the MusciEes AND FASCI# IN THE REGION OF THE URETHRA. He has
dissected the perineum in 210 subjects, and eighty specimens of
perineal organs after removal. His conclusions are as follows: The
M. constrictor urethre membranacee seu Wilsoni encircles the pars
membranosa, arises from the walls of the venous plexus of Santorini,
lies on both sides of the urethra, and ends in the superior process of
the septum perineale. The upper part of the muscle reaches the
sides of the prostate. It acts as a constrictor of the urethra, and,
from its attachment to the walls of the veins, as a relaxator penis.
It is separated from the inner layer of the levator ani by a definite
membrane. Three Mm. transversi perinei are situated between the
inner wall of the pelvis and the perineal septum ; the superficialis
occurs only as a rare anomaly in 7°‘74 per cent. of the subjects
examined; the medius is wanting only in exceptional cases, it is
the superficial transverse muscle of authors; the profundus is the
most constant. They tighten the perineal fascia, and the profundus
acts indeed as a dilatator urethre and compressor of Cowper’s glands.
The JL transversus urethre is placed in front of the urethra; it
arises from the inner surface of the descending pubic ramus, and
ends in front of the middle of the urethra: one fasciculus commonly
passes above the vena dorsalis penis to the fascia penis. It acts as a
dilatator urethra, and contributes, by compressing the dorsal vein,
and by tightening the fascia penis, to erection. The caput acces-
sorium M, bulbo-cavernosus is an anomalous head of the J. bulbo-
cavernosus, which arises from the tuber or ramus ischil: it les in
the same layer as the Jf. transv. perinet medius. The pelvic fascia
stretched between the pelvic walls, the rectum and the bladder, gives
off on each side two lateral descending processes, an internal and an-
external, and a median process. The internal obturator muscle
lies between the side wall of the pelvis and the external descending
process. The ano-perineal fascia is a prolongation of the fascia.
REPORT ON THE PROGRESS OF ANATOMY. 161
glutea: it subdivides behind the border of the m. transversus medius
into a superficial Jamina, which passes forwards into the fascia penis,
and a lamina profunda, which ends in the pubic arch. The pelvic
fascia, the anterior part of its internal descending process, its median
process, the lamina profunda of the perineal fascia, and the lower
inner portion of the synchondrosis pubis, form a capsula prostato-
urethralis, in which the prostate, the membranous part of the
urethra, Cowper’s glands, the colliculus bulbi intermedius, the
Mm. constrictores, the Mm. transversi perinei profundi, and trans-
versi urethre, the internal pudendal nerves and vessels, the pro3-
tatico-urethral, bulbous and bulbo-urethral veins, are enclosed.
In the same Archiv, p. 126, A. v. Brunn describes a variety of the
MM. interosseous dorsal. manus II, which possesses a third head
arising from the dorsum of the unciform bone. Henry 8. Williams
compares (Trans. Connecticut Acad. of Arts, and Se. II., Part 2
1873) the Muscres or THE Human AND CHELONIAN SHOULDER-
GIRDLES. His object has been to show the importance of the posi-
tions and relations to each other and to the axes of the bones of the
areas of origin and insertion of muscles. The teres major and
scapular part of the latissimus in man he regards as homologous with
the chelonian teres major; the teres minor with the m. scapulo-
acromio-humeralis; the deltoid with a muscle arising from near the
scapulo-acromial angle to near the medial extremity of the acromion ;
the supra-spinatus with the M. acromio-humeralis secundus, and
M. coraco-humeralis secundus; the coraco-brachialis with the M.
coraco-humeralis primus; the biceps with the M. coraco-ulnaris and
M. coraco-radialis; the subscapularis with the MM. scapulo-humeralis
secundus and tertius: the long head of the triceps with the chelonian
muscle which arises from the rim of the glenoid cavity on the outer
side, and which, after joining a bundle which possesses a humeral
origin, is inserted into the dorsal side of the proximal head of the
ulna. An abstract of a Memoir on the MrecuanicaL ConpbiTIons
OF THE ResprravoRY Movements in Man, by A. Ransome, is in
Proc. Roy. Soc. Lond., Noy. 21, 1872.——G. R. Wagener concludes,
from his observations on the SrRUCTURE OF TRANSVERSELY STRIPED
Muscie (Schultze’s Archiv, 1x. 712), that the fibrilla is the ultimate
constituent of the fibre. That all the forms of transverse discs take
their origin only out of the subdivision of the contractile substance
in different parts of the fibrilla, and that the intermediate discs
(Zwischenscheiben) are not definite structures——An_ elaborate
memoir by Rudolf Arndt, on the ENDING or NERVES IN TRANSVERSELY
STRIPED MUSCLE-FIBRES, occupies upwards of 100 pages of Schultze’s
Archiv, 1x. 481, and is illustrated by three large plates. He believes
that he can distinguish between the mode of termination of the motor
and sensitive nerves in a muscle. The chief difference between them
being this: the motor-nerves are broad, medullated fibres which ter-
minate within the sarcolemma, and are therefore intra-muscular,
whilst the sensory fibres are narrow fibres, poor in medulla, and in
part pale and non-medullated, which end outside the sarcolemma
either in sensitive ‘plaques, or in excessively delicate penicillated
VOL. VIII. VE
162 PROFESSOR TURNER.
filaments. These are extra-muscular, and not unfrequently the
sensitive nerves encircle the muscular fibre.
ConNnECcTIVE TissuE.—Arnold Spina relates Observations on the
structure of Tendons (Med. Jahrb. 11. Heft 1873). After referring to
the more recent researches on the subject, he describes his method,
and then proceeds to give an account of his own observations, which
may be summarized as follows :—The development of tendon may be
considered in three stages. First Stage. Tendon consists of three
fibrillary fascicles, between which run rows of solid cells. The cells
are embraced by a substance that does not swell with acids, and
refracts light strongly. When teased it appears in the form of
thorn-shaped processes. A transverse section shows it in the form
of a ring, which surrounds the ce!l and sends out membrane-like
processes which separate the connective-tissue fascicles from each
other. Longitudinal sections show this substance as a ring which
surrounds the ellipsoid cells. Second stage. The connective-tissue
fascicles have become thicker, and the rows of cells have become
longer. The cells have become flattened to a domino shape, and
towards the ends of the row they are seen to be gradually narrower,
so that the row is pointed in shape at its extremities. The substance
enclosing the cells has acquired a ladder-shape, the sides of which, at
the extremities of the cell-row, are continued in a single line. But,
besides, threads of the same substance are seen stretching along the
middle line of the cellxow. ‘These cannot be considered as being
anything else than the thickened cell-wall. These terminal and
median threads are to be identified as elastic fibres. The rows of
cells are no longer perfectly parallel, but come into contact with
each other; the threads of one cell-row often becoming continuous
with those of another. The sheaths of the connective-tissue bundles,
visible only in the transverse sections, appear in this stage also as
membranous processes of the cell-wall. Third stage. The rows of
cells have changed to flat bands, some of which are striated longi-
tudinally and transversely; others are non-striated. The striated
appearance is produced by elastic matter which persists in the
above-mentioned ladder-form. ‘These threads can be traced a long
way into the fibrillary tissue, and can be diagnosed as elastic fibres.
They form a wide-meshed net which penetrates the connective tissue,
the meshes being the cause of the known bulgings of the tendon
fascicles. Hn résumé.—(1) The elastic tissue of tendon takes its origin
on the surface of the cells. (2) The fine sheaths in tendon are formed
by membrane-like processes of the cell-wall. (8) The cellrows
become striated and non-striated bands.
EriraeLium.—P. Langerhans communicates (Virchow’s Archiv,
Lvl. 83) some observations on stratified epithelium, more especially
with reference to the distribution of epithelial cells possessing den-
ticulated processes. He figures various examples from the cornea,
rete Malpighii, ceesophageal and vesical epithelium.
Pracenta.—F. Winkler communicates observations on the Human
Placenta (Archiv fiir Gynaekologie, tv. 238). He describes his methods
REPORT ON THE PROGRESS OF ANATOMY. 163
of injecting and hardening the organ. He considers that the feetal
part of the placenta is of no architektonic importance, and applies the
term Netto-placenta to the ideal organ as it would appear after the
elimination of the entire foetal portion, whilst to the complete organ
he gives the name Brutto-placenta. The Netto-placenta has a cavern-
ous structure, bounded on the uterine aspect by the Basa/platte, on
the feetal aspect by the Schlussplatte, between which the wide-meshed
cavernous trabecular structure is situated. The uterine basalplatte
or serotina consists of two layers, an external small-celled and an
internal large-celled. The small-celled layer consists of a preponde-
rating portion of intercellular substance, mostly homogeneous, but
here and there streaked. It is through this layer that the separation
of the mature placenta takes place. The large-celled layer contains
the well-known colossal decidual cells imbedded in a sparse intercel-
lular tissue. The intercellular substance increases In importance on
the feetal aspect of this layer, and the trabecule, which are formed of
homogeneous intercellular substance, with small, round, spindle, rarely
star-shaped, connective-tissue corpuscles scattered in it, are continuous
with and derived from this layer. Except in the cotyledonary septa,
in which for short distances the large cells penetrate, the whole of the
rest of the Wetio-placenta consists of this kind of connective tissue.
The Sch/ussplatte is a subchorionic layer, and bounds the blood-caverns
on the feetal aspect of the placenta; like the inner layer of the Basal-
platte, it consists of homogeneous intercellular substance with cells
scattered in it. Vertical trabecule extend through the placenta
from the basal to the schlussplatte, from these vertical bands trans-
verse or oblique trabecles proceed, to trace which outwards is almost
impossible on account of their delicacy. The basalplatte, the schluas-
platte, and the trabecles, bound the maternal blood-spaces, which are
every where, except where large villi have broken through the wall, lined
by anendothelium. The placenta possesses in its basalplatte, especially
at the places of intersection of the cotyledonary septa, apertures
through which the cavernous spaces within the placenta communicate
with the venous sinuses of the uterus, which apertures open either
directly or by a sort of vascular tube three or four J/m. long into the
blood-caverns. Winkler believes that the histological detail supports
the opinion of Virchow, that all the vascular spaces of the Wetto-pla-
centa consist of ectatic capillaries with consecutive cavernous for-
mations. He believes that muscle-elements exist in the layer of
colossal cells of the serotina. He distinguishes three kinds of chori-
onic villi: a. those which become obliterated and end in the schluss-
platte, without ever penetrating into the blood-caverns: 0b. short
vascular villi which end directly in connection with the blood-caverns
lying immediately under the schlussplatte; only that portion of these
villi which hangs free in the blood-caverns possesses a recogniz-
able epithelial envelope: c. long and many-branched vascular villi
which penetrate deeply into the placenta, some reaching as far as the
basalplatte. These villi are either imbedded in the maternal tissue of
the placental trabecule, in which case they are without epithelium;
or they lie free in the cavernous spaces, and then possess the well-
11—2
164 PROFESSOR TURNER.
known epithelial investment. In their whole extent the stems of
the villi are intimately connected with the placental trabecule.
He believes that a system of juice-canals (Saftcandlchen) permeates
the connective tissue of the chorion, the gelatine of Wharton, and even
the amnion, the epithelial layer of which possesses apertures like the
serous investment of the diaphragm. He finds these juice-canals to
be connected with the blood-vessels, as Carter (see this Journal,
Iv. p. 97) had experimentally shown in other tissues. Winkler
opposes the statement of Reitz that the blood-vessels of the villi lie
in perivascular spaces. From the examination of a nine-weeks old
abortion he describes the maternal part as consisting of a small-celled
tissue in which many spaces exist, lined by a single layer of epi-
thelium, which is often cylindriform in shape. These spaces he
regards as utricular glands, and into them villi, presenting the usual
epithelial investment, project, though without filling the space. Other
villi projected simply into the maternal tissue ‘without being con-
tained in a space lined by epithelium; he considers that these villi
have burst through the walls of the glands in the process of develop-
ment and growth of their various branches. By a continuance of
this process of growth they then press against the walls of the mater-
nal blood-spaces, and parting asunder the endothelial cells, project into
the blood-caverns. As the villi in the mature placenta which are
invested by the maternal tissues are naked, the pressure of growth
must have made to disappear both the epithelium proper to the villus,
and that of the utricular glands. He entirely disagrees with Fried-
linder’s view, that utricular glands exist in the placental area, and has
never seen a trace of a gland either in the serotina or placental trabecule,
neither has he seen the double layer of cells on the villi which Jassinsky
(Report 111. p.- 203) has described. C. Hennig publishes’ researches
on the Human Placenta (pamphlet, Leipzig, 1872). He makes a care-
ful examination of a uterus in the third month of gestation, in which
he found no sharp line of demarcation between the placental and
non-placental part of the fatal membranes. The uterine mucous
membrane has considerable thickness, and in addition to a cavernous
venous reticulation possesses glands both in the decidua vera and
serotina. They disappear from the reflexa in the earlier months, and
in the later months they are no longer recognisable with certainty
beyond the placental area, the only indications of their existence
being the cribriform apertures of the decidua; their ciliated epithe-
lium is much more difficult to be recognised than on the free surface
of the mucous membrane, and appears to be converted both in the
serotina and vera into tessellated scaly epithelium. The chorionic
villi consist of a gelatinous connective tissue containing corpuscles, of
the fcetal vessels, and of a foetal epithelium consisting of a nucleated
pavement epithelium, which sometimes peels off like a glove from a
finger: this epithelium is the internal cells of Goodsir, the inner epi-
thelium of van der Kolk. Further, in many localities, though he
admits not universally, one or two other coverings are found: a
maternal epithelium of serotina cells, the outer cells of Goodsir and
van der Kolk, the gland-organ of Ercolani; and the wall of the
hee SA erway 6.
.
REPORT ON THE PROGRESS OF ANATOMY. 165
maternal vessel—the external membrane of Goodsir and van der
Kolk. The villi of the chorion grow through the reflexa and vera,
and penetrate in the later months of pregnancy into the serotina.
At the end of pregnancy the serotina forms the thin layer which
covers the placental spot and the covering of the uterine aspect of the
placenta, from which latter proceed the processes which penetrate
between the cotyledons. Although on the inner surface of a uterus
from which the placenta has just been separated the blind ends of
young utricular glands are situated, the tubes of these glands cannot
be seen in the placental part of the serotina. For not only the glands
themselves, but the interglandular tissue, are converted in the latter
half of pregnancy into a framework of delicate columns and tubular
spaces between, and in which epithelial-like and often nucleated
colossal cells are situated. These columns and tubular spaces arise
partly from the basis tissue (Grundgewebe) of the placenta, indeed
from the connective tissue of the serotina and partly from new-formed
glands, which cannot however be recognised as such, as they are
imbedded in the growing cells, and their epithelium is completely
modified. The serotina cells arise partly from the endothelium of
the young glands, partly from the connective-tissue corpuscle of the
intermediate tissue, and probably from cells which have migrated
out of the blood-vessels. The largest of these cells have several nuclei,
as many as nine have been seen, which nuclei are disposed like a
cylindrical. epithelium: other modifications of these cells are also
described. Hennig concludes his account of the structure of the
placenta with the following statement: ‘The chorionic villi reach
in the second and third months the decidua reflexa, through which
they grow, then they penetrate the vera, as Spiegelberg had already
shown, and in the last months pass through the most inferior
parts of the serotina, not only into the mouths of the glands, but
into spaces in the intermediate tissue: finally, they penetrate
the blood-vessels as when they enter the marginal sinus of the
placenta.” In consequence of the observations of W. Turner on
the placenta of Orea gladiator (Abstract in Report, v. 383 and vi. 439),
and of Winkler and Hennig on the human placenta (Abstracts as
above), G. B. Ercolani has been led to re-investigate the structure of the
Placenta in the Mammalia (Mem. dell’ Accad. d. Scienze di Bologna.
T. 111. 1873), more especially with reference to the determination of the
part which the utricular glands take in the formation of the maternal
part of the placenta and in the nutrition of the fetus. In his pre-
vious very elaborate memoirs (Abstract in Report, v1. 439), Ercolani
had maintained that the utricular glands furnish materials for the
nutrition of the embryo only in the early period of its development,
and that the villi do not penetrate into their mouths; but that sub-
sequently a new maternal glandular organ is formed, consisting of
secreting follicles, lined by a pavement epithelium, into which the villi
of the chorion fit, and from the secretion formed by which the foetus
is nourished. In his new memoir he describes additional researches
on the sow, and summarises the differences between the non-gravid
and gravid uterus of that animal as follows:—Ist. A want in the
166 PROFESSOR TURNER.
gravid of the well-marked rug of mucous membrane with numerous
folds such as are seen in the non-gravid. 2nd. A remarkable increase
in the gravid of the muscular layer and of the blood-vessels, especially
those of the mucosa. 3rd. A diminution in thickness in the
gravid of the glandular layer. 4th. A change in the internal super-
ficies of the mucosa, which is smooth in the non-gravid, uniformly
subdivided into fine ridges and furrows in the gravid uterus. It is
into these furrows, which constitute the new-formed glandular organ,
that the villi of the chorion are lodged. 5th. The utricular glands
are more developed in the gravid, which he attributes to being more
distended with contents, and to an increased thickness of the epithe-
lium. In the non-gravid the cylindrical epithelium of the glands is
twice the thickness “of that covering the free surface of the mucosa.
In the gravid uterus the epithelium covering the free surface of the
mucosa “is pavement-like when seen on its free surface. The glandular
organ in the sow is not so completely subdivided into distinct crypts
as was described by Turner in Orca, for the dissepiments or ridges
between the crypt-like furrows are less perfectly decekpad Scat-
tered over the surface of the gravid mucosa are circular areas circum-
scribed by a raised border formed by the rounded ends of the minute
ridges of the mucosa: in the centre of each of these areas the mouth
of an utricular gland opens; the cylindrical epithelium of the gland
contrasts with the pavement-like epithelial covering of the area itself.
He considers that the mouths of these glands are plugged up during
the early period of pregnancy by rounded growths such as Eschricht
had described, which are not villi, from the surface of that part of the
chorion which lies opposite the mouths of the glands; whilst in the
later periods the hyperplasy of the connective tissue of the mucosa
around the mouths of the glands completely obliterates their apertures.
Hence he concludes that natural obstacles exist to the flow of the
secretion of these glands out of their mouths, so that it cannot be
absorbed by the villi, and does not take a part in the nutrition of the
foetus. He admits however that in the mare and ass the utricular
glands do secrete freely during the whole period of pregnancy. He
meets the objection raised by Turner to the secreting function of the
follicles or crypts of the new-formed organ, because they are lined by
a pavement epithelium, by the statement that the objection is “eli-
minated by the fact of the constant and complete change in the physi-
cal and anatomical qualities of the epithelium in the uterus of the
sow, which, from being cylindrical as it is in the unimpregnated
uterus, becomes pavement-like in pregnancy,” and by the further
statement, that there are other instances in the vertebrata of secreting
organs with pavement epithelium, as the sebaceous, sudoriparous and
ceruminous glands. As an additional illustration that the utricular
glands are not concerned in placental formation Ercolani adduces the
structure and mode of growth of the cotyledons in the sheep, in which
in the non-gravid state the position of the future cotyledons is mapped
out on the mucosa by areas in which the utricular glands are absent,
though they exist in abundance at the borders of these areas. In the
changes which take place during pregnancy a new formed glandular
REPORT ON THE PROGRESS OF ANATOMY. 167
organ .takes place in the cotyledon by an abundant cell-production,
producing follicles into which the villi of the chorion fit. He states
that his previous observations on the production of w new-formed
maternal cellular organ in the early and later stages have been con-
firmed by Romiti in the intermediate stages. In opposition to Leydig,
who holds that utricular glands do not exist in the Muridz, he states
that he has seen them, though few in number, in the uterine mucosa
of Mus musculus. They are simple, slightly sinuous, not very long,
and have a cylindrical epithelium. The epithelium is shed from the
areas to which the ova are attached, and a rich new formation of cells
takes place, by which the ova are embraced: this formation is not
derived from the utricular glands, because in these localities they lose
their internal epithelium during pregnancy, are augmented in volume,
and show in their interior an amorphous transparent substance. In
the non-placental parts of the mucosa the glands undergo no change.
The Hydroperion in the human female possesses a nutritive function,
and is secreted by the utricular glands in the earliest period of gesta-
tion. The disappearance of the hydroperion, and the impossibility of
demonstrating utricular glands in the later period, he considers as
correlated conditions. He believes, in opposition to Hennig, that the
utricular glands in the human female take no part in the formation of
the placenta. A, H. Garrod notes (Proc. Zool. Soc. Lond., Nov.
19, 1872) some points on the Placenta of the Hippopotamus. The
placenta is a strong cylindrical bag 34 feet long. The bag had rup-
tured during parturition at the end opposite the os uteri. The outer
surface of the bag is uniformly covered with villi of a bright red
colour, at the closed end there is no reduction in their number, but
at the ruptured end they are paler and more scattered. The umbili-
cal cord is attached to the placenta about halfway between its two
ends. Itis 1} foot long and 1} inch in diameter in the middle, and gets
larger as it approaches its attachment, near which there are many
spherical bodies, as big as peas, yellow in colour, supported on short
amniotic pedicles. In the course of a description of an early Human
Embryo in the vesicular stage of development, C. B. Reichert (Archiv,
1873, 127) states some facts about the Decidua. The embryo was
believed to be at the 12th or 13th day. The ovum is completely
enclosed by the decidua reflexa. The decidua vera is developed from
the decidua menstrualis by the formation of cotyledonary elevations
or islands in the uterine mucous membrane, and by a remarkable
growth of primary and secondary papillary processes on the surface of
these islands. At the margin of these islands the uterine mucosa is
smooth, and shows most distinctly the widened orifices of the uterine
glands. The cotyledonary walls of the d. vera form an equilateral
triangle with the apex at the cervix uteri, and are separated by a
median cleft into two halves; and a bilaterally symmetrical arrange-
ment is indicated in the other islands. In the posterior wall, which
alone was preserved, eight islands, more or less irregularly polyhedral
and separated by furrows, were seen. The embryo was imbedded in
the parenchyma of one of the islands. On separating it several small
villi were drawn out of the ducts of the utricular glands. The decidua
168 PROFESSOR TURNER.
reflexa possessed on its inner concave surface utricular glands, although
the villi had especially grown from the marginal zone of the ovum;
likewise a short cylindrical epithelium, free from cilia, was found
here which passed into the epithelium of the utricular glands. -The
reflexa has only one independent wall in which the scar indicative of
the closing in of the ovigerous chamber could be recognised, but in it
no utricular glands could be seen: the remaining part in which the
mouths of the glands were seen could not be separated by any line of
demarcation from the parenchyma of the islands. The wall of the
chamber next the embryo he names the basilar wall, and although in
it utricular glands open, yet here, in the region of the embryonal spot
of M. Coste, no villi pass into them. The view generally entertained
that the reflexa is formed around the ovum resting on the vera by the
elevation of a circular wall is untenable. It would appear from this
specimen that the spot on the island, to which the ovum attaches
itself, does not increase at the same rate as the rest of the island, in
consequence of which a cup-shaped depression is formed which con-
stitutes the basilar wall and marginal zone of the reflexa; a general
growth of the free edge of the cup over the free surface of the ovum
then takes place, which closes in the chamber and forms the free
wall of the reflexa. The paper concludes with some observations on the
stage of development of the embryo itself.
Evpryotocy.—F. M. Balfour has recently published three inte-
resting embryological memoirs (Quart. Journ. Mic. Sc. July, 1873).
In one he relates the DEVELOPMENT AND GROWTH OF THE LAYERS OF
THE BLAsTODERM OF THE HeEn’s Kec. He first describes the unin-
cubated blastoderm as consisting of two layers of cells; a superficial
single row of nucleated columnar cells, and a deeper layer of several
rows of rounded non-nucleated granular cells, varying from ;4,5 to
2009 Wech in diameter ; whilst quite at the bottom of the segmenta-
tion cavity “formative cells” ~*- inch in diameter, and containing
highly refracting spherules, are situated. 6 to 8 hours after incubation
a hypoblast and a mesoblast are formed, whilst the superficial
columnar layer forms the epiblast. The hypoblast is formed by a
metamorphosis of a number of the cells of the deeper layer from their
originally spherical form into flattened nucleated cells. Enclosed
between the epi- and hypoblast are found numerous cells of the
originally deeper layer, together with the “ formative cells,” which, as
Paremeschko, Oellacher and Klein had previously shown, begin to
travel towards the circumference and then pass in between the epi-
and hypoblast. Contrary to the opinion of the above observers,
Balfour believes that the cells of the mesoblast are derived, not merely
from the “formative cells,” but both from them and the enclosed cells
of the deeper layer, by a process of conversion into new cells, though
as to the exact manner of conversion he is not quite sure, but thinks
that the spherules of the original cells develope into the nuclei, whilst
the protoplasm of the new cells is formed from that of the original
cells. The mesoblast in the chick is formed coincidentally with the
hypoblast out of apparently similar segmentation cells, A kind of
REPORT ON THE PROGRESS OF ANATOMY. 169
fusion, as development goes on, takes place between it and the epiblast
along the line of the primitive streak, to form the axis-string of His.
Its growth is effected by means of the “ formative cells,” which rapidly
increase, probably by division, and act as carriers of food from the
white yolk to the mesoblast, till the formation of the vascular area,
when they are no longer necessary. The growth of the hypoblast is
by direct conversion, cell for cell, of the white yolk spheres into hypo-
blast cells. The epiblast cells increase entirely by division, and the
new material is most probably derived directly from the white yolk.
In his second memoir Balfour supports, in opposition to the opinion
generally entertained by embryologists, the view originally propounded
by Dursy (Der Primitivstreif des Hiihnchens, Lahr, 1866), that THe
PRIMITIVE GROOVE Is A TEMPORARY STRUCTURE, and has no connection
with the development of the neural canal, or indeed with any part of
the future chick. This groove is distinguished in transverse sections
by the epiblast and mesoblast being fused together beneath it, by the
epiblast not becoming thinner where it lines the groove, and by the
mesoblast beneath it never showing any sign of being differentiated
into a chorda dorsalis or other organ. Near the time when the
primitive groove has reached its full length, there appears at about
the 16th hour, or a little later, altogether in front of, and non-
continuous with it, a thickening of the mesoblast, which forms an
opaque streak, the medullary streak ; and near the anterior extremity
of the area pellucida arises a semicircular fold, against which the
medullary streak ends abruptly. This fold is the head fold, and a
groove which extends along the central line of the streak is the
medullary groove, which subsequently forms the cavity of the neural
canal. As the medullary groove increases in size, the primitive
groove diminishes and is pushed backwards: at about 36 hours only
a small and curved remnant is to be seen behind the sinus rhomboti-
dalis, though Dursy has been able to distinguish it up to the 49th
hour. The medullary groove widens and deepens, its margins, by a
thickening of the mesoblast, are elevated into the medullary folds,
which broaden and finally close in the neural canal. The epiblast
becomes thinner where it lines the canal ; there is no fusion between
it and the mesoblast as beneath the primitive groove; and the noto-
chord begins to be differentiated out of the cells of the mesoblast.
In his third memoir Balfour describes the DEVELOPMENT OF THE BLoop-
VESSELS OF THE Cuick. He investigates the development of the
blood-vessels in the area pellucida. He agrees with Klein in the
fundamental fact that some of the cells of the mesoblast send out
processes which unite with processes from other cells and form a
network. The nuclei of the original cells divide, and at the points
from which the processes start their division is especially rapid. Some
nuclei acquire, especially at these points, a red colour and become red
blood-corpuscles, others, together with that part of the protoplasm in
which they are imbedded, become converted into an endothelium
both for the processes and the masses of blood-corpuscles ; the remain-
ing protoplasm becomes fluid, and thus the original network of cells
is converted into a network of hollow vessels, filled with fluid, in
170 PROFESSOR TURNER.
which corpuscles float. From their mode of formation, the blood-
corpuscles of the Sauropsida are, he believes, to be looked upon as
nuclei containing nucleoli, rather than as cells containing nuclei,
which would make them to be morphologically as well as functionally
homologous with the mammalian red-corpuscles. This view of the
formation of capillaries, by the hollowing out of an anastomosing cell
network, agrees closely with the observations of Julius Arnold, of
which the Reporter has given an abstract in Vol. vr. of this Journal,
p. 437. Balfour also agrees with Klein in the mode of formation
of.a secondary investment of the capillaries from cells of the meso-
blast, situated in the meshes of the capillary system, which flatten
and send out processes, and look spindle-shaped on section. The
cavity of the heart is produced by a splitting or absorption of central
cells of the thickened mesoblast of the splanchnopleure, while its
muscular walls are formed from the remaining cells of this thickened
portion. M. Balbiani communicates (Ann. des Se. Nat. 1873) an
elaborate and beautifully illustrated memoir on the EmpryoLocy OF
THE ARANEINA. E. Ray Lankester discusses (Ann. Nat. Hist.,
May, 1873) the Primitive CELL-LAYERS OF THE Empryo as the basis
of a genealogical Classification of Animals. Regarded from the stand-
point of their development, animal organisms may be grouped in
three classes: A. the Homoblastica, in which there is no arrangement
of the cell-units into definite layers. This class coincides with the
Protozoa, the Sponges being excluded. B. the Diploblastica, in which
two layers of cells forming the ectoderm and endoderm remain
throughout life, as the basis of histological differentiation, To this
class belong the Coelenterata, including the Sponges. C. the Triplo-
blastica possess, like B, an ectoderm or epiblast, and an endoderm or
hypoblast, but a third layer of cells, or mesoblast, appears between
the two. This class comprises Vermes, Echinodermata, Mollusca,
Vertebrata, Arthropoda. A. Gotte makes some contributions
(Schultze’s Archiv, 1x. 679) to the DEVELOPMENT OF THE VERTEBRATA.
This paper contains the results of his observations on the blastoderm
of the trout’s egg. K. Slavjansky describes (Ludwiy’s Arbetten,
1872) the RetrocressivE CHANGES which occur in the EPITHELIAL
CELLS of the Serous Layer or THE Ovum oF THE Rassit. The
protoplasm of the cells in the first instance begins to disappear, holes
form in the cells which gradually enlarge, and what remains of the
protoplasm forms a branched and anastomosing network within
the cell, the branches seeming to radiate from a ceutral mass of
protoplasm. The spaces between the meshes of the network are
empty. Ina later stage the individuality of the cells is lost by the
disappearance of their houndaries, and a protoplasmic network is
formed by the junction of the meshes of adjacent cells with each
other. He calls the degeneration reticular, and considers it to be a
physiological prototype of the pathological changes in epithelium seen
by Wagner in cases of croup and diphtheria,
Matrormations.—M. Flesch describes (Virchow’s Archiv, Lvit.
289) a malformation of the thorax, in which there is a depression
REPORT ON THE PROGRESS OF ANATOMY. 171
of the sternal wall, and a diminution of the sterno-vertebral dia-
meter of the chest, the transverse diameter is, however, increased.
C. C. Th. Litzmann gives an account (Archiv fiir Gynackologie,
Iv. 266) of a woman, aged 24, who died in child-bed, and in whom
there was extroversio vesicae, and non-union of the pubic bones at the
symphysis. P. Gervais in his Journal, u. 1873, gives an account
of Polygnatic and Heterognathic Monsters. —— R. Hein describes
(Virchow’s Archiv, Lv. 326) a foetus in which through defect of the
anterior-wall of the abdomen there was ectopia viscerum and imperfect
development of the extremities. R. Jaensch describes a specimen
of pregnancy in a rudimentary uterine cornu (Virchow’s Archiv,
Lyi. 185). The woman died about the 4th month from rupture
and internal hemorrhage, and on post-mortem examination the left
uterine cornu was found to be pregnant. In this case no communica-
tion was found between the pregnant horn and the right uterus,
so that it agrees closely with the two cases described by W. Turner in
Edinburgh Med. Jowrn. May, 1866. The author adopts the hy-
pothesis, supported by Turner, of the mode in which impregnation
took place, viz. by an extra-uterine transmigration of the seminal
fluid.
Nervous System.—V. Butzke investigates (Archiv f. Psychiatrie,
1872, ut.) the Minute Structure oF THE CEREBRAL CONVOLUTIONS.
The nerve-cells possess a longitudinal striation, a character which
the author considers to be distinctive of nerve-cells. The axile
prolongation of Deiters either joins the cell directly, or through the
intermediation of an offshoot from the base of the cell. He has
never seen any of the other processes of the cell continued into
nerve-fibres. He thinks that these processes divide, and form very fine
fibrille, which constitute a terminal network. He also investigates
the cell-forms of the Neuroglia.——Gerlach and E. Rindfleisch dis-
cuss (Centralblatt, 1872, 273 and 277) the mode of TERMINATION of the
NERVES in the THE GREY CorTEX OF THE CEREBRUM. Gerlach holds
that the non-medullated nerve-fibres of the cortex, which possess a
transverse direction, form a wide-meshed network along with the
radiating fibres situated in the plane of the pyramidal cells. These
meshes are occupied by an extremely fine secondary network of
delicate non-medullated fibrille. In this network the protoplasmic
prolongations of the nerve-cells also end; so that it forms the
medium of connection between the nerve-fibres and ganglion cells.
Rindfleisch does not admit the very fine network of fibrille which
Gerlach describes, but considers that the nerve-fibres, after a series
of dichotomous divisions, terminate in the granular amorphous
material of the grey matter in a penicillated ramification of very
minute fibrille. The ends of the branched processes of the pyramidal
cells also terminate, he believes, in the same amorphous material,
which acts therefore as the medium of communication between
nerve-cells and nerve-fibres. Franz Boll has published (Archiv
fiir Psychiatrie, tv. and separate pamphlet, Berlin, 1873) an elaborate
memoir on the HistoLocy AND HIsToGENESIS OF THE CENTRAL
172 PROFESSOR TURNER.
OrGANS OF THE Nervous System. In his first chapter he discusses
the nature of the connective tissue of these organs, and in succeeding
chapters the minute structure of the nerve-elements in the spinal
cord, the white substance of the brain, and the grey matter of the
cerebrum and cerebellum. He examines also the perivascular and
epicerebral spaces, and the development of the central organs of
the nervous system. He considers that he has established two
general conclusions as to the connective tissue: first, that the view
propounded originally by Henle and supported by Rindfleisch of the
nervous nature of the molecular material, in which the nerve-cells,
fibres and blood-vessels are imbedded, cannot be substantiated; that
the “granules” (Kérner) in that material are not, as Henle and
Merkel contended, indifferent elements capable of being developed
either into connective tissue or nerve-corpuscles, but are in the white
substance distinct connective tissue-cells, and in the grey matter
distinct nuclei, mostly with a characteristic double contour, Secondly,
that there is a general unity of structure of the connective tissue in
the different central organs of the nervous system. The apparent
differences arise from this: that in the conversion of the protoplasm
of the embryonic cells into connective tissue, a greater or less pro-
portion of granular albuminous substance remains between the
fibrille which arise out of the protoplasm. As there are differences
in the relative proportion of the fibrillee and the granular albuminous
material in various parts, so different appearances are occasioned.
The cells of Deiters which occur in the white matter of the spinal
marrow of the ox, &c., are obviously only embryonal cells, which
in a preponderating manner are converted into connective-tissue
fibrille, and very slightly into granular albuminous substance. In
smaller animals again these substances occur in more eqnal propor-
tions. The white substance of the cerebrum is very analogous to
that of the spinal marrow. Cells of Deiters, however, occur in regions
such as the grey matter of the cerebrum, where the connective tissue
exhibits a preponderance of the granular albuminous substance. These
cells in their distribution follow closely that of the blood-vessels. He
believes that under the term perivascular lymph-spaces, observers
have hitherto mingled together two different kinds of hollow spaces;
one adventitious lymph-spaces which have a physiological import,
and communicate with the lymph-vessels of the pia; the other, the
perivascular spaces which are not lymph-vessels, but artificial produc-
tions. He considers that one common anatomical principle regulates
the connection of the nerve-fibres with the nerve-cells in the grey
matter of the spinal cord, of the cerebral convolutions, and of the
cerebellum. In each case a cell is connected with a well-marked
medullated fibre, by the “ axis-cylinder process,” but in addition,
the “ branched processes” break up into delicate threads, which be-
come continuous with a very fine nervous network, from which
medullated nerve-fibres spring. He has never seen an anastomosis
between ganglion cells in the sense usually expressed, viz. by a direct
junction between a strong process of one cell and a corresponding
process of an adjacent cell——In Virchow’s Archiv, Lvmi. p. 259,
REPORT ON THE PROGRESS OF ANATOMY. 173
M. Roth communicates observations On Varicose HyprertropHy
OF THE NERVE-FIBRES OF THE BRAIN; on p. 323, O. Obermeier
refers to VARICOSE AXIAL CYLINDERS in the central nervous system ;
and on p. 310, T. Simon describes a NEw Formation or Brain-
SuBSTANCE, in the form of tumours on the surface of the cerebral
conyolutions—— W. Turner considers the CONVOLUTIONS OF THE
Human Brain IN RELATION TO THE INTELLIGENCE (West Riding
Asylum Reports, 1873). He arranges his remarks under the heads
mass and weight, external configuration, internal structure, vascular
supply, and concludes with some observations on the organology of
the convolutions. In the same feports, p. 97, H. C. Major makes
observations on the HistoLocy OF THE BRAIN IN THE INSANE, and on
p- 285, W. C. 8. Clapham communicates the results of the WEIGHINGS
OF THE Brain in 716 cases. L. Gerlach gives an account (Ludwig's
Arbeiten, 1872) of the Myrenteric PLexus or AverRBAcH. He de-
scribes ganglia in it as well as larger and smaller bands of nerve-fibres
which form primary and secondary networks. From the secondary
network very fine nerves arise, each of which ends in a corpuscle,
which may give off one or two processes; these processes end between
the smooth muscular fibres. The ganglia are very vascular.
E. Klein contributes (Quart. Mic. Journ, Oct. 1873) a short paper
on AUERBACH’S PiLeExus in the Jntestine of the Krog and Toad. In
addition to the isolated groups of ganglion-cells, he describes other
ganglion-cells between the circular and longitudinal muscular coats.
Eyr-patt.—A. T. Norton investigates the anatomy of the
Citiary Bopy, and the accommodation of the eye to vision (P.
toy. Soc. L., June 19, 1873), His paper is to show that the increase
in the convexity of the lens, when accommodated for near vision, is
effected by compression of its equator by the ciliary processes
turged with blood, the ciliary muscle being the motor agent: the iris
aids accommodation by increasing its rapidity, but accommodation
can be effected slowly without the aid of the iris. R. J. Lee
communicates P. Roy. Soc. Z., Jan. 9, 1873, some further remarks
on the Sense or Sicut 1x Birps.——A. V. Gruenhagen gives in
Schultze’s Archiv, 1x. an account of the posterior surface of the Iris.
Bioop Aynp Lympa VascuLar System.—Paul Langerhans con-
tributes (Virchow’s Archiv, Lyi. 65) to the HistoLoGy OF THE
Heart. He first describes the characters of the muscular fibre.
He figures the branched muscle-cells from the heart of the Sala-
mander, also the appearance of the transverse striation in Leu-
ciscus, the frog and the human foetus. He points out also the
relation of the nerves to the muscle-cells. George Rolleston in
the Harveian Oration (Brit. Med. Journ. July 5 and 12, 1873, and
separate pamphlet) describes a ‘ MoDERaTOR BanpD’ in the HEART of
the Cassowary, which stretches across the cavity of the right ven-
tricle from the septum to the moveable wall of the ventricle. He
points out its morphological relations to a similar band in the
heart of the sheep, and to a band in the human right ventricle,
174 PROFESSOR TURNER.
which when well developed extends from the base of the muse.
papillaris arising from the outer or moveable wall of the ventricle
towards the conus arteriosus. C. Giacomini communicates a
memoir on the VEINS oF THE [INFERIOR Extremity (Giornale della
Accad. di Med. di Torino, 1873). Not only does he consider the
usual arrangement of the veins, but he also describes cases of
variation from the normal mode of disposition. He refers also to
their comparative anatomy, and describes the venous system in the
hind-lmbs of Cercopitheci. He concludes with a chapter on some
general deductions from his observations. F. Schmuziger figures
and describes (Schulize’s Archiv, 1x. 709) the Micration of the Rep
AND WuHiTE Bioop-corpuscLtres from the mesenteric vessels of the
froy——A. Heller (Ludwig's Arbeiten, 1872) investigates the
BLcop-VESSELS OF THE SMALL INTESTINE in man and several mam-
mals. Each villus as a rule contains an unbranched artery extending
to the tip of the villus. In man only does it begin to form a
capillary network about the middle of the villus. The vein begins
either at the tip of the villus (rabbit, man), or near the tip (rat),
and passes as a rule without lateral twigs directly into the sub-
mucosa; or it arises near the base of the villus and receives late-
ral twigs out of the glandular layer (dog, cat, pig, hedgehog). In
none of the animals examined was the plan usually described
seen, of an ascending artery, a descending vein, and of a complex
capillary network connecting together these vessels. In the
same Arbeiten, J. Michel enquires into the BLoop anp LymMPH-CHAN-
NELS of the CEREBRAL Dura Mater. The peculiarity of the anatomical
arrangement of the blood-channels of the dura consists in this: that
the arterial capillary net opens into two venous systems, of which
the stronger is found on the outer surface, the feebler on the inner,
but which communicate with each other by branches proceeding from
the network on the inner surface which traverse the tissue of the
dura. The space between the dura and arachnoid (arachnoid cavity)
does not communicate with the vessels of the dura, specially not
with the blood-vascular net on its inner surface. A system of
inter-communicating spaces exists throughout the entire thickness
of the dura which is in connection with the subdural space, as
well as with a number of larger and smaller spaces situated between
the dura and the bone, which may be called epidural; both on the
inner and outer surfaces of the dura an endothelium exists, which
forms the inner boundary of the epi-dural and the outer boundary
of the sub-dural space, whilst the spaces within the dura are also
clothed by an endothelium. This system of spaces he believes to
be employed in the passage of lymph. To the same Arbeiten
also Schwalbe communicates a short note on the LyMPH-CHANNELS
of the Retina Anp Virreous Bopy.
ReEsPrrATORY System.—Sam. Pozzi records (Revue d’Anthropolo-
gie, 1872, 443) the case of a right human lung which possesses a
lobus impar. It adheres to the inferior lobe of the lung by the
inner part of its upper face, whilst the external part is free: its
REPORT ON THE PROGRESS OF ANATOMY. 175
inferior face is moulded on the diaphragm. It is triangular in
form, and its volume is about one quarter of that of the middle
lobe. He regards this lobe as a true dobus impar such as is seen
in the generality of mammals, and as exhibiting a tendency to
reversion in man to a lower type.
Skin anp AppenpAces. — P. Langerhans gives an account
(Schulize’s Archiv, 1x. 730) of the Toucu-corpuscLes AND REeETE
Mauricuit. He considers the touch-corpuscle to be composed of
a number of individual cells, which are characterized by the deli-
cacy and sparing quantity of cell-substance, and between these
cells throughout the entire organ the nervous elements are ar-
ranged. He disbelieves therefore in a central core, as distinguished
from a peripheral substance, as well as in an investing membrane,
for the cells of the periphery abut directly on the surrounding
connective tissue. L. Stieda adversely criticises (Schultze’s
Archiv, 1x. 795) Schobl’s researches on the NERVE-corLs AT THE Roots
oF THE Hairs of certain animals (Report, vu. p. 331). Stieda re-
gards the appearance as indicative merely of different stages which
mark the changes of the hair.
ALIMENTARY CanaL.—Hugo Crampe communicates (Reichert u.
du Bois Reymond’s Archiv, 1872, 569) aseries of comparative obser-
vations on the Leners oF THE INTESTINE and extent of its mu-
cous surface in different animals. He does not consider that the
nature, z.e. the chemical composition, of the food exercises much
influence on the structure of the digestive apparatus, but that the
form in which the food is administered, by causing the organs to
accommodsute themselves to its volume, occasions modifications.
Again, he finds variations in length even in animals born of the
same mother; and that cats fed on vegetables have possessed rela-
tively shorter intestines than cats fed exclusively on flesh. He does
not believe that any constant relation exists between the weight of
the body and the extent of the intestinal mucous surface.
Kipyey.—Th. Egli gives a short account (Schultze’s Archiv, rx.
653) of the GLANDs IN THE PELViIs oF THE KipNey. He was induced
to examine into this subject owing to the discrepancies in the
writings of anatomists, and has studied the pelvis of the kidney
of the ox, pig, horse and man. The ox and pig have no glands.
The mucous lining of the pelvis of the horse is studded with simple
and compound tubular glands possessing a single layer of goblet
and cylindrical cells. In man compound glands, intermediate in
form between tubular and racemose, filled with cylinder and spindle-
shaped cells, exist. The duct is very short.
CoMPARATIVE ANATOMY.
QuapRUMANA. —Jas. Maurie continues (P. Zool. Soc. June 18,
1872) his observations on the Macaques, and gives an account of
Bélanger’s Monkey (l/. arctoides), of the Formosan or round-faced
monkey (J. cyclopis), and of the Japanese Monkey (J. speciosus).
176 PROFESSOR TURNER.
Carnivora.—A. H. Garrod notes (Proc. Zool. Soc. Feb. 18, 1873)
certain points in the anatomy of Arctictis binturong. The alimentary
canal, liver, lungs, genito-urinary organs and brain are described, and
in some instances figured. J. Chatin in Ann. des Sc. Nat. 1873,
Art. 12, describes the visceral anatomy of Viverra civetta.
PiynepepiA.—B. Dybowski gives in Feichert. u. du Bois Rey-
mond’s Archiv, 1873, 109, a detailed description with figures of the
skull of Phoca baicalensis.
PerissopactyLa.—A. H. Garrod gives an account (Proc. Zool. Soc.
Jan. 21, 1873) of the visceral anatomy of the Sumatran Rhinoceros.
He figures the tongue, stomach, colon and liver.——J. E. Gray
records (Ann. Nat. Hist. May, 1873) observations on the Dentition
of Rhinoceroses, and on the characters afforded by their skulls.
PacuypErRMATA.—J. E. Gray (Ann. Nat. Hist. June, 1873)
makes some observations on Pigs and their skulls, more especially in
connection with the classification of these animals. An abstract
of a memoir by W. K. Parker on the structure and development
of the skull of Sus scrofa, is in P. Roy. Soc. L. June 19, 1873.
CrTacea.—A catalogue of the Whales and Dolphins of the New
Zealand Seas has recently been drawn up by Jas. Hector (Trans.
New Zealand Inst. v. 1872). It is illustrated with six octavo plates.
H. 8. Wilson describes in Cambridge Univ. Reporter, June 3,
1873, the Rete mirabile of the Narwhal. He showed that the rete
was divisible into halves, the vessels of which were derived from two
sources, and presented the same calibre at their origin. He divided
arterial retia into two classes, bilateral and axial, and subdivided the
axial into terminal and mediate, and each of these into complete and
incomplete. The axial system he regards as fulfilling the office of
supplying a large amount of blood to parts, as a means of avoiding
injury from compression of vessels, and to check sudden pressure on
nerve centres, whilst he believes the bilateral to act not only as a
storehouse for oxygenated blood, but as a diverticulum protective
against over-pressure. J. E. Gray (Ann. Nat. Hist. Aug. 1873)
notes some points in the anatomy of two skeletons of Kogia Mac-
leaywt recently received at the British Museum from Australia.
M. du Bus describes (Gervais’ Journ. de Zoologie, 11. 97, 1873) the
remains of the Delphinide found in the Antwerp Crag. He refers
them to the genera Hurhino-delphis, Prisco-delphinus, Platydelphis,
Champso-delphis, Phocenopsis, Eudelphis, Hoplocetus, Paleo-delphis
and Scaldicetus. P. J. Van Beneden gives in Acad. Roy. de Belgique,
July, 1873, xxxv1., two coloured drawings of cetaceans from the Cape
of Good Hope ; one he refers to the Orca capensis of Gray, the other
he names Lagenorhynchus de Castelnau. Rt. Walker in Scottish
Naturalist, Oct. 1873, figures the lower jaw of a common porpoise in
which conical-shaped bodies 7; to }inch in height were attached to
the gum alternately between the normal teeth. A similar arrange-
ment exists in the upper jaw. He also states that he has examined
REPORT ON THE PROGRESS OF ANATOMY. VTE
eight or nine specimens of the porpoise caught on the coast of Scotland,
all of which had tubercles on the dorsal fin. He does not regard these
tubercles as Dr Gray does as specific, but only as indicating possibly
a variety of porpoise.
Brrps.—Jas. Murie describes (Zbis, April, 1873) and illustrates
with numerous figures the Upuprp# and their relationships. AOE
Garrod (Proc. Zool. Soc. June 18, 1872) notes the CHARACTERS OF THE
ToneuE of the psittacine genus WVestor. The peculiarity consists in
the anterior edge of the unguis being prolonged forwards beyond the
tip of the tongue for about one-tenth of an inch as a delicate crescentic
fringe of hairs, which seem to result from the breaking up into fibres
of the forward growing plate. The structure of the tongue leads to
the placing of Nestor among the typical parrots and not with the
Tricho-glossine. A. H. Garrod points out (Proc. Zool. Soc, Jan. 7,
1873) the value in CuassiricaTion of a peculiarity in the ANTERIOR
Marcin or tHE Nasau Bones of certain birds. In most birds the
anterior margin of the nasal bone is concave with the two cornua
directed forwards, which become continuous behind with the body of
the bone and with one another. Birds possessing this form of nasal
bone the author terms holorhinal. In other birds the posterior
contour of the osseous external nares instead of being rounded, as in
holorhinal birds, is apparently formed by the divergence of two
straight bars of bone, which enclose an angular space between them.
These birds he terms schizorhinal. Nearly all the schizorhinal birds
are included among the Schizognathe of Huxley. M. G, Duchamp
makes (Ann. des Se. Nat. 1873, Art. 11) some observations on the
Awatomy or Dromaius Nov#-Hotianpim. He describes the diges-
tive, respiratory, urinary and genital organs, also the spleen and
thyroid body. M. Coughtrey notes (Ann. Nat. Hist. Sep. 1873)
some peculiarities in the TRacHEAL Poucu or AN Emeu which he had
dissected. V. Mihalkovics relates his investigations on the PECTEN
IN THE EYE oF THE Brirp (Schultze’s Archiv, 1x. 591). Histologically
it consists of a convoluted mass of capillaries, the spaces between
which are filled with gelatinous material containing pigment. He
considers that it plays an important part in the nutrition of the lens
and partially of the retina.
RepritiA.—F, Leydig describes (Schultze’s Archiv, 1x. 598) the
GLANDS OF THE Heap in the following Opuipians: Z’ropidonotus
natric and tessellatus, Coronella levis, Coluber viridiflavus, Vipera
verus and ammodytes; and on p, 753 Leydig gives a description of
the STRUCTURE OF THE SKIN of the same Ophidians. J. E. Gray
makes additional notes on the rorm of the Bones in the STERNUM of
Be Les Torroises and thei development (Ann. Nat. Hist. Oct.
73).
Ampuipians.—P. Langerhans describes (Schultze’s Archiv, 1x. 745)
the Skin of the Larva or SALAMANDRA Macutosa.
Fisnes.— P. Harting, by the invention of an instrument, which
he calls a physometer, has been able to make observations on the
VOL. VIII. 12
178 PROF. TURNER. REPORT ON THE PROGRESS OF ANATOMY.
Functions OF THE Swimmine Biapper (Abstract in Gervais’ Journ.
de Zool. 11. 116).° He discusses the various theories which have been
propounded as to its use; as an accessory respiratory organ, or as a
hydrostatic apparatus by which the fish preserves its equilibrium in
the water, and refers to the experiments of Moreau, who esta-
blished that the air contained in it possessed an excess of oxygen. By
his instrument the dilatations and contractions of the air in the swim-
ming bladder can be measured. G. Gulliver (Proc. Z. Soe. Nov.
19, 1872) records measurements of the BLoop CorPUSCLES OF THE
Satmonip#. The average sizes are as follows: Salmo Sn
Long Bi sizz» Short Diam. zeeo3 ; Salmo sags i; D:
Ss. D. Salmo te wo, L. D. 54; 8. D. oe i Salmo Sao,
ia): sore nl sa00 3 dee vulgaris, L, Do ee ¥a00>
_ Osmerus eperlanus, L. D. 53-,, 8. D. = 355. He compares their size
with that of several other osseous fishes and states that amongst
osseous fishes the Salmonide have the largest blood-dises. P. D.
Handyside communicated to Roy. Soe. Edinburgh, Jan. 20, 1873, an
account of the external characters of a new SPECIES OF PoLyopon
from the river Yang-tsze-kiang, and on June 2nd a description of its
nervous and muscular systems.
Tea
Crustacea.—E. T, Newton gives a description (Quart. Mic. Journ.
Oct. 1873) of the EYE or THE Lopsrer. Tt is illustrated with two
oct. plates. The Anatomy of the American King Crab (LiruLus
POLYPHEMUS) is described by R. Owen in 7rans. Linn. Soc. XXVUI.
Mo.tivsca.—R. Bergh describes (Jowrnal des Museum Godefroy,
Heft 11.) several forms of Vudibranchs, and illustrates them by four
large plates. The specimens belong to the genera Phyllidia, Plako-
branchus, Llysia, Cyerce, Fiona and Cerberilla.
REPORT ON PHYSIOLOGY. By Wituiam Sriruine, D. Sc,
M.B., C.M., Demonstrator of Practical Natural History in the
Tniversity of Edinburgh’.
Nervous System.
EXPERIMENTAL RESEARCHES IN CEREBRAL PHYSIOLOGY AND Pa-
THOLOGY.—Dr Ferrier (rit. Med. Journ., April, 1873, West Riding
Asylum Reports, 111.) has uncovered particular parts of the encepha-
lar and Faradised them, observing the movements which follow, with
a view to obtaining information regarding the source of the volun-
tary movements in certain parts of the surface of the brain, The
results of Hitzig, obtained in an almost similar manner, have already
been noticed in this Jowrnal, vu. 175. Ferrier’s experiments were
practised upon dogs, guinea-pigs, cats, and rabbits. For the results
see paper by Dr Ferrier, p. 152. Nothnagel, vide this Jowrnal, vil.
177, (Virchow’s Archiv, 1873, Lyi. 184,) finds that on making an
injury, by injection of solution of chromic acid, of the diameter of
2mm., and to the depth of 1—13mm. on the upper surface of the
hemisphere 12—16 mm., according to the size of the animal, from the
point of the hemisphere after removal of the olfactory lobes, and 2 mm.
from the longitudinal fissure, one observes that the fore-leg of the
opposite side, and less clearly the hind-limb, is extended forwards and
outwards. This condition lasts 6—12 days, and then becomes less and
less obvious, and ultimately disappears altogether. The author attri-
butes it to a partial paralysis of the muscular sense. By irritating a
particular point of the corpus striatum (nucleus caudatus), the animal
begins without the least irritation from without to hop, then to rest,
in a short time to hop again, always quicker, and with shorter inter-
vals, and lastly to rush forward with great speed. The animal soon
tires, and falls over, but still the legs continue to move violently.
The animal could not be preserved more than eighteen hours. In
Centralblatt, No. 35, 1873, the same author finds that in 23 cases
by injuring the cortical portion of the brain with a fine needle
(without injection of chromic acid), at a circumscribed spot lying near
to the hinder end of the hemispheres, immediately there ensued
violent spasmodic movements. The animal makes powerful jumps,
and often rises 3—1 metre from the floor, tle extremities then be-
come rigid, After 1, 2, 5 minutes this passes off, the animal sits
still a minute, and then hops away as if nothing had happened.
Fournié also communicates experimental researches (pampilet),
Paris, 4 plates. A portion of the skull is removed and by means of
a Pravaz syringe a small quantity of a saturated solution of chloride
of zine coloured blue by aniline is injected into the brain substance.
We must be careful not to confound the results of small extravasa-
1 To assist in rendering this report more complete, authors are invited to
send copies of their papers to Dr Stirling, Edinburgh University.
12—2
180 DR STIRLING.
tions into the brain substance, caused by the needle of the syringe
puncturing a vessel, with those of the injection. The dog was the
animal operated upon. In 5 experiments out of 7 a complete abo-
lition of the sense of touch was produced, which coincided with the
complete destruction of the thalamus opticus. The author seeks to
explain this by the relatively large commissure which unites the
two thal. optici in the dog. ‘The thalamus opticus, the author thinks,
furnishes the conditions for a ‘perception simple’ (he does not say
‘avee connaissance’). With regard to lesions affecting motion, by far
the most salient and pronounced is the ‘mouvement de galop,’ which
is produced in one and the same animal incessantly till just before
death, when, according to the intensity or seat of the lesion, these
movements are followed by paralysis more or less complete. These
movements the author attributes to the effect of the caustie upon
the fibres which pass between the optic thalamus and the corpus
striatum. The paralysis appears because the sensitive parts of the
thalamus are destroyed by the injection, and, in consequence, cannot
excite the motor cells of the corpus striatum. Lesion of the white
substance below and to the imner side of the thalamus, and which
connects it with the corpus striatum, is followed by paralysis of the
hind extremities. The respiration is very peculiar, being rapid and.
deep. Corpus striatum ; as in the thalamus, lesion of this organ on one
side produces the same phenomena as if both had been injured. As
a general rule, no affection of sensibility was observed, while para-
lysis of motion had a constant occurrence. The tendency presented
by the animal to move in a circle, or to turn to the injured side,
the author observes, are not characteristic of lesion of the corpus
striatum, for they also follow injury to the white substance of the
brain, and of the convolutions. | Lesions of the cervical portion of the
brain at the anterior, middle, or posterior parts, did not extinguish
simple perceptions, while, on the contrary, the absence of intelligence
(‘‘ connaissance”) and memory was constantly observed. Regarding
motions, two periods were constantly observed, the one a period of
excitation when the animal ran, or moved in a circle &c., and the
other a period of prostration, and sometimes of paralysis. The first
period the author thinks corresponds to the simple excitant action of
the caustic, while the latter is referable to a destruction of the tissues
by the same. Injury to the white matter of the brain was followed
by the same phenomena which accompany lesion of the convolutions,
optic thalamus, and corpus striatum, and this because the fibres
which compose the white part of the brain serve to bind, on the one
hand, the thalamus with the peripheral part of the brain, and on the
other, the peripheral part of the brain with the corpus striatum.
Lesion of the cerebellum produced phenomena of excitement ; the
animals ran about wildly, assumed ridiculous positions, while the eye-
balls were always turned from below upwards, &c. This state was suc-
ceeded by a period of collapse, characterised by paralysis. More or
less complete sensibility was profoundly affected in the greater number
of these lesions. Then follow the results of a series of complex lesions,
not referable distinctly to injury of any one part of the brain, be-
REPORT ON PHYSIOLOGY. 181i
cause several parts of the brain had sustained injury. Injury of
the cornu ammonis was followed by the same phenomena as lesion
of the other convolutions, but, what was very peculiar, the
animal without being paralysed was unable to stand upon its four
feet, being only able to crawl along on its belly. It had lost the
sense of equilibrium. See also a paper communicated by the author
to the Académie de Médicine, of which an abstract is given in the
Lond. Med. Rec., No. 35.
CoMPOSITION OF THE GREY AND WHITE MATTER OF THE BRAIN.—
Phliiger’s Archiv, 1873, vu. 367. Petrowsky, under Hoppe-Seyler’s
direction, has investigated the chemical composition of the grey and
white matter of the brain. The following represents the composition
of the brain of the Ox.
Water. Solids.
100 grms. of grey substance contain 81:6042 18-3958
100 grms. of white substance contain 68°3508 31°6492,
100 grms. ef Solids. Gray. White.
Albuminous compounds and Glutin 95°3733 24-7252
Lecithin : : ; 5 : 17-2402 9-9045
Cholestrine and Fats. ; : 18:°6845 51:9088
Cerebrin : : ” , : 0:5331 9-5472
Substances insoluble in pure ether 6:7135 3°3421
Salts . P ¥ - 4 is 1:4552 O79:
CuemicaAL Reaction oF THE CrnTRAL Nervous SysTem.—
R. Gscheidlen (Pjliig. Archiv, vut, 171) experimented upon the
brains of horses, dogs, rabbits, &c. An incision was made into the brain,
and its reaction tested by means of Liebreich’s plates (Berichte d.
Chem Gesell. zu Berlin, 1868, p. 48). The grey substance always _
showed an acid reaction, while the white reacted always neutral,
er slightly alkaline. These results were constant in more than
70 animals, and it was immaterial whether the animals were poisoned
with morphia, or curara, or whether they had served for a longer
or shorter time for experiments, or had been killed at once. These
results are not due to post-mortem changes. The same results were
obtained on the ganglia of the sympathetic, which exhibited an acid
reaction, while the connecting nerve-fibres were neutral or slightly
alkaline. The same also is true of the grey and white substance of
the spinal cord. The author thinks that the ganglion cells contain
as a constituent a free acid, which is in all probability lactic acid.
After death the acid reaction of the grey substance increases to a
certain degree, while in the white this reaction occurs quite as little
as in peripheral nerves (vide Heidenhain’s observations). If the
white substance shows an acid reaction, it is due to the passage of
the acid into it from the grey matter, for if in a large animal the
gray and white matter are kept apart, the reaction of the white
remains alkaline or neutral, the grey of course acid.
EmpryonaL DEVELOPMENT OF NervE-CeLLs.—Lubimoff (Central-
dlatt, No. 41, 1873) has studied the development of the nerve-cells
182 DR STIRLING.
in the human embryo. He found that the cells of the sympathetic
system reach perfection sooner than those of the central part of the
cerebro-spinal nervous system. In the sympathetic system itself
these cells, which are found in the branches of the cerebro-spinal
nervous system, reach perfection sooner than those of the pre-verte-
bral cords and the ganglion cceliacum. In the central cerebro-spinal
system the cells of the spinal cord (earliest the nerve-cells of the
anterior horn) are sooner perfected than those of the grey sub-
stance of the cerebrum and cerebellum.
TRANSMISSION OF REFLEX IMPRESSIONS IN THE SprnaL Corp.—
Rosenthal (Bericht der K. Akad. der Wissensch. Berlin, February,
1873. Abstract by Dr Brunton, in London Med. Record, No. 29)
finds that a certain time is required for the transmission of a sensory
impression to a motor nerve.’ This time (reflex time) depends on the
strength of the irritation—the stronger the irritation the shorter the
time required. A longer time is required for the transmission of a
sensory impression made on one side of the body to the muscles of
the other side, than to the muscles on the same side. This difference
also depends upon the strength of the irritation—the stronger the
irritation the shorter the time. In peripheral motor nerves no de-
pendence of the rapidity of conduction on the strength of the irrita-
tion can be observed. The nearer the irritated spot is to the spinal
cord, the more easily is the reflex time diminished. The reflex time
is altered by the exhaustion of the spinal cord. When an adequate
irritation is applied to an exposed sensory nerve, at two points as far
apart as possible, the reflex time for the point furthest away from the
cord is greater than for the nearer one.
ReEFLex RELATIONS OF THE StoMAcd To THE NERVE CENTRES OF
THE CiRCULATION.—Sittz.-berich. der Wien Akad. der Wissensch, LXVI.
1872. Centralblatt, No. 13, 1873. Mayer and Pribram have ex-
perimented upon dogs and cats. The experiments of Goltz on irri-
tating the walls of the stomach have shown that this is followed
by slowing of the pulse, but these authors have found that there
is an increase in the arterial pressure. The results were the same
both by mechanical and electrical irritation, or by insufflation of the
stomach by means of an india-rubber bag. In opposition to the
experiments of Hermann and Ganz, these authors state that neither
by injection of cold water, nor by pieces of ice introduced into the
stomach, could they observe any effect. It seems, from these experi-
ments, that irritation of the mucous membrane alone has no effect
on the pulse or blood-pressure ; but the irritation to produce these
effects must extend to the serous and muscular layers.
ReFLex PARALYSIS OF VESSELS AND AFFECTIONS OF THE SPINAL
CorRD AFTER SUPPRESSION OF THE PERSPIRATION BY VARNISHING
ANIMALS.—Feinberg (Centralblatt, No. 35, 1873) finds that in ani-
mals coated with varnish the greater part of the morbid phenomena
ave spinal-cord affections. To be observed are tremor, hyperzsthesia,
REPORT ON PHYSIOLOGY. 183
later partial anzesthesia, enacted reflex sensibility, reflex and tetanic
spasms, rotation round the axis (observed three times), paralysis of
the bladder, &e. Frequency of respiration decreased, action of the
heart lessened, and the temperature diminished. AI] these phe-
nomena are of different duration and different intensity. If the
animal dies rapidly the period of irritation is very short, hyperzs-
thesia and reflex irritability are not very pronounced, and merge
quickly into partial anesthesia and paralysis. Spasms never fail.
In fact, all the phenomena of depression are present. When death
takes place more slowly, hyperesthesia and reflex irritability are so
intense that the slightest contact with the skin, a stamp of the foot
on the ground, is followed by a succession of powerful reflex spasms ;
the other phenomena appear later, some time before death. The
temperature depends upon the unhindered or hindered radiation of heat.
In the first case it sinks quickly and progressively till 19 — 20°C.
In the second case it exceeds or differs little from the normal, and
then just before death it begins quickly or slowly to sink, then death
quickly sets in. The post-mortem phenomena observed were: Dila-
tation of the sub-cutaneous vessels, which were injected; large extra-
vasations from the pulmonary capillaries, not seldom also under the
pleura, heart-chambers over filled with blood, often extravasations in
the heart-substance, distension of the roots of the portal veins, and
extravasations in the liver-substance; the mucous membrane of the
stomach constantly exhibited extravasations; strong injection of all
the capillaries of the serous layer of the intestine, with catarrh of its
mucous lining ; kidneys filled with blood, with commencing parenchy-
matous inflammation, capillaries of the peripheral nerves strongly i in-
jected, often with extravasations, the same in the voluntary muscles.
Membranes of brain injected. Gray substance of the cervical spinal
cord dark red, with small extravasations. Dorsal and lumbar
regions of the cord less injected. Microscopical examination shows
in the cervical part of the spinal cord numerous capillary extravasa-
tions with partial destruction of its substance, proliferation of neuro-
glia less obvious in those animals which died quickly, but strongly
developed in those which died slowly. The nerve-fibres were com-
pressed and atrophied. All these phenomena were more pronounced
in the cervical part of the spinal cord than lower down, These phe-
nomena can only be explained by a paralysis of all the vaso-motor
nerves in the cervical part of the spinal cord or their centre in the
med, oblongata.
INFLUENCE DE L’ Activité REFLEXE DES Centres NERVEUX VASs-
CULAIRES SUR LA DILATATION DES ARTERES PERIPHERIQUES ET SUR
LA SECRETION DES GLANDES sous-MAXILLAIRESs.—Owsjannikow and
Tschiriew (Lull. de [T Acad, Imp. des Se. de St Pétersbourg, Xvut.
18, Mai 1872. Abst. in Arch. de Phys. 1873, No. 1, p. 90) find
that irritation of the central end of the N. ischiadicus with in-
tact chorda tympani is followed by secretion of saliva from the sub-
maxillary gland. These authors explain this as due to the increased
blood-pressure (from irritation of a sensory nerve) in the body gene-
184 DR STIRLING.
rally and in the gland. Irritation of the N. ischiadicus when the chorda
is cut diminishes the secretion. Irritation of the ganglion sub-maxillare
can diminish a rapid secretion, These authors suppose that there is in
the chorda tympani fibres which widen the vessels (secretion fibres),
and from the ganglion sub-maxillare proceed fibres which narrow the
vessels (fibres which diminish the secretion). Irritation of the peri-
pheral end of the N. splanchnicus (which is followed by increased
blood-pressure) produced the same result on the secretion as irritation
of the N. ischiadicus, Griitzner and Chtapowski (Pjliig. Arch. vu.
p. 522) have repeated the above experiments, and find exactly as the
previous authors, that on irritating the central end of the N. ischia-
dieus the secretion from the canula fell drop for drop with a fre-
quency scarcely inferior to that obtained by directly irritating the
chorda. The saliva also was the so-called chorda saliva. On irri-
tating the left N. ischiadicus and cutting the left chorda (canula in
both ducts) the right sub-maxillary glands secreted in the old way,
while the left yielded very little or almost no secretion. The blood-
pressure was measured in the art. cruralis. These authors explain
the fact why irritation of a sensory nerve, with cut chorda, produces
now and again a small outflow of saliva thus. They say it is owing
to the pressure of the contraction of the muscles which pass over
these glands. These muscles can contract (in an animal not too
powerfully poisoned with curara) quite as well as the other muscles
of the body. Heidenhain has shown that atropia paralyses the
secretory fibres of the chorda, while the inhibitory fibres are intact,
and Ludwig has shown that the increase of blood-pressure and in-
crease of secretion do not stand in the relation of cause and effect. To
determine whether the increase of secretion observed on irritating the
N. ischiadicus was really due to increase of the blood-pressure, or was
only a reflex secretion, these authors injected atropia (0-001—0-005)
into a vein in a-curarised dog, and found that on strong irritation of
the N. ischiadicus no secretion was obtained from the canula, but red
blood often at rhythmical intervals flowed from the vein of the gland,
as ought to happen on peripheral irritation of the chorda. The
vaso-motor nerves of this nerve were here reflexly stimulated, and
their section brought the whole to an end, for then there is in-
crease of blood-pressure in the vessels of the gland, and no secretion.
The increased salivary secretion on irritation of the N. ischiadicus is
therefore produced by a reflex action on the secreting fibres of the
chorda. From these experiments these authors think that the fibres
of the chorda, as well as of the sympathetic, have their origin in the
medulla oblongata.
INNERVATION OF THE Ear oF THE Raspit.—Moreau states in
Arch. de Phys. 1v. 667, that section of the sympathetic alone pro-
duces redness of the under half of the ear due to widening of the
vessels; while section of the N. auriculus magnus (cervical plexus)
produces a similar effect on the upper half. It is therefore obvious
that both nerves are concerned in the innervation of the vessels of
the ear, and that the contraction of the vessels of the under half of
REPORT ON PHYSIOLOGY. 185
the ear depends upon the sympathetic, while that of the upper
depends on the N, auriculus. “ Functions of the Lingual Nerve.”
Prévost, in Arch. de Phys. v. 1873, 253 and 375. ‘“‘ Contributions
to Physiology of the Vagus.” Arloing and Trissier in Arch. de
Phys. Vv. 1873, p. 157.
REASONS FOR THE PATHOLOGICAL CHANGES IN THE LUNGS AFTER
SECTION OF BOTH VaGi.—AlIf. Genzmer in Pfliig. Archiv, vit. 101.
The author concludes from his experiments (1) that the altered
action of the heart produced by paralysis of the vagus is without
influence on the lung-tissue. (2) Hindering of the supply of blood.
(3) Forced passage of saliva does not produce disease, as was found
after section of both vagi. (4) Paralysis of the vagus can remain
without visible consequences on the condition of the lung-tissues, it
however increases their tendency to disease, and that more so on the
paralysed side. (5) Paralysis of both pulmonary vagi produces a
neuro-paralytic hyperemia of the lungs. (6) If saliva is forced into
the lungs, rendered hyperzemic (cedematous) through section of the
vagus, it produces inflammation.
INNERVATION OF THE SPLEEN AND ITS RELATION TO LeEucocy-
THAMIA.—Tarchanoff in Pfliig. Archiv, vul. 97 says, that in a cura-
rised dog with artificial respiration kept up and_ blood-pressure
measured in carotid, he obtained the following results. (1) Inri-
tation of the central end of the vagus (medium and strong) for
a minute or more, produced, besides the increase of blood-pressure,
a powerful contraction of the spleen, about 1—2 ¢.m., while irritation
of the peripheral end produced a scarcely perceptible contraction, or
no effect at all. (2) Irritation of the central end of the N. ischia-
dicus produced, besides the increase of blood-pressure, a contraction
of the spleen, but less than in the above case. (3) Irritation of the
medulla produced a very powerful contraction of the spleen about
2$cm., but this only when the N. splanchn. are intact, for these
nerves contain the centrifugal nerves for the vessels of the spleen.
This contraction does not disappear on cessation of the irritation, but
the spleen returns gradually to its former size, whilst the blood-
pressure has already returned to its original height. (4) Gradual
section of the nerves of the spleen, as is known, is followed by
swelling of the organ to one-and-half times the original size. After
section of the nerves of the spleen the blood taken from the spleen in
the 2nd, 3rd, and 4th days after the operation showed a quantity of
white blood-corpuscles in the field of the microscope, varying from
60 to 70, whilst the blood taken from the ear before the experiment
showed only from 6 to 15. Leucocythzemia so produced however
gradually disappears, and at the end of a week the number of white
blood-corpuscles is normal. Parallel with this phenomenon is the
diminution of the size of the spleen. Vide also “Physiology of
the Spleen,” by Bochefontaine in Arch. de Phys. v. 558. “ Tnner-
vation of Blood-Vessels,” by E. Pick in Reich. und Du Bois Rey,
Arch. 1873, 1. Abstract in Centralblatt, No. 46, 1873.
186 DR STIRLING.
On THE CENTRE OF THE NerveES or Erection.—Goltz, Pflig.
Archiv, vit. 582. Eckhard says that the nervi erigentes can be fol-
lowed in the interior of the spinal cord to the pons varolii and
cerebrum. The author, from experiments upon dogs, arrives at
a different conclusion, viz. that the nearest centre from which the
nervi erigentes spring is placed in the lumbar portion of the spinal
cord. Upon section of the posterior part of the thoracic part of the
spinal cord erection could be produced in a reflex manner. Further,
that the condition of the reflex can be hindered by irritation of
sensory nerves. He has observed peculiar reflex rhythmical contrac-
tions of the sphincter ani in animals with divided spinal cord. The
lumbar spinal cord is a more important centre for reflex movements
and reflex inhibition than was formerly supposed.
Hermann, “ Action of galvanic currents on Muscles and Nerves.”
Phliig. Archiv, vi. 312 and 561. Abstract by Rosenthal, Centralblatt,
Notas 1873: Griinhagen, “On the secondary Contractions in
Muscle.” Pfliig. Archiv, v1. 119, 1872. Abstract in Centralblatt, No.
LSP LS7 3: Griinhagen and Englemann, “ Intermittent Irritation
of Nerves and Muscles.” Pfliig. Archiv, v1. 157, 1872. Abstract in
Centralblatt, Nos. 19 and 20, 1873. Vulpian, “ Influence of trau-
matic Lesions of Nerves on the physiological properties and the struc-
ture of Muscles.” Arch. de Phys. tv. 245, Abstract in Centralblatt,
No. 15, 1873. “Transverse Conduction in Frog’s Nerves.” Hitzig
in Pfliig. Archiv, vu. 263. “On the Form of the Contraction in
the so-called transverse Conduction in Frog’s Nerves.” Wm. Filehne
in Pfliig. Archiv, vit. 71. “ Influence of Section of a Nerve on the
nutrition and regeneration of the Tissues.” Schulz in Centralblatt,
No. 45, 1873.——“ Alterations of the Spinal Cord after extraction of
the Sciatic Nerve in the Rabbit.” G. Hayem in Arch. de Phys. v.
504, “Union of the cut end of the Lingual and Hypoglossal
Nerves.” Vulpianin Arch. de Phys. v. 597. “ Wlectrical Irritation
of Nerves.” Fick in Arbeiten aus d. Physiol. Lab. d. Wurzbiirger
Hochschule, Pt. 1. 65. Ranvier, “Degeneration of Nerves after
Section” in. Comptes Rendus, txxv. 1831; and “ Researches on the
Histology and Physiology of Nerves,’ in Archiv. de Phys. et Pathol.
Iv. 427, 1872. “ Trritation of Nerves by electric currents.” Zeitsch.
F. Biolog. vit. 71 and 100, Schiff, Fuchs. Englemann, Vederl.
Arch. S.A. 5. Abstract in Centralblatt, No. 33, 1873. “« Action
of very gradual thermal irritation on the sensibility of Nerves.”
Heinzmann in Pfliig. Archiv, vi. 222. “‘ Regeneration of Nerves
in Paraplegic Animals.” »Prévost and Waller in Gaz. Méd. de Paris,
1873, No. 10. These authors find in rats and guinea-pigs, rendered
paraplegic by section of the spinal cord, that degeneration and regene-
ration of the cut or pinched sciatic nerve takes place just as in the
sound animals.
CoNDITION OF THE SKIN UNDER SLIGHT MECHANICAL IRRITATION.
—Centralblatt, No. 26, 1873. Petrowsky remarks that the pheno-
menon of white stripes in the skin, observed by Bouchert in Scarla-
tina, and by Baumler in other febrile disorders, Exanthematic Typhus,
REPORT ON PHYSIOLOGY. 187
Pneumonia, Traumatic fever (Centralblatt, No. 12, 1873), he has
found also to take place in the normal skin. Make a line with the
fingernail gently upon the skin, at first at the irritated spot nothing
is observed, but after the lapse of }—} min. the irritated line begins
by degrees to grow pale, the paleness rapidly reaches its maximum,
persists for a short time, and then disappears by degrees. The dura-
tion of the phenomenon, until it has completely disappeared, varies
from 4—6 minutes, and depends upon the strength of the irritant
employed, and also on the individual. This condition can be pro-
duced on a dry skin as well as one wet with perspiration. The
author believes that the phenomenon is probably due to contraction
of the cutaneous capillaries, and that other histological elements of
the skin are also active.
Cuorpa Tympani.—Vulpian has continued his researches on this
nerve, and has communicated his results to the Société de Biologie.
The observation of Heidenhain is confirmed, that in a dog brought
under the influence of atropia, excitation of this nerve is no longer
followed by increased sub-maxillary secretion, but further dilatation
of the vessels of the glands, and that in excitation of the lingual
nerve the vessels dilated just as if no atropia had been administered.
Vaso-MoTor ACTION OF THE Sprancunic Nerve.—Vulpian (Gaz.
hebdom. 1873, No. 21). Section of this nerve 3 cm. above the left
suprarenal body, as performed ona curarised dog. The volume of
the kidney on the same side and the quantity of blood it contained
was increased. The urine albuminous and diminished in quantity,
but not containing blood or epithelial cells from the tubuli. By
irritating the peripheral end of the nerve by an induced current
the kidney became pale, the veins contracted and the secretion of
urine ceased.
Eyr.—‘ Keratitis after Section of the Trigeminus,” Eberth, in
Centralblatt, No. 32, 1873.——“ Section of the Optic Nerve,” Berlin,
in Centralblatt, No. 28, 1873. “Influence of the Sympathetic
upon the Eye,” Eckhard in Centralblatt, No. 35, 1873. “ Visible
Direction,” J. Jago, in Proc. Roy. Soc. Lond. xxi. 213. “On the
Physiological Action of Light,” by Jas. Dewar and J. G. M. Kendrick,
in Journ. of Anat. and Phys. June, 1873.
Eax.—< Sensibility to Sound,” Miiller, in Ludwig’s Arbeiten, vi.
pel. H. Buck and Burnett, “ Experiments on the Mechanism of
the Auditory Ossicles,’ in Centralblatt, No. 17, 1873. Rumbold,
* On Functions of the Eustachian Tube,” in Lond. Med. Rec. No. 29.
Riidinger, on “ Closure of the Eustachian Tube,” bid. No. 18.
“ Perception of high Musical Notes,” in Boston Med. and Surg.
Journ. 1872, x. 20. “ Accommodation of the Ear,’ Mach and
Kessel, Abstract in Lond. Med. Rec. No. 37, 1873. “On the
Movements following Section of the Semicircular Canals.” Bottcher
(Dorpat. Med. Zeitschr. 1872, ut. 97. Abstract in Centralblatt, No.
5, 1873) in his experiments has used frogs, because he found that by
188 DR STIRLING.
careful dissection he could expose these canals without loss of blood,
and without injury or disturbance to any of the surrounding impor-
tant parts. In opposition to the results of Flourens and Goltz, B. finds
that no disturbance of equilibrium was produced by this injury,
whether it was performed on one or both sides. These results are in
direct opposition to those of Lowenberg (Arch. f. Augen- u. Ohrenheilk.
1872, 11. Abstract in Centralblatt, No. 18, 1873), who operated upon
pigeons, and by a modification in the method of operating, the author
wisued to determine whether the movements were caused by pain as
supposed by Flourens, and whether the presence of consciousness was
necessary for these disturbed movements, and whether the cause
consists in an irritation or paralysis of nerves. The results of his
experiments are: (1) The disturbance of movement recurring after
section of the semicircular canals of the ear depends only upon this
injury and not from any injury to the brain. (2) The vomiting
observed by Czermak in his experiments depends upon injury of the
cerebellum. (3) The disturbance of movement is the consequence
of irritation of the membranous canals and not of paralysis of the
same. (4) The irritation of the canals produces the spasmodic move-
ments in a reflex manner without the cooperation of consciousness.
(5) The transference of this reflex irritation of the nerves of the
membranous canals to the motor nerves takes place in the thalamus
opticus. With regard to the statement of Brown-Séquard, that
section of the N. acusticus produces these movements, the author,
operating upon rabbits, finds that section of this nerve from the
tympanum yields the same results, but in this case the semicircular
canals are at the same time partly pinched and partly pierced.
Shin.
Excretion oF CO, By THE Skriv 1n Man.—Aubert (Pjliig. Archiv,
vi. Hf. ii.) has determined by a special apparatus the amount of CO,
excreted by the skin of a healthy man per diem. The subject
experimented upon was made to sit in a box which fits tightly round
the neck, and through which air is gently passed. In 24 hours a
maximum amount of 63grms. and a minimum of 2-3 grms. was
eliminated by the skin of the whole body minus the head. This
gives a mean of 3°87 grms., while the quantity discharged by the
lungs daily is 14,000 grms.
Bacrerta IN Sweat.—Eberth (Cenéralblatt, No. 20, 1873) finds
that in ordinary as well as in yellow sweat there are multitudes of
bacteria. These bacteria are small, oval, united in strings of twos
and threes and endowed with living movements. In spots covered
with hair they attach themselves to the hair and even penetrate into
the hair, which then splits and breaks.
Circulatory System.
“Circulation and its Disturbances in the Frog’s Lung.’”—Hiiter
in Centralblatt, Nos. 5 and 6, 1873. “Physiological Action of
REPORT ON PHYSIOLOGY. 189
Digitalis on the Circulation and Temperature.” Abstract in Cenéral-
blatt, No. 13, also Brunton and Meyer, Journal of Anat. and Phys.
vu. Nov. 134, 1872. “On the Physiology of the Circulation in
Plants, in the Lower Animals and in Man.” Lectures by Pettigrew,
Edinb. Med. Journ. 1872—73.——*“ Law which Regulates the Fre-
quency of the Pulse.” Garrod in Journ. Anat. and Phys. vir. 219.——
“On the Work of the Frog’s Heart under varying Blood Pressure.”
Blasius in Arb. aus d. Phys. Lab. d. Wiirzburger Hochschule, Pt. 1. p. 1
and ‘On the Variation of the Blood Pressure in different parts of the
Circulatory System,” by Fick, in cbed. Pt. 11. p. 183.
PRESSURE IN THE PeRicaRDIUM.—Adam Kiewiez and Jacobson
(Centralblatt, No. 31, 1873) have measured in living animals the
intra-pericardial pressure by means of a small trocar with a short
stiletto point introduced through the 4th intercostal space at the left
margin of the sternum into the pericardium. On sheep, dogs and
rabbits, when the experiment succeeded, i.e. when the pericardium
was perforated with no loss of blood and no injury to the cardiac
muscles, they found without exception a negative pressure within the
pericardium. The negative pressure varied between — 3 and —5 mm.
Hg. By gentle respiration they never saw this bound exceeded, but
in the rabbit with artificially induced dyspneea, they observed it to
fall to -9 mm. Hg. The force with which the venous blood is
drawn to the heart is less important than is generally believed.
Donders estimated it in the respiratory pause at about 7} m., after
ordinary inspiration at least 9, after deepest 30 mm. Hg. These
authors’ results are a half lower.
CIRCULATION IN THE Coronary ARTERY.—Archives de Médicine,
Feb. 1873. M. Rabatel has ascertained that in the horse the current
of the blood in the coronary artery undergoes a double variation.
The first increase strongly pronounced, and coincident with the
ventricular systole, is due to the contraction of the ventricle; the
second coincident with the diastole is explained by the facility with
which the blood passes through the peripheral capillaries.
On THE ConDITION OF THE Bioop STREAM AFTER LIGATURE OF
THE VENA Porta#.—H. Tappeier (Ludwig’s Arbeiten, vu. p. 11).
It has long been known that rabbits die very soon after ligature of
the vena porte, and with a great decrease in the arterial pressure.
The decrease of arterial pressure and the death of the animal have
been explained by the previous authors (C. Ludwig and Thiry) upon
this subject as due to the accumulation of such a large quantity of
blood in the rootlets of the vena port so that there is not sufficient
blood in the other vessels and parts of the body to carry on the
circulation. F. Hofmann and the author have found that after ligature
of the vena porte and death had resulted, in a rabbit the quantity of
blood which had accumulated in the rootlets of the vena porte was
only 0-8 per cent. of the weight of the animal. By performing
experiments with blood-letting the author has found that the abstrac-
tion of a quantity of blood equal to 1°3 per cent. of the weight of the
190 DR STIRLING.
body does not cause the death of the animal, and gives quite other
changes in the arterial pressure. The author thus shows that the
explanation of the previous authors is not tenable. Studying the
curves of decreasing blood-pressure during the time of ligature of the
vena porte, the author has found that the decrease takes place regularly
in cases where the animal remains quite quiet. This decrease however
is not regular when muscular contractions and irritation in the region
of the vaso-motor nerves are present. The variations by which the
heart-beats and respiration are represented in the arterial curves
become after ligature of the vena porte at once flatter, and then
disappear altogether when the arterial pressure has already clearly
decreased. After removal of the ligature in some cases, the increase
in arterial pressure begins and rises ‘until the tension that was present
earlier is reached. In other cases the arterial pressure may remain
low for some seconds or even minutes before it begins to rise, which
it does at first gradually, then more quickly. ‘Active or passive
movements during the ascent accelerate clearly the increase of the
pressure already present. At other times removal of the ligature is
not followed by a spontaneous increase in the arterial pressure.
With regard to the pulse-beats, in all cases the initial diminished
frequence increases with the duration of the experiment, so that when
by the continued occlusion the animal is brought near to death, there
is a period in the middle between the initial slowing and that induced
later by dying, in which the pulse is accelerated. Section and irrita-
tion of the spinal cord during ligature of the vena porte produces an
increase of the arterial pressure, and the waves of the pulse are to be
seen more clearly. Studying the rapidity of the movements of the
blood in the carotid when the vena porte is ligatured, the author
found that it is always decreased. For a description of the apparatus
by which the above results were obtained, we must refer to the
original.
Unirormity or THE Hearts WoRK WHEN THE ORGAN IS NOT
SUBJECT TO ANY Exterior Nervous InFrLuence.—Marey (Comptes
ftendus, Aug. 4, 1873, p. 367) seeks to establish the theory that (the
innervation remaining constant) the heart which never rests executes
an amount of work sensibly uniform, its beats being rare when each
of them has to overcome a considerable resistance ; they are frequent,
on the contrary, when that resistance diminishes. The resistance to
the effort of the heart is the pressure of the blood already contained
in the arteries. He then cites a number of facts by which he supports
his theory. With regard to the experiments of Cyon with the
depressor nerve, the heart is in this case influenced by the nervous
system. Irritation of the central end of this nerve produces a retar-
dation of the heart’s beats and consecutive diminution of the arterial
pressure. Certain facts, the author remarks, seem to stand in opposi-
tion to his theory, viz. that even when in three rabbits Cyon had
destroyed all the nerves which run along the vessels, so as to isolate
the heart from all external nervous influence ; on the depressor nerve
being excised under these conditions, retardation of the heart-beats
REPORT ON PHYSIOLOGY, 191
followed. The author thinks that some nerve-fibres of the pneumo-
gastric may not have escaped the scalpel. Ludwig has shown that
the excised heart of a frog (and even the heart of a rabbit) when filled
with serum, will continue its movements for a long time, and Bow-
ditch, Coats and Cyon have measured with a manometer the energy
of its movements. Such an excised heart (Tortoise), free from all
external nervous influence, was fitted with an artificial circulatory
apparatus formed of caoutchouc tubes, in which circulated fresh calf’s -
blood. Blood was introduced into the veins and auricles from a
raised reservoir by meaus of a syphon, passing from the ventricles to
the arteries, the blood was forced into the elastic tubes, and was again
poured into the reservoir. Jn spite of a high temperature, this circu-
lation was maintained over five hours, and the following experiment
was repeated a great number of times. Every time upon contracting
the orifice of outflow of the arterial blood, only raising it to a greater
or less height, the pressure in the arteries thereby being increased,
the heart’s movements were retarded. Every time on the contrary on
diminution of the pressure, the heart-beats were accelerated. In the
absence of all communication with the nervous centre, the heart beats
more quickly, in proportion as it performs less work in each of its
beats.
A Perriopic Function or THE IsoLatep Froe@’s HEart.—
L. Luciani (Ludwig's Arbeiten, vu. 115). When the author ligatured
the auricles of a frog’s heart, into whose ventricle a canula had been
introduced through the sinus venosus, and thereafter the excised
heart filled with serum and brought into connection with a small
registering manometer, he observed, instead of the long pause in
which the heart generally remains after the method of ligaturing of
Stannius, that a long series of pulsations followed in a very charac-
teristic, intermittent manner. To study more exactly this function
the author employed the method of Bowditch (Ludwig’s Arbeiten,
vi. 139), with certain modifications, for a description of which we
must refer to the original. By means of a canula, which had at one
end five small rings adjoining each other (each ring 2mm. broad),
the author could ligature the ventricle at any height he wished. This
intermittent pulse-rhythm shows itself in every frog’s heart filled
with serum, by a paroxysm (“ Anfall”) of many frequent beats,
which is then followed soon, or after a longer pause, by a series of
slower beats. Then the groups (“Gruppen”) first appear, consisting
of two and more seldom sixty contractions (when the serum contains
many blood corpuscles), which are more frequent towards the middle
of the group than at the beginning or end of the same. Pauses of
from 20 seconds to 10 minutes separate the groups. In one and the
same heart however there is often a remarkable regularity in the
proportion of the groups and pauses. When such periodic work has
lasted for a long time (often hours) the groups, which are already
chiefly composed of lower beats, become dissolved, and merge into
isolated contractions (“ Krisis”). The act of ligature produces often
(especially on ligaturing near the sulcus) a very high and long
192 DR STIRLING.
(seldom more than 9 minutes) continued tonic contraction of the
heart, upon whose curve often single beats are marked, sometimes
arranged in groups separated by pauses. On loosening the ligature
the tonus disappears immediately. Ligature of the places tied can-
not again produce these results; the periods therefore following the
tonic paroxysm are not to be regarded as phenomena of irritation,
but as consequences of the separation of the centres regulating the
- beats. The influence of the serum is shown in that upon its renewal
it makes the beats in the groups higher and more frequent and the
pauses shorter. Warming of the heart from 24° to 30° Cels. makes
the groups and pauses shorter, the pulsation higher and much
quicker, so that the rapidly widening heart causcs the mercurial
column in the manometer to oscillate under the abscissa ; cooling
to 3° Cels. produces an opposite effect. The single beat lasts often
more than four seconds. The height of the beats following each
other diminishes very gradually. The absolute number of beats
in a given time under all the above conditions is smaller when the
beats are arranged in groups than when they follow each other in
the normal manner, but are equal (to those of the uninjured heart)
with regard to height and duration. The pauses are longer in a heart
beating in oil than in one filled and bathed with serum. One can
therefore conclude that during the pause the irritation is actually
absent, not, however, that its manifestation is only suppressed by the
increased resistance. In the course of a normal pause, if contrac-
tions are produced either by mechanical or electrical stimulation the
pause is thereby increased. The easily changed rhythm of the
periodic acting heart is easily affected by poisons. Nicotine and
atropia make the groups longer, the pauses shorter. The latter,
however, in small doses causes death of the heart without dissolution
of the groups in the period of ‘ Krisis.” Large doses cause at first
along series of frequent then more seldom beats, whilst nicotine
makes the pauses shorter, and produces “ Krisis” more rapidly with-
out disturbing the energy of the heart. Whilst new serum could
not reproduce the periods after atropia poisoning, this happened in
most instances after strong poisoning with nicotine. A heart poisoned
with nicotine that had already passed through all the phases from
“Anfall” to “Krisis” could be brought to a new periodic action
when it was either filled with new serum or again poisoned with
nicotine. After:a strong dose of extract of muscarin (5 milligr.) the
heart remained motionless (as already known) after it had made
‘a short group (sometimes peristaltic) of beats, but it reacted upon
mechanical and electrical irritation. Careful washing out of the
heart with fresh serum produced no spontaneous beats. A new
ligature produced tetanus, upon whose curve single beats were
marked, but immediately after the heart again relapsed into rest.
Medium doses (2 milligr. ext. musc.) make the groups shorter, the
beats more seldom and smaller, and the pauses longer. Minimum
doses of muscarin (0-05 milligr. of the sulphate), after a stage of
excitation, in which the height and frequency of the beats are in-
creased, appear to produce the depressing effects of medium doses.
REPORT ON PHYSICLOGY. 193
Atrepin prodaced its effect regularly upon the muscarin poisoned
heart as well as on the normal one. On the contrary, nicotine ean-
not annul the action of muscarin. All the facts seem to the author
to render the idea untenable, that there are present in the heart sepa-
rate inhibitory and motor centres.
CHEMICAL IRRITATION OF THE NERVES OF THE Herart.—Mosso
(Lo Sperimentale Anno, xxiv. 1872. Abstract in Centralblatt, No. 14,
1873) lends some confirmation to Schiff’s old view that the vagus is
the probable or only motor nerve of the heart. He chemically irri-
tated the cardiac nerves in dogs poisoned with atropine. In these
animals the heart-beats were rendered independent of blood-pressure
by the sub-cutaneous injection of atropine, and the nervi vagi recur-
rentes were irritated by the careful application of a drop of solution
of caustic potash. From his experiments he deduces the following
conclusions: (1) Irritation of the vagi increases the frequency of the
pulse in consequence of the excito-motor fibres running in the trunk
of the vagus. (2) If the sheath of the vagi-sympathetic be opened,
and the sympathetic be separated from the vagus, chemical irritation
of the latter constantly produces an increase of the pulse frequency,
whilst excitation of the sympathetic is without perceptible effect on
the rhythm of the heart. (3) Mechanical irritation of the inferior
laryngeal nerves by simple section is sufficient to increase the pulse
frequency, and (4) this quite independent of an increase in the blood-
pressure.
AcTION OF THE VaGus ON THE Heart.—-Metschnikoff and
Setschenow (Centralblatt, No. 11, 1873) from their experiments are
led to the opposite conclusion from that of Mosso. When in a
turtle (Emys Europea) the irritation of the vagus is carried beyond
the bounds, which are known as the phenomena of exhaustion, a
very striking picture of the periodicity of the inhibitory action of
the vagus is obtained. The phenomena were much the same whether
the animal had the brain destroyed or not (with intact vagus on the
other side), as well as after section of the branch of the sympathetic
from the vagus. They draw the two following provisional conclu-
sions: (1) The inhibitory action of the vagus on the heart of the
turtle is periodic. (2) In the periodicity of this action is a new
proof that the inhibitory fibres of the heart end in a sort of nerve
centre. In Centralblatt (No. 19, 1873) Setschenow finds also that in
the heart of the frog with brain and spinal cord destroyed the action
of the vagus is periodic. In frogs the pulse-frequence never rises
during the negative phase over the normal. Repechoff investigated
the reflex inhibition of the lymph heart by the vagus, and the cor-
responding phenomena in the blood heart by the sympathetic. In
both cases by a continued stimulation of the nerve the effect was
certainly periodic. “ Cause of the retardation of the Pulse follow-
ing artificial or voluntary closure of the nostrils in the Rabbit,” by
W. Rutherford, in Journ. of Anat. and Phys. vu. 283.
On THE INTERFERENCE OF THE RETARDING AND ACCELERATING
Nerves or THE Heart.—H. P. Bowditch (Ludwig’s Arbeiten, vu.
VOL. VIII. 13
194 DR STIRLING.
259). Ina short history of the N. accelerans cordis, whose discovery
by the brothers Cyon and A. v. Bezold gave full confirmation to the
idea of the latter, upon the necessary connection of the cardiac nerves
with the spinal cord, the author gives the results of the researches of
M. and E. Cyon upon this nerve in rabbits and dogs, and shows how
A. v. Bezold, Beven and Schmiedeberg have increased our knowledge
of its function. The latter showed that it is also present in the frog.
The branches of the accelerans and those of the vagus are united into
one stem before they reach the heart, so that it is possible to irritate
both equally strongly, and at the same time. By doing so, we see at
first only the inhibitory action of the vagus, and by continuing the
irritation there is consequent exhaustion of the vagus, and its action
becomes weaker and weaker. After interruption of the irritating
stream, the complete action of the accelerating nerve appears. Simi-
lar results are obtained when the isolated vagus of the one side, and
the N. accelerans of the other, are irritated by currents of the same
strength.
It is now necessary to ascertain if both nerves acting oppositely,
when these are irritated, yield a medium value of heart-beats, which
each would have given if irritated by itself. For this purpose the
author used dogs, poisoned with curara, and isolated the N. accelerans
by the method of O. Schmiedeberg (Journal of Anat. and Phys.
Vol. vir. 180). The pulse was written partly with a mercurial and
partly with the spring manometer. For a special apparatus for
recording the time, we must refer to the original. At first the effect
of the maximum irritation of the carefully isolated N. accelerans was
studied. No exact proportion could be made out between the form
of the curve and the duration, strength, and consecutiveness of the
irritation. But probably a nicer gradation of the irritation might
yield some results. The latent period varied (from 1 to 22 sec.) in
the same and different animals, and was independent of the strength
of the induction stream. The only indication of regularity exhibited
was that the latent period was shorter when a second irritation fol-
lowed the previous one before the acceleration of the pulse which it
had produced had disappeared. The maximum value which the num-
ber of pulse-beats reaches in a given time in the same animal, within
narrow limits, unmistakably depends upon the strength of the
irritation. .
This value, however, is very unequal in different animals. It is
a rule without exception, that the time of the duration of the maxi-
mum of heart-beats is proportionally short. It is of importance
to know how many beats more the heart produces in a given time,
when the N. accelerans is irritated, over the number obtained in the
same time without irritation. The result is, that the increase of the
pulse-beats is greater with the duration of the irritation, but not at
all proportionally. In addition to acceleration of the pulse, the
irritation of the nerves which are contained in the path of the N.
accelerans often produces an increase of the mean arterial pressure.
But even where the pressure and the pulse increase at the same time,
there is a complete independence the one of the other. Without
REPORT ON PHYSIOLOGY. 195
exception, after irritation the arterial pressure reaches earlier its
maximum than the acceleration of the pulse, and still further, when
the pulse-beats have reached their highest value they sink regularly
to that which they had before the irritation. The arterial pressure,
on the contrary, sinks with wave-like variations under its normal
value, and without exception the waves of pressure become lower and
shorter the further they are from the beginning of the irritation.
The independence of the acceleration-wave, and of the variation in
the pressure, are best explained by supposing that there are in the
irritated stem two nerves of different functions, one of which acts
upon the heart, the other upon blood-vessels elsewhere, in the same
way as O. Schmiedeberg has demonstrated that, by irritating a branch
proceeding from the ganglion stellatim only, the pressure-wave is
produced without at the same time any acceleration of the heart’s
beats. The author also observes that in some dogs, but not in all,
as Fick has already shown, a dicrotic pulse is produced. In those
animals in which it was observed it always disappeared when the
pulse-beats reached from 210 to 220 in a minute.
By observing the changes of the rhythm of the pulse following
irritation of the N. accelerans, the author compares them with those
that are obvious in a frog’s heart exhausted but warmed, or in the
exhausted frog’s heart filled with fresh blood serum. The antagonism
between the N. accelerans and N. retardans is at least not a complete
one. Maximum induction currents, acting at different times upon
the isolated accelerans, gave an approximately equal frequence of
pulse-beats ; the regular tetanisation of the same part of the N. vagus
yields not so satisfactor y results, e.g. the distance of the coils necessary
to produce the slightest perceptible retardation of the pulse must be
altered for consecutive irritations. The cause of this irregularity is
very probably not to be sought for in exhaustion of the nerve-trunk,
but in a changing susceptibility of the automatic apparatus to the
irritations of the vagus. It is necessary to find the minimum irrita-
-tion of the vagus, by which it does not lose its effect on the N.
accelerans, to employ this by itself, and then at the same time
to employ the maximum of the N. accelerans. From the curves
figured at the end of the paper, it is shown that a very weak irrita-
tion of the vagus is sufficient completely to overcome the effects
of even a maximum iuritation of the N. accelerans. From the study
of the curves obtained from different animals, it is possible that very
weak degrees of irritation of the vagus may not be able to overcome
the maximum irritation of the N. accelerans. The vagus has not the
power to abolish the condition produced in the heart by the N. ac-
celerans. When the pulse, rendered quicker by the accelerans, was
rendered slower by the vagus, the increased pulse-beats returned
when the vagus was removed from the induction circuit.
By an increase of arterial pressure, causing irritation of the
central ends of the vagus, the author made further investigations
to see if this natural irritation would disappear under maximum
irritation of the accelerans. For these experiments of course only
those animals could be employed whose pulse is retarded by the
13—2
196 DR STIRLING,
increased pressure. The result was, that the irritation of the vagus,
effected by increased arterial pressure, was strong enough to overcome
a maximum one of the N. accelerans. But, as it has been shown
that irritation of the N. accelerans by itself alone can produce an
increase of the arterial pressure, so it may happen that by increase
of the arterial tension the effects of irritation of the accelerator nerve
may be balanced.
So in this manner it can be understood why sometimes in the
course of an acceleration, caused by irritation of a very carefully iso-
lated N. accelerans, there appear suddenly some heart-beats with
long pauses. In curarised dogs, dying from asphyxia, and in which
the blood had already become of a deep dark colour, the author found
that the N. accelerans had not lost its irritability but was very
active, and it seems to be an important fact for the condition
produced in the heart by irritation of the N. accelerans, that oxyge-
nated blood is not necessary for the same.
VARIATIONS OF Ha2MOGLOBIN IN THE ZooLocicaAL SERIES.—
Quinquand, Comptes Rendus, August 18th, 1873, p. 487. The
method consists in determining by aid of a liquid triturated with
hyposulphite, the maximum quantity of oxygen, absorbed by the
blood,which ought to be effected in five minutes, using 2 ce. of blood
(for details, Comptes Rendus, Lxxvi., 1489, or Lond. Med. Rec., No.
30). (1) The progressive diminution of the quantity of hemoglobin
contained in the same volume of blood follows in general the degrees
in the animal scale. At the same time the blood of the primates
is not that which contains most. (2) The blood of young animals
is less rich in hemoglobin than that of adults; in many species the
placental blood contains as much as the blood of the general circula-
tion. In old age the quantity diminishes. The curve of variations
would be represented by a slight fall at first corresponding to the
first days of uterine life, the curve then rises in the infant, and
remains horizontal during adult life (25 to 50 in man), after which
it slowly falls in old age.
EsTIMATION OF THE MrnERALS OF BiLoop-SeruM By Direct PRE-
crpiraTion.—L, Gerlach (Ludwig's Arbeiten, vii. 99). Dr Pribram’s
experiments showed that it is practicable to precipitate from fresh
blood-serum all the lime and a part of the phosphoric acid which
formerly could only be obtained from the ashes of the serum (Jowrn.
of Anat. and Phys. vu. 190). Dr Pribram, however, in his ex-
periments took no notice of the magnesia. The author, from his
experiments, finds that magnesia as well as lime can be precipitated
directly from the serum, but to obtain reliable results it is necessary
to precipitate the lime in an acetic acid solution to prevent the
contamination of the precipitate by ammoniaco-imagnesian phosphate.
Wuat Constituents oF ASPHYXIATED BLOOD SERVE TO BIND THE
DirFusisLE OxycEn!—Afonassiew (Ludwig's Arbeiten, vu. 71).
Alex. Schmidt has shown that asphyxiated blood possesses the pro-
perty of changing a portion of the oxygen which is added to it, and
REPORT ON PHYSIOLOGY. 197
that the whole of the oxygen cannot be recovered from blood so
treated. In place of an aliquot part of the oxygen, CO, is obtained.
The author, under Ludwig's direction, wished to know “whether the
stuff which holds this oxygen is contained in the serum or in the
blood-corpuscles. He began with the investigation of the compo-
sition of the serum. The serum was obtained free from blood-
corpuscles by means of the centrifugal apparatus. He mixed equal
volumes of this serum and asphyxiated blood (taken from an as-
phyxiated animal nearly dead). The quantity of CO,, oxygen and
nitrogen contained in the serum as well as in the blood added, was
estimated so that the contents of the mixture in gases could be
easily derived therefrom. The gases were pumped out of the mix-
ture and analysed. In five experiments the quantity of CO,, oxygen
and nitrogen obtained was so near the quantity calculated, that it
is quite certain that the serum of asphyxiated blood does not con-
tain any substance which could cause the formation of CO, from
the oxygen of the blood-discs. In one experiment the quantity of
CO, actually obtained was 32°23 per cent., while the quantity reck-
oned was 32°40 per cent., and the actual quantity of oxygen obtained
7°24 per cent., the calculated amount 7°31 per cent. ; small variations
quite within the limits of analytical errors. The author now added
a given quantity of oxygen to a known quantity of asphyxiated
blood whose gaseous contents were known, and then ascertained the
amount of gases in the same. Thus in one of the experiments the
asphyxiated blood contained 1:48 volumes per cent. of oxygen, 11:12
per cent. of oxygen was added, 11°86 per cent. of oxygen was ob-
tained. So that 0-74 volumes per cent. of oxygen had disappeared
and 0°37 per cent. of CO, more was obtained. The serum of the
same blood, on the contrary, absorbed only 0:21 volumes per cent.
of oxygen and yielded 0:16 volumes per cent. more of CO, Ina
second experiment, after the addition of oxygen, 1:04 volumes of
oxygen disappeared from the asphyxiated blood and oxygen, ‘93
volumes CO, more was obtained. From this it appears that the
substance in asphyxiated blood, which allows the finding of the
oxygen and the formation of CO,, i is present in the floating ‘elements
of the blood (red and white corpuscles).
Expunsion oF Nirric Oxipe FRoM THE Broop.—Zmitz and
Donders have shown that CO can be expelled by other gases, e.g.
oxygen, from its hemoglobin compounds. Podolinski (Pfliig.
Archiv, vi. 1872 , 953) has also succeeded in driving out completely
CO from blood containing it, by means of oxygen (atmospheric air),
and also by hydrogen, though hydrogen acts a little slower. Nitric
oxide can also be expelled by hydrogen, though here also longer time
is required than for the CO.
EstiMATION OF THE ABSOLUTE QuaANTITY OF BLoop.—Steinberg
(Phiig. Archiv, 1872, vit. 101) in his researches has adopted the
method of Preyer, viz. the giving of a green light in the spectrum
by a certain estimated solution of heemoglobin, which serves as a
test object for the washings. The proportion of blood to the body
198 DR STIRLING.
weight in the rabbit was 1 : 12°3 to 13:3. In guinea-pig, 1 : 12
to 12°3. Adult dog, 1:11:2 to 12°5. Young dog, 1: 16-2 to
17°8. Adult cat, 1: 10-4 to 11:9. Young cat, 1:17°3 to 18-4.
The colorimetric method of Welcker adopted by Bischoff, Heidenhain,
Ranke, &e. yields very similar results.
Minute Movine Particies as Constant Constituents oF Nor-
MAL Bioop.—Nedsvetski (Centralblatt, 1873, No. 10) finds that in
perfectly fresh human blood, examined with a power of not less than
900 to 1000 diams., there is to be observed in addition to the red
and white corpuscles very small spherical bodies without cilia of the
size of the granules of the white blood-corpuscles, which exhibit move-
ments of rotation round their axis, and move from place to place.
No structure is to be observed in them, Nedsvetski has named these
bodies harmo-cocci. They attach themselves to the filaments of fibrin
when it is found under the microscope. Other bodies, whose source
seems to be the white blood-corpuscles, are also described by this
author. Still other granules are to be distinguished ; they resemble
the white blood-corpuscles, and are to be distinguished from them
by being more rounded, and by being devoid of movement. They
remain longer without change than the other elements of the blood.
BAcTERIA-FORMING MAssES PRESENT IN Bxroop.—Osler and
Schifer (Centralblatt, 1873, No. 37). In many diseases there is
present in the blood a greater or lesser number of colourless granular
bodies, of the same size, or often larger than the white blood-cor-
puscles, and consisting of small pale particles) When a drop of
blood is diluted with solution of NaCl (2 per cent.) and kept at the
temperature of the body, small fibres are observed to be pushed out
from the surface of these masses which soon exhibit a violent vibra-
tile movement, while at the last they separate themselves from the
mass and float free in the fluid. Every fibre exhibits either in its
middle or at one of its ends a swelling, which, according to the
position of the fibre, appears either circular or linear. The swelling
is not globular, but discoid. These masses are present sometimes in
apparently normal human blood as well as in that of animals, but
they do not exist previously as such in the blood-vessels. They are
formed after extraction of the blood by the flowing together of the
pale particles from which they are made up.
Action oF Quinine on Buioop.—Binz (Arch. f. Expt. Path. 1.)
pointed out some years ago that Quinine arrested the movements
of the white blood-corpuscles, and this he explains by the drug di-
minishing the oxydizing power of the red blood-corpuscles, for the
white corpuscles are only active when supplied with oxygen, which
is yielded up to them by the red corpuscles as they pass. Binz has
further found that when oxygen is withheld from them they do not
penetrate the walls of the blood-vessels, and his observation has been
confirmed by Heller and Zahn.
CoLourine Marrers or tHE Bioop.—H. Struve (Virchow’s
Archiv, uv. 423) has succeeded in discovering two colouring mat-
REPORT ON PHYSIOLOGY. 199
ters in the blood. The one, probably identical with Virchow’s heema-
toidin, is crystalline, dark blue in colour, insoluble in water, alcohol,
ether, chloroform, and acids, but dissolving in weak alkalies, giving
the solution a brownish tinge. On heating it, ammoniacal fumes are
evolved. The other is soluble in water and in alcohol, but with
difficulty in ether, and is probably identical with Von Wittich’s
heematin.
ON THE SUGAR-PRODUCING FERMENT OF Bioop.—P. Plész and
E. Tiegel (Pfliig. Archiv, 1873, vi. 391). When defibrinated blood
is mixed with a weak solution of NaCl, and the blood-corpuscles
allowed to separate, the supernatant fluid contains the ferment in
large quantity, while the blood-corpuscles contain none, or only a
very small quantity. The authors conclude that the Na Cl solution
has extracted the ferment from the blood-corpuscles, and quote the
analogous phenomenon that it is possible by a 3 per cent. solution
of NaCl to extract from fibrin a sugar-producing ferment nearly
allied to globulin. The authors, in repeating Bock-Hoffman’s experi-
ments, found in the excreted urine a sugar-producing ferment ; also
in diabetic urine a substance which, when freed from sugar, possessed
the same property. The authors’ experiments render it probable
that in the Bock-Hoffman experiments the formation of sugar takes
place not in the kidneys, but in the blood or liver. With regard
to V. Wittich’s experiments, by which he sought to prove the exis-
tence of a special liver-ferment which changes glycogen into sugar,
V. Wittich has stated (Pjliig. Archiv, 1873, 28, abstract in Lond.
Med, Rec. 1873) that in a liver quite free from sugar and from which
the blood has been completely washed out, after some time the sugar
returns, and the sugar-producing ferment can be demonstrated. The
authors think it more probable, however, that by the washing out of
the liver with water, the ferment contained in the blood was dis-
solved, and fixed in the coagulated liver-cells.
“ Presence of Soluble Earths and Phosphoric Acid in Alkaline
Blood,” Fokker, in Pfliig. Arch., 1873, vir. 274. “Absorption of
Oxygen by Blood,” Gréhaut, in Comptes Rendus, txxv. 495, abstract
in Lond. Med. Kec., No. 17, ‘ Physical Nature of the Coagulation
of the Blood,” A. H. Smee, in Journ. of Anat. and Phys. vu. 210.
“Observations on Welcker’s Method of Estimating the Quantity of
Blood,” Gscheidlen, in Pfliig. Arch. vit. 530.
Lymph.
SzcreTIoN oF Lympn IN THE Fore-times or THE Doc.—(Lud-
wig’s Arbeiten, v1. 198.) Paschutin has selected by preference the
fore-limbs of the dog as the source of the lymph, because the roots of
the brachial-lymph stem arise only in the skin and muscles, and the
lymph is not mixed with that of other tissues, as would be the case
if it were obtained from the truncus thoracicus, and, further, because
the circulation in the limb is easily modified, and its nerves excited.
On the living animal, after cutting the skin at the outer margin of
200 DR STIRLING.
the vena jugularis, all small arteries must be tied, and alt blood re-
moved. The A. transversa colli is next sought for by aid of a blunt
instrument, and two ligatures are placed upon it, and the artery cut
between them. Immediately in the neighbourhood of this vessel,
but nearer the spinal column, the lymph stem is to be found. By
pressing the lymph from the limb upwards towards the head, the
trunk is easily rendered visible. Place a ligature on the vessel as
near as possible to its place of opening into the truncus colli; but
before tying the ligature see that no small branches join the vessel
from the scapular or pectoral regions, if so these must be ligatured.
The brachial stem must be carefally isolated for 5 mm., a ligature
placed round it, and a canula bound in it. The best material for
making the canula is glass. The observations of Generisch and
Lesser have shown that the lymph only flows regularly from the
limbs, when these are actively or passively moved; and that further,
the rapidity of the out-flow probably depends upon the intensity of
the movements and the tact displayed. It was therefore necessary
to have the limb moved regularly, and the method by which this was
obtained, by means of a machine, is to be found in the original. In
order to vary the experiments it was necessary to divide the plexus
brachialis, the cervical spinal cord, or both. The plexus brachialis is
easily reached through the original wound in the skin. In dividing
the spinal cord in the neck, the best spot is at the lower margin of
the second cervical vertebra; but the loss of a small quantity of
blood causes the death of the animal. It is therefore better, in lay-
ing bare the vertebra, to ligature the large branch of the A. profunda
cervicis, which runs backwards on the middle layer of the cervical
muscles. After section of the spinal cord artificial respiration is kept
up, and the electrodes for irritation of the spinal cord applied. The
blood-pressure was measured in the A. carotis.
I. The medium rapidity of outflow of Lymph.
1. ATI experiments show that with the duration of the experi-
ment the out-flow of lymph is diminished, i.e. when the circumstances,
under which the animal is operated on, remain the same. But it is
different whether the animal is poisoned or not, or placed under the
ordinary temperature of the air or if the temperature is higher, or
whether the spinal cord is injured or remains intact. With regard
to the source of the lymph it is easy to show that one has not to do
merely with an emptying of the lymph vessels, for when the lymph
is carefully pressed out of the limb with the hand, and this is
again repeated immediately thereafter, employing very powerful pres-
sure, no greater quantity will be obtained. The assumption that
during the time of the experiment a store of tissue juice is expelled,
the author cannot refute, considering the striking proofs which Ham-
marsten has had, on account of the large quantity of lymph which
he obtained. But not less important is the observation that in one
experiment, after the weak movements of the limb caused by the
machine had been followed by the powerful one caused by the hand,
the rapidity of out-flow rose suddenly, and lasted 25 minutes, without,
REPORT ON PHYSIOLOGY. 201
however, sinking to the small quantity which was obtained before
the commencement of the powerful pumping. From this, and spe-
cially from the composition of the lymph, every portion of which has
a different value, it seems to the author very probable that the out-
flowing lymph is actually secreted during the period of movement.
And again, in several experiments the out-flow of lymph during the
period of rest either ceased entirely, or only a very small quantity
could be obtained on pressing the limbs; but as the pumping move-
ment again increased, no more lymph flowed than was dropped by
the same movement before the rest. It is to be observed that these
results were obtained also on animals whose cervical spinal cord and
plexus brachialis were cut. The observation that the secretion of
lymph ceases during the period of rest is not valid for all cases.
2. Action of Curara on the Secretion of Lymph.—The experi-
ments of Lesser rendered it very probable that curara modified the
secretion of lymph. The cervical spinal cord and plexus brachialis
were cut, and then the lymph pumped out for a certain time; the
animal was then rapidly poisoned with a sufficient quantity of curara,
and the lymph obtained as before. After curara poisoning the ra-
pidity of secretion increases, reaches its maximum in 40 to 50 minutes
afterwards, and then by degrees gradually decreases. This accelera-
tion takes place not only during the period of movement, but also
during rest. The increase and decrease of the secretion does not run
parallel with the arterial blood-pressure. Curara produces redness of
the skin. It also modifies the composition of the secreted lymph.
3. Change of Lymph Secretion in consequence of an increased
supply of Blood.—The simplest method to produce arterial congestion
in the fore-limb of a curarised dog is to lay bare carefully the brachial
plexus, then pump out the lymph under these conditions, and
then, after the section of the plexus, to proceed with gaining the
lymph. After this the foot became warmer, and blood flowed from
the foot, after puncture with a needle, much more freely than be-
fore. Tables of the results so obtained show very clearly that in-
creased supply of blood is without influence on the secretion of
lymph, that its rapidity fell regularly after section of the nerves, not-
withstanding that the blood-pressure after the same was higher than
before, and notwithstanding that the foot and upper part of the ex-
tremity was more richly supplied with blood. To be without doubt
on this point, the cervical spinal cord was cut and electrodes applied,
and at the same time the plexus brachialis was divided. One could
therefore change, after irritation of the spinal cord, the lymph caught
during high and during low blood-pressure, and therefore be sure
that the change of the blood-pressure measured in the carotid also
took place in the fore extremity, because the vaso-motor nerves of the
same were paralysed. From numerous results it is impossible to
doubt that in the dog increased blood-supply is without influence on
the secretion of lymph. At no time was there produced by the ap-
pointed irritation even a stoppage in the sinking of the rapidity of
secretion, to say nothing of an increase of the latter. These results,
therefore, stand in open contradiction to the idea of the dependence
202 DR STIRLING.
between the blood-pressure and the lymph secretion. Till now it
was believed that the cause of the movement of fluid from the blood-
vessels in the tissue spaces was founded in the difference of pressure
which was active on both sides of the wall of the vessels.
4. Change of the Secretion of Lymph by increased temperature of
the body.—TYo test whether the cooling which the animal undergoes
when it has lain for a long time on the operating table is the cause
of the secretion becoming smaller and smaller with the duration of
the time of observation, the animal was aliowed to cool at the ordi-
nary temperature while the lymph was obtained, and whilst the
lymph was being obtained the animal was warmed from without.
For this purpose the double walled box (described at p. 203) was em-
ployed. An increased temperature accelerated the out-flow of the
lymph.
II. Per-centage composition of Lymph Serum in fixed residue.
1. To free the lymph serum from fibrin and corpuscles the lymph,
which during the catching was shaken before evaporation, was placed
in closed vessels in the centrifugal machine, until the clear serum
could be poured off from the sediment. A weighed quantity of the
latter was carefully dried. In 84 different examples of lymph, taken
from different dogs or from the same dog at different times, the per-
centage composition of lymph serum in fixed residue clearly changed.
The variation was from 2-61 to 6:55 per cent. From numerous ex-
periments it is shown that the per-centage increases with crease in
the time of experiment. Similar results were obtained by Generisch
when he collected the lymph from surviving limbs whilst an artificial
blood stream was passed through the same. The formation of the
lymph is not due merely to transudation,
2. In the course of an experiment when the initial slowing of
the rapidity of the lymph-secretion again increases, the per-centage of
its serum very often decreases again, and this independent of the cause
which produces the acceleration. The relation which exists between
the lymph which flows out in a certain time, and its per-centage in
fixed constituents, is here very obvious. For when the rapidity of
secretion was accelerated by new means, in spite of the continuation
of these means, the rapidity becomes by degrees less and less, the
per-centage of the residue increases.
3. In one experiment, in the course of which the animal was
poisoned with curara, the contents in fixed constituents rose 1-2 per
cent. It is therefore to be expected that all the animals narcotised with
this poison before yielded a concentrated lymph. When in seven
cases the animals were curarised before the beginning of the experi-
ment, the per-centage of the first portion of lymph varied from 4-4
and 6-5 per cent., i.e. it had already reached in the beginning of the
experiment the worth which is only reached towards the end in
unpoisoned animals. In short, when in the curarised animals, in
consequence of a cause, the rapidity of out-flow of the lymph in-
creases, exactly the same proportions in the per-centage of the lymph
in fixed constituents occur as in the non-curarised animals, but not
REPORT ON PHYSIOLOGY. 203
with the same regularity. In other experiments the rapidity of
out-flow of lymph was increased by warming the animal. In these
experiments the contents of the lymph in fixed stuffs decreased,
although the rapidity of secretion had already begun to decrease. If
these results are confirmed, there would be proved a specific action of
increased temperature on the composition of lymph serum which was
secreted in animals with uninjured spinal cord. With regard to the
question whether the lymph is obtained by the passive movements of
the limbs, or is formed from a previously prepared store during the
period of the experiment, the different composition of the obtained
fluid speaks in favour of the latter. And further, this remains
scarcely doubtful when, the movement remaining the same, the com-
position of the serum changes after this more or less rich out-flow.
It further shows how insufficient it is to observe solely the volume
of the fiuid, for its composition must also be investigated.
It is not possible to explain the results obtained simply by the
categories of a propelling force and a resistance, for two conditions
which equally increase the out-flowing volume act quite in an
exactly opposite manner on the quantity of albumen of the serum,
as is the case with curara in opposition to the active movements of
the limbs and the warming of an injured animal.
Respiratory System.
Quinquand, “ Respiration of Fishes,” in Lond. Med. Ree. No.
22. Moin, “On Cubic Space and Volume of Air,” in Comptes
Rtendus, Aug., 1873, Abstract in Lond. Med. Rec. No. 33.
Schroetter, “‘ Movements of the Trachea.” Abstract in Lond. Med,
tec. No. 27.——“ Graphic Representation of the Respiratory Move-
ments,” Deutsch. Archiv f. Klin. Med., 1872, x. 124, and 1873, x1.
379. “Erectile Action of the Blood-Pressure in Inspiration,”
Hoggan, in Edinb. Med. Jour., 1872, ccvut. “ Functions of the
Nerves and Muscles of the Larynx,” Schech, in Zeit. f. Biolog., 1x.
Hft 2, 258. “Movements of Respiration,’ E. Lockenberg, in
Arb. aus d. Phys. Lab. d. Wiirzburger Hochschule, Pt. 1. 199.
“ Dyspnea from Warmth,” Goldstein, in ed. Pt. 1. 77. “On Res-
piration,” Nussbaum, in Pfliger’s Archiv, vii. 296.
Action oF InrercostaL Muscres.—From experiments on a
guillotined criminal, Onimus (Gaz. Hebr. de Méd. et de Chir., Jan.
24, 1873) has shown by means of electricity, that the external
intercostals raise the ribs and are inspirators, while the internal
ones depress them and are expirators, thus confirming Hamberger’s
theory: muscular contractility was further observed- not to dis-
appear in all the muscles at once. The diaphragm and muscles of the
tongue are the first to lose their excitability, while the masseters
retain theirs longer than any muscle of the face. (Abstract in Lond.
Med. Ree., No. 9.)
INFLUENCE OF ARTIFICIAL RESPIRATION IN STRYCHNIA POISONING.
Rossbach, Centralblatt, 1873, No. 24. The author concludes from
204 DR STIRLING.
his experiments on rabbits, that artificial respiration has no influence
either upon the preservation of the life of the strychnia-poisoned
animal or upon the intensity or duration of the spasms.
Apnaa.—Ewald (Pjliiger’s Archiv, vu. 575) used always the same
animals for comparing the oxygen contained in the blood. A quan-
tity of blood was obtained during ordinary respiration, and its gases
obtained. Apnoea was then produced by inflation, another quantity of
blood was obtained when the animal had completely recovered, and
so forth, and the quantity of oxygen was estimated at the different
periods. The chief results of these experiments are in the condition
of apneea ; the oxygen of the artificial blood is increased almost to
complete saturation, that of the venous blood is diminished, while the
CO, was clearly diminished in both cases. If the animal again
breathed naturally, the quantity of oxygen of the arterial blood sank
to the normal, the quantity of CO, also increased. By natural forced
respiration almost complete saturation of the blood by oxygen can be
produced.
That the diminution of oxygen in venous blood is not produced
by a diminution of temperature through the blood-lettings, whereby
the tissues would have increased need for oxygen, is shown in the
experiments where the animals were kept at a constant temperature.
Withdrawal of air during apneea causes asphyxiation much later
than during natural respiration. In apneea, after cessation of arti-
ficial respiration, after a long time (40 seconds), at first a scarcely per-
ceptible diminution of oxygen appeared, which, as well as the mentioned
retardation of asphyxiation, speaks for a diminution of the consumption
of oxygen. Experiments of Pfliiger, not yet published, however, speak
against this ; he observes neither increase nor diminution in the con-
sumption of oxygen in apneea. The author would explain the fact
mentioned through a diminution of the rapidity of the blood stream,
and he found that the blood-pressure during apneea was diminished
about one-third of the normal. During natural respiration the
oxygen contained in arterial blood was about 5-5 per cent. higher
than that of venous, during apnea about 7:1 per cent.
Alimentation.
“ Physiological Studies on the Action of Flesh-broth, Extract of
Meat, Alkaline Salts, and Kreatin.” Bogoslowsky in Arch. f. Anat. u.
Phys. 1872, 347. Seegen and Nowak, “ Estimation of Nitrogen in
Albumates,” Pyliig. Archiv, vit. 1873, 284. Pettenkofer and Voit,
* Feeding with Flesh and Fat,” Zeitschr. f. Biolog. 1x. 1873, 1.
Nasse, “Studies on Albumen,” Pjliig. Archiv, vi. and vi. 589.
Wolffhiigel, “Digestion of Fibrin without Pepsin,” in Pfliig. Archiv,
vir. 188. Sanson, “On a Mechanical Coefficient of Food,” in
Comptes Rendus, No. 24, 1490—1873. Abstract in Lond. Med. Ree.
No. 30. Mohlenfield, ““On Peptones,” Pjliig. Archiv, Vol. v.
Abstract in Lond. Med. Rec. No. 27. Von Wittich, “ Action of
Pepsin on Fibrin,” in did. “ Contribution to the Physiology of
REPORT ON PHYSIOLOGY. 205
Water,” Falck in Zeitschr. f. Biolog. vit. 398.——“ Absorption of
Fat,” Radziejewski in Virch. Archiv, 1872, v1. “On the import-
ance of Chloride of Sodium and its relation to Potassium Salts in the
human economy,” G. Bunge, Zeitschr. f. Biolog. 1x. Hft 1, 104.
AssIMILATION OF Fats.—Hoffman, Zeitschr. f. Biolog. vi. Ab-
stract in Lond. Med. Rec. No. 17, The animals were starved till all the
fat was supposed to have disappeared, when they were fed on nearly
pure fat, in order to determine whether fat is deposited in the tissues
from the food or not, or whether it first undergoes conversion. On
post-mortem examination it was found that the deposit is chiefly in
the liver and mesentery. Upon analysis it was shown that a con-
siderable quantity was assimilated and deposited in the tissues.
Brunyer’s GLanps.—Krolow (Jnaug. Dissert. Berlin, 1872) finds
that the fluid secreted by these glands has the power of converting
starch into sugar, and completely dissolves raw fibrin, but has no
effect upon fat and coagulated albumen.
RESEARCHES ON THE FUNCTION OF THE GLANDS OF THE INTESTINAL
Mucous Memprane.—Costa (abstract in Centralblatt, No. 20, 1873).
The experiments were made upon horses, which were killed during
digestion. The glands of Brunner and Lieberkiihn were specially
prepared after Von Wittich’s method (extracted by glycerine), and
with the fluid so obtained digestion experiments were made. His
results are: (1) That the glands of Brunner possess strongly the power
of converting starch into sugar, but have no effect on albumen or
fat; thus confirming Krolow’s observations. (2) Extract of Lieber-
kiihn’s glands of the small intestine immediately changes starch into
sugar, but is also without effect on the albumen and fat. (3) The
extract of Lieberkiihn’s glands of the great intestine has no diastatic
effect, and is neutral with regard to albumen and fat. (4) The ex-
tract of Brunner’s is in the horse, and especially in the dog, very
thick and stringy, and on the addition of acetic acid deposits flakes
of mucus. (5) The extract of Lieberkiihn’s glands is less thick and
more fluid, but besides its diastatic function it also serves to keep the
contents of the small intestine fluid. (6) The intestinal juice, the
mixture of the separately observed secretions, has no other effect
than its constituents.
THE NUTRITIVE VALUE OF PEAS AND FLESH, AND THE QUANTITA-
TIVE RELATION BETWEEN THE NITROGEN INGESTED AND THAT EX-
CRETED IN THE Urine.—Woroschiloff in Berlin. Klin. Wochensch.
1873, No.8. Abstract in Lond. Med. Rec. No. 18. The experiments
were made upon himself. Both peas and flesh yielded complete
nourishment when the requisite quantity of carbo-hydrates in the
form of bread and sugar and a small quantity of NaCl were added.
The power of assimilation for the albuminous constituents of flesh is
higher than for those of peas, for the amount of nitrogen in the feces
in the first case was only 6—10 per cent. of the ingested nitrogen,
in the latter 10—17 per cent, During violent muscular exercise (lifting
206 DR STIRLING.
a heavy weight) the quantity of flesh and peas must be increased to
keep the body-weight unchanged. The quantity of urea excreted
under these circumstances was not increased. A quantity of the assimi-
lated nitrogen was retained in the body to be employed for repairing
the muscle substance. From this and other observations it seems
probable that the source of muscular power is to be sought for in
the non-nitrogenous constituents of muscle.
ON THE FERMENTATIVE ACTION OF PANCREATIC JUICE AND
PaROTIDEAN SALIVA OF NEWLY-BORN CHILDREN AND INFANTS UPON
Srarcu.—Korowin, Centralblatt, No. 17, 1873. The pancreas was
taken from children which had died of various affections, chiefly
intestinal and pulmonary, and at different times after death. In all
cases where possible the glucose was estimated quantitatively. The
results obtained are: Pancreatic juice in the first month showed no
effect in changing starch into sugar. In the second month it begins
to exert the fermentative change, and at the end of the third month
the action was so strong that in several cases the quantity of sugar
could be estimated quantitatively. The complete effect is fully esta-
blished at the end of the first year. The parotid secretion converts
starch into sugar in the first day of life, and the action is so strong
that the quantity can be easily ascertained. In Centralblatt, No. 20,
1873, Korowin details his method of obtaining the saliva. The
4 children are allowed to suck small pieces of meerschaum, from which
the saliva is pressed out. It is possible to collect the saliva in the
first minutes after birth. The results are corroborated by Schiffer
(Arch. f. Anat. u. Phys. 1872, 464. “Effects of Stimuli on the
secretion of the Parotid Gland,” by P. B. Stoney, in Journ. Anat.
and Phys. vu. 161.
PLACE OF DESTRUCTION OF ALBUMEN—THE ANIMAL ORGANISM.—
Hoppe-Seyler, Pfltig. Archiv, vit. 1873, 399. This paper is of a cri-
tical nature, and is partly directed against the idea of an organic and
circulating albumen, introduced into physiology by Voit. Lehmann,
Frerichs, and then Bidder and Schmidt, believed that from the great
rapidity with which increased nitrogenous diet was followed by an
increased excretion of urea, that this urea was produced directly from
the albumen of the food without the food having previously become
a constituent of the tissues of the body. Liebig imagined that occa-
sionally a part of the nitrogenous material employed as nutriment
might form urea without having previously been a constituent of the
body. Then Voit from his observations concluded that there was an
organic and a circulating albumen. According to his view only the
albumen taken in as food, and which is flowing in the organs, is dis-
posed to be broken up, while, on the contrary, the albumen which com-
poses the organs of the body is so only when it is again rendered fluid
and again returned to the general juice-stream. This view, however,
is in opposition to a great number of facts. Shortly, the deductions
of the author are that the blood and the lymph vessels possess neither
a ferment nor the peculiarities for oxydation which would lead one
to believe that the place of breaking-up of the nutriment is to be-
REPORT ON PHYSIOLOGY. 207
sought for either in the blood or lymph; on the contrary, we know
chemical changes in the composition of the glands and muscles, which
show that the albuminous matter of the organ can be decomposed
comparatively quickly by fermentation and oxydation. The idea
that urea is formed in the blood directly from the surplus nutriment
is untenable, but much more is the idea of Voit of an organic and
circulating albumen to be rejected.
STOMACH FERMENT OF CoLD-BLOODED ANIMALS.—A. Fick, Verh.
d. Wiirzb. Phys. 1873, 1v. 120. The lowest temperature at which
the stomach ferment of the mammalia is active was fixed by Schiff at
+ 13°, and by Kiihne at + 5°. If digestion in the cold-blooded animals
does not cease when the temperature sinks to 5°, then in this case
we have to do with another ferment. The watery acid extract (HCl)
of the mucous membrane of the stomach of the dog and pig exhibits
digestive powers even under 10°, but at 0° never, while, on the con-
trary, a similar extract prepared from the frog, trout, and pike digests
fibrin at 0°, and at 40° was quite as active as that of the dog. The
stomach ferment of cold-blooded animals is therefore not quite iden-
tical with that of the warm-blooded.
ForMATION OF PrEpsIN IN THE StromAcu.-—Ebstein and Griitzner,
Pfliig. Archiv, vit. 122. This paper is a reply to that of Von Wittich
(Pjliig. Archiv, vit. 19), in which he states that the pyloric glands
have no pectic action. They find that an extract of the pyloric made
with HCl possesses digestive action, but an extract of the same with
glycerine does not possess this property, as Von Wittich found. They
find that HCl alone does not dissolve so much albuminate as HCl
with only the smallest trace of pepsin added to it. They conclude
that the pyloric glands do possess digestive powers, and that they do
not obtain their pepsin by infiltration from the fundus; and further,
from their experiments, they think it in the highest degree probable
that the chief-cells (“‘ Hauptzellen”) of the fundus and the glandular
cells of the pylorus prepare a secretion, in which pepsin is present
not yet free, or not completely developed. It is first freed or com-
pletely developed by contact with NaCl, or HCl (or probably, in
general, with all combinations of chlorine).
METAMORPHOSIS OF Foop AND TissuE IN FLESH AND Fat Drer.
—Pettenkofer and Voit (Zeitschrift f. Biolog. 1x. Heft 1), experi-
menting on their famous dog, determined what becomes of fat when
taken together with flesh. They give tables which show the effects
of dieting with various proportions of fat and flesh. They find that
fat is taken up in large quantities from the intestine. In one expe-
riment, lasting over fifty days, the animal was fed with 500 grms. of
flesh and 200 of fat; 14°7 grms. of dry feces were evacuated, con-
taining 4°6 of fat, so that in every 24 hours 195-4 grms. of fat out
of the 200 were absorbed. Although when a moderate amount of
fat is given some part of it is discharged by the feces, yet this is not
because it cannot be absorbed, for if larger quantities are given
nearly all is taken up. Fat it seems splits up in the animal economy
208 3 DR STIRLING.
into simpler compounds with greater difficulty than albumen; indeed
fat resulting from the decomposition of albumen is more easily oxy-
dized than that in the food.
PHYSIOLOGY @F THE MOVEMENTS OF THE IntTEstINES.— Horvath,
Centralblatt, Nos. 38, 39, 40, 41, and 42. In 1869 the author showed
that in artificially cooled animals the intestines became completely
motionless at a certain degree of cooling, and did not contract when
electrically stimulated, but that upon warming the same piece of
imtestine resumed its normal movements and its sensibility to elec-
trical irritation. When a piece of excised is changed from cold to
warm water its movements decrease very obviously with the time,
so that at the 3rd or 4th change from the cold to the warm water the
movements are scarcely perceptible. Excised living intestine when
placed in warm water always everts its edges, while the edges of
dead intestine never show such an eversion. The abdomen of an
animal was opened, and through a loop of its intestine well supplied
with blood, and in connection with the general circulation, by means
of the mesentery, water of different degrees of temperature was
passed. For irritating it a Du Bois Reymond’s induction machine
and one or two Daniell’s elements were employed. By the passage
of cold water the loop of intestine remained motionless (10 to 15 mins.
at most), till the cold was compensated by the heat, which produced
the peristaltic movement. At all temperatures between + 19°C and
0° C the intestines remained motionless and insensible, and that for
a long time; but between + 19°C and + 41°C, above which no expe-
riments were made, they exhibited spontaneous movements, and con- .
tracted when stimulated electrically. The peristaltic movements (be-
tween + 19°C and +41°C) rose somewhat proportionately with the
increase of temperature. Similar results were obtained from the great
intestine, coecum, and different parts of the small intestine in cats,
dogs, guinea-pigs, and rabbits, and in the stomach of the frog. A
pale-coloured piece of intestine, i.e. with little blood, in spite of the
warming, contracts less energetically than a piece well supplied with
blood. Heat is the only till now known means of producing peri-
stalsis In a piece of intestine rendered for a long time motionless by
cold. On warming a piece of cooled intestine a second before the
movements begin it suddenly widens. For the movements of the
intestines in addition to heat an ample supply of blood is necessary,
without which the peristalsis is very weak, or ceases altogether. The
author thinks that the diarrhcea of abdominal typhus is perhaps alone
to be ascribed to the elevated temperature and the thereby increased
peristalsis. The diarrhoea and constipation in different affections
may be explained by a combination of the temperature and the
addition of blood to the intestines. The intestines may remain
motionless for weeks or months during hybernation, and they doubt-
less play an important part in hybernating animals. From experi-
ments upon artificially cooled animals the author is of opinion that
there is a connection between the death of such a cooled animal and
the absence of motion in its intestines. “ Peristaltic movements
REPORT ON PHYSIOLOGY. 209
of the Stomach and Intestines.” Van Braam Honckgeest, Pfliig.
Archiv, vi. 266. (Abstract in Cenéralblatt, No. 29, 1873, and
vir. 163.)
Liver.
“Cholic Acid.” Baumstark in Berlin Klin. Wochensch. 1873,
No. 4. “Source of Glycogen in the Liver.” Weiss in Sitz.-ber. d.
Wien. Acad. d. Wiss. LXVIl. “On the Albuminous Substances of
the Liver-cells,” in Pfliig. Archiv, 1873, vu. 371. “ Absorption
Spectrum of Hydrobilirubin.” C. Vierordt in Zeit. f Biol. 1x. 160.
“ Wistinghausen’s Endosmotic Experiment on the Action of the
Bile in the Absorption of Neutral Fat.” Steiner in Reich. and Du Bois
Rey. Arch. 1873, 137. “ Formation of Colouring Matter of Bile.”
Steiner in ibid. -160. “Lectures on Diabetes and the Glycogenic
Function of the Liver” (Translation of, from Revue des Cours Scienti-
Jiques), Lond. Med. Rec. No. 40, 1873.
Secretion or Bite.—Réhrig in Medicin. Jahrbiicher, Heft 11.
1873. A bent glass tube was introduced into the duct. communis
choledochus of a curarised rabbit, in which artificial respiration was
kept up. At the outset of the experiment one drop of bile was
obtained every 8 secs., but towards its close one drop in 17 secs.,
when the bile was more glutinous and deeply coloured. Compression
of either the vena porte or the hepatic artery or both together caused
a diminution of the quantity of bile, though after a time the liver
recovered to some extent. In the dog ligature of the thoracic aorta
above the diaphragm diminished the flow of bile largely from 1 drop
in 7 to 1 in 50 secs. Ligature below the cceliac anis in the rabbit
raised it from 1 in 8 tol in 3. Injection of water into the intestine
and digestion always increased it. Section of the splanchnics in-
creases it. Division of the spinal cord is followed at first by increase
and then by diminution of the flow. Compare also I. Munk, “On
the Excretion of Bile by irritation of Sensory Nerves.” Pjliig.
Archi, vil. 151.
ACTION OF THE BILE IN CONVERTING STARCH INTO SUGAR.—
Pjliig. Archiv, v1. Von Wittich states in opposition to Ranke that abso-
lutely fresh bile has the power of converting starch into sugar. Also
Pjliig. Archiv, vu. Heft 1, 1873. Von Wittich from his experiments
thinks that the liver, even when washed free from blood, contains
a ferment capable of converting starch into sugar, and that there is
good evidence to show that this ferment is generated in the liver-
cells,
Milk.
“ Organic particles in Milk.” Béchamp in Comptes Rendus, 1873,
x. 654. “Cause of Coagulation of Caseine by Rennet,” in Journ.
SJ. Pract. Chem. 1872, vi. 174.
Composition oF Human Mitx.—Th. Brunner, Pjliig. Archiv,
vit. 440, The author undertook this investigation after seeing an
VOL. VIII. 14
210 ---> DR STIRLING.
analysis by Sourdat (Comptes Rendus, Lxx1. 87, 1870), in which
the milk from the right breast difiered in its composition from
that of the left. The author estimated the albuminous substances
together. These substances (casein and albumen) are easily and com-
pletely separated on adding dilute acetic acid till the alkaline reaction
disappears. The precipitate when washed, and so freed from any
sulphate of soda that may be in it, was then dried and weighed, and
represents the casein + albumen + fat. The fat was then estimated in
another portion of milk by Trommer’s method. By subtraction the
weight of the casein and albumen was easily obtained. The sugar
was determined by suturation with Febling’s solution in the usual
way. From a table containing 14 double analyses (milk from right
and left breasts), with remarks upon the age of the woman, and
number of children, the author gives the following as the mean
composition of the collective analyses in 100 pts.: 0°63 albuminous
bodies (casein and albumen), 1-73 fat, 6°23 sugar, 90-00 water, and
1:41 soluble salts and extractives. The quantity of albumen in
these analyses is far under that cited by Vernois and Becquerel, viz.
3°92. The author thinks that this is to be accounted for in that
these authors did not estimate the albumen directly, but all the other
constituents, and regarded the remainder as casein, and that they did
not completely dry the milk. The author further remarks that,
while other experimenters have analysed milk soon after delivery, his
analyses were made on milk obtained several mouths after delivery,
when it seems that the milk becomes with the time poorer in albu-
men and fat, whilst the other constituents remain tolerably un-
changed. A table is then given showing the composition at different
periods after delivery. The milk of the cow is poorer in water,
sugar, soluble salts, and extractives, than human milk, but much
richer in albuminous stuff and fat. From a table, indicating the
composition of the milk taken from the right and left breasts at the
same time, it is shown that the milk in each can have a different
composition.
Muscle.
On THE FaticUE AND RECOVERY OF TRANSVERSELY STRIPED
Muscrtes.—H. Kronecker, Ludwig's Arbeiten, v1. 177. This commu-
nication is specially interesting in that the results from a very large
number of carefully conducted experiments, seem to stand in oppo-
sition to the generally accepted view of the influence of work upon
the consumption of material. According to the laws of the change
of material one would expect that excised muscles, which have only
a distinct quantity of material at their disposal, would consume this
by a corresponding quantity of work; thus, by a single great effort
quickly, by a small one slowly. Heidenhain (Mechanische Leistung,
Warmeentwicklung und Stoffumsatz bei der Muskelthdtigkeit, 1864)
has found that the warming of a muscle is always greater during its
contraction the greater the weight it lifts, and also that with the
principal mechanical the secondary thermal expenditure increases.
REPORT ON PHYSIOLOGY. 211
One has therefore more ground for the belief that a muscle when it
raises a large weight must exhaust its foree much sooner than when
it lifts none. The author has found that the fatigue is quite inde-
pendent of the work, and follows very simple laws, and really depends
upon the frequency with which the single acts follow each other.
To estimate exactly by comparison the dependence of the fatigue
on the period of rest, on the weight, on the strength of the irritation,
and also the influence of substances circulating in the muscles, the
author wished to irritate the muscles (triceps femoris) of a frog
loaded or overloaded (‘‘belastet oder iiberlastet”) with selected weights,
by opening and closing induction-strokes of selected intensity, in
opposite directions and in easily changed intervals of time, and to
have the height of the contractions written on paper in equal small
(1mm.) distances. The following arrangement of apparatus was
employed.
The two corresponding muscles (triceps femoris) of a frog were
placed by means of a string in connection with two levers for writing,
which marked the height of the contractions (enlarged twice) upon
the paper on the cylinder of a large kymographion. For irritating,
opening, or shutting, induction-strokes, which affected directly both
muscles (after the one pole was placed to the lower end of one muscle
and the other pole to the lower end of the other), were employed.
The irritation was increased till it produced maximum contractions
before the proper experiment commenced. By means of a metro-
nome, which closed the primary circuit of a du Bois Reymond’s
induction-apparatus, induction-strokes were discharged in equal in-
tervals of time (whose length could be easily changed within wide
limits), one sort of the induction-strokes (opening or shutting stroke)
could be rendered ineffective by a Pfliiger’s apparatus. Generally
after every contraction the direction of the current was inverted.
After every contraction the electro-magnet, indirectly dependent
on the metronome, allowed the regulator of the clock-work of the
myograph to make a haif-revolution, and therewith the cylinder
rotated a short distance. The heights of the contractions were drawn
about 1m.m. from each other. The course of the work of the
muscles, which often wrote many hundred contractions before com-
plete exhaustion, could thus be easily seen.
Having arranged this apparatus, the muscle, in unchanged in-
tervals of time (2—12 secs.) lifts a weight (which must not exceed
that of an entire frog, 50 grms.) from a support (“ Ueberlastung”),
then the difference between the heights of two contractions fol-
lowing each other—the difference of fatigue (‘“‘Ermiidungsdifferenz”)
—is constant, the line connecting ull the highest points of the equi-
distant series of contractions is straight ( Ermiidungscurve”). The
line of fatigue falls more rapidly towards the abscissa the smaller the
interval between the irritations ( Ruhezeiten”), i.e. the difference
of fatigue diminishes when the intervals between the irritations in-
creases, A striking peculiarity of muscle is this, that the height
of the contraction, observed at a given moment during the experi-
ment, is quite independent of the quantity of work which it has
14—2
22 DR STIRLING.
previously accomplished, only the nwmber of irritations (maximal)
which the muscle has previously received regulates the height of
its present work. When the interval between the irritations is
changed, the fatigue progresses during this tempo of the contrac-
tions, in the same manner as if all the contractions from the beginning
onwards had been made with the same interval.
Let the interval be constant, but change the weight (‘‘ Ueber-
lastung”), the smaller weight is of course lifted higher than the larger
one, but the terminal points of all the heights to which the muscle
has raised the equal weights lie in a straight line, or, in other words,
the difference of fatigue during unchanged intervals of irritation
remains constant even when the overweightings (‘‘ Ueberlastungen”)
of the working muscle are changed. 'he lines of fatigue of different
overweights run parallel.
If the weight is attached to the muscle so that it can stretch the
muscle, even when the latter is at rest (“ Belastung”), the line of
fatigue remains, with equal intervals and weight, a straight one only
to the point where it cuts the abscissa drawn by the inactive non-
weighted muscle. From this point onwards—where the shortening of
the active muscle is equal to the elongation of the passive one carrying
the same weight (“Dehnungslinge”)—the line of fatigue is a hyperbole
in an algebraical formula: thus from where the shortening is equal to
the above elongation (6) the difference of fatigue (D) becomes always
smaller with the number (7) of contractions, and is represented thus:
Dp’? in other words, the line of fatigue straight to this point
becomes thence nearly a hyperbola whose asymptote is the abscissa of
the passive weighted muscle. This relation is easily explained when
one assumes that the elasticity of an active muscle is quite as great as
that of a muscle at rest. For the confirmation of this we must refer
to p. 239—245. It is obvious that the line of fatigue of a stretched
muscle (“belastet”) cannot remain straight to the end of the work,
because of the weight which is held in equilibrium by the elastic
force of the passive stretched muscle, a smaller fraction requires to be
lifted by the contractions, the smaller the shortenings are, so that the
muscle in a manner contracts always with a smaller weight. There
is thus explained the at first sight somewhat paradoxical phenomenon
that a fatigued muscle (which can contract very little) can raise great
weights the same relative height as light ones (the latter of course
from an absolute higher position), the muscle being less stretched by a
light weight. .
We have not space to mention many other remarkable facts that
are contained in the original, to which we must refer for further infor-
mation. We can only remark that the author has found both im the
muscles of dogs, and in those of living intact frogs, that the funda-
mental law of the constant difference of fatigue is valid, and by means
of transfusion of different liquids through the vessels of the muscles has
shown, that not only arterial blood of the same or different classes
(rabbits’ blood in frogs) can partly restore the fatigue of muscles
without their requiring to rest, but that also a 0:5 per cent. solution
REPORT ON PHYSIOLOGY. 273
of NaCl, which contains a very small quantity (0-01 per cent.) of
permanganate of potash can produce the same effect. The author
cites an experiment in which he weighted a very exhausted gastrocne-
mius of a frog with 40 grms., and found after injection of this solution
that 190 contractions were produced, which at the beginning were
four times as high as those made by the muscle before the injection.
There was at most in this case 0:018 milligr. of ozonized oxygen
which the muscle could have extracted from the reduced outflowing
fluid, an increase of work of 0°01176 kilogrammeter.
Pure solution of common salt (0-5 per cent.) had little or no effect.
Muscles.
Post-Mortem Ricipiry or Muscies.—Michelsohn (Jnaug. Dissert.,
Dorpat, 1872) has arrived at the conclusion from experiments con-
ducted under Schmidt’s direction, that the cougulation of myosin post-
mortem is brought about by a ferment.
CuemicaL Reaction of Active AND Inactive Muscie.—Griitz-
ner (Pjliig. Archiv, 1873, vu. 254) tetanized the gastrocnemius
muscle of a frog till it was exhausted, while the opposite muscle
remained at rest under similar conditions with the active one. Hach
of the muscles was now triturated with 5 cm. of a half per cent
solution of pyrogallic acid, and the product filtered. The active
muscle yielded a clear or yellowish filtrate, the inactive one a brown
one. In the one purpurogallin was formed, in the other none, or
only a trace. The latter had been able to oxydise the pyrogallic acid,
the former not. The browning in the second case is not to be
ascribed to the strong alkaline reaction of the inactive muscle. When
a mixture of pyrogallic acid, and chloride of iron is used, which is a
reducing as well as an oxydising mixture, itself of a brownish red
colour—with the inactive muscle a red-brown colour is obtained, and
with the active a bright violet. This change the author ascribes to
the large quantity of lactate of an alkali present in the active muscle,
which the author found yielded a violet colour when treated with
the above mixture.
“ Electrotonus,” Hermann, in Pfliig. Arch. vu. 301, 323 and
497,——“The Conditions of Equilibrium for Stimulated and Non-
stimulated Muscles,” Fuchs in ibid. 421. “Contraction of Muscu-
lar Fibre,” Krause in Pfliig. Arch. vir. 508. “ Action of Veratria on
Muscular Fibre,” Fick and Bohm in Arb. aus d. Phys. Lab. d. Wirz.
Hochschule, u1. 142. ‘¢ Rlectrotonus,” Bernstein in Pyliig. Arch,
vur. 40,——“ Action of Electricity on Muscle and Nerve,” Bernheim,
in ibid. 60.
Muscutar IRRITABILITY AFTER SysTEMIC DeEatTH.—Croonian
Lecture. B. W. Richardson. (Proc. Roy. Soc. Lond. 339.) The
principal results obtained by the author are: 1, There are three
degrees of muscular irritability—the active efficient, passive efficient,
negative inefficient. The muscle after death may be suspended in
214 DR STIRLING.
any of these conditions for action ; but, as a rule, it is the last con-
dition only that is maintained long after death. 2. Muscular irrita-
bility may be suspended or stopped altogether under three different
conditions, having reference to its connection with nervous activity :—
(a) the nervous muscular activities may be suspended equally, on
which there may follow spontaneous return of motion. (b) The
muscular irritability may outlive the nervous function, on which
the phenomenon of irritability may be induced by the application
of the motor forces, but there is no return of spontaneous irritability,
i.e. of irritability belonging to the animal as a distinct agent.
(c) The nervous function may outlive the muscular irritability, under
which circumstances irritability is invariably stopped by the pro-
duction of persistent contractility of the muscular fibres. 3. Nervous
activity exciting muscular action is identical with all the motor forces,
and particular to none. It is equivalent to mechanical, calorific or
electrical force, and equally susceptible of manifestation through
either. 4. Muscular irritability is possible after death. Cold, in
certain defined degrees, suspends without destroying it. The motor
forces strike it into rest. Blood sustains it or stops it according to
the balance of powers existing between the muscular and nervous
systems. Some chemical agents suspend it independently, others
suspend it together and equally, with suspension of the nervous
function. When suspension is equal there may be spontaneous re-
turn ; when it is unequal there is no return.
ON THE SO-CALLED SHORTENING OF TenpDoNSs.—Hermann (Pjiiig.
Arch, vu. 417) cites an experiment to show the connection between
this condition and the coagulation of the albuminous substances
contained in the tendon when a piece of tendon of a known length
is placed in water; when the water is at 65° the length of the tendon
is completely unchanged, but just at this point there begins a marked
shortening, which is complete at 75°. This condition of the tendon
‘takes place at exactly the same temperature at which ordinary al-
bumen coagulates. (See reply to the above, Englemann in Pflig,
Arch, vit. 95).
Miscellanzous.
RerLex Movements oF THE Urerus.—Schlesinger (Wien Med.
Jahrb., 1873, 1. Hft. 4). Electrical irritation of the central end of
a spinal nerve produces after 5—15 seconds powerful general move
ments of the uterus. In a curarised rabbit, upon which tracheotomy
has been performed and artificial respiration kept up, if the artificial
respiration is suspended, the uterus exhibits energetic contractions.
The same results when the central end of the median or crural
nerve is electrically stimulated. The path of the reflex action
is not through the spinal cord, for after section of the medulla,
between the occipital bone and atlas, a continued irritation of the
stump of a nerve for 40 seconds was without effect. So that the
author believes that the path by which irritations going from the
=
REPORT ON PHYSIOLOGY. 215
brain reach the uterus, is at least partly through the nervous
plexus of the aorta, but still this is not the only route by which the
motor influence travels.
Influence of Changes in the Barometric Pressure.
On THE PuHeNomEeNA oF Lire.—Bert. (Comptes Rendus, No. 8,
Aug. 25, 1873, p. 531.) Twelfth note on this subject :—Ist. When
oxygen, reaches the proportion of 28 to 30 vols. for 100 vols. of
arterial blood in a dog, the animal is seized with convulsions, while a
proportion of about 35 vols. proves fatal. 2nd. All these convulsions,
however varied their type, are due to direct irritation of the spinal
cord. From the outset of the convulsive attack, the temperature falls
several degrees. There is therefore a profound alteration in nutri-
tion, which does not take place in poisoning by convulsive substances,
e.g. strychnine. These convulsions are to be regarded as epipheno-
mena manifestations of the spinal cord of general disturbance of the
organism, such as happens in asphyxia or rapidly fatal hemorrhages.
To what alterations in the blood are we to attribute these strange
results ? Is it to be supposed that under a little more pressure oxygen
forms with the blood-corpuscles a more stable combination than with
ordinary hemoglobin? a combination from which the tissues are
not able to remove the oxygen of which they have need? This is
not so, for scarcely had the animal been placed under normal pressure,
when the excess of oxygen disappeared from the blood, whilst the
convulsions often continued many hours and the temperature con-
tinued to fall. Nor is it likely that a substance so formed by the
super-oxydation of the blood would persist after the return of air,
the blood having thus become a toxic substance ; for the author has
injected with impunity considerable quantities of blood, charged with
oxygen to the fatal extent, into dogs rendered nearly bloodless. All
tends to show that the blood is an intermediary, carrying the blood |
to the tissues. The author believes that it is the excess of oxygen
in the tissues themselves which alters the chemical phenomena of
nutrition ; the central nervous system is the first part affected by the
sudden change in nutrition, hence the epiphenomenon of convulsions,
Fishes die with convulsions when the water contains more than
10 vols. of oxygen. The toxic action appears also in the invertebrata.
Insects die more rapidly in compressed oxygen than Arachnida and
Myriapoda, and the latter more rapidly than Mollusca and Earth-
worms.
Regarding the general nature of the alteration of the nutritive
phenomena, the most evident is a diminution in the intensity of the
phenomena of oxydation, 1. If an animal is made to respire in a
certain volume of air, in the normal state, and afterwards in air
poisoned with oxygen, it is found to absorb much less oxygen in a
given time during the second period than during the first. 2. On
analysis of the arterial blood of a dog which had convulsions due
to oxygen, and which had respired some time in open air, the
216 DR STIRLING. REPORT ON PHYSIOLOGY.
quantity of CO, is very small (25, 20, 15 vols. for 100 vols. blood.).
3. The proportion of urea falls considerably under the influence of
compressed air. In one case it fell 12 grms. to 4 after seven
hours, at 8 atmospheres. Similar results were obtained in Vitro.
A fragment of muscle or other tissue separated from the body,
absorbs in a given time less oxygen and forms less CO, in compressed
air (the same is true of seeds), This diminution in oxydation is the
cause and consequence of a retardation, and even remarkable arrest of
the numerous chemical actions which are closely allied to those which
take place in living beings. Fragments of muscle had not com-
menced to putrefy after eight days in oxygen, compressed to twenty-
four atmospheres, while putrefaction was complete in four days under
identical conditions. Glucose added to blood is destroyed much more
slowly in compressed oxygen than at normal pressure. The same,
though less marked, is the case in the transformation of starch into
glucose by the saliva. Milk undergoes the lactic fermentation more
slowly. It is therefore not astonishing that the nutritive actions
in animals and vegetables are similarly arrested and death ensues.
But diminution in the intensity of the nutritive acts does not
explain all. Slow asphyxia as well as low barometic pressure also
diminish it, and yet do not give convulsions lasting for several hours,
or disturbances that persist even when the quantity of oxygen
absorbed in a given time is reduced to the normal. Seeds of barley
arrested in their evolutions by a vacuum do not die, but they die in
compressed air. There is then, in the physico-chemical acts of nu-
trition, not only a diminution in quantity, but also a modification in
. quality.
REPORT ON PHARMACOLOGY. By Dr Tuomas R. Fraser.
Puospuorus.—In a research on the action of phosphorus
(Virchow’s Archiv, tv., June, 1872; and abstract, Glasgow Medical
Journal, November, 1872), Wegner examined its effects on the
stomach and liver, and on the osseous system. Moderate doses
were found to produce in rabbits a catarrhal condition of the
stomach, followed by great thickening and pigmentation of the mucous
coat, and the formation of characteristic flat ulcers. By similar
doses, the connective tissue of the liver was increased, and by its
subsequent contraction the organ became cirrhotic. The observations
on the effects of this substance on the osseous system are of great
interest. It does not appear to have any selective influence on the
bones of the jaws, for when the tibia was exposed to the action of
phosphorus vapour, changes soon occurred in it. These changes
appear to be the result of irritation of the periosteum, leading to
increased formation of bone and to suppuration; and they are induced
by administering phosphorus by the stomach as well as by bringing
its vapour into direct contact with the periosteum. In growing
animals to whom small doses of phosphorus were given by the
stomach, its peculiar action on the osseous system was manifested
by an unusual density and compactness of the bones. The cartilage
of the young bones was quickly changed into true osseous tissue,
and the periosteum formed new dense bone with unwonted rapidity.
This action on the osseous system is not produced by phosphoric acid
nor by the phosphates.
SILVER-SALTS.—From several experiments on animals of many
different classes, Rouget (Archives de Physiologie, No. 4, 1873,
p- 333) finds that, without exception, the first symptoms produced
by the absorption of silver are due to disturbances in the functions
of the nervous and muscular systems, ranging from feebleness of the
extremities, torpor, and somnolence, to complete loss of voluntary
movement, convulsions, persistent spasm, and paralysis. The respi-
ration is afterwards affected, its rate being gradually diminished
until, in fatal cases, it ceases; and, at the same time, there occurs
in many animals hyper-secretion of bronchial mucus, with an
appearance after death of retraction, or diminution in the volume,
of the lungs. The voluntary muscles are also directly affected,
and in frogs and other animals their contractility may be destroyed
before complete death. The circulatory system resists the action for
the longest time, as the heart continues to contract after the function
of the cerebro-spinal nervous system has been abolished, and even
after the voluntary muscles are no longer contractile. In opposition
to the opinion of Krahmer, Rabuteau and Mourier (and also to
that of Bogolowsky, with whose elaborate research the author seems
218 DR FRASER.
unacquainted), Rouget maintains that silver does not alter either the
composition or the properties of the blood. The greater number
of the experiments were made with nitrate or chloride of silver
dissolved in hyposulphite of soda.
ARSENIC.—F. Schifer and Rudolf Boehm (Verhandlungen der
Physikal.-Med. Gesellschaft in Wiirzburg, 11. 3) believe that previous
experiments have established (1) that pure metallic arsenic is not
poisonous; (2) that no compound is formed between arsenious acid
and albumen; (3) that arsenious acid exists as such in the urine -
and coagulum, not serum, of the blood of poisoned animals, in the
latter case perhaps in combination with potash; and (4) that
arsenic is able to prevent putrefaction. In order to determine if
arsenic interferes with digestion, they compared the action upon
white of egg of natural and artificial gastric juice to which no
arsenious acid had been added, with portions of the same specimens
of gastric juice to which small quantities of this poison had been
added. They arrive at the important conclusion that arsenic is not
able to prevent the digestion of albumen. From similar experi-
ments, they arrive at the conclusion that the pancreatic digestion of
albumen is also unchecked by this substance.—From an experiment
on a dog to which arsenic was administered for several days, and
chemical analysis made of the urime and feces previously to, coinci-
dently with, and subsequently to the administration, H. Von Boeck
(see Vew York Medical Journal, xvi. 1872, p. 301) concludes that
the administration of arsenic exerts no essential influence upon the
amount of nitrogen excreted, nor upon the decomposition of albu-
minoid substances.
Bismuta.—On undertaking a series of researches that have yielded
many important results, on the presence of various metals in the
tissues, Mayencon and Bergeret have found it necessary to plan a
new method of procedure, those already existing being too tedious
and difficult of application (Robin’s Journal del Anatomie et de la
Physiologie, No. 3, 1873). They simply treat the substance or
tissue with a strong acid, and insert into the product a voltaic pair,
with platinum as one of its constituents. In this way, the metal
is deposited on the platinum, from which it may be removed and
tested by converting it into a chloride. Applying this electrolytic
method to determine if bismuth is absorbed into the tissues of man
and the lower animals to whom subnitrate was administered by the
stomach, Mayengon and Bergeret conclude (1) that subnitrate of
bismuth is promptly absorbed, and deeply impregnates the tissues ;
(2) that it is slowly eliminated; and also (3) that it seems to increase
the plasticity of the blood.
CatciuM-Satts.— On numerous occasions, Rabutreau has enun-
ciated the law, based on his own experiments and those of others,
that the activity of metallic compounds increases according as the
atomic weight of the metal is more elevated. He has recently made
some experiments to determine if the calcium-salts adhere to this
REPORT ON PHARMACOLOGY. 219
law (Comptes Rendus, 10 Février, 1873). When the chloride in
solution was injected into the circulation of a dog, it caused death
if the dose was such that the calcium present in the salt was
about the same in weight as the potassium in a dose of chloride of
potassium previously found sufficient to cause the same effect.
Calcium, therefore, forms no exception to the above law, its atomic
weight being 40, while that of potassium is 39. When administered
in this way, calcium causes death by arresting the heart’s contrac-
tions in diastole, just as potassium does. Further, it resembles all
other metallic substances, excepting sodium and lithium, in acting
as a muscular poison.
Mercury.—Adopting with mercury the method of research
described in the abstract of their paper on bismuth (see ante),
Mayengon and Bergeret (Robin’s Journal de l Anatomie et de la
Physiologie, No. 1, 1873, p. 81, and No. 3, p. 233) have determined:
—(1) That mercury and its saits are absorbed by the skin as well
as by the stomach. (2) That of the absorbed mercury, the greater
part is immediately eliminated, while only a small portion is deposited
in the tissues, from which it is very slowly eliminated. (3) That
elimination seems to occur through all the excretory fluids (urine,
feeces, saliva, milk, spermatic fluid), but chiefly through the urine
and intestinal juices. (4) That iodide of potassium has a marked
action in freeing the system of deposited mercury. In the course
of their research, they found that perchloride of mercury excites an
abundant secretion of bile. In a case of acute poisoning by per-
chloride of mercury, A. Olivier (Archives de Physiologie, No. 5,
1873, p. 547) observed a remarkable lowering of temperature at the
commencement of the poisoning, and the occurrence of albuminous
nephritis indicated by the presence in the urine of albumen, tumefied
renal cells, and granular, fatty, and hyaline cylinders. Contrary to
the reported observations of Salkowsky and Bouchard, the urine
was altogether free from sugar.
Leap, Gotp.—Mayengon and Bergeret have applied with some
modifications their method of analysis (see Bismuth) to a research
on the absorption and elimination of lead, with the following results:
—(1) Salts of lead are not absorbed by the skin. (2) They are
absorbed slowly and in small quantity by the intestine. (3) Ab-
sorbed lead seems to be deposited principally in the liver and
spleen. (4) Sometimes, after long-continued administration, the
kidneys and urine contain traces of this metal. (5) The elimination
is prompt and complete, and is chiefly effected through the liver.
A similar research on gold has yielded the same observers only
doubtful results (Op. cit., p. 226).
CuLoriDE oF Soprum.—Falck (Virchow’s Archiv, Vol. tvt. p. 315)
finds that chloride of sodium is a more active poison for mammals than
is generally supposed, from three to five parts to every thousand of
the animal’s weight producing death in dogs, when injected into the
veins. It is more rapidly eliminated when injected into a vein than
220 DR FRASER.
when introduced into the stomach, and during its elimination it
greatly increases the quantity of urine and so probably causes thirst.
In fatal doses, frothy liquid escapes from the mouth and nose, the
heart’s contractions are weakened, and tremors and convulsions
precede death. The most conspicuous post-mortem appearances are
pallor of the muscles, dilatation of the heart, dark loosely coagulated
blood, frothy liquid in the bronchi, edema of the lungs, congestion
of the liver and spleen, and mucous fluid in the stomach. Sedium-
phosphate was found to be less poisonous than chloride; it acts less
powerfully on the heart, and does not produce exudation into the
bronchi, nor edema of the lungs.
ALCOHOL AND ABsINTH.—The comparative effects of alcohol and
absinth have for some time been carefully studied in France. In a
recent contribution, Magnan (Archives de Physiologie, No. 2, 1873,
p. 145, and No. 3, p. 281) shows that the prolonged administration
to dogs of somewhat large doses of alcohol produces symptoms closely
resembling those of chronic alcoholism in man. On about the fifteenth
day of intoxication, irritability and excitability occur; two days
afterwards, there are illusions and hallucinations at night; and at
the end of the month, delirium both day and night. Should the
administration be prolonged into the second month, tremors make
their appearance in the posterior extremities, and extend from them
to the anterior extremities and then to all parts of the body. The
animal finally succumbs to disorders of digestion and other complica-
tions, which recall the accidents that terminate alcoholic poisoning
in man. After death, dogs present, Im various degrees, steatosis of
the liver, kidneys, and heart, and results of chronic irritation of the
meninges, cord, and pericardium. Magnan finds that the symptoms
of poisoning by absinth are quite distinguishable from those of
poisoning by alcohol. Even in small single doses, the former sub-
stance causes, in dogs, vertigo and muscular twitchings in the ante-
rior parts of the body; while in large doses, it causes epilepsy and
delirium. By anumber of careful experiments, he has determined
that during the stage of tonic spasm in absinth-epilepsy, the pupils
dilate, the retina becomes injected, and congestion occurs in the
brain—phenomena that do not accord with the generally received
theories of the mechanism of epilepsy.—Accepting the opinion of
Beale, Binz, and others that alcohol checks the movements of white
blood-corpuscles and other masses of protoplasm, Ross endeavours
to show how on this opinion a theory of the action of alcohol may
be founded, sufficient to explain its effects in health and disease
(British Medical Journal, October 4 and 25, 1873).
CuioraL.—Arndt (Archiv fiir Psych. und Nervenkrank, Bd. 111.,
Hft. 3, 1872) and Gubler (Journal de Pharmacie et de Chimie, July
and August, 1873) refer the poisonous symptoms that have been
observed in many cases of chloral-administration, to an injurious
action on the vaso-motor nerves and the heart. Gubler also advances
many arguments against Liebreich’s theory of the action of chloral
being due to the production from it of chloroform in the blood.
REPORT ON PHARMACOLOGY. 221
Nitrous Oxipe.—The effects of nitrous oxide on plants and
animals have been carefully studied by Jolyet and Blanche (Archives
de Physiologie, No. 4, 1873, p. 364). They find that when chemically
pure it prevents the germination of seeds, as those of barley and
cress; and acts as an asphyxiating gas on animals, death being
produced with all’the signs of asphyxia by strangulation or by the
respiration of inert gases such as hydrogen or nitrogen, and in
about the same time. If it produce anesthesia when respired pure,
it is by bringing about an insufficiency of oxygen in the blood; and
insensibility commences to show itself only when the arterial blood
contains no more than 2 or 3 per cent. of oxygen. When this
occurs, the arterial blood is very dark in colour, and it contains from
30 to 40 per cent. of nitrous oxide. Animals may, however, live
if they are made to respire a mixture of nitrous oxide and oxygen in
the proportion of atmospheric air, the nitrogen being replaced by
nitrous oxide: but they do not then exhibit any symptoms of dis-
turbed sensibility, although the arterial blood may contain from 30
to 35 per cent. of nitrous oxide. The absence of oxygen seems
to be a necessary condition before anesthesia results, nitrous oxide
being itself quite irrespirable.
Nitrite or Amyu.—In a long paper on the action of nitrite of
amyl (Archives de Physiologie, No. 5, 1873, p. 467), which contains an
account of numerous experiments in which this substance was ad-
ministered to various animals by inhalation, subcutaneous injection,
and injection into veins, Amez-Droz arrives at conclusions that are
generally in accordance with the opinions usually held. General
uneasiness, great increase in the rate of the heart’s contractions,
dilatation of blood-vessels with reduced blood-tension, irregularity
of respiration, lowering of temperature, and paralysis, when the
doses are comparatively small; with the addition of convulsive move-
ments, passage of feces and urine, reduction following increase in
the rate of the heart’s contractions, and finally, coma and death,
when the doses are large—are the chief symptoms to which he draws
attention. His experiments do not support Richardson’s statement
that the paralysis results from an action on motor-nerves, but they
lead him to refer this effect to a stupefying action on the nerve-centres.
Amez-Droz also differs from Brunton in his explanation of the
remarkable dilatation of blood-vessels produced by nitrate of amyl.
He argues that it cannot be a result of paralysis of the muscles of
the minute vessels, because after dilatation has been produced he
has frequently observed that the vessels contract momentarily
during movements of the animal, and again dilate when these
movements have ceased—changes of calibre that could not occur
were the muscular sheath of the blood-vessels paralysed. He be-
lieves it to be more in accordance with observation to suppose
that by increasing the quantity of carbonic acid in the blood, nitrite
of amyl renders this fluid sufficiently irritating to enable it to
stimulate the peripheral ramifications of the vaso-motor nerves, and
thus to produce a reflex dilatation of the vessels. (No sufficient
222 DR FRASER.
experimental proof of this theory, however, is advanced, and, be-
sides, it is well known that the changes in the minute blood-
vessels during asphyxia are not similar to those produced by nitrite
of amyl. Rep).——F. A. Hoffmann (Reichert und Du Bois Rey-
mond’s Archiv, 1872, Heft 6, p. 746; and Centralblatt, 1873, No. 36,
p- 374) has made an important addition to our knowledge of the
effects of nitrite of amyl. He injected under the skin of rabbits
doses varying from 0:45 to 0°66 gramme (7 to 10 grains). The
quantity of urine voided was increased, and in 24 hours it was
double the normal amount, and contained from 1:0 to 2°5 per
cent. of sugar, which entirely disappeared in from 12 to 30 hours,
Diabetes could be produced several times in the same animal, but
if the injections followed each other too quickly, the animal died.
Urea.—From experiments made on himself, Rabuteau (L’Union
Médicale, Tom. xtv., 1872, p. 841) has found that when urea is intro-
duced into the organism it is entirely eliminated by the urine,
without undergoing any change, within twenty-four hours. It may
also be detected in the saliva, which, however, normally contains
a small quantity. Contrary to the usually held opinion, Rabuteau
maintains that urea has only a very feeble diuretic action.
BenzinE, Nirro-GLYCERINE, Nitric AND SuLtpHuric Acips.—In
an interesting contribution to the toxicology of these substances,
Starkow (Virchows Archiv, uu. No. 4) remarks that by replacing the
hydrogen in carbo-hydrogens by the radicle NO,, their action is
modified and their toxicity increased. At the same time, the nitrous
products confer upon the blood a new modification which is not
produced by the carbo-hydrogens. Thus, in all animals poisoned by
binitro-benzine, the blood invariably exhibits, besides the two oxyhe-
moglobine lines, an absorption line at the boundary of the orange
and red of the spectrum, and corresponding to Frauenhofer’s line C.
The same line is produced in acid hematine by nitro-benzine, nitro-
aniline, and nitro-naphthaline, in all of which a single H is replaced
by NO,, but not markedly in the blood of animals poisoned by these
substances. Corresponding to these characters, the toxic action of
the latter substances is much less powerful than that of binitro-ben-
zine. Chloro-benzine and benzine are greatly exceeded in the in-
tensity of their action by nitro benzine, and they have not the above
action on the blood. They, however, are able to separate crystals of
hemoglobine from the blood to which they are added, and as this
property is especially possessed by chloro-benzine, it may be employed
in place of ether to separate blood-crystals. Binitro-benzine presents
the peculiarity of having a toxic power which is not related to its
solubility. The action of nitro-glycerine on the blood-pigment is ana-
logous to that of nitro-benzine, and the two substances are nearly
equal in toxicity. Nitric and sulphuric acids have only partly this
action on the blood. Spectrum-analysis of the blood of animals
poisoned by them shows the line of acid hematine, which is not
produced by hydro-chloric, phosphoric, and other acids. Absorbed
by the stomach, they seriously modify the blood, and act as poisons
REPORT ON PHARMACOLOGY. 223
as much by this property as by their local caustic action. Starkow
is inclined to disagree with Letheby’s opinion of the possibility of
nitro-benzine becoming transformed in the organism into aniline,
—The occurrence of this transformation is also denied by Lehmann
(Journal de Pharmacie et de Chemie, Octobre, 1873, p. 335) in some
observations appended to an account of a case of poisoning by nitro-
benzine. The chief symptoms observed were cephalgia, vertigo,
unsteady movements, cyanosis, and a strong odour of bitter almonds.
AconiT1A.— Rudolf Boehm and L. Wartmann have published a
valuable research on the physiological action of Merck’s amorphous
aconitia (Verhandl. der Physikal-Medic. Gesellschaft in Wirzburg,
mi., Hft. 1, 1872). They did not succeed in confirming Achscharu-
mow’s statement, that aconitia paralyses the terminations of the
motor nerves (see Journal of Anatomy and Physiology, Vol. 1., 1867,
p. 154), nor the same statement in reference to crystalline aconitia more
recently made by Gréhaut and Duquesnel (see Jowrnal of Anatomy and
Physiology, Vol. v1., 1872, p. 496), On the contrary, their experiments
have led them to conclude that paralysis is produced by this substance,
by an action on the cord, whereby diminution in the activity of the
sensory ganglia, with resulting impairment of reflex sensibility, is in
the first place induced, and is succeeded by diminished excitability
of the motor ganglia with total paralysis of voluntary movement.
The only effect observed by them on the peripheral terminations
of the motor nerves, was an irritative one, resulting in fibrillary
muscular contractions. The muscles were otherwise unaffected. A
large portion of their research is devoted to the action of aconitia on
the circulation. They found that in large doses it diminished the
frequence of the heart’s contractions, and finally caused complete
stoppage of the heart’s action in diastole, before which, however, a
temporary acceleration occurred in the rate of contraction. The
mean pressure of the blood was a little increased at the commence-
ment of the poisoning in rabbits, but it was always lessened in dogs
and cats; and at a later stage, in all animals, it was progressively
lessened. Their experiments lead them to conclude that the effects
on the heart are due to the excitation of the inhibitory apparatus
situated in the heart, and not of the central origin of the vagi as
Achscharumow holds. The lowered blood-tension is due in the first
instance to paralysis of the afferent vascular nerves, and afterwards
to a further paralysis of the vaso-motor centre.
ApomorPHiA.—Quehl (Ueber die Physioloyischen Wirkungen des
Apomorphins. Inaug. Dissert., Halle, 1872; Abstract in Lond.
Med. Rec., January 22, 1873, p. 44), under the direction of Kohler,
confirms many of Siebert’s previous observations regarding the emetic
action of apomorphia (see Journal of Anatomy and Physiology,
November, 1872, p. 194). He differs from Siebert in stating that divi-
sion of the vagi prevents this action. . During five weeks, he produced
vomiting at least once a day in a dog, by the administration of
apomorphia, and found that no tolerance occurred, and that the health
was not injured, the dog having actually gained 23lbs. at the end
224 DR FRASER.
of that time. It would appear that only small doses produce vomiting
in dogs, when it is administered subcutaneously. Somewhat large
doses (3 grs. or more) do not produce vomiting, but symptoms of
general poisoning such as staggering, weakness of the posterior ex-
tremities, great salivation, and partial paralysis, during which the
animal lies extended and executes natatory movements. After death
from a fatal dose, no special appearances are seen. Quehl further
states that apomorphia has no action on sensory or motor nerves,
voluntary or involuntary muscles, nor on the blood-pressure. Chloro-
form-narcosis prevents its emetic action.
Hyprocorarnin.—Hydrocotarnin is one of the opium alkaloids
discovered by O. Hesse (Ann. der Chem. u. Pharm., Supplbd. v111.),
who assigned to it the formula C,, H,, NO,. A somewhat frag-
mentary investigation on its action has been published by F. A. Falck
(Viertel-Jahrschrift f. Gericht. Med., Januar., 1873, 49, and Jnau-
gural Dissertation, Marburg, 1872). With the chlorhydrate, this
observer has made two series of experiments ; one on frogs, and the
other on rabbits. He defines the fatal dose for the latter as three
grains per kilogramme of animal, and for the former, about ten times
this dose per kilogramme. This alkaloid has therefore a toxic
activity less than that of thebaia and codeia, and greater than that
of morphia. It produces, like codeia and other opium alkaloids, two
classes of symptoms in rabbits; a tetanic and a narcotic. In frogs,
the symptoms belonging to the tetanic variety predominate, but be-
fore death a stage occurs, in which the motor nerves are paralysed,
and the heart alone remains active. Falck has made the interesting
observation, that hydrocotarnin possesses, in common with atropia
and codeia, the power of causing the heart in frogs to resume its
contractions after it has been brought to rest in diastole by muscaria.
Ruvus VENENATA AND R. Toxicopenpron.—J. C. White publishes
a valuable paper on the action of Rhus Venenata and LR. Toxicodendron
upon the human skin (Wew York Medical Journal, March 1873, 225).
He describes minutely a number of cases, including his own, in which
he observed the effects on the skin produced by contact with these
plants. His opinion is, that the eruption is of an eczematous, and not
of an erysipelatous nature, that it is not capable of being transferred
by contact or otherwise from one person to another, and that it is pro-
duced by a volatile poison (probably the toxicodendric acid of Maiseh),
which may be conveyed to the skin by exhalations from the plant as
well as by direct contact with it.
Quin1a.—The numerous and valuable papers of Binz on the
action of quinia have been increased by a recent important addition
(Ueber Chinin und Blut: Archiv fiir eaperimentale Pathologie und
Pharmakologie, Bd. 1., Heft 1., 1873, p. 18). This paper treats
chiefly of the effects of the alkaloid upon the oxidation changes
of the blood. It is shown to arrest the formation of acid in the
blood-clot, to diminish the power of blood to carry ozone, and to
lessen oxidation. The previous experiments of this observer had led
REPORT ON PHARMACOLOGY. IOS
him to conclude that quinia retards, and even prevents, the move-
ments of white blood-corpuscles. In the present paper this is at-
tributed to the diminished power of’ oxidation by the red corpuscles
produced by the action of quinia on hemoglobin, a liberal supply of
oxygen being necessary for the migratory movements of the white
cells. The principal action of quinia would therefore appear to be a
lessening cf oxidation ; in virtue of which the temperature is lowered,
the quantity of nitrogen excreted diminished, and the movements of
the colourless blood-corpuscles retarded or paralysed. Jt has besides
a remarkable power in preventing putrefaction and destroying low
forms of organism, regarding which Binz has made many valuable
observations.—This parasiticide action of quinia is treated of in the
first part of a paper by Rochefontaine (Archives de Physiologie, No. 4.
1873, p. 389); but the report of this paper will be deferred until its
publication is completed.—The absorption of quinia, and the trans-
formation undergone by it in the system, are discussed by Guyochin
(These de Paris, 1872; and Revue de Sciences Médicales, Tome 1.,
No. 1, 1873, p. 283). By means of the double iodide of potassium
and mercury, he has discovered the presence of quinia not only
in the urine, but also in the saliva, blood, and intestinal contents.
He differs from Kerner’s opinion that this alkaloid is transformed in
the blood into a new base, and maintains that it is easy to estab-
lish, by applying suitable reactions, that the supposed new base is
quinidine.
THEIN, CAFFEIN, GUARANIN, Cocaiy, THEoBromin.—The action
of these substances has been carefully investigated by numerous ex-
periments on warm- and cold-blooded animals, by A. Bennett (Hdin-
burgh Medical Journal, October, 1873, p. 323). His investigation
was mainly directed to determine the symptoms produced by their
administration, and the causation of their effects on the muscular and
nervous systems. He finds that these five principles are to all
appearance identical in their physiological action, They cause, in
small doses, cerebral excitement, not succeeded by coma, with partial
loss of sensibility ; and in large doses, cerebral excitement, complete
paralysis of sensibility, and tetanic spasms and convulsions. The
influence on sensibility is explained by an action on the posterior
columns of the cord, and on the entire system of peripheral sensory
nerves, both of which are paralysed by large doses. The anterior
columns of the cord, the peripheral motor nerves, and the striped
muscles do not seem to have their functional activity diminished ;
but an irritant action is produced on the cord, which shows itself by
the occurrence of spontaneous spasms. ‘The vaso-motor system is
decidedly affected, contraction of the vessels being first produced, and
afterwards dilatation with stasis of the blocd. Further, these substances
contract the pupils, lower and then elevate the temperature, increase
salivary secretion, and induce a peculiar form of tenesmus accom-
panied with a copious discharge of clear mucus from the bowels.
AtroriA.—From the results of several experiments in which
atropia caused great increase of the pulse-rate, after the vagi had
VOR. VL 15
226 DR FRASER.
been divided, H. C. Wood (American Journal of the Medical Sciences,
April, 1873, p. 332) is inclined to think that it possesses a stimulant
power upon either the accelerator nerves of the heart or their centres,
and that the increase of the pulse-rate is not therefore entirely due
to paralysis of the peripheral vagi. Ina subsequent part of the same
communication, Wood states his decided conviction of the medical
value of atropia in opium poisoning. Although its action is not
completely antagonistic to that of morphia, still it exerts valuable
counter-effects, and notably upon the respiration, which it directly
and powerfully stimulates.—Schiff’s experiments (Za Nazione, No.
235, Augosto, 1872) have established that the sensibility of the heart
is lessened by doses of atropia, rather greater than are necessary to
dilate the pupils, to such an extent, that the pulse-rate is neither
accelerated nor diminished by increasing the blood-pressure to three
or four times its normal amount, or by lowering it greatly.
Nicotra, Topacco-Smoxe. — Basch and Oser (Wiener Med.
Jahrbuch, 1872, p. 367) have investigated the effects of nicotia upon
the intestinal movements, and the relations of these effects to the
changes produced in the circulation, They find that nicotia produces
movements of three kinds. The first two, already made known by
Nasse, occur soon after the administration, consist at the commence-
ment of only slight movements lasting a very short time, and then
pass into tetanic contractions. Then occurs a period of repose ex-
tending over four or seven minutes, after which the intestinal
peristalsis becomes more and more powerful, but retains its normal
characters. Coincidently with the first and second kinds of move-
ment, the pulse-rate is slowed by excitation of the vagi, and the
blood-pressure is lowered, and the vessels dilated. During the third
kind of movement the vagi continue to be excited, but the vaso-
motor system is stimulated, and the vessels of the intestines contracted
and pale. These intestinal movements are followed by a period of
repose, during which the excitation of the vagi is diminished, the
pulse-rate increased, and the intestinal vessels again dilated. These
effects on the intestinal movements were found to be caused by an
action of nicotia on the walls of the intestines aided by the changes
produced upon the circulation. The latter are the combined result
of its influence on the vaso-motor centre, and on peripheral vaso-moter
nerves. In a preliminary communication (Centralblatt, No. 41,
1872, p. 641), E. Heubel describes the method followed, and some
_ of the results obtained, in an investigation on the compocnran of
tobacco-smoke, which he has carried on under the direction of
Rosenthal. Since the researches of Vohl, Eulenburg and others,
it has been generally supposed that tobacco-smoke is nearly or alto-
gether devoid of nicotia, the temperature of the burning tobacco
being sufficient te dissipate or decompose this alkaloid. Heubel
imitated the process of smoking by drawing air through burning
tobacco by means of an aspirator. Twenty-five cigars were employed
in one experiment, and the smoke was passed through a Liebig’s
condensor, and then through water, and dilute solutions of sulphuric
REPORT ON PHARMACOLOGY. aE
acid and caustic potash. The products were submitted to physio-
logical and chemical examination, with the result that a large quan-
tity of nicotia was proved to be present in them. ‘To explain the
difference thus shown to occur between the results of submitting
nicotia and tobacco to elevated temperatures, the author maintains
that in the tobacco nicotia is present not in the free state but as a
salt. This salt is able to be conveyed in tobacco-smoke without
undergoing much change, although free nicotia is decomposed when
subjected to an equally elevated temperature.
Dieiratin.—The researches of numerous observers seem now to
have established that digitalis and digitalin produce on the heart,
first a slowing of its contractions, then an acceleration, and finall
again a slowing terminating in paralysis of this organ if the dose
be a sufficiently large one. Coincidently with these effects, there
occurs an increase of blood-tension, which gives place to a diminution
when, at the end of the last stage, the heart’s contractions are slow
and irregular. These phenomena have received various interpretations.
Traube refers the primary slowing of the heart to an excitation of
the vagi nerves, and this view is supported in an elaborate paper by
Ackermann (Deutsches Archiv fiir Klinische Medicin, Vol. Xt.
December, 1872, p. 125), who shows that if the vagi be paralysed by
atropia, digitalin no longer slows the pulse. This observer has also
found that during the stage of cardiac acceleration following slowing
under the action of digitalis, the vagi nerves have completely lost
their inhibitory power over the heart, and in this respect also he
supports Traube’s view that the acceleration produced by digitalis
is a result of paralysis of the vagi nerves. He maintains, however,
that it is likewise in all probability a result of a stimulation of the
accelerating cardiac nerves. The slowing and arrest of the heart,
which subsequently occur, are brought about by a totally different
mechanism, namely, a direct action of digitalin on the muscular
substance of the heart. In investigating the cause of the increase
of blood-tension, Ackermann divided the spinal cord in the neck
before administering digitalin, and found that the blood-pressure
nevertheless increased. This substance, therefore, seems to produce
contraction of the arteries independently of any influence on the
vaso-motor centre, and the probabilities are that its action on the
blood-vessels is due to excitation of the peripheral terminations of
the vaso-motor nerves. It cannot be caused by an action on the
heart, because blood-tension remains increased alike during accelera-
tion and slowing of the contractions of that organ. Besides these
effects, the temperature of the interior of the body falls, while
that of the exterior rises, for a short time, coinciding with the
increase of blood-tension. Ackermann suggests that this is due to
the accelerated circulation causing an increase of temperature at the
surface, and a cooling of the interior of the body. Ackermann’s
views are opposed in a very ingenious and valuable paper by Boehm
(Dorpater medicinische Zeitschrift, 1v.1, 1873) who maintains that the
effects of digitalin on the circulation are the result of a direct para-
928 DR FRASER.
lysing action upon the heart, and a stimulating action upon the vaso-
motor centre in the medulla. Mégevand (Action de la Digitale,
Paris, 1872), among many other experiments, administered to a rab-
bit a small dose of Nativelle’s crystalline digitalin, and then divided
the vagi, and afterwards the right cervical sympathetic. Proof
was thus obtained that after the administration of digitalin see-
tion of the vagi greatly increases the cardiac pulsations without
materially affecting the blood-tension, and that division of one cer-
vical sympathetic causes a considerable diminution of blood-pressure,
without influencing the heart’s contractions to any important extent.
He, therefore, concludes that digitalin directly excites the central
origin of the vaso-motor nerves, and affects the vagi nerve-centres.
The acceleration of the heart’s beats and lowering of blood-tension,
which follow large doses, are referred to paralysis of the vagi and
vaso-motor nerves. In a subsequent portion of his paper, Mégevand
advances evidence to show that digitalin diminishes the excretion
of urea by lessening denutrition and organic combustion, and lowers
the respiratory activity; and he believes that the diminution of
temperature results from these two actions.
VerATRIA. — Claus (Haperimentelle Studien ueber die Tempe-
raturverhdlinisse bei einigen Intoxicationen, Marburg, 1872) finds
that a toxic dose of veratria causes, first, a slight fall in the tem-
perature, then a rise to about the normal point, and finally a fall
immediately before death. Sabadillin, on the contrary, produced
effects nearly opposite to these, and even at the moment of death
the temperature was above the normal.——The report of a valuable
paper by Fick and Boehm, on the action of veratria on muscle, must
be postponed.
Ercotin.—The researches of Holmes (see Journal of Anatomy
and Physiology, Vol. v. 1871, p. 206) and others have established that
ergotin contracts the minute vessels by a direct action on their
walls; and it has been supposed by many pharmacologists that uterine
contractions are merely the secondary result of the diminished calibre
of the vessels of the uterus. A recent investigation by A. Wernich
(Virchow’s Archiv, Vol. tv1. 1872, p. 505) throws considerable doubt
on this supposition. Having injected ergotin into the veins of female
cats and dogs, he found that the first effect was contraction of the
arterial capillaries, accompanied with venous congestion of the skin,
muscles, intestines, bladder, and vascular covering of the brain and
cord. As this effect was not prevented by section of the sympa-
thetics, Wernich’s experiments confirm the opinion of Holmes that it
is the result of a direct action of ergotin on the blood-vessels. The
blood-vessels of the uterus, however, were not observed to become
contracted at the time when the other blood-vessels became so.
Indeed, no change occurred in the uterus until the blood-vessels of
other parts of the body were already diminished in calibre; it then
itself began to contract, and coincidently with its contractions,
anemia occurred in its substance, as a result of obliteration of
the lumen of its vessels. Section of the spinal cord between the
REPORT ON PHARMACOLOGY. 229
third and fifth dorsal vertebree completely prevented the action of
ergotin on the uterus. It is, therefore, probably due to stimulation
(anaemic !) of the centre of uterine movement in the brain or upper
part of the cord, and not to a direct action upon the nerves or
vessels in the uterus, nor upon its muscular fibres.——In a later com-
munication (Centralblatt, No. 15, 1873, p. 353), Wernich draws
attention to a matter of considerable practical importance relative
to the administration of ergot in labour. In the autopsies of indi-
viduals who have perished under its influence, the bladder has
habitually been found distended. This has been regarded as the
result of retention caused by spasm of the sphincter, and the sup-
position has supplied an indication for the use of .ergot in inconti-
nence due to paralysis of this muscle. Wernich, however, finds
that in animals poisoned by ergot the bladder quickly becomes re-
filled after being emptied with the catheter. The secretion of urine
is therefore augmented by this substance, doubtless on account
of its action on blood-pressure. The distended bladder which is
thereby caused has frequently been accountable for retarded parturi-
tion. Wernich advises repeated catheterization in all cases where
ergot is used during labour.
INFLUENCE OF CERTAIN SUBSTANCES UPON REFLEX EXCITABILITY.—
Meihwizeen (Pjliiger’s Archiv, Vol. vu. 1873, p. 201) has made some
experiments on the above subject. The spinal cord was divided
behind the ears in frogs, the animals were then suspended by the
lower jaw, the reflex excitability was tested by measuring the time that
elapsed between the dipping of one foot into 0.2 per cent. solution of
sulphuric acid and the subsequent movement that occurred, the sub-
stance to be tested was injected under the skin of the back, and the
reflex excitability again tested at intervals. The results were as
follows :—Bromide of potassium, in doses of from 5 to 10 milligrammes,
lessened the excitability, and doses greater than 20 mgm. killed the
animal. Bromide of sodium seemed to have no effect on spinal reflex
excitability, even when given in doses of 60 mgm. Acetate of zine,
in doses of from 10 to 20 mgm., almost always proved fatal, and its
action was a central one. Chloral, in large doses, reduces reflex ex-
citability, but in small doses, 0.0005 grm., it has no effect, or produced
only a slight diminution, which was not seen to be preceded by any
increase as Radziejewski affirms. Strychnia increases the reflex exci-
tability for mechanical but not for chemical excitants. Caffein, in
doses of from 5 to 10 mgm., reduces the excitability for chemical stimu-
lation, Doses larger than the above produce death in a few hours.
The reflex centres in the spinal cord are paralysed; but the author
has not observed the tetanus of the extremities described by Loven.
Morphia, in doses of from 5 to 8 mgm. at first. diminishes reflex
excitability, which, after about three hours, returns to. its normal
state; and afterwards it increases it so that it becomes markedly
exaggerated during from 12 to 24 hours after the injection. This
increase is rendered apparent by chemical but not by mechanical
stimulation, and when it has become developed, spasms occur. On
230 DR FRASER.
the increase giving place to diminution, however, the spasms still
continue, with unabated frequency and strength. Quinia seemed to
affect reflex excitability through its influence on the circulation.
Doses of from 4 to 6 mgm. slowed and weakened the heart in about
ten minutes, and altogether stopped it a few minutes afterwards; but
the reflex excitability was not modified until from fifteen to thirty
minutes after the heart’s contractions had become feeble. Alcohol,
in a dose of 1 ec. of a 10 per cent. solution, for a long period pro-
duced great diminution of excitability, but afterwards great increase,
by a central action. With digitalin, Meihwizeen observed the same
phenomena as Weil (see Journal of Anatomy and Physiology, Vol. vt.
1872, p. 500). In frogs, which the day previously had had the
cerebral hemispheres removed, 1 mgm. of digitalin produced a marked
decrease of reflex excitability, aud this before any obvious effect upon
the circulation. After section of the cord behind the tympanum
the excitability returns to its former state. In frogs, with both brain
and medulla removed, no direct depression is produced, only an
indirect one after the circulation is weakened. The author does not
adhere to Weil’s explanation of the depression being produced by
excitation of an inhibitory centre for reflex activity. He thinks it
may be satisfactorily explained as a result of the action of digitalin
on the circulation.
AcTIon oF ConvuLsants.—Several substanees appear to combine
a paralysing with a convulsant action. It occurred to H. C. Wood
(An investigation into the Action of Convulsants, pamphlet) that the
latter action may have a cerebral origin. Selecting aconite, prussic
acid, veratrum viride, and impure veratria, as examples of such
substances, he administered them to frogs and rabbits in which the
spinal cord had been divided; and found that, with the exception of
the impure veratria, they all failed to produce convulsions in the
parts connected with the lower portion of the divided cord. Their
convulsant action, therefore, is not of spinal origin. These various
substances further are powerful modifiers of the circulation, and they
probably diminish the quantity of blood in the brain, Wood describes
several ingenious experiments which lead him to believe that the
convulsant action is of brain origin, occurring only after the cireula-
tion is profoundly affected, and probably resulting from deranged
cerebral circulation.
AcTION OF CERTAIN Emetics.—An interesting paper by A. E.
D’Ornellas (Bulletin Général de Thérapeutique, 15 Mars 1873, p. 193)
on the physiology of vomiting includes an examination of the emetic
action of emetia and tartar emetic. The conclusion is arrived at
that when these substances are injected under the skin, the vomiting
they produce is consecutive to their elimination by the stomach.
This is founded on the observations, that they require about three
times longer time to produce vomiting when they are administered by
. subcutaneous injection than by the stomach; that the time of vomiting
coincides with that of elimination by the stomach; that such doses as
cause vomiting when subcutaneously injected also cause inflammatory
REPORT ON PHARMACOLOGY. 231
lesions of the stomach and intestines, while such as are too small to
cause vomiting produce no gastric nor intestinal lesions; and that, as
determined by Kleiman and Simonowitsch, when tartar emetic is
injected into a vein, vomiting is produced in a longer time than when
it is introduced into the stomach, while in the former case antimony
may be detected in the matters: first vomited. D’Ornellas further
believes that, in general, the excitation to vomit caused by emetics is
the result of a direct action upon the peripheral terminations of the
vagi in the stomach. (However the last statement may accord with
observations referring to emetia and tartar emetic, it cannot be ex-
tended to the whole class of emetics, Of the other members of this
class of which we have suthiicient knowledge, there are several, and
notably apomorphia (see ante), regarding which statements cannot be
advanced in harmony with those on which D’Ornellas’ last general
conclusion is founded—Leporter.)
Action oF Pureatives.—Moreau, in oppositien to Thiry and
Radziejewski, was led by experiment to maintain (see Journal of
‘Anatomy and Physiology, Vol. v. 1871, p. 201) that saline purgatives
do not increase intestinal peristalsis, but modify osmosis in such a
manner as to cause a great increase in the fluid contents of the intes-
tines. This opinion is supported by Vulpian, who has experimented
on curarised and morphised dogs, by injecting solutions of several
purgative substances into the small intestine, and noting the changes
during life and after death (Bulletin Général de Thérapeutique, No.
11, Vol. txxxiv. 1873, p. 522). Sulphate of magnesia was found
to rapidly increase the intestinal secretion, producing distension and
redness of the gut, but no increased peristalsis; and after death, the
production of a true intesinal catarrh was substantiated. A small
portion of the injected sulphate was absorbed, and its presence could
be detected in the urine soon after purgation occurred. Jalap pro-
duced a more intense catarrhal condition, chiefly apparent, however,
in the large intestine; but it alse slightly increased peristalsis.
ANTAGONISM.—(Between Saponin and Digitalin.) From a pre-
vious elaborate and valuable research (Die lokale Anaesthesirung
durch Saponin, 1873) H. Koehler had been led to conclude that
saponin, (@) paralyses the respiratory centre in the medulla, (6) pro-
duces temporary stimulation followed by paralysis of the vaso-motor
centre, and (c) paralyses completely the cardiac nerves and muscular
substance. Before its muscular contractility is destroyed, the heart,
under the influence of this substance, acts in the same way as after
section of the cardiac branches of the vagi and the sympathetics, and
cannot be excited except through its intracardiac nerves. In many
respects, therefore, the action of saponin is opposite to that of digita-
lin; and Koehler has been led, on this account, to make a special
investigation on the antagonism between these two substances (Archiv
fiir experimentelle Pathologie und Pharmakologie, BA 1, Hft 2, 1873,
p- 138). His paper contains many facts of the greatest interest and
importance, but we must content ourselves by briefly mentioning a
few of the more prominent results of his experiments. The animals
pAS DR FRASER. REPORT ON PHARMACOLOGY.
employed were frogs, rabbits, and dogs. It was found that digitalin
increases ‘or reproduces the heart’s contractions after they have been
retarded or suspended by saponin, and that, likewise, saponin in-
creases or reproduces the heart’s contractions after they have been
retarded or suspended by digitalin, Digitalin is able to prevent, for
a considerable time, the paralysing action of saponin upon the regu-
lator-nerves of the heart. It also prevents, for a somewhat longer.
time, the action of saponin in reducing blood-pressure and paralysing
the respiratory centre; but it does not seem to antagonise the lower-
ing of temperature. As might have been expected from the identity,
or at least similarity, of some of their important actions, the antago-
nism between them is, however, incomplete. “Thus, although digitalin
can retard for some time the paralysing action of saponin upon the
regulating nerves of the heart, it afterwards aids the latter substance
in paralysing these nerves, and it joims with it in destroying the
contractility of the cardiac muscle. The one substance cannot there-
fore act as an antidote to the other, and prevent death when a lethal
dose of either has been administered.—( Between Atropia and Physo*
stigmia.)) Qsler has made a number ef observations to determine if
any antagonism exists between the actions of atropia and physostigmia
upon the colourless blood-corpuscles of newts, frogs, and human beings
(Lhe Monthly Microscopic Journal, August, 1873, p. 102). He
employed one per cent. solutions ef sulphate of atropia and sulphate of
physostigmia, dissolved im half per cent. solutien of common salt;
- and applied them in the proportion of four times as much reagent as
blood in the case of newt’s and frog’s blood, and in the proportion of
five to one in the case of human blood. The observations were made
on a Stricker’s stage, at a temperature of 39° Cent. Atropia caused
all motion to cease in the corpuscles ; sooner in the blood of the newt
and frog than in that of man, and sooner with strong than with weak
solutions. At the same time, the blood of frogs and newts poisoned
by the internal administration of atropia showed normal ameeboid
movements, without any modification whatever. The action of phy-
sostigmia was found to be somewhat different. A solution of the
strength of 1 to 800 of water did not modify the movements of the
white corpuscles; but a solution of 1 to 300 greatly impeded the
formation of processes, and caused the movements to be of an undu-
lating and heaving character; while a stronger solution produced the
same changes as atropia. One or two per cent. solutions of either
substance acted distinctly on the red corpuscles, producing irregulari-
ties of surface from involutions and cuppings; but scarcely two
corpuscles were affected alike. The result of applications of both
substances was that no antagonism exists between atropia and phy-
sostigmia, so far as their topical action on blood-corpuscles is concerned.
The special changes produced by each substance could be recognised
in blood to which the two had been together added.
Hournal of Anatomp and Phpstology.
ON SOME EFFECTS OF ALCOHOL ON WARM-
BLOODED ANIMALS*. By C. Binz, M.D., Professor
of Pharmacology at the Bonn University.
In the medical world alcohol is considered by some a stimulant,
and by others a narcotic. Both designations are correct under
certain circumstances ; but they do not embrace the whole sub-
ject. A precise experimental analysis is necessary in order to
obtain a definite view.
It seems to me that the question concerning the warmth of
the body under the influence of alcohol is one of the most im-
portant. As faras I know it was first touched upon by Professor
Nasse, in Marburg, 1846’; then again in 1848 by Duméril and
Demarquay in Paris’, and by Lichtenfels and Frohlich, 1852, in
Vienna*. They all had proved, in contradiction to the old and
still much received opinion, that the warmth of the body slightly
decreases after alcohol. These results were forgotten. Ringer
and Rickards published, 1866, similar observations* ; and Parkes
and Wollowicz°® refuted the former prejudice, which, by the
way, Shakspeare represents very merrily®. But still, in the
very last year, the old assertion of alcohol elevating the com-
bustion of our body, with apparently great precision turned up
again in Germany’.
Whilst with us almost every one was convinced of the cor-
rectness of the view that using alcohol in fever. would only feed
the flames, I was reminded of the question by reading the
accounts of Todd and his school. His communications could
* After a paper read in the Section of Biology of the British Association,
Bradford, 1873.
VOL. VIII. 16
23 PROFESSOR BINZ.
not be right, if alcohol heats ; his method must on the whole be
called correct, if alcohol does not heat.
The experiments which one of my scholars, Dr Bouvier,
made turned out as I expected®. They proved on healthy men
and animals what the majority of authors have said since 1846.
I print here a curve belonging to a later series of experiments,
because all precautions in this case were observed with especial
precision®. It is taken from a strong healthy man of twenty
Time P.M.
“30
37,0 -
centigrades
........ Normal curve, average of 7 observations.
—— Curve under the action of alcohol, average of 7 observations
in the case of the same healthy subject.
years of age, who was suffering from a joint disease without in-
flammation, and who had abstained from all use of alcohol for a
long time. Those hours were chosen as the time of measuring,
in which, according to experience, the heat of the body never
decreases by itself, in which, on the contrary, there is a ten-
dency to an increase. The experiment is thus rendered more
difficult, but in case of success it is made more evident. The
person was measured seven days without, and in the meantime
seven days after the administration of alcohol. The quantity of
alcohol changing from thirty to ninety cubic centimeters (pure
\
alcohol mixed only with tepid water and some sugar) was
SS ———
EFFECTS OF ALCOHOL ON WARM-BLOODED ANIMALS. 235
always so regulated, that no trace of intoxication occurred. It
is of course to be understood that all the outward circumstances
during both series of days remained exactly the same. The
comparison of each single curve of alcohol with the normal
curves shows us clearly the difference. In order, however, to
avoid every arbitrary explanation I have designed the average
curve, and have drawn it in the graphic form.
My opinion, based on such experiments, may be summed up
as follows: The pretended heat of the organism does not exist.
The subjective impression is, at least partially, the consequence
of an irritation of the nerves of the stomach and of the en-
largement of the vessels arising in the skin. When given in
small doses the thermometer shows no extraordinary increase
or decrease of the temperature of the blood. Moderate doses,
which lead by no means to drunkenness, show a distinct de-
crease of about half-an-hour’s duration or more; and strong
inebriating quantities evince a still more decided lowering of
3°5 to 5 F., which lasts several hours. The decrease in the
temperature after moderate doses takes place most distinctly in
warm-blooded animals, which have not for some time previously
had alcohol administered. When inured to it, the organism
does not answer to such doses by any measurable cooling or by
the reverse *.
This is the result of more than one hundred measurings,
which lasted generally from three to five hours. Only the
rectum was used on these occasions, as the axilla is perfectly
impracticable for ascertaining small variations. Another of my
scholars has proved, that even the entrance of the thermometer
into the contents of the rectum may entirely disturb the ther-
mometrical result and especially obscure the obtained decrease.
Good results are yielded more easily by a feverish than by
a healthy animal. I generally used for these experiments
strong rabbits or dogs of the same origin and of the same
quality, and injected under their skin some cubic centimeters of
ichor or putrefying blood. As is well known, after thus pro-
* As to the great influence of habit I need only refer to tobacco. People
who have been habitual smokers, and who smoke again occasionally, resist
the toxic effects of the nicotine, etc., perfectly well. I speak here from my
own experience.
16—2
236 PROFESSOR BINZ.
ceeding, the warmth of the animal rises several degrees, and
all the symptoms appear which are to be observed in human
beings suffering from putrid fever. Particularly the severe
symptoms in the intestinal canal remind us distinctly of our
enteric typhoid fever. If the quality of the poisonous sub-
stance is right the animal expires in a few days. Not so,
however, if simultaneously with the commencement of the ex-
periment alcohol diluted with water is administered, either by
means of the stomach or the skin. The temperature then
remains lower from the beginning, the intestinal catarrh is
slighter, the animal is more lively, and the one may be seen
gradually to die, whilst the other takes its food kindly. The
same effect takes place if we allow in both animals the fever
iy,
1 day. 2 day.
post meridiem. : ante meridiem.
aaa =) SS
3 4 5 6 7 8 9 10 9 10
40,
39,
WETANDDOHP NWR OOD
39,
38,
AOnNwooo re bw
A large dog.
At 3.30, Injection of 3 cem. ichor under the skin;
6 o’clock, 5 ecm. alcohol (96 per cent.) with some water;
8.30, again 5 cem, alcohol, and the following day
9 o'clock again 5 cem.—The dog untouched the next day
died.
(The injection of the ichor is marked by +, that of alcohol by *).
EFFECTS OF ALCOHOL ON WARM-BLOODED ANIMALS. 237
to make its way. I have drawn also the curve from such a
case*, The analysis of this experiment shews me a threefold
action produced by alcohol, (1) the diminution of the heat of
the body, (2) reduction of the putrid processes, and (3) rising of
the action of the heart. We here clearly see alcohol emerging
from the small sphere allotted to it by many practitioners. It
is much more than a simple stimulant in these circumstances,
though it is no doubt sometimes a stimulant for the heart and
the nerves.
By poisoning animals with ichor one can see clearly that
even strong doses of alcohol need not be a narcotic. On the
contrary, the ichor-poisoned animal which cowered down, sleepy
and exhausted, becomes, after alcohol, lively, and runs about.
Todd and his followers made similar observations in man,
Such experiments are easily repeated. It is only the ques-
tion of procuring such ichor as will with certainty produce
fever, two animals of the same quality, and then a careful in-
jection of the pyretic matter; for it is clear, that if we go too
far with the latter, the use of alcohol will not be able to save
life. But in such cases it is always perceptible, even if the
animal treated with alcohol dies, that there was some healing
influence. The alcoholized animal either dies later or under
less severe symptoms.
Space does not permit me to enter into any farther details
of these experiments. If we wish to gain the knowledge of
their foundation and their rational use for hfe, it seemed sug-
gestive to me, to search for the causes which lead us to the
above-described results. It thus became a necessity to analyse
the synthetic experiments in their separate parts.
What are the causes of the decrease of the temperature
after the reception of alcohol? How does it work in the ani-
mal’s economy? At first one would think an increase in heat
must take place. Alcohol becomes easily oxydised. In the
tissues of the animal body alcohol burns, if not given in exces-
sive quantities, into carbonic acid and water, and thus warmth
is set free. But the same occurs to a far greater extent if we
consume grease or oil, and yet in this case no rise of the ther-
* For the sake of clearness I give only the curve of one animal, as it implies
the other.
238 PROFESSOR BINZ.
mometer is caused. The regulation of our whole system works
in sufficient perfection to retain the bodily heat on the same
scale. There exist other causes which counterbalance the little
warmth in the circulation produced by the burning alcohol.
First of all one would search for these causes in the nervous
system ; but that which has hitherto been discussed under the
name of thermic nerves, is still up to the present day entirely
without proof, and also the centre for moderating the heat is
doubted by many. If one separates the spinal cord of large
animals at a certain height, one finds under some circumstances
an excessive increase in the heat of the body, as Sir B. Brodie
has first described in a man, who in consequence of an accident
was thus situated. However the case may be, I have found in
a series of experiments, that alcohol has the effect of lessening
the temperature also of such feverish animals. Its influence,
therefore, is independent of all those nerves which, having their
issue in the brain, traverse the spine’®.
It can further be caused by the action on all the striped
muscles of the body. According to Zuntz and Rohrig the mus-
cles are the organs in which, with the assistance of the nerves,
the greatest part of tissue metamorphosis, and especially oxyda-
tion, occurs. As one knows that after the reception of alcohol
a feeling of relaxation in the striped muscles appears, so the
connection between the two factors seems quite probable.
How far it goes, further experiments must decide.
We have a clearer evidence of this from another point of
view. The weaker the action of the heart the less quantity of
blood is it able to throw to the periphery of the body, and
therefore the cooling of the blood is diminished. Now, if we
introduce, where the action of the heart is weak, an agent
which drives more blood to the cool surface, the contrary oe-
curs. The greater quantity of blood in the skin will irradiate
a greater portion of warmth. In the same time the perceptible
as well as the imperceptible perspiration becomes augmented.
It is needless for me to explain, how, according to a simple
physical law, a diminution of warmth must ensue.
Finally, one of the chief causes is the direct impediment of
the activity of our cells. The great many microscopical ele-
ments of which our glands are composed, and through whose
EFFECTS OF ALCOHOL ON WARM-BLOODED ANIMALS. 239
action the albumen of food is decomposed, become slightly pa-
ralysed by alcohol. We have a very clear example of this
fact in the action of alcohol upon yeast. It can be asserted
from all points of view that the cells of the organism answer
on the whole in the same manner to the reagents, as those of
the mykoderma vini. The higher the percentage of alcohol in
a fluid the less able is the protoplasm of the cells to work and
to produce warmth. And not only does common yeast show
us this, but in every other form of fermentation, particularly in
the oxydation, commonly called putrefaction, the impeding in-
fluence on the protoplasm becomes most apparent; and even
the highest and most complicated issue of the protoplasm—
the hemoglabin—is affected by it, when it is about to pass
over its oxygen to other substances. It imparts its oxygen to
combustible substances in a slower manner when but a small
quantity of alcohol is present”.
Of all these described effects which alcohol produces upon
the animal organism each single one may be very slight, but
summed up they mark a decided decrease in the thermometer.
I will here only briefly say, that another series of experiments
has proved to me, that also the post-mortem temperature is
lowered by previous injection of alcohol”. As is known, this
warmth after the death of almost every warm-blooded animal
not only lasts for a shorter or longer time, but even rises by
several degrees Fahrenheit. The fact, that this warmth also is
under the influence of alcohol, proves to us the direct action
on the chemical processes of the animal body more than any-
thing else.
We thus see from all points that the pretended rise of tem-
perature of the blood by means of alcohol exists as little as
does the extraordinary cold in a fever patient whose bed is
shaking under him. In both cases we are deceived by sub-
jective impression, and only the instrumental observation gives
us the truth.
It is @ priori to be expected that alcohol will be not with-
out influence on the metamorphosis of tissues. An agent that,
consumed in somewhat larger doses, so clearly lowers the com-
bustion, must also be supposed to decrease the urea and the
carbonic acid, both the most important excretions of the organ-
240 PROFESSOR BINZ.
ism. And in fact the researches of several authors prove to
us that this is the case”, although none of them has considered
the great amount of CO,, formed by the consumed alcohol
itself. The gradual accumulation of fat in the tissues of
drinkers is thus most probably explained, and also other occur-
rences are illustrated by these facts.
We feel in winter or in damp cold weather the want of
consuming alcohol in small quantities, and we see further, that
the abuse of it, especially in hot climates, leads to serious dis-
turbances of the health.
Both of these occurrences can be explained by physiological
facts. In a cold atmosphere our metamorphosis of tissues is
accelerated, we consume more of them. This is agreed to by
all physiologists, and proved by the experience of common life.
The colder the climate the more substantial our food must be.
Thus, when we imbibe a fluid which, without sensibly cooling
our blood slowly, lowers the consumption of our tissues, we
apply to the flaming furnace a kind of moderator. Exactly the
contrary is to be remarked in India or in the Tropics. Here
our tissue-metamorphosis is of itself sluggish. If we there con-
tinue to consume the same amount of alcohol, as we are in the
habit of taking in any stimulating atmosphere, the consequence
will be that to the already existing inertness of transformation
in tissues and blood an artificial increase of this inertness will
be added, which can only result in an accumulation of delete-
rious dross within our organism. The same, of course, is also
observable when, in our moderate climates, we indulge freely in
the use of alcohol ; though greater quantities are admissible here.
With regard to the application of alcohol to fever patients we
must not overlook, that its effect is not as strong and lasting as
that of other antipyretic methods. Large and repeated doses
are necessary to retain the lowering of the temperature. On
the other hand, there are certain cases, for instance, where the
heart is very weak, in which alcohol acts as an antipyretic,
whilst quinine is powerless. Here alcohol revives the circula-
tion, like subcutaneous injections of camphor, and thence arises
the cooling at the periphery. Also camphor, as by me and
one of my scholars has been proved, acts coolingly in putrid
fevers”.
EFFECTS OF ALCOHOL ON WARM-BLOODED ANIMALS. 241
In the treatment of all feverish illnesses, the first thing
wanted is to suppress high temperatures. We thus remove the
greatest danger, moderate the process of the disease, and give
to the organism the possibility of resisting with success the
internal cause of the illness. Further thermometrical obser-
vation is necessary in order to decide, in which cases alcohol
may help to this effect, and in which other cases its administra-
tion may be useless or even injurious.
The question has been vehemently discussed, whether alco-
hol is a food or not. The answer depends quite upon circum-
stances. It is certain that we do not require alcohol under
regular conditions to sustain life. Should, however, through
any cause, such as cold air or feverish excitement, an increase
of our tissue-metamorphosis arise, then the matter changes.
Alcohol—according to the different basis—becomes a direct
food; for burning without heating it yields to the body warmth
and power of tension. It becomes an indirect food ; for acting
as we saw before, it moderates the wasting of the body.
As to the excretion of alcohol through the kidneys I agree,
after repeated researches with Geisler’s vaporimeter, thoroughly
with those who declare, that under ordinary circumstances
especially in fever, only traces of alcohol appear in the urine. ;
In speaking of alcohol I mean the absolutely pure prepa-
ration of about 98 per cent., which I gave diluted with much
water and a little sugar. All good alcoholic liquids, the per-
centage of which one knows, may serve for common use. But
I protest decidedly against all those bad artificial and unpuri-
fied mixtures which are so generally given. The nauseous
smell from the breath of the consumer indicates their injurious
composition. Pure alcohol gives no taint to the breath, and
good alcoholic liquids only leave that smell which belongs to
their ethers.
Science has to solve the problem as to what alcohol ean ac-
complish on the healthy and on the diseased frame”, and in
which form and way it ought to be administered. Such re-
searches can be as little disturbed by the lamentable abuse of
alcohol as by the somewhat immoderate reaction and agitation
against that abuse.
242 EFFECTS OF ALCOHOL ON WARM-BLOODED ANIMALS.
LITERATURE.
1H. Nasse, in Heller’s Archiv, 1846, p, 178.
2 Duméril and Demarquay, Arch. gén. de médic. 1848, p. 189 and p. 332.
% Lichtenfels and Frohlich, Denkschriften der Wiener Akademie, 1852, p. 131.
Besides these: Jacobi in Germany (Marburg) 1857, Setschenoff, Sulzynski and
Walther 1860—1865, in Russia (Cf. Manassein, p. xurx. and p. 59), Anstie and
others in England.
4 Ringer and Rickards, Lancet, 25 Aug. 1866, p. 208.
5 Parkes and Wollowicz, Proceedings of the Roy. Soc. 1870, No. 120, p. 391.
—No. 123, p. 89.
6 Shakspeare, Henry IV. Part u. Act iv. Se. 3.
7 8. Rabow (Strassburg), rep. in the Centralblatt, Berlin, 1873, No. 21,
refuted by P. Daub (Bonn), in the same periodical 1873, No. 30.
8 C, Bouvier, Pharmakologische Studien tiber den Alkohol. Berlin, A. Hirsch-
wald, 1872,64 pp. (In this book the greater part of the former literature is
indicated. )
9 C. Binz, Ueber die antipyretische Wirkung von Chinin und Alkohol.
Virchow’s Archiv, Bd. 51, p. 6. On the antipyretic action of quinine and
alcohol in paralytic fever. Practitioner, Voi. v. 1870, pp. 1—7.
10 W. Manassein (St Petersburg), Ueber die Dimensionen der rothen Blut-
korperchen unter verschiedenen LEinfliissen (Kélte, Chinin, Alkohol, etc.).
Berlin, A. Hirschwald, 1872. A most valuable experimental and literary con-
tribution to the knowledge of fever and its treatment.
11 In Bouvier’s book, p. 20 to 31.
12 KE. Berg, Ueber Ausscheidung der Kohlensiure durch die Lungen. Archiv
fiir klin. Med. Bd. 6, p. 291.
13 J. Baum, in the Centralblatt, Berlin, 1870, p. 467; and his doctor’s disser-
tation, Bonn, 1872.
14 F, Riegel, Ueber den Hinfiuss des Alkohols auf die Kérperwirme. Archiv
fiir klin. Med. Bd. 12, p. 79 (August, 1873). An experimental paper with the
same results as mine and those of my pupils.
ON THE ACTION OF INORGANIC SUBSTANCES
WHEN INTRODUCED DIRECTLY INTO THE
BLOOD. By James Buaxeg, M.D., F.R.CS. San Fran-
cisco, California.
THE experiments I am about to describe will shew the action
of the salts of Lime, Strontia, Baryta and Lead, when intro-
duced directly into the blood of living animals. Since my last
communication in connection with these researches the subject
has been invested with additional interest from a discovery I
have made of the connection between the atomic weight, and
the physiological action of these salts of metallic bases. I find
that in the same isomorphous group the intensity of physiolo-
gical action increases as the atomic weight of the elements,
shewing a new and important connection between physiological
action and the molecular properties of these inorganic com-
pounds. I have published a paper on this subject in the
American Journal of Science for March, 1874. For the man-
ner in which the following experiments were performed, I
would refer to the May Number of the Journal of Anatomy
and Physiology, for 1870.
Salts of Lime.
The experiment was made on a dog, weight 17lbs. The
manometer was connected with the arteries. A solution con-
taining 4egrs. of chloride of calcium was injected into the
jugular. Action of the heart slightly quickened; expiration
performed by two or three spasmodic efforts; inspiration nor-
mal.—lInject 10grs., no effect on the circulation; expiration
more spasmodic. Inject 30grs. In 12” the action of the
heart was quickened; the pressure in the arteries not
affected except by the struggles of the animal; respiration
spasmodic. Inject 60 grs. In 12” the heart stopped beating :
30”, violent spasmodic respiratory efforts; head drawn down
forcibly towards the chest; rigidity of abdominal muscles: 2’
after the injection, all movements ceased with the exception
244 DR BLAKE.
of a strong quivering of the whole of the voluntary muscles
which lasted 45”. On opening the thorax, auricles contracting,
ventricles still: each cavity contained blood, right dark, left
bright scarlet: blood coagulated firmly. The muscles that were
exposed on opening the thorax contracted some time after death.
Exp. 2. Injection of nitrate of lime into the veins ; animal
not confined. Inject 3grs. into the jugular; no aponreciable
effect. Inject 6 grs.; expression of pain as the injection was
made. Inject 14grs.; same expression of pain; expiration
spasmodic: animal jumped about apparently from uncontrollable
action of the muscles. Inject 14grs.; 12” after injection,
efforts to vomit; animal seems weaker. Inject 14 grs.; animal
jumped about with involuntary action of the muscles: 45”,
lay down apparently from weakness. Inject 20grs.; animal
kicked about with irregular choreic movements; no expression
of pain; after the general movements had stopped, the fore-
legs were in continual motion. Inject 40 grs.; 10”, heart’s
action arrested: 40”, respiration stopped in a general tonic
spasm; this relaxed in 1’ 30”, and three or four respiratory
movements took place. On opening the thorax, the heart was
still; left cavities contained scarlet blood’.
Salts of Strontva.
Exp. 3. A solution containing 4 grs. of chloride of strontium
was injected into the jugular: 10”, the pressure in the arteries
diminished 1}in.; 15”, pressure again at its former height 6 to
1 Tn the last number of this Journal (p. 218) there is a notice of a communica-
tion made by Rabutreau to the Académie des Sciences, in which he states that the
quantities of the salts of lime and of potash which, when introduced directly into
the blood, destroy life, are about equal; and as the atomic weights of these sub-
stances are nearly the same he adduces this fact in support of the statement he
had made as to the connection between the atomic weight of metals and the
intensity of their physiological action. The result of my own experiments with
these two classes of salts is, that they afford the most striking proof against the
existence of such a connection as he has stated. In the many experiments I
have performed with the salts of potash I have found them six or eight times as
poisonous as the salts of lime, ten or twelve grains of nitrate of potash being
sufficient to arrest the action of the heart, while it requires about one hundred
and twenty grains of nitrate of lime to produce the same effect. It is only
between members of the same isomorphous group that the intensity of physio-
logical action increases with atomic weight.
ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 245
8in., oscillation greater, action of the heart slower. Inject
12 ers.: respiration affected in 10”, pressure diminished: 15”,
pressure rising: 45”, 9 in., oscillations 3 to 3Lin., heart’s action
slower. Inject 20grs.: 18”, the action of the heart arrested,
respiration continued uninterruptedly a minute after the heart
had stopped, it then became intermittent but full inspiratory
movements took place three minutes after the arrest of the
circulation. On opening the thorax the ventricles were still,
nor did they contract on irritating them with the scalpel;
auricles contracting rhythmically, and continued contracting for
five minutes. Left cavities contained scarlet blood which
coagulated firmly. Shortly after the thorax had been opened
the whole of the muscles of the trunk began to contract, and
contracted for more than a quarter of an hour. When the
intercostals had been exposed in opening the thorax they con-
tracted so as to move the ribs forty minutes after death,
although the temperature of the room was 42”.
Exp. 4. Injection of chloride of strontium into the arteries.
Pressure in arteries, 4 to 7in. Inject 12 ers. of the salt into the
axillary artery: 5”, action of the heart quickened, oscillation
less: 45”, the action of the heart as before the injection. Inject
24 ers.: the same action on the heart 5” after the injection ; no
other marked symptoms. Inject 36 grs.: immediate expression
of pain. The direct effect on the heart could not be ascertained
on account of the movements of the animal: 20”, it was stopped,
probably earlier: 30”, respiration suspended for 1’ 30"; the
muscles of trachea and larynx then began to contract, and in
a few seconds the movement extended to the whole of the
respiratory muscles ; fourteen respiratory movements were made,
they were short and spasmodic, lasting about 1’ 35’. On open-
ing the thorax, the auricles were found contracting, ventricles
still ; left cavities contained scarlet blood. Four minutes after
respiration had ceased the whole of the voluntary muscles
commenced contracting so as to produce general movement of
the body. The muscles of the leg contracted with sufficient
force to move the body on the table when a point d'appui was
furnished to the feet. These contractions continued more than
a quarter of an hour.
246 DR BLAKE.
Salts of Baryta.
Exp. 5. A solution containing 0°25 grs. of chloride of barium
was injected into the jugular: 12”, the arterial pressure slightly
increased: 1’, action of the heart slower. Inject 0°50 grs.: 10”,
slight diminution of the pressure, heart’s action fluttering: 14”,
pressure increased lin. above the level before the injection.
Inject 1 gr.: 11”, pressure diminishing with fluttering action
of the heart: 14”, pressure again increased, heart’s action slower
and very irregular, two or three quick beats and then a number
of slow ones. Inject 2 grs.: 12”, action of heart arrested ; re-
spiration continued irregular for 1’ 30”, it then became inter-
mittent and ceased 2°45” after the heart had stopped. On
opening the thorax, the auricles were contracting vigorously,
and continued contracting for some minutes, The ventricles
were still, and did not contract whenirritated. The left cavities
contained scarlet blood. Five minutes after the thorax had
been opened the muscles commenced contracting and continued
in motion for fifteen minutes. When injected into the arteries,
the salts of baryta are exactly analogous in their action to the
salts of strontia. The quantity required to arrest the action of
the heart is about 3grs. When the salts of baryta are injected
into the veins and the animal is left at liberty, no marked
effects are produced until a sufficient quantity has been used
to arrest the action of the heart.
Salts of Lead.
Exp. 6. Pressure on arteries 5-5 to 6in. Inject a solution
containing 3 grs. of acetate of lead into the jugular: 7”, pressure
diminished: 11”, pressure increased to 74 74in. Inject 14 grs.:
7”, pressure diminishing: 11”, pressure 5in.: 15”, pressure 34 10in.
Inject 30 grs.: 7”, pressure diminishing; 14”, pressure, 3in.,
heart’s action slow, oscillations 1 to 14in. at each pulsation;
violent struggles 20” after the injection.—Inject 60 grs.: 10”,
the action of the heart apparently arrested, pressure fell to 1in.,
no oscillations: 1’, respiration stopped. The animal lay as if
dead for two minutes, the heart then commenced beating and
respiration was renewed; the pressure in the arteries increased,
so that in two minutes it was up to 8 in.—Inject 60 grs.: 10”,
ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 247
heart stopped: 25”, respiration arrested in a violent spasm ;
when this relaxed the animal lay as if dead for 2’ 30”, when
respiration again commenced and continued about 1’; the heart
could also be felt pulsating, but no effect was produced on the
manometer. On opening the thorax the right auricle only was
contracting. The trachea and bronchial tubes were filled with
frothy fluid tinged with blood; lungs oedematous. The frothy
fluid had escaped from the mouth before death in considerable
quantities. Shortly after the thorax had been opened muscular
movements took place over the whole body, and one of the fore
legs was moved strongly for many minutes.
Injection into Arteries.
Exp. 7. Pressure 5 to Gin. A solution containing 6 grs. of
acetate of lead was injected into the axillary artery: 4”, pressure
rising: 80”, pressure 12 in. 1’: 15”, respiration arrested, pressure
fallmg slowly; two minutes after respiration arrested, pressure
10in.; three minutes pressure 6 in., heart’s action regular, oscil-
lations 0°5in. At seven minutes after respiration had ceased
the pressure had fallen to 2in., heart’s action weak. At this
time one deep respiratory movement was made; its immediate
effect was to increase the pressure to six inches, There was
no other respiratory movement, and the action of the heart
gradually became weaker as in asphyxia, apparently stopping
in about three minutes. On opening the thorax the heart was
found to be still contracting feebly, lungs natural.
The above experiments shew that the more strictly iso-
morphous members of this group Lime, Strontia and Baryta are
essentially heart poisons, that is, they kill by arresting the action
of the heart. Their apparently exclusive action on the heart is
most evident when they are injected into the arteries, as then
no marked effects are produced until they are introduced in
such quantities as to reach the heart in a sufficiently concen-
trated state to destroy its irritability, the quantities required
being much larger than when introduced into the veins. ‘The
action of these substances in paralysing the heart is owing to
their direct contact with its tissues, although some action is
exerted on the organ through the nervous system, as is shewn
248 DR BLAKE.
by the change caused in its movement five or six seconds after
they are injected into the arteries, and before they have had
time to reach the coronary arteries. The action of the salts of
Lead differs in many respects from that of the other members of
the group. In its action on the lungs, and on the muscular
coats of the arteries, it agrees with the salts of silver, with which
substance it has certain isomorphous relations; but in the pe-
culiar action on the voluntary muscles, which. distinguishes the
elements of the Baryta group from all other inorganic com-
pounds with which I have experimented, the salts of lead agree
with the other members of the group. This action on the
muscles is most strongly marked after the injection of the salts
of Strontia and Baryta, and shews itself sometimes in a curious
manner. In one instance in an animal killed by chloride of
strontium, there were no movements until ten minutes after
death. The muscles of the ear then commenced contracting, so
as to cause it to move; and from this point the muscular con-
tractions spread until all the muscles of the trunk and limbs
were in motion, and they continued contracting more than a
quarter of an hour. The longest time after death m which I
have observed these spontaneous contractions of muscles has
been forty-five minutes; this was in the muscles of the penis and
scrotum in a dog that had been killed by the injection of chlo-
ride of barium into the veins. It is probable that the occurrence
of respiratory movements so long after apparent death (im one
instance seven minutes) is connected with the action of these
substances on the muscles, as the contractions they cause are
not simply contractions of individual muscles, but coordinated
movements. This action on the voluntary muscles is curiously
contrasted with their action on the muscular tissue of the heart,
at least on the ventricles, as I believe the irritability of the
auricles is not affected, the action of the Baryta group on the
heart being in this respect the reverse of the action of the chlo-
rine group, which, as I have already shewn, destroy the irrita-
bility of the auricles, whilst increasing that of the ventricles.
The salts of Lime, Strontia and Baryta exert but little action
on the muscular coats of the arteries, either pulmonic or sys-
temic; whilst the salts of Lead cause contraction in each set_of
vessels, agreeing in this respect with the salts of silver. I shall
ACTION OF INORGANIC SUBSTANCES IN THE BLOOD. 249
not however at present discuss the connection of these facts
with the general question of muscular irritability, or do more
than allude to the pathological changes caused by lead in the
muscular tissue, although they afford a new point of view from
which to investigate these questions. I would state that my
experiments prove that these muscular contractions are caused
by the direct contact of the substance with the muscular fibre.
The physiological action of these substances furnishes a well-
marked example of the increase of the intensity of physiological
action in the same isomorphous group as the atomic weight is
greater. Thus we find:
Atomic weight. Quantity required to kill.
[DDC aps ae ae ROMS ene ce eee wars 100 grs.
BSURUTECIA. ccs esness), s2 =o (aeeeeaters Sac Ae BOE 36 grs.
1 3 UG ee i 6s gee OL EN ae ee 4 grs,
With the salts of lead this connection between the atomic
weight and intensity of physiological action does not exist, but,
as has already been shewn, they also differ in their physiological
action in many respects from the other members of the group.
That this difference should be in the direction of assimilating
their action to that of the salts of silver, with which they have
isomorphous relations, is interesting, as when molecular physics
shall be able to point out in what consist the differences and
the resemblances of lead to the baryta and soda groups, we
shall probably attain more definite notions as to the causes of
the physiological phenomena to which it gives rise,
1 The quantities of the different salts used in these experiments are undoubt-
edly greater than would have been required had they been injected at one time.
My object has been to ascertain the general physiological action of the substances,
and not the smallest quantity that would arrest the vital reactions. As, how-
ever, the experiments were all performed in the same manner the data they
furnished are fairly comparable.
VOL. VIIt. Li
ON DOUBLE-BODIED MONSTERS, AND THE DEVE-—
LOPMENT OF THE TONGUE. By PRoFEssoR CLE-
LAND, M.D., F.R.S., Galway. (Pl. VIII. Figs. 3 and 4.)
SoME time ago I received from my colleague, Dr Browne, a
kitten with four additional limbs attached to its breast (Fig. 3).
Otherwise it appeared to be normal; but when I opened its
mouth I was much surprised to find that it had a cleft palate,
and that the well-developed tongue lay completely in the nose,
supported by the incomplete halves of the palate. This is the
point which is most remarkable; but as the monstrosity is one
which to my mind presents a considerable degree of interest,
and is less easily explained in detail than most others, I shall
shortly describe the more important peculiarities of the speci-
men, before discussing further the position of the tongue.
The mother of the little monster, instead of biting the um-
bilical cord neatly away, appears to have been so discomposed
as to have torn her unsightly offspring, and removed the whole
intestine as far up as the middle of the stomach, and down as
far as the rectum; but with this exception and the removal of
part of the liver, there was little damage done.
The appended hinder-limbs lay in the middle line, while
the fore pair inclined to the right side. The left appended fore-
limb had the head of its humerus placed between two portions
of the right sterno-mastoid of the supporting foetus; and ex-
tending upwards from this point was a lamina of tissue passing
inwards in front of the prevertebral muscles, and contaiing a
cord which was attached superiorly to the base of the skull in
the sphenoidal region, in the middle line, in intimate connexion
with the back of the pharynx (Fig. 4,q). The anterior division
of the right sterno-mastoid muscle was partly attached to the
head of the humerus of the right appended fore-limb, and
partly to the end of the cartilaginous left half of the divided
sternum, while its posterior division was attached to the right
half of the sternum. The scapule of the two appended fore-
limbs were directed towards one another, with their dorsal
DOUBLE-BODIED MONSTERS. 251
aspects toward the surface, and had their bases united together;
and immediately beneath these scapule, united to them by
fibrous tissue and muscle, was a pelvis reduced to the simplicity
of the piscine form, composed of two bones, which met in the
middle line and each supported on the superficial aspect the
head of a femur. Thus the four appended limbs diverged from
a central table formed by the union of scapule and pelvic
bones. All of them were smaller than those which belonged
to the developed kitten; and the left fore-limb had only three
toes, while the others had the proper number.
The sternum, as has been said, was in two parts, which had
not begun to ossify. These were separated one from the other
by the table of origin of the appended limbs, and, toward the
hinder part of each, a long pointed process projected inwards
towards its fellow on the deep side of the table referred to
(s, s). At first, I thought that these were the elongated
xiphoid extremities of the half sterna; but a more careful ex-
amination showed that the xiphoid extremities were in their
proper positions, and that these were separate elements. They
are probably to be looked on as rudimentary half sterna be-
longing to the appended embryo.
A deeper dissection exhibited two trachee with one pharynx
and cesophagus between them. The left trachea belonged to
the developed kitten; the other was situated with the properly
vertebral aspect foremost, that is to say, the incomplete parts
of its rings looking forwards; and it was surmounted with a
larynx smaller than the other, opening on the occipital wall of
the pharynx, with its epiglottis next the skull, and without any
tongue connected with it.
The thorax contained two completely s2parate hearts; and
in connection with each was a well-developed pair of lungs ;
those, however, which lay in the middle of the cavity, between
the two hearts, being more compressed than the others. The
left heart was that of the developed kitten. It had two ven-
tricles and two auricles: its aorta and pulmonary artery were
normal, and its right auricle received an anterior and posterior
vena cava normally. The left heart was slightly smaller; it
had no pericardial sac, and had only one auricle and ventricle.
Its auricle had two rudimentary appendages directed to the
17—2
JaZ - PROFESSOR CLELAND.
vertebral aspect, and received a vein (t), formed by union of
the azygos vein with an anterior trunk into which fell the left
subclavian and a deep vein from the left side of the neck.
The arterial trunk of this heart, springing from between the
auricular appendages, formed an arch directed towards the
left, and approaching the sternal wall of the thorax to be con-
tinued into a descending aorta which supplied the appended
limbs and gave off a hypogastric artery; but before coming to
the surface it communicated by a large branch with the ante-
rior end of the descending aorta of the developed kitten.
On the visceral side of the appended pelvic bones was a
little round body, apparently filled with meconium (n); and
from this a structure extended to the damaged umbilicus. I
believe that this was a vestige of intestine which had separated,
near the umbilicus, from that of the developed kitten, and
ended in a blind dilated rectum.
Concerning monsters consisting of a perfect foetus, with an
appendix which is “an acephalus with four extremities,’ Vrolik
states that, in the instances described, genital organs existed in
the appendix, but the anus was closed. Also, “in many cases
the evacuation of urine has been observed; the appendix showed
circulation of blood, it had its own temperature, and was de-
pendent for nutrition on the chief or perfect body: in the inte-
rior were found uropoietic organs, vessels connected with those
of the chief body, and an imperfect intestinal canal (Otto,
Serres). In the supporting foetus are sometimes found traces
of double organs (Otto, Serres, Rosenstiel) }.”
The nearest approach, which has come under my notice, to
the monster now described, is one of those described by Serres’,
and alluded to by Vrolik. It also was a kitten with the ap-
pended extremities very similar to those in my case; but the
hinder-limbs were about an inch anda half removed from the
fore-limbs. The rectum of the appended foetus was prolonged,
after a very short course, up to a cecum, whence an ileum was
continued to join that of the developed kitten im its lower
third. Thus, it will be perceived, an intestine, single above,
1 Vrolik, article Teratology, in Cyclopedia of Anatomy and Physiology.
2 Sur Vorganisation anatomique des monstres Andee Reprinted
from the Mémoires du Muséum.
DOUBLE-BODIED MONSTERS. 953
was double below, in the fashion common among double mon-
sters which have duplicity of the lower part of the trunk.
Further, in Serres’ monster the appended foetus had a urachus
and a pair of kidneys; and the representation of these latter
enables me to pronounce as a renal rudiment, in my specimen,
a small mass of substance near the lower end of the additional
aorta, which could not otherwise have been recognised. The
fore and hinder appended extremities were united by a thread,
which Serres describes as nervous and ending at its extremities
in ganglia to which converged the nerves of the limbs. There
were two pairs of lungs, and two traches opening into a single
gullet, exactly as in my specimen; but, curiously enough, there
was one respect in which the appended foetus was less deve-
loped than in my specimen; for in Serres’ kitten there was
only one heart. The artery of the appended fcetus came off
from the aorta in front of the trachez, and turned to the right,
while the arch of the aorta turned to the left.
Within my reach for consultation are the descriptions of
two other monstrous kittens, in both of which there was a
single head and two fully developed bodies. The first of these
is by the second Monro’. In his specimen the cerebro-spinal
axes were distinct; that of the right foetus presented a com-
plete encephalon, while the other had a much smaller “sphe-
rical body supplying the place of brain and cerebellum.”
Unfortunately no description is given of the osseous surround-
ings of this spherical body, and it is not apparent how it was
related to the cranium. Jn this monster there was only one
heart, from the left ventricle of which one aorta was sent out,
which soon divided into right and left aortz proper to each
body. There were, however, two pairs of lungs; one pair con-
nected with a normal trachea, and the other opening by sepa-
rate bronchi into the lower part of the cesophagus. Another
monstrous kitten has been described by Dr M*Intosh in the
pages of this Journal (May, 1868). In this specimen bifurcation
began in the back part of the skull, and there was a single
cerebrum continued into a double cerebellum, pons and me-
dulla. Two trachesee were disposed in a manner evidently
_ 1 Structure and Functions of the Nervous system, 1783. Table x11.
254 _ PROFESSOR CLELAND.
exactly similar to the disposition in my specimen; and there
were two distinct hearts, one much larger than the other, and
supplying both sides of the head.
From a comparison of the different monstrosities referred to,
it may, in the first place, be manifestly gathered that monsters
with appended limbs have fundamentally the same origin as
monsters with one head and two bodies. The nomenclature
which speaks of the appended limbs as belonging to an “ace-
phalus” is as misleading as could well be conceived. It is
intended to convey the idea of a headless embryo being added
to an otherwise normal foetus; whereas the real state of matters
is, that the head is the common property of the developed and
the appended body. This is evident from two considerations :
first, the resemblance of monsters with appended lmbs to
double-bodied monsters, and secondly, from double-bodied mon-
sters being always the result of fissiparous division of a single
embryo. In Dr M°Intosh’s specimen the fore-part of the head
manifestly was the only undivided part of an embryo which
had undergone fission from behind forwards at a period prior to
the laying down of the primitive groove; and, if one of the bodies
so produced had suffered arrest of development in the spimal
region, a specimen like that of Serres or the one in my posses-
sion would have been the result.
Yet there is this to be said in favour of the term “acephalus”
for the appended fcetus, that by far the most probable hypo-
thesis to account for the production of completely separate
acephali is that they have become, in process of early growth,
detached by fissiparous division from the developed feetus which
always accompanies such a monstrosity, and are therefore to be
considered as a third variety of result from posterior fission of
an embryo. In this view the head of the developed foetus is no
doubt originally the common property of it and the acephalus,
exactly as the head of a double-bodied monster is common to
both bodies; but the acephalus may be said to have forfeited
its claim to property in the head when it parted company com-
pletely from it. It would be of the highest interest, when oppor-
tunity occurs, to institute careful dissection of the body of the
developed foetus born along with a separate acephalus.
It, may be further suggested that, while in one set of in-
DOUBLE-BODIED MONSTERS. 255
stances of posterior fission, the two bodies resulting therefrom
grow nearly equally, in the other set the degree of imperfection
of the less developed body is dependent on the effects of being
stretched out, set on the rack, by the more rapid growth of the
other. It will doubtless be granted that the tough string ex-
tending, in my specimen, from the base of the skull to the fore-
part of the appendix, was really the vestige of the spinal column
of the appended embryo. Thus, taking the view suggested, it
might be said that the spinal column of the less developed body
was reduced to the condition of a string, by being attached at
one end to the skull, while a part beyond was, by its connexions
with the fore-limbs of the more developed body, pulled out at a
rate more rapid than that at which it had been able to grow;
and that the continued existence of the string prevented the
visceral parts from being as well developed as in the specimen
described by Serres. So also an early rupture of the connexion
with the head, and with the body of the perfect foetus, in the
case of a completely separate acephalus, would give freedom for
growth of the part separated. Reasoning in the same way, the
continuity of the spinal column with the head is seen to be a
reason why the column should be liable to suffer by stretching,
while the limbs, appearing in disconnected portions of blastema,
are only carried to unexpected positions, remaining uninjured
and developing freely.
In my specimen the appendix was the right division of the
original embryo, and had been carried round to the front by the
growth of the right thoracic parietes of the left or developed
division. In fact, in all double monsters the divisions of the
embryo lay originally side by side, and therefore, even in those
in which two bodies most completely face one another, there is
a ventral aspect originally turned towards the yelk, to be dis-
tinguished from a dorsal aspect. In this connexion it is inter-
~ esting to note that appendices, or so-called acephali, connected
with the sacral end of the column, remain attached to the
sacrum, while those adherent to the thorax are carried to the
front; and it is not quite obvious whether this is to be explained
by supposing that sacral appendices result from a later and less
perfect fission, or whether the earlier and more perfect growth
of thoracic parietes enables them better to carry to the front
256 PROFESSOR CLELAND.
appendices connected with them. 1 should incline to the first
supposition.
The presence of two pairs of lungs in all the four instances
of monstrosity recited, with formation of two tracheze in three
out of the four, while, on the other hand, in two cases there
were two hearts, and in the other two only one heart, is very
remarkable, and is even more so when it is considered that one
of the bicardial monsters was double-bodied, and that the other,
the one which had the smaller heart most nearly approaching
In size to the larger, was. the kitten in my possession, which
had the other parts belonging to the appended body less de-
veloped than they were in Serres’ specimen.
In trying to discover the explanation of these phenomena
one is led to consider the mode in which fissiparous division in
the embryo takes place. We have no evidence that complete
fission of an impregnated ovum prior to the appearance of the
embryo ever occurs; and the phenomenon which has been met
with, and which explains the formation of double monstrosities,
is fission more or less perfect, not of the ovum, but of the
embryo placed thereon. The embryo and the ovum are two
very different things; indeed to my mind it is obvious, as I
have stated elsewhere’, that the development of the embryo
from the ovum is a process of gemmation; and, especially when
multiplication of the embryo by division is attempted, I can see
no difference between the plan of reproduction in vertebrata
and the alternate generation of meduse. But what interests
us at present is that fissiparous division, when it occurs, being
confined to the embryo, the remainder of the ovum presents the
obstacle which most interferes with its completion, and the part
most intimately connected with the yelk or cavity of the ovum
exhibits least tendency to divide. That part is the alimentary
tube, particularly the portion neighbouring to the umbilical
vesicle. Indeed the tract of intestine usually found single in -
double monsters extends from the duodenum to the place of
connexion with the umbilical vesicle; but. the reason why it
reaches so far up I judge to be connected with the very early
appearance of the liver; having found in a dissection of a
1 Animal Physiology, p. 276. 1874.
DOUBLE-BODIED MONSTERS. 257
double monster, made many years ago for Sir James Simpson,
and still in the Edinburgh University Museum, a common
duct, from two pancreases, and a liver with two gall-bladders,
opening at the angle of junction of the two duodena.
We see, then, that division of the embryo by fission, whether
extending from before backwards or behind forwards, meets with
least obstruction on the dorsal aspect; and the things which
strike me as the interesting points to be learned from the du-
plicity of the air-passages and occasional duplicity of the heart
in double-bodied monsters with single head are, first, that in a
part already separated from the ovum, the tendency to duplicity
exhibits itself on the ventral as well as the dorsal aspect, while
the gullet situated in the centre between the two aspeets remains
single. Secondly, it is interesting to note that while the tend-
ency to duplicity is interrupted on the ventral aspect by the
connexion with the yelk, the direction in which the duplicity
has travelled continues the same in front of the interruption as
behind, the fore part of the head being in every case single,
‘That the heart is in some cases single, when the trachea, which
must be considered as a deeper structure, is double, is perhaps
to be explained by the earliness of the heart’s first appear-
ance; it may in one instance be already laid down as a single
tube, and in another be not yet begun to form, at the time
when the tendency to fission has travelled so far forwards on the
ventral aspect.
The still earlier period at which the gullet appears may also
be taken into consideration in connexion with the singleness of
that structure. But I confess that it is a subject of marvel tome
to find, in my own specimen, as seems also to have been the case
in that described by Dr M‘Intosh, the right trachea, the smaller
of the two, turned completely round, and the larynx opening on
the cranial side of the pharynx. That the right heart was so
turned is easily understood: it was carried round with the body
to which it belonged. But applying this explanation to the
larynx and trachea, the axis of revolution must be considered
as passing along the centre of the gullet; and it is simply im-
possible to conceive a larynx and trachea, originally placed to
the right, rotating to the left, and continuing their revolution
till the larynx touched the vertebral column of the body to which
258 PROFESSOR CLELAND.
it was opposed, and the trachea regained its position on the right
side. Therefore, either the larynx revolved toward the right side
of the gullet after the body to which it belonged had revolved
from right to left, which seems impossible, or the revolution of
the body took place before the windpipe began to be developed.
Even on that supposition it might well puzzle a metaphysician
to apportion the parts of the gullet between the bedies to which
the windpipes belong. Explain it as we may, there seems to be
an interlocking of the arez in which the formative forces of the
two bodies dominated. Matters are not made simpler by the
consideration that in my specimen the vestiges of the spinal cord
of the right body were adherent to the top of the gullet, close
to where the larynx opened.
These remarks have become much fuller than I had intended.
Let us revert now to the peculiar position of the tongue in the
cavity of the nose, folded round the vomer, and above the cleft
palate. In all the three cases which I have cited to illustrate
the kitten in my possession the palate was cleft, and apparently
so widely, that nose and mouth made one cavity, and no in-
formation was to be gained as to the relation of tongue to palate,
But the circumstance which led me to think of writing this
paper was, that a few weeks ago my colleague, Dr Doherty,
placed in my hands a spirit preparation of a two-headed kitten,
belonging to the Montgomery Museum, and, finding it neces-
sary to change the spirit, I took the opportunity to examine the
mouths of the two heads of the monster, and found a very re-
markable state of matters. In the right head the palate was
cleft, and the tongue folded round the vomer and placed com-
pletely in the nose, exactly as in the double-bodied kitten. In
the left head the palate was perfect, but the mouth contained
no tongue; a pillar of substance, however, ascended in the back
of the throat, completely filling the pharynx, so that at once’
it could be perceived without dissection that, in this head also,
the tongue was impacted in the nose. The question however
occurred, how is the tongue disposed? With the complete de-
velopment of the palate there is ordinarily associated complete
formation of the nasal septum; but in this instance there must
needs be either deficient development of the septum, or
deficient development of the tongue. To determine this point
DOUBLE-BODIED. MONSTERS. 259
I obtained permission from Dr Doherty, and laid open the nasal
cavity on one side, so as to display the relations of parts. It
was then found that the tongue was less than half its proper
size; the vomer failed to pass so far backwards as it ought, and
the tongue was rendered bifid at its extremity, by its attempt
to grow forwards into the nasal fossze on each side of the vomer.
Let us explain the matter as we may, it is certainly very
remarkable that, in six heads of monstrous kittens, the only
examples of monstrosity in those animals coming, by observa-
tion or otherwise, under my notice, there should not be a
normally developed mouth; that in five out of the six the
palate should be cleft, and that in the three cases in which the
palate existed, in whole or in part, the tongue lay above it. Nor
does the matter seem less curious when it is considered that
two of the heads were attached to a single body, while the
other four were each in connexion with two bodies more or
less completely developed. In all instances, however, the con-
nexion of double structures with single was calculated to lead
to crowding; and it is possible to imagine that this crowding led
to the tongue being forced unduly upwards, and that the altered
position of the tongue occasioned the cleft palate by interfering
with the closure of that structure.
This displacement of the tongue is particularly interesting
as throwing light on a question of development which has not,
so far as I know, been settled by direct observation. The mouth,
as is well known, makes its appearance originally as a depres-
sion of the integument; and afterwards, at a very early period,
a cleft of communication takes place between this depression
and the anterior extremity of the alimentary tube. The point
to be determined is, whether the tongue is to be considered as
connected with the integumentary depression, or with the mu-
cous membrane. These monstrosities would seem to show that
it belongs to the cul-de-sac of mucous membrane. The original
roof of the integumentary depression is, indeed, a totally differ-
ent thing from the ledges which, long after its rupture, appear
and unite to form the palate; it is of similar nature to the
membranes which originally occlude all the cervical clefts; and,
as I have said, the position of the tongue above or below the
palate might well depend on the amount of space left by the
260 PROFESSOR CLELAND. DOUBLE-BODIED MONSTERS.
growth of parts below. But while the normal tongue may be
said to arise from the floor of the mouth forwards to the sym-
physis of the jaw, the arrangement in these cases of monstrosity
shows that the root of the tongue makes its appearance first
behind and afterwards grows forwards,
EXPLANATION OF PLATE.
Fig. 3. Kitten with eight limbs described in the text; some-
what reduced. Fig. 4. Dissection of the same; somewhat enlarged.
a, cleft palate ; 6, tongue drawn aside from its position, grooved so
as to fit round the vomer; c, c, lower jaw cut across; d, e, left. and
right larynx, surmounting their trachee ; f, g, anterior and posterior
vena cava of left heart; h, left heart; 7, aorta from the same;
k, esophagus; /, aorta of right heart; m, hypogastric branch of the
same; 7, rectum of appendix; o, kidney of the same; p, right heart;
g, cord, cut across, which attached appendix to base of skull; 7,
sterno-mastoid muscle; s, s, appendices to the two halves of the
sternum; ¢, vein entering right heart; wu, symphysis of the pelvis of
the appended limbs.
ON THE CARTILAGES AND SYNOVIAL MEMBRANES
OF THE JOINTS. By Cart Revuer, M.D., Privat-
docent of Surgery in the University of Dorpat. (Plate IX.)
IN a previous publication’ I have given the results of an
experimental study, which I had undertaken, of the changes
which the joints undergo under conditions of prolonged inac-
tivity. It was there shewn that, both in the young and in
the adult, beyond those parts of the articular surfaces which
are habitually in contact, the large, flattened, round cartilage-
cells pass, by the gradual acquisition of cell-processes, into the
spindle-shaped or stellate connective tissue corpuscles of the
synovial membrane. In the same communication also I drew
attention to the great similarity observable between this tran-
sition and that seen in the development of the so-called mar-
ginal zone of the synovial membrane, or, in other words, the
portion of that membrane which extends over those parts
of the articular surfaces which are not ordinarily in contact,
and started the question as to whether this marginal zone were
really to be regarded as a growth from and production of the
synovial membrane or not. The generally received opinion is,
that at the time when the cartilages corresponding to the several
bones that are to be formed become separated from one another
by fissuring of the cartilaginous substance accompanied by
increased growth of the cells and liquefaction of the inter-
cellular substance, at the same time a corresponding portion
of perichondrium developes into the capsule of the joint; and
finally, that those parts of the cartilaginous ends which are not
constantly subjected to mutual contact become covered with
an ingrowth of the synovial membrane. Hiiter, in accordance
with these views, conceives the marginal zone of the synovial
membrane to be derived, not from the rapidly growing layer
of embryonic cells by the dehiscence of which the cavity of the
joint is formed, but, as a true ingrowth of the synovial mem-
brane considered as a tissue the matrix of which is situated in
1 Deutsch. Zeitsch. f. Chirurgie, Bd. 111. p. 189.
262 DR REYHER.
the innermost layer of the capsule, and from which it becomes
infolded over those surfaces of the joint which are not habitually
in contact. If this opinion be correct, immediately after the
formation of the joint each cartilaginous end should possess the
peculiar form seen in the future bone, e.g. in the hip, the one
the concave form, the other the convex: the latter (the head)
will then have portions of the surface in constant contact or not,
according to the movement which is permitted to the limb in
utero. This, however, is not in accordance with our ex-
perience ; on the contrary, we are forced to consider that at first
the joints are very simple and incompletely developed, possess-
ing gliding and rotatory movements around one axis only; and
it is only in the course of development, by aid of the constant
attempts which are continually being made to move the limb
around other axes, that the stereometric forms of the articular
ends, which determine the typical movements of the limbs after
birth, become developed. In accordance, also, with the manner
in which subsequent movements take place, marked changes
are found to occur’.
Whichever view of the formation of the articular surfaces is
the correct one the question of the origin of the “marginal zone”
of the synovial membrane still remains open. The investiga-
tion of foetal joints can alone definitely solve the problem: the
results of observations on these form the basis of the present
article.
The investigation of the surfaces of embryonic joints presents
considerable difficulty, the parts composing them being too
small and soft to enable good sections to be made: the latter
also are difficult to obtain on account of the sharp curves of
the surfaces. For this reason I have not been able to subject
the earliest stages to the microscope, but have been obliged to
be contented with such as are to be found in embryo sheep of
about two inches long; I hope, however, that the facts arrived
at will appear sufficient to yield a correct notion of the develop-
ment of the so-called “synovial processes*.” I have carried on
1 Tn illustration of the same principle I need only refer to the physiological
changes which occur in the bones (particularly in the head of the astragalus) in
the different forms of club-foot.
2 The synovial ingrowths or processes referred to in the text are not to be
eonfounded with the well-known Hayersian fringes, which project freely in the
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 263
similar investigations in the embryos of guinea-pigs and dogs,
and in new-born puppies, rabbits, and cats; besides studying
the structure of the parts in full-grown sheep, oxen, and
dogs, and in the human joints at different ages, In order to
demonstrate the structure of these synovial processes, I have
employed, as was done by Hiiter, the method of treatment
with nitrate of silver.
Before, however, the observations are considered in detail, it
may not be out of place to say a few words as to the value of the
silver treatment in the investigation of the synovial membrane. As
is well known, the usual interpretation of the images obtained by
means of the silver treatment has been called in question by Schweig-
ger-Seidel, and doubt has been thrown upon the cellular nature of
the figures and appearances which are produced in the synovial mem-
brane by means of this reagent. His objections have been fully
disproved as regards other organs (e.g. the cornea), in which, with
different methods of treatment, corresponding outlines are always ob-
tainable. Whilst firmly convinced that the same holds good with
regard to the synovial membrane (having looked upon it in this
light in the work which I have before referred to), it must be remem-
bered that so long as neither the treatment with chloride of gold nor
combined methods had been employed in the investigation of this
tissue, a gap was left in the evidence as to the nature of the siiver
outlines. I anticipated that these would render service, more espe-
cially in bringing to light the exact meaning of the large white
stellate fields’, apparently belonging to the same category as those
demonstrable in the cornea, butas to which it was uncertain whether
they belonged to groups of cells or only to single ones. My in-
vestigations with respect to this point have been principally made
on the joints of full-grown sheep and oxen, the tarso-metatarsal
joints of which, and especially of the last-named, yield marginal
zones a finger’s-breadth wide. The sections were always made sub-
sequently to the occurrence of the silver precipitation; in this way
the clearest images are obtained, and there is no fear of cutting
sections of cartilage from which the marginal zone has been acci-
dentally rubbed off.
I had so often attempted to combine the staining with other
reagents, such as carmine and aniline, with that obtained by the
silver method, but without any great measure of success, that I was
extremely pleased to find that hematoxylin, which I made trial of at
Prof. Sanderson’s suggestion, furnished a perfectly reliable means of
articular cavity, and the microscopic structure of which has been carefully
described by Rainey and others, but are, as already mentioned, merely those
portions of the synovial membrane which lie over the borders of the cartilage,
and the connective-tissue corpuscles of which pass by gradual transition into the
cartilage-cells,
? Comp. Hiiter, Virch. Arch. xxxvt. Plate 1. fig. 10,
264 DR REYHER.,
staining the cell-nuclei. By the employment of sections which are
sufticiently thin to obviate any sources of fallacy arising from the
presence of the nuclei of the more deeply-seated cartilage-cells, it is
not difficult to be convinced that the white fields on the brown
ground of the silver preparation, from the more circular spaces of the
cartilage to the stellate and epithelioid forms of the inner layer of
the capsule, each contain either one or several (violet-coloured) nuclei.
By this method then it is demonstrable that in each of the white
fields of the silver preparation there lie, according to the size of the
fields, one or several cellular elements. It is, however, impossible by
this means to say whether the cells entirely fill the cavities, and by
means of their processes extend into the lymphatic canaliculi, form-
ing a complete anastomosing network, or not ; for the elucidation of
these points the treatment with chloride of gold is necessary. With-
out going farther into the description, I need only recommend a
comparison of Figs. 3 and 4, which are taken from two prepara-
tions of the tarso-metatarsal joint of the ox, the one representing
a preparation treated by the combined silver and hematoxylin
method, the other with gold. In both kinds of preparations appear-
ances are to be met with, which, in general form and in mode of
branching of their processes, are more or less similar.
As the hematoxylin shews the presence in the silver prepara-
tions of cell-nuclei corresponding to the white spaces, so the treat-
ment with gold shews that these nuclei belong to protoplasmic
bodies, which—the conclusion will hardly be assailed—correspond on
the whole to the spaces in the silver preparations. It is quite
another question whether these masses of protoplasm completely fill
the spaces or not. Proof of this could only be obtained were it
possible to produce both the gold and the silver appearances in the
same preparation; as is well known, however, if a preparation be
treated first with silver then with gold, the only effect is to produce
a reduction of the latter in the parts impregnated with silver, whilst
the converse mode of treatment altogether fails to yield the silver
spaces. The question must, therefore, so far remain unsettled. All
one can say is, that in both silver and gold preparations appearances
are frequently obtained, which, as regards form, are precisely similar,
apparently even to the minutest details, although it is not everywhere
possible to trace the same exact resemblance. or instance, the pro-
toplasmic masses of the one might be said to be smaller relatively
than the spaces of the other: the sizes, however, in both are so
varied that it is difficult to compare them. If the forms obtained by
the gold treatment differ from those obtained by the silver treat-
ment, in one point more than in another, it is in the diameter of the
processes, which here and there appear somewhat smaller and more
tapering than those proceeding from the spaces of the silver pre-
paration. On the other hand, the appearances presented in silver
preparations, which have been placed in spirit, are in favour of the
idea that the spaces are completely filled by protoplasm. In these
it may here and there be seen, although, it must be admitted, not as
a rule, that both the nucleus and the protoplasm are to be made out,
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 265.
_the latter appearing as a finely granular substance, which is separated ;
from the brown intercellular substance by a crescentic, clear zone
or space (perhaps caused by a shrinking of the protoplasm).
We may conclude, therefore, that the white spaces and canaliculi
-shewn to exist in the synovial membrane by treatment with nitrate
of silver, correspond generally to a network of protoplasm (made
evident by chloride of gold) consisting of connective-tissue corpuscles.
A similar statement may be made with regard to the cartilage-
‘cells of the surface, which appear after treatment with silver as
round white spaces, in which hematoxylin brings the nuclei into view,
_whilst on the other hand chloride of gold colours the protoplasm of
the cells,
The appearances produced by the silver method having
been thus explained, we may proceed by its aid to investi-
gate the structure of the joints. If the head of the femur
of a sheep's embryo 1?inch long be brought under observa-
tion, it is easy to satisfy oneself that the layer of cells described
as an epithelium by Todd and Bowman, and by Reichert, does
not exist. The appearances presented by the surface of the
cartilage after treatment by this method are precisely similar
to those seen within its substance. The cells at this early stage
jie so close to one another, that there is no matrix apparent,
and it is difficult even to make out their outline. All that is
visible is an apparently homogeneous substance with nuclei
imbedded in it of varying form, which might easily be taken
for the cells themselves imbedded in an intercellular substance.
Teazed preparations shew, however, that they are really nuclei
surrounded with a variable amount of protoplasm. This obser-
vation coincides in the main with that made by Toynbee, who
found (p. 163) that in the calf foetus of 12 lines long there
-was no difference between the cells on the surface and those in
the substance of the cartilage. The small processes which
Luschka describes as projecting into the articular cavity I have
not succeeded in coming across, either in these young ones or
in older embryos: I have however only investigated the larger
jomts. The appearances seen in sections from embryos a
little more advanced (sheep’s embryo about 24 inches long) are
somewhat different. Besides places in which the above de-
scribed condition obtains, others are also found in which the
nuclei on the surface are rather more separated, brown lines
appearing here and there between them. In some places the
VOL. VIII. 18
266 DR REYHER.
irregular cartilage-cells are still closely juxtaposed, whilst in
others they are somewhat larger and flatter, and fine brown
lines shoot out between them, and in some parts completely
bound the cells. Later on, the brown lines become more fre-
quent and broader, and in certain places form an elegant net-
work like that on the surfaces of the serous membranes, dif-
fering only in the fact that the territories marked off are com-
monly smaller and more irregular: the appearance is quite
that of an epithelium (fig. 1, a). In older embryos this epi-
thelioid appearance is still more common and complete, so that
the larger part of the articular surfaces is covered with these
flat epithelium-like cells. This is particularly well seen in the
head and condyles of the femur of new-born rabbits and cats,
and it is doubtless this that Todd and Bowman refer to in
stating that (p. 127) “in the feetus it (the epithelium) is con-
tinued over the whole cartilage.” In this, however, I cannot
concur, for the transition is gradual from the parenchymatous
€ondition (without matrix) of the surface-cartilage to the ap-
pearance of a slight amount of intercellular substance, the
latter becoming gradually more and more increased in amount,
so that the white fields become separated from one another by
broad bands of matrix, and assume an irregularly angular
form; in fact, the appearances obtained are quite similar to
these seen on the so-called “synovial process” of the adult’.
In preparations taken from the shoulder-joint the surface-cells
are roundish and connected here and there to one another ; in
those from the patella, more spindle-shaped and trailing off into
processes (fig. 2,¢, f), their long axis being parallel with that of the
limb. These differences appear to me to be produced mechani-
cally. The all-round movements of the shoulder-jomt would
naturally not influence the growth of the cells in one direction
more than in another, whilst the single to-and-fro movement of
the knee-joint might be conceived to have the effect of pro-
ducing elongation of them. But be that as it may, it is certain
that with the growth of the cartilage the intercellular substance,
- 1 These forms of cells have been termed by Hiiter epithelioid and keratoid,
meaning thereby to imply a resemblance in form to pavement epithelium and to
the connective tissue corpuscles of the cornea respectively. The term ‘tkeratoid”
is however somewhat ambiguous, and has therefore not been adopted in this
paper.
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 267
as well on the surface as in its depth, increases in amount and
causes the cells to become more separated from one another.
To recapitulate:—The surface-cells in the earliest stages
lie close together without intercellular substance ; later on, the
latter becomes developed as fine lines between the cells, pro-
ducing an epithelium-like appearance (fig. 1, a); stall later, by
a further development of intercellular substance, the cells
become more separated and irregular (as in fig. 1, 6). The
intercellular substance (matrix) becomes increased over, as well
as between, the cells, so that in the adult there is a distinct
layer of hyaline matrix covering the surface of the cartilage.
While this is occurring the irregular stellate and angular
cells on the one hand, and the elongated cells with trailing
processes (fig. 2, f) on the other hand, become gradually trans-
formed, in parts where the articular surfaces are constantly in
contact, with loss of their processes into the round scattered
cells of ordinary cartilage (fig. 2, d).
Thus much having been said as to the development of the
parts of the surface which are constantly in contact, we have next
to consider the condition of the articular surface in the neigh-
bourhood of the capsule. At the time that the proper articular
surface has the epithelium-like appearance before described, the
same can be traced over its margin as far as the insertion of
the capsule. The parts in which, in the adult, vessels and
irregularly disposed cells are to be found, are covered in the
foetus merely by an epithelioid layer of cells similar, as just
remarked, to that on the remainder of the surface of the car-
tilage ; and it is only in places which are more affected by the
movements of the joint that other forms are to be found. In
size the cells are quite similar to those covering the cartilage, but
differ from them somewhat in shape, being less polygonal and
separated by rather more intercellular substance, into which they
send processes which are, however, very short and knobbed, not
long and tapering, and extending far into the intercellular sub-
stance, as is the case in the adult (figs. 3, 4). Fig. 1 is a correct
representation of a portion of the peripheral surface of the
glenoid cavity of the shoulder-joint from a sheep’s embryo five
inches long. The larger part of the surface presented a beau-
tiful epithelioid appearance. In the figure, which corresponds
18—2
268 - DR REYHER.
to the outer raised margin bounding the glenoid hollow, two
parts are to be discerned, a vascular (c) and a non-vascular (a, 6).
The last-named must necessarily have been in contact with the
head of the humerus, whilst the vascular part corresponds with
the line of insertion of the capsule, and represents the circulus
articuli vasculosus of W. Hunter. On the glenoid part (fig. 1, a)
we see the regular epithelioid arrangement, on the synovial part
(c) the irregularly disposed cells of the inner layer of the capsule,
and between these every transition. The appearances seen
give the impression of a change being produced in the pre-
viously more regular cells by the tension to which the synovial
membrane is subjected in the neighbourhood of its insertion.
In the knee-joint we find the different stages we have described
still earlier. In sections from the surface of the patella of an
embryo sheep, four inches long, comprising both the cartilage
surface and the synovial margin, I have been able to see in
silvered preparations how the cartilage-cells, as we approach
the synovial membrane, begin to exhibit processes which get con-
tinually more numerous, and the cells more irregular, until they
take on an appearance similar to those on the inner layer of
the capsule, and indeed come into connection with them by
freely anastomosing branches. Part of such a preparation is
represented in fig. 2. At (d) the round white fields corresponding
to the cartilage-cells are to be seen; at (e) these possess processes,
at (f) they come to be still more branched and irregular. This
part corresponds to the beginning of the capsule, the inner
layer of which is to be seen in another part of the same prepa-
ration as a beautiful layer of epithelioid spaces. .
Whilst in fig. 1 the cells of the cartilage-surface (qa) still
present an epithelioid arrangement, and those of the inner layer
of the capsule may also be termed epithelium-like, in a further
stage of development the appearances are quite the reverse.
Over the cartilage the cells (fig. 2, e) are not only far apart, but
present most irregular forms, whilst in the neighbourhood of,
and upon the capsule they may be termed epithelioid. These
differences get more and more marked as development proceeds.
In full-grown animals the round spaces become so common,
and the irregular ones so rare, that it is now no longer easy to
find the latter, and it is only in particular localities that they
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 269
present themselves, and then in no great amount. Instances of
such places are to be found in man at the lower border of the
patella and the anterior border of the tibial surface of the astra-
galus (Hiiter). Larger animals would naturally be expected to
shew this more markedly. In accordance with this in oxen, in
sections of the articular surface, including the marginal zone,
which had been treated with nitrate of silver, and subsequently
with hematoxylin, 1 have observed almost every transition,
from the irregularly stellate cells, ike those found in the cornea,
to the round cell-spaces of the cartilage.
It rarely but occasionally happens that a network of irregular
cells is seen in the immediate neighbourhood of round ones:
here also, however, some of the cartilage-cells are met with, pos-
sessing a single process and representing rudimentary stellate
cells, as pointed out already by Hiiter and Béhm. The meaning
of these silver images needs but little further explanation. As-
suming the white fields to represent masses of protoplasm with
nuclei, and the round ones in the homogeneous ground-substance
to correspond with the cartilage-cells, the rest, whether pos-
sessing processes or arranged like an epithelium, must be looked
upon as connective-tissue corpuscles.
The Synovial Membrane—In spite of the difficulties pre-
sented by the softness of the embryonic tissue to subjecting
silvered portions of the synovial membrane uninjured to the
microscope, I have nevertheless been so far successful, and I
am able to confirm, in every respect, in the embryonic tissue,
the observations of Hitter upon the adult; and I must distinctly
deny the existence of an epithelium, or at least of a distinct
layer of cells, such as Landzert and Albert have lately described
as covering the vessels and the sublying cell-spaces. It is true
that places are to be found on the inner surface of the capsule
in which the cells are regularly arranged, like those upon the
surface of the cartilage, represented in fig. 1, a: such patches
are, however, never very extensive, and always shew at their
margins a transition to the more separated cell-spaces. The
layer is in reality characterised by the variety of the cell-forms,
and the differences in width and regularity of the intercellular
substance; and it is only upon the surface of the cartilage itself
that, as before seen, any appearance similar to that of the serous
270 DR REYHER.
epithelium is observed: as we approach the capsule this dis-
appears, and the irregular arrangement is seen (fig. 1, b, ¢, c’).
As regards the relations of the vessels, also, 1 can confirm
Hiiter. In the same manner as in the marginal zone they le
between the epithelioid cells, so here on the inner surface of the
capsule. It is true that they are very often seen covered by
cell-spaces (Saftcandlchen); some, however, rise to the surface.
As we have seen then, in the earliest stages of feetal life the
form and arrangement of the cells on the surface of the cartilage
are precisely similar to those in its substance. They then become
flattened, and form a layer of cells somewhat similar to a serous
epithelium. At the borders of the cartilage these become jagged,
and pass into forms which are more and more like those of the
“Saftcanilchen” of the serous membranes. This arrangement
obtains throughout life on the inner surface of the capsule. A
change, however, takes place in the cells on the surface of the
cartilage, and this change depends, firstly, on the growth of the
articular surfaces; secondly, on the varying conditions of contact
and pressure to which they are exposed.
With respect to the first cause it may be remarked that the
cells become more or less evenly or unevenly disposed, accord-
ing as the general articular surface becomes evenly developed
or not. Thus on the concave articular surfaces, the growth of
which is more or less uniform both near the centre and at the
periphery, the cells, whether epithelioid, stellate or rounded,
are more or less similar throughout both in form and dis-
position, whereas on the convex surfaces such as the head of the
humerus or femur, the superficial cells in the parts m the
neighbourhood of the neck are far less separated than those
nearer the centre of the articular surface, where growth and
development are more rapid.
With regard to the second, it is to be noticed that in parts
of the surface which are always in contact, the epithelioid cells
become, as development proceeds, irregularly stellate, finally
losing their processes and becoming round, so that by the time
of birth the epithelioid arrangement has in most animals dis-
appeared at these parts.
The fact of the conversion of the epithelioid layer on the
articular surface of the foetal cartilages into the large and round
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 271
cartilage-cells, met with on the surfaces of joints which are no
longer embryonic, such conversion occurring not only on parts
of the surface which are not constantly in contact, but also,
although at an earlier stage, on parts which are, se not, I
Hetiene! been previously pointed out.
One is at first inclined to ascribe this conversion of the
epithelioid cells into stellate corpuscles, and those again into
round cartilage cells, to the rubbing together of the articular
surfaces. JI am, however, rather disposed to believe most of it
to be due to the mutual pressure of the opposed cartilages, and
this I am led to think because new canal-systems become formed
below the superficial network of protoplasm (fig. 4), long pro-
cesses passing down from this into the substance of the cartilage,
where, of course, there is no exposure to friction. Such a one is
seen in fig. 3, g, the body of the cell being quite on the surface,
the process, as may be seen by focussing the microscope, dipping
down into the matrix.
As the pressure and counter-pressure exercised on the dif-
ferent parts of the articular surfaces vary with the direction and
frequency of the movements of the joint, so we find correspond-
ing differences in the cells, such as have already been more than
once alluded to. The converse fact I had previously demonstrated
experimentally in the most striking manner by keeping the
joints of dogs at rest. Under circumstances such as these, in
which all the effects of pressure and movement are removed,
the cells on the articular surfaces again take a more or less
epithelioid arrangement. This change is accompanied by an
absorption of intercellular substance, and extends also to the
deeper layers of the cartilage.
These facts shew, I think, that the synovial process, so
called, is not to be looked upon as an ingrowth of the synovial
membrane, as some have asserted, but rather as being formed
in situ as the development of the joint proceeds, its cells being
intimately related both by the history of their development and
by the presence of intermediate forms with the cartilage-cells
of the articular surface. Whether it is derived from the
dehiscing substance or not I am unable to say, not being suf-
ficiently familiar with the changes which the newly-formed cells
here undergo; but this at least is certain that, at a later period,
Dia DR REYHER.
both the cells on the inner surface of the capsule and those on
the surface of the cartilage form a continuous layer, and the
so-called synovial process—the marginal zone of later periods of
development —is seen to be derived from part of this layer.
The existence of vessels in the synovial process is not, in
my opinion, any argument for its origin from the inner layer of
the capsule, for, as Toynbee’s researches shew, these parts are
at the time of the fissuring to form the joint all devoid of ves-
sels, which only appear later, and may as readily be conceived
to develope in one part as in another. But however that may
be, the fact that the epithelioid layer of cells may be in ex-
istence, and even disappear long before the appearance of
vessels, and without any development of vessels occurring in it
(e.g. on the concave surfaces), shews that the vessels are of no
importance in deciding the question at issue. That they are con-
tinuous with those of the synovial membrane, and play an im-
portant part in the secreting and absorbing functions of the
parts supplied by them, is however undoubted: and, as already
remarked, at the margins of the articular surfaces the cells are
of an irregular form, and, although derived from cartilage-cells,
are similar in appearance to those on the inner surface of the
capsule. The function also of the tissue here is doubtless similar
to that, and for this reason possesses vessels.. So far, then, it
may not altogether incorrectly be considered an offshoot of the
synovial membrane. It is not, however, merely a membranous
layer of cells spread like a carpet over the marginal surface of
the cartilage, but is, on the contrary, to be considered an integral
part of the latter, for, as already shewn, the processes of the rami-
fied cells actually dip down into the matrix of the cartilage:
this may be especially well’ seen near the inner margin of the
“synovial processes,” where its cells are disposed in a single layer
only. Nearer the synovial membrane the ramified cells are dis-
posed in more than one layer, being placed one over the other,
somewhat as in the cornea (fig. 4). The dipping down of the
processes is represented in fig. 3: they are often 20 or 30 times
as long as the diameter of the cell-body, and pass obliquely
inwards into the intercellular substance of the cartilage. The
cells, therefore, of the “synovial process” would seem to serve as
a medium for the conveyance of fluid, not only between the
CARTILAGES AND SYNOVIAL MEMBRANES OF THE JOINTS. 273
blood-vessels and the articular cavities, but also, to some extent,
between the vessels and the subjacent cartilage itself.
The investigations which form the basis of this article have been
mainly conducted in the Physiological Laboratory of University
College, and I would take this opportunity of expressing my thanks
to Professor Sanderson for the abundant materials placed at my
disposal, as also to Professor Sharpey and Mr Schiifer, to the latter
of whom I am indebted for the drawings from which the figures in
the Plate have been executed.
LITERATURE.
Rathke, Lntwickelungsgeschichte der Natter, Konigsberg, 1839.
Rainey, Proc. Roy. Soc., 1846.
Luschka, MWiiller’s Archiv, 1855, p. 487.
Toynbee, Philosophical Transactions, 1841, p. 163.
Todd and Bowman, Physiological Anatomy and Physiology of
Man, London, 1843, pp. 90, 93, 127.
Henle, Handbuch der systematischen Anatomie, Biinderlehre, p. 11
and p. 2.
Reichert, Miiller’s Archiv, 1849, p. 16.
Hiiter, Virchow’s Archiv, Bd 36, p. 65.
—— Clinik der Gelenkkrankheiten, p. 49.
Landzert, Centralblatt der med. Wissenschaft. 1867, p. 370.
Albert, Stricker’s Handbuch der Gewebelehre, p. 1232.
Bohm, Jnauguraldissertation, Wiirzburg, 1868.
Schweigger-Seidel, Berichte der Kon, Scichsisch. Gesellschaft d.
Wiss., Math. Phys. Klasse, 1866.
DESCRIPTION OF PLATE IX.
Ficure 1. Surface of peripheral part of glenoid cavity of 5 inch
sheep’s embryo (silver preparation).
a. Epithelioid arrangement of superficial cartilage-cells.
b. Transitions from these to
¢, ¢’. Saft-caniilchen of synovial membrane. Opposite ¢ the Cir-
culus articuli vasculosus c is taken from the part of the
membrane lining the capsule.
Ficure 2. From the patella of 4 inch sheep’s embryo, shewing
transitions of cartilage-cells into connective-tissue corpuscles of syno-
vial membrane (silver preparation).
d. Cartilage-cell-spaces.
e. The same acquiring processes.
J. Elongated and irregular spaces of the so-called synovial
process.
Ficures 3 and 4 are from preparations of the marginal zone of the
tarso-metatarsal joint of the ox. In figure 3 (nitrate of silver pre-
paration) the cell-spaces, in figure 4 (chloride of gold preparation)
the cells themselves, are shewn.
THE ACIDITY OF GASTRIC JUICE. By Jas. Reoca,
M.A., M.B. (Edin.), Demonstrator of Practical Phystology,
College of Medicine, Newcastle-upon-Tyne,
SomE time ago I began a series of experiments on the Sulpho-
cyanide of Potassium, with a view of determining its use in the
saliva. Commencing, then, with its reactions on the various
Pharmacopeeial preparations of iron, I found that while the
ordinary ferric salts, such as the Perchloride and Nitrate, gave
the blood-red coloration of sulphoeyanide of iron readily, the
compound iron salts with organic acids did-not do so at all; but
when a drop of dilute hydrochloric acid was added, the reaction
came out at once. There are three salts which in solution ex-
hibit this peculiarity—Ferrum Tartaratum, Ferri et Ammonis
Citras, and Ferri et Quiniz Citras; but as the two former are
red themselves, while the latter is golden yellow, I did not con-
sider the reaction so delicate or so free from suspicion of error
in the former as in the latter, and therefore confined my experi-
ments chiefly to the Ferr. et Quin. Citr., which for shortness I
shall call Ferr. Citr. Now, when a solution of this salt, of the
strength of 1 gr. or 2 grs. to the ounce, is mixed with a solution
of the Sulphocyanide of Pot. of similar strength, no reaction
takes place, but the addition of a single drop of dilute HCl
develops the red colour of sulphocyanide of iron at once; and
the delicacy of this reaction is so great that, conversely, when it
is required to know if free HCl exists in a liquid, the addition
of this mixed solution will detect it when it amounts to only
sdsath part of the total liquid, and where the acid is only 54th
part the red colour is so dark as to appear almost black by
reflected light; as therefore the amount of HCl in the gastric
juice is ordinarily estimated at ;4jth part, or ‘2 per cent., it is
evident that this method might give valuable results, because
by it an approximate analysis of the gastric juice, in regard to
its free HCl, may be made in less than a minute, whereas for-
merly, by estimating the amount of acid and base separately,
and then calculating the excess of acid, the best part of a day
THE ACIDITY OF GASTRIC JUICE. 275
was spent over one analysis, and, after all, the results were not
so satisfactory.
But it must first be inquired whether any free acids other
than HCl produce this reaction, and also if any substances can
destroy or modify it. I found, then, that dilute Nitric, Sulphuric
and Tartaric acids produced the reaction as well as HCl; that
Carbonic, Lactic, Acetic, Citric, Benzoiec and Uric acids had no
effect on it, while Phosphoric and Oxalic acids not only did not
produce it, but destroyed it when produced. As for the alka-
lies they, of course, destroy the reaction; but as it is impossible
that they should exist in a solution containing free HCl, they
may be dismissed at once. Now, the fact that Tartaric Acid
produces this reaction, while Ferrum Tartaratum does not do
so, seems to shew that what happens is, that the free tartaric
acid unites with the potassium, and liberates the sulphocy-
anogen which in its nascent state attacks the ferric citrate or
potassic tartrate, and forms the sulphocyanide of iron; this
appears also proved by the conduct of HCl, which, when added
to a solution of Ferr. Citr., does not change the colour at all, as
it should do if it formed perchloride of iron; and moreover it
makes no difference to the reaction whether the Sulphocyanide of
Pot. be added first or last; and yet it seems scarcely probable that,
when added before the HCl, the latter should form perchloride of
iron, and then change the iron for potassium, to enable the
sulphocyanide of iron to be formed, I believe it, therefore, to
be most probable that this reaction is caused by the liberation
of nascent sulphocyanogen. In regard to phosphoric and ox-
alice acids, which I have described as destroying this reaction, I
think the word destroy is hardly applicable, but that the re-
action is really produced, as I have satisfied myself by careful
experiment, and then dissolved, probably by the neutral phos-
phate and oxalate of Potassium formed from the Sulph., for, on
adding strong HCl to the solution effected by oxalic acid, the
colour is restored to a great extent.
While, therefore, this method fails in distinguishing free
HCl from free Nitric, Sulphuric and Tartaric Acids, and while
phosphoric and oxalic acids in a free state negative the reaction,
yet, as these acids are easily tested for separately, and elimi-
nated from an inquiry, the value of this method is scarcely
276 DR REOCH.
diminished, more particularly as it holds in the very point in
which it is most required, the distinction between free HCl and
free lactic acid; for while it is not yet absolutely settled to
which acid the gastric juice of man owes its acidity, it is gene-
rally believed that in man it is due to free HCl, and in the dog
to free lactic acid. Now, during my first experiments on this
subject, I failed to find HCl, while I took especiai care to prove
not only the absence of free HCl, but, by adding an equal
quantity of a solution of HCl of the strength ‘2 per cent., and
obtaining the reaction, I proved that there was nothing in the
fluid to prevent the reaction coming out, and that therefore, if
free HCl had been present in anything like the quantity *2 per
cent., it could not fail to be detected. These experiments were,
however, defective in that they were made on mice, cats, and
vomited matter from the human subject; but on establishing a
gastric fistula in a dog I obtained, by many scores of experi-
ments, distinct evidences of free HCl. Before, however, de-
scribing these later researches, I shall refer to some results which
I obtained from the former class of experiments. It appeared
to me at that time that a result so much at variance with com-
mon opinions must be supported by irrefragable evidence, and
therefore I undertook a series of experiments to ascertain if any
foreign substance, such as albumen, &c., might interfere with
the reaction in any way. It was not probable that such was the
case, for 1 have already mentioned that the reaction came out
at once on adding free HCl to the original liquid; but still ex-
periments on that point required to be multiplied. Now, on
adding my mixed solution to urine in a test-tube, I observed
that no reaction took place, while, if two or three drops of dilute
HCl had been previously added, the reaction at once developed
itself. The conclusion appeared, therefore, that there is no free
HCl in urine, and that if there was, it would shew itself; but
on performing the experiment with the greatest care, I found
that the reaction did not come out with urine to the same ex-
tent as with distilled water if the same quantity of HCI had
been added to each. If, for example, six test-tubes be arranged
with 2 cc. urine in each, and six test-tubes with 2 cc. distilled
water, and if 1, 2, 3, 4, 5, and 6 drops of dilute HCl be dropped _
into each series, it will be found that in the first test-tube of the
THE ACIDITY OF GASTRIC JUICE. Qt
urine series there will be no reaction developed on adding the
mixed solution, while the 2nd, 3rd, 4th, 5th and 6th will corre-
spond in depth of reaction more or less with the Ist, 2nd, 3rd,
4th and 5th of the water series, or perhaps the first two of the
urine series will give no reaction, and the others correspond
with the first four of the water series; obviously, therefore, there
is nothing in urine to destroy the reaction, but there is some-
thing which delays it and slightly dissolves it; and this some-
thing varies within certain limits. The only explanation of this
peculiarity seems to me to be that there is something in the
urine which absorbs or combines with HCl, and that thus when
free HCl is added to urine, though the already acid urine is ren-
dered more acid, yet there is no free HCl till a certain point is
reached ; now every one knows that when HCl is added to urine
it precipitates uric acid by assuming to itself the sodium which
the latter is united with; and it is evident, therefore, that when
free HCl is added to urine there is formed chloride of sodium
and uric acid, and to that extent, therefore, free HCl, when added
to urine, will disappear; but it is evident that, the quantity of
uric acid in the urine being only 8 grs. per diem, this is alto-
gether insufficient to account for the disappearance of 1 or 2
drops of dilute HCl in 2cc. urme. I therefore examined care-
fully the relations of the alkaline and acid phosphates of soda
to the sulphocyanide reaction, with the following results :—
When a solution of the ordinary or neutral, or rather alkaline,
phosphate of soda Na, H PO, is added to a mixed solution of
the Ferr. Citr. and Sulph. of Pot., no reaction of sulphocyanide
of iron takes place when HCl is dropped into the mixed solu-
tion until a quantity of HCl is added, exactly sufficient to con-
vert the alkaline into the acid phosphate, and then it comes out
at once, allowing only for a slight solubility of the sulph. in the
acid phosphate.
I shall not trouble the reader with an account of the experi-
ments by which I proved this important conclusion, that Na,
H PO,+ HCl does not remain so, but becomes at once arranged
as Na H, PO,+ Na Cl, because there is a fact known to every
physiologist, which, had it been duly considered, might have
proved this long ago. Neubauer and Vogel in their work on
the urine say that when uric acid is heated with alkaline phos-
278 DR REOCH.
phate it assumes an atom of sodium, becomes urate of soda, and
converts the alkaline into the acid phosphate: now every one
knows that when HCl is added to urine it appropriates an-atom
of sodium from the urate of soda, forms chloride of sodium, and
precipitates the uric acid. If then such a feeble acid as uric
acid can take sodium from the alkaline phosphate and convert
it into acid phosphate, is it not probable (and to this I shall
afterwards refer) that other acids can do the same, and is it not
demonstrably certain that HCl will do so when it can make
uric acid itself give up the sodium which it has taken from the
phosphate? The dilution of the HCl makes no difference, for
though it is added to urine in the concentrated form when uric
acid is wanted, that is only because the urine itself dilutes it so
much that a teaspoonful of strong HCl in twenty ounces of
urine is not any stronger a solution of HCl than the gastric
juice at ‘2 per cent. Moreover, every one knows that you never
get as much uric acid as you add of HCl, simply because, as I
maintain, a large quantity of the HCl is required to convert the
alkaline phosphate which has not already been converted into
acid phosphate. The same reasoning applies to phosphate of
lime. The ordinary phosphate of calcium is Ca, P, O,, and this
is usually said to be insoluble in water, but soluble in dilute
acids; but, according to my experiments, it is not soluble in HCI
except in so far as the latter is added in sufficient quantity to
convert it into acid phosphate, which is then dissolved, and thus
when HCl is added to Ca, P, O, the reaction is Ca, P, O,+
4HCl] = 2Ca Cl,+ Ca H, P, O,, therefore the HCl disappears
until this point is reached, and accordingly I found that it gave
no sulphocyanide reaction till this point, and beyond that it
gave it in an increasing ratio with the amount of HCl added,
excepting only for a very slight solubility of the colouring
matter in the acid phosphate. When these facts are fully con-
sidered it is evident that Schmidt’s results will not hold ina
quantitative analysis of gastric juice; for unless you allow that
any free HCl present in that juice converts the alkaline phos-
phates present into the acid phosphates, you will infallibly
make the quantity of free HCl much too large, and in Schmidt's
analyses the relation of the phosphates to the free HCl is evi-
dent, while no relation of the latter to the ehlorides is apparent.
THE ACIDITY OF GASTRIC JUICE. 279
Human G. J. Sheep’s G, J. Dog's G. J.
HCl 20 1:50 2°70
Chlorides 2°07 5:98 5:87
Phosphates ‘12 2:09 273
To return, however, to my first experiments: I examined more
than a dozen mice, and found that the reaction of the stomach
and its contents was generally acid, and even in some cases
post-mortem digestion of the stomach itself was almost com-
plete, and yet in every case I failed to find HCl, though on
adding a drop of ‘2 per cent. solution to the contents of the
stomach the reaction came out. It appears to me therefore a
very doubtful matter whether these animals have free HCl in
their gastric juice, and probably they have some acid analogous
to that which gives them their peculiar smell. In regard to
cats also I failed to find free HCl. In one cat I made a gastric
fistula; the animal lived for ten days, and I examined its gastric
juice several times without detecting HCl; as however it did
not live long enough to leave its milk-diet perhaps that might
have accounted for the failure. JI examined also several speci-
mens of vomited matter from the human subject; after filtration
the fluid was generally acid, though in one case neutral, and
yet there was no evidence of free HCl; these however might be
regarded as abnormal specimens, and I therefore made a gastric
fistula in a dog*. I was unable to keep a tube in the opening,
as the dog injured himself by constantly trying to get it out,
and moreover it keeps the part in much better condition to
allow the dog to lick the sore. If the opening be not too large
the muscular movements of the stomach and abdomen will not
allow the food to escape, while a director can be passed and the
1 My first dog died in three days, and I am persuaded his death resulted
from the operation being performed in the way generally recommended, by
giving the animal a full meal to distend his stomach prior to the operation,
and thus give facility in seizing it. I did so, and found when I opened the
abdomen that his stomach was highly vascular. I attached it to the wall of
the abdomen by silver sutures, and then opened it, and on his death three days
afterwards I found the coats of the stomach highly inflamed, and almost gan-
grenous. It appeared to me therefore that the danger of exposing the peri-
toneum too much while searching for the stomach when empty, was imaginary
compared with the danger of opening the latter when the coats were highly
vascular, and in the full performance of the digestive process. Accordingly, in
my second operation, I did not feed the animal for some hours previously, and
the result was in every way satisfactory.
280 DR REOCH.
fluid: will flow along the curve in a much better way than by
tickling the inside of the stomach with a feather; for by using
the director you can apply it to any part of the stomach at
once, and thus compare the results. I kept the dog for a week
on milk and bread, and then gradually returned him to a mixed
diet. The result of my experiments was that for the first fort-
night no HCl was procurable from the juice. I did not examine
the juice for a week after the operation to allow the animal
time to recover, but during the next week I examined his juice
frequently, and failed to find HCl; it then began to appear,
and continued more or less during the whole period of my ex-
arnination.
I found, however, that Schmidt’s estimate was much too
high for the free acid in the dog. I never found it to rise
much above *2 per cent. under any circumstances, and I there-
fore conclude that his analyses which gave 2 per cent. did not
take sufficient account of the acidity of the phosphates. It
appears, however, that free HCl is a nearly constant product
in the gastric juice of the dog, varying, however, with the state
of the animal; after long fasting it appears in greater quan-
tity than after a limited fast, and its quantity varies also with
the kind of food supplied. The question therefore arises, to
what is the free HCl due? There can be only three sources
of it, from the decomposition of chloride of sodium, chloride
of potassium, or chloride of calcium. Many chemists say that
it is caused by lactic acid decomposing calcic chloride, but that
this is not so is evident, for calcic chloride exists neither in the
food nor in the blood, which is a result of the food : to say there-
fore that HCl arises from the decomposition of calcie chloride
just removes the difficulty one step, for what causes the decom-
position of sodic or potassic chloride to give rise to calcic chloride
in the stomach? Moreover, according to my views the calcic
ehloride in the gastric juice is easily explained, for it is a result
of the free HCl and the phosphate of lime forming acid phos-
phate and calcic chloride, and, finally, to upset this theory
entirely, it is only necessary to deny that lactic acid can take
calcium from calcic chloride, much less form HCl from sodie
or potassic chloride. I have in my possession a bottle of so-
called lactic acid, which undoubtedly decomposes calcic chloride,
THE ACIDITY OF GASTRIC JUICE. 281
but I have satisfied myself that this is from the presence of
oxalic acid as an impurity, probably from higher oxidation of
lactic acid by defect in the process of preparation, and that this
is not a reaction of lactic acid itself. No doubt, this very fact
that lactic acid may be oxidized to oxalic acid, may account for
the presence of the latter in the urine as a frequent consti-
tuent, but that this is the ordinary method of production of HCl
is wholly improbable; and I may add, from numerous examina-
tions of the gastric juice mieroscopically, that I. never detected
the presence of oxalate of lime. And though oxalic acid were
proved present, yet it cannot decompose sodic or potassic chlo-
ride, to which I believe the presence of free HCl in the gastric
juice is due. Tartaric acid as an agent in the production of
HCl, by precipitating the potassium as tartrate of potash, is
equally objectionable, for there is no evidence of the presence
of tartaric acid in the gastric juice. There is another theory
of the production of free HCl which seems to be more popular
than the lactic acid theory I have referred to, and that attri-
butes the decomposition to electrolysis; but this theory appears
to me even more unsatisfactory than the other, for where is the
source of the electricity? Who has proved its existence? and
if it does exist, why does it decompose chloride of sodium in
particular ? why not decompose every other. salt? The theory
in fact is little better than attributing the origin of HCl toa
vital principle. I have made numerous experiments on the
effect of the introduction of sodic chloride into the stomach,
and uniformly found that it greatly increased the flow of gastric
juice, but did not increase the proportion of free HCl above
"2 per cent. In fact it stimulated but did not alter the secre-
tion. It is evident, therefore, that the sodic chloride introduced
into the stomach is not decomposed in it by electrolysis, or at
least this conclusion is nearly certain, for though I introduced
the salt not into the blood, but into the cavity of the stomach,
yet it must have passed by diffusion into the blood supplying
the glands of the latter; and as I examined the fluid coming
from the same part of the stomach, the quantity of HCl should
have been increased. The only objection to these experiments
is that while adding the electrolyte, I did not add additional
electromotive force to decompose it, but still, that being im-
VOL. VIII. 19
282 DR REOCH.
possible, I yet think it forms a strong argument against the
theory of electrolysis.
It appears to me that the true theory will best be found by
recurring to elementary chemical facts. Albumen contains
sulphur, and this is without doubt oxidized in the blood into
sulphuric acid, for the nitrogen, &c. of albumen being removed
as urea and uric acid, the sulphur is acknowledged by all to be
removed by oxidation, conversion into sulphates, and excretion
with the urine. Now sulphuric acid is almost the only acid
which is able to decompose sodic chloride; and it appears to me
therefore highly probable that the oxidation of the sulphur of
the albumen takes place in the walls of the stomach during the
production of its peculiar secretion, and that this in its nascent
state seizes on the sodium of sodic chloride, and allows the free
HCl to be eliminated in the gastric juice. It is no argument
against this theory that sulphates are not found in the juice,
for if the decomposition took place in the walls of the stomach,
there is no more reason why sulphates should appear in that
secretion than why sodic phosphate, which exists so largely in
the blood, should appear in it also. This theory then fully
accounts for the free HCl in the gastric juice, and the latter
fully accounts for the presence of calcic chloride, as I have
already explained*. In regard to the phosphate of lime so con-
stantly present in the gastric juice, I do not think that it isa
secretory product of the gastric glands themselves, unless in the
upper part where they are lined by columnar epithelium, but
rather that it is a product of those other glands which secrete
the mucus of the stomach. Phosphate of lime is a very constant
1 As a possible objection to my theory that there is not enough H,SO,
in the urine to account for all the HCl in the gastric juice, I may reply, that
allowing with Parkes the H,SO, in urine to be 31-11 grs., and only one-third of
this to be derived from the food, then since 3—4 grs. of oxidized sulphur are
excreted by the urine independently of sulphuric acid, it is evident that even
supposing the sulphates produced in the stomach not to be again decomposed,
there will be S oxidized equal to about 32 grs. of H,SO,, and this would be
equal to 12 grs. of HCl, or to 14 ounces of -2 per cent. solution of HCl. Now
though the gastric juice secreted is 10 or 20 times this amount, yet it must be
remembered that it is only after long fasting that you can get even ‘2 per cent.
solution of HCl; ordinarily, as I have remarked, in analyzing vomited matters
the acid chyme shews no trace of HCl, which is only secreted for the purpose of
digesting flesh-meat, or something similar, and therefore it is quite a mistake to
estimate the gastric juice as if each drop of it contained ‘2 per cent. HCl. Eyen
in the dog this is rarely the case, for it is not found while giving the animal
milk, and little or none when giving it bread.
THE ACIDITY OF GASTRIC JUICE. 283
constituent of mucus, and I think therefore that this is its
origin in the gastric juice, though no doubt it is largely in-
creased from the peculiarly irritating nature of the gastric
juice which contains so much free acid. But is not lactic acid
present in the gastric juice of the dog? I believe it 1s. Ber-
nard has found it to be so, and I have found in evaporating the
filtered juice crystals analogous to those which Scherer describes
as those of lactate of lime; but the very fact that I obtained
these without adding any calcic carbonate to the solution, shews
that lactic acid possesses the property in regard to phosphate
of lime which uric acid is known to have with regard to phos-
phate of soda. It forms lactate of lime and acid phosphate;
and this appears to me to be the true function of lactic acid in
the gastric juice, and also one reason why starchy matter
such as potatoes is so agreeable a mouthful along with flesh-
meat, for the starch is converted by the saliva into glucose,
and that by the secretion or ferment of the stomach is converted
into lactic acid, and thus the alkaline phosphates are converted
into acid phosphates without using up HCl, which is so valuable
in digestion, and the production of which must task the stomach
so highly. But into this subject I will not enter, as experiment
is here mixed with theory, and will therefore pass on to remark
that numerous experiments which J made shew that the cardiac
and not the pyloric end of the stomach is that in which the
greatest secretion of HCl goes on; in the latter part, indeed, I
scarcely found any HCl at all, but that might possibly be due
to the fact that the fistula was near the pylorus, and therefore
its presence and some slight inflammatory reaction might have
vitiated results. I will add further, that the introduction of
sapid substances stimulated the secretion of the juice much
more than mere mechanical irritation, and that in such cases
the amount of HCl was almost always kept up to nearly ‘2 per
cent. along with the increased amount of juice, shewing evi-
dently that true stimulation of the stomach is not effected by
tickling its inside, but by the substance entering the walls of
the stomach by diffusion and stimulating its glands, or perhaps
the ultimate filaments of the vagus which control their secretion.
The relation of these facts to the important question of why
the stomach does not digest itself is evident. It is not simply
19—2
284 DR REOCH. THE ACIDITY OF GASTRIC JUICE.
from the alkalinity of the blood, but from the fact that doubt-
less the nervous system possesses a regulating power over the
production of HCl, and that even if the latter were formed in
greater excess, its force would be rapidly utilized by union with
the alkali of the alkaline phosphate. What chance could HCl
have to dissolve a stomach in whose walls alkaline phosphate of
soda was constantly circulating? If it attempted to do so it
would form simply sodic chloride and acid phosphate, and these
would immediately be swept on by the current of blood and
excreted by the urine. Under ordinary circumstances, indeed,
there is no necessity for this, as the acid phosphate of lime
intervening between the gastric juice and the proper coats of
the stomach will form a sufficient protection ; but should this
balance be destroyed there would still not be digestion of the
stomach itself, for the reason I have indicated.
Notre. In the foregoing paper I have constantly spoken of the free HCl in
the gastric juice as ‘2 per cent. Properly speaking, I should have said ‘02 per
cent., as thatis the proportion found by Schmidt, but though the ordinary books
nominally give °02 per cent., they yet speak of -2 per cent. in relating experi-
ments on artificial gastric juice, and admit that no less proportion than °2 or ‘1
per cent. will digest fibrin. I therefore assumed that Schmidt’s results were
wrongly referred to 1000 parts instead of 100, as I had no access to his original
paper. If however it be correct to refer them to 1000, then it seems clear that
the theory of the origin of the free HCl in the gastric juice which I have pro-
posed by the production of H,SO, from oxidized sulphur will fully account for
the free HCl of 7 pints of gastric juice at ‘02 per cent.
ADDITIONAL OBSERVATIONS ON THE ANATOMY
OF THE GREENLAND SHARK (LHZMARGUS
BOREALIS). By PRoresson TURNER.
In June, 1873, I published in this Journal some observations
on the visceral anatomy of the Greenland Shark, based upon
a dissection of two well-grown females, captured in the preced-
ing February off the Bell Rock. Early in March of the present
year a young male was hooked on a deep-sea-line off the Isle
of May at the mouth of the Firth of Forth. I purchased the
specimen from Mr Anderson the fishmonger, and am now able
to render my description of the visceral anatomy of this shark
more complete, not only by giving an account of the male
sexual organs, but in some other particulars.
As the published figures of this shark by Scoresby, Yarrell
and Couch, are, in my opinion, unsatisfactory, I requested
Mr C. Berjeau, whose skill as a draughtsman is so well known
to anatomists, to prepare a drawing of the fish, which he has
reproduced on wood to illustrate this paper. The figure is on
the scale of one inch to the foot, and has been drawn under my
supervision. The colour of the skin was, as in the former
specimens, bluish-gray, but the sides were not, as in those fish,
barred with faint stripes.
Spiracle placed behind and above the eye, mouth on the
ventral surface immediately below the eye. The base of the
1st dorsal fin was a short distance in front of a point midway
between the tip of the snout and the tip of the tail. The centre
of the base of the 1st dorsal fin was about the mid-distance
between the anterior borders of the pectoral and ventral fins.
The 2nd dorsal fin was immediately posterior to a line drawn
vertically from the cloaca. The 2nd dorsal fin was not only
smaller but more pointed at its upper and posterior angle than
the Ist dorsal.
In my former specimens, owing to the liver having broken
away by its great weight, I was not able to determine if a
286 PROFESSOR TURNER.
gall-bladder existed. In this specimen, which, on account of
its smaller size, was much more manageable to dissect, I found
a gall-bladder, about twice as large as the corresponding organ
in man. It was lodged in a fossa in the liver, and its surface
of attachment was overlapped by liver substance. Though it
projected from the liver in a well-marked manner, it was not
pyriform in shape; its sides were flattened, and it had no
neck. Its duct lay for some distance in contact with that part
of its wall which was next the liver-substance, and opened into
the bladder by a wide orifice. Lying in the fold of peritoneum,
which enclosed the bile-duct on its way to the duodenum, was
an oval body about the size of a small almond. It had a red-
dish colour, and possessed the general appearance of a lymphatic
gland.
The dimensions were as follows:
From tip of snout to iip%of axis of tail <<... ..or<ssseeee Oil
to back of base of 1st dorsal fin... 2 114
to back of base of 2nd dorsal fin... 4 7
to anterior border of root of pec-
Gopal fi “westse .ssccceccoss sansa oe
to anterior border of root of ven-
Brel Tania eoe cee tens 2 cie05 sea eee 3 10
Lo Spiraclet./...-0lr ase -oseseeoe eee 84
LO CVO secaes Ger awe cosehcy 2-0 ne eee 54
TO TOWED Saose ss cc sae see onc seat 6
See aw, vane... DOM MOMDIM ES eth cas ocean ass seen 24
From cloaca to tip of axis of tail .......0....--c...000e0e 1. ‘94
Between upper and lower tips of tail .................. Eig iat
Antero-post. diameter of base of Ist dorsal fin ...... 4h
(EUS VS TMG) Geabhaatbrbanenn ay Adapbormtissci scm uo67 3t
Antero-post. diameter of base of 2nd dorsal fin ...... 4
Pets e Ole). .0c sete cae actncs oecasee eee 24
Antero-post. diameter of pectoral fin .................. 8
SOL Wembrealuliineen sestsemmnasre cs -/ae 64
An injecting pipe was introduced into the conus arteriosus
and some coloured gelatine was thrown into the branchial
vessels. From the sides of the aorta three pairs of branchial
ANATOMY OF THE GREENLAND SHARK, 287
ne
Ith
Wide
A specimen of the Lerneopoda
ingall,
ipl
iV
rH
inch to a foot,
), scale 1
ight cornea. Engraved by Mr W, Ball
4s
elongata is attached to the r
Profile yiew of a young male Greenland Shark (Lemargus boreal
288 PROFESSOR TURNER. =
arteries arose, and at its anterior end, immediately behind the
basi-hyal, the aorta ended by bifurcating into a terminal pair,
each of which, after a course of about an inch, again bifurcated,
so that: five pairs of branchial arteries resulted. The first or
most anterior pair extended outwards along the line of origin
of the 1st gill from the hyoid cartilage; the 2nd, 3rd, 4th and
5th pairs ran outwards to their respective gill-arches; each
artery lay on the distal side (i.e. on the side furthest removed
fromthe heart) of the cartilaginous styles which supported the
more posterior of the two gill-lamine attached to a given
branchial arch. A well-defined layer of striped muscle inter-
vened between the trunk of the branchial artery and the
more anterior of the two gill-lamine attached to a branchial
arch. The 1st branchial vein at the dorsal angle of the gill-
cleft bifurcated, one branch ran forward in a groove on the
under surface of the palate, entered a foramen, and assisted in
the supply of blood to the head, the other branch joined the
more anterior of the two veins proceeding from the gill-lamina
lying behind the 1st branchial cleft, and formed the first or
most anterior of the roots of the dorsal aorta. The 2nd, 3rd
and 4th roots of the aorta were formed in a similar manner by
the junction at the dorsal angle of the gill-cleft of a vein from
the gill-lamina in front of the cleft with a vein from the lamina
behind. In addition to the transverse branches which arose
from the dorsal aorta in this region, it gave off two branches at
its anterior end, which, diverging from each other, entered fora-
mina in the occipital cartilage.
The testicles were two narrow elongated glands situated
at the anterior part of the abdominal cavity. Each gland was
connected by a mes-orchium to the ventral surface of the cor-
responding kidney ; the two testicles were separated from each
other by the meso-gastric fold of the peritoneum. Each testicle
was between 10 and 11 inches in length, 3ths broad, and 4th
inch in thickness. It terminated in a narrow pointed extremity
at each end. The surface of the testicle was quite smooth, but
a narrew band extended along that border which was opposite
the line of attachment of the mes-orchium; sections through
this band were made, but no trace of a tube was seen within it.
From the immature growth of the fish it is probable that the
ANATOMY OF THE GREENLAND SHARK. 289
testicles were not sexually ripe. I carefully examined both the
surface of each gland and its mes-orchial fold for an epididymis
and a vas deferens. No trace of an excretory duct could be
seen. The mes-orchium was so translucent that if a duct had
been present I could not but have seen one; nothing indeed
was visible between its two layers but a few thin-walled vessels
and a little fat. When the mes-orchium was dissected off the
kidney a large, long blood-vessel was opened into, the canal of
which was subdivided by slender bands. _
In the next place I carefully examined the cloaca with the
view of determining the number and relation of its openings.
The rectum opened by a large orifice into its anterior part. At
the bottom of a shallow fossa, immediately behind this orifice,
was a small papilla, at the summit of which a minute opening
was seen. Into this opening I introduced the fine nozzle of an
injecting syringe and forced a coloured fluid into it. This fluid
ran freely into two long ducts, one situated on the ventral
surface of each kidney, and from the ducts into the kidney
substance, but no injection passed into the testicles. The con-
clusion I came to therefore was, that the minute orifice at the
summit of the papilla was common to the two ureters. In this
respect the male differs from the female fish previously de-
scribed, in which each ureter opened independently into the
back wall of the cloaca. Where the cloaca approached the
surface of the integument two transversely elongated abdo-
minal pores were situated, which freely communicated with
the general peritoneal cavity. From this dissection, therefore,
it would appear that in the male, as in the female Lemargus
borealis, the genital ducts are not developed, so that the
products of both the male and female genital glands are shed.
directly into the general peritoneal cavity. As I discussed
the importance of this arrangement in connection with the
classification of this fish in my previous paper, I need not
further refer to it on this occasion.
A cleft 14inch in length was situated at the posterior border
of the ventral fin. It separated the innermost style-shaped ap-
pendage of the basal cartilage of the fin from the other append-
ages of that structure. The innermost appendage, which was
only 123 ths inch long, formed with its tegumentary covering a
290 PROF. TURNER. ANATOMY OF THE GREENLAND SHARK.
short rudimentary clasper. A longitudinal groove was situated
on the dorsal surface of the clasper, which passed into a deep
cleft lined by a prolongation of soft, smooth integument, No
gland was found at the root of the clasper.
As an illustration of the omnivorous character of this shark,
I may state that its stomach contained a large haddock, the
remains of several herring, a portion of a cuttle-fish, the shell
of a dog-whelk containing a hermit crab, a partially digested
biscuit, and a boy’s playing marble.
ON THE USE OF THE LIGAMENTUM TERES OF THE
HIP-JOINT. By W.S. Savory, F.R.S., Surgeon and
Lecturer on Surgery, St Bartholomew's Hospital, late Pro-
fessor of Comp. Anat. & Physiol., R.C.S.E.
Wuaat is the function of the Ligamentum Teres? Many authors
who describe this structure are silent on the question, while of
those who answer it the general conclusion is that it has, for its
chief function, to limit adduction of the thigh, or when the
thigh is fixed, to limit lateral movement of the pelvis on the
femur—to prevent the pelvis from rolling toward the opposite
side. Authors, of course, are not fully agreed in explanation
of its use, and other less prominent functions are by many
assigned to it, such as to limit rotation of the thigh; but the
conclusion substantially arrived at is the one given above.
I cannot, and need not, here quote from the several authors
who are entitled to speak with authority on the subject, but
neither in them nor elsewhere can I find any allusion to what
appears to me to be the prime purpose of this ligament’. Its
strength is very great; its attachments are remarkable; its situ-
ation is peculiar. I submit the following explanation of its use.
When the person is erect the ligament is vertical and tight.
This statement, although generally accepted, has been chal-
lenged. I am satisfied of its accuracy. By removing the bottom
of the acetabulum from the pelvis with a trephine the state of
1 Since this paper has been in type I have learnt that Prof. Partridge in his lec-
tures on anatomy at King’s College was accustomed to compare the Ligamentum
Teres, in its function, to the leathern straps by which the body of a carriage is
suspended on © springs; and my attention has been called to the following
passages published by Prof. Turner in 1857 (Hwman Anatomy and Physiology,
Edinburgh): ‘In the interior of the joint (hip) is a strong band of fibres called
the inter-articular or suspensory ligament. When a person is standing erect or
with the body slightly bent, a portion of the weight of the trunk is borne directly
by the heads of both thigh-bones, or of one thigh-bone, according as he stands
upon one or both legs, owing to the direct pressure of the acetabulum upon the
heads of those bones. Now as the end of this ligament, which is connected to
the lower margin of the acetabulum, is much lower than the end connected to
the thigh-bone, it of necessity suspends that portion of the weight of the body
which is thrown upon it. The effect of this is to distribute over the head of
the thigh-bone that weight which, supposing the suspensory ligament had not
been present, would have been sustained by that portion merely which is in
direct contact with the upper part of the acetabulum.” (p. 42.)
292 PROFESSOR SAVORY.
the ligament may be demonstrated. But I think the discre-
pancy of observation is due to the fact that the degree of tension
of the ligament is dependent on the line of direction of the
femur. The ligament is moderately tight when a person stands
evenly upon both legs. It is tighter when the femur is slightly
flexed as it more usually is. But when resting upon one leg,
inasmuch as the pelvis is then raised on that side, which of
course affects the ligament in the same way as adduction of the
femur would do, then the hgament becomes extremely tense.
In other words, it becomes tightest when the hip-joint has to
sustain the greatest weight.
When therefore the pelvis is borne down upon the femur,
or when the femur is forced upwards—that is, when the pressure
would be greatest between the upper part of the acetabulum
and the opposite surface of the head of the femur—it is put
directly on the stretch. More precisely, its great purpose is to
prevent undue pressure between the upper portion of the aceta-
bulum, just within the margin, and the corresponding part of
the head of the femur. But for this ligament such undue pres-
sure must inevitably occur. Suppose the Ligamentum Teres
absent, and the person standing upright : owing to the obliquity
of the acetabulum and the head of the femur—of the axis of the
joint—pressure between the two could not be equally, or nearly
equally, diffused over their opposing surfaces, but it would be
concentrated on a spot in the upper part of the socket through.
which a line drawn down the body, through the joint into the
leg, would pass. When the thigh is straight, when the femur is
in a line with the body, as when one stands upright, then is the
Ligamentum Teres in the same line too, and consequently any
force which drives the femur and pelvis together must tell at
once upon the ligament, and be directly checked by it.
Owing, therefore, to the shape and obliquity of the hip-joint,
and the weight of the body, the Ligamentum Teres is neces-
sary to prevent concentration of pressure at a particular point
above it.
The line through which the weight or force acts between the
upper part of the acetabulum and the opposed surface of the
head of the femur forms, with the line of weight or force which
passes through the Ligamentum Teres, an obtuse angle; and the
USE OF THE LIGAMENTUM TERES OF THE HIP-JOINT. 293
resultant of these forces is in a line which passes through the
long axis of the head of the femur.
When the person is erect the body partly hangs upon the
Ligamentum Teres.
I submit that this is the prime function of the Ligamentum
Teres. Other purposes I do not deny, but would maintain that
they only occasionally come into play, and are altogether sub-
ordinate to this one, which is especially called into action when-
ever the weight of the body is thrown upon one leg.
Now this view may be tested by the facts of comparative
anatomy.
It has often been remarked that the Ligamentum Teres is
apparently distributed among animals in a very arbitrary
manner.
In most of the mammalia it is present, e.g. in ruminants,
rodents, and terrestrial carnivora. In many other absent, e.g.
in the elephant, sloth, seal, walrus, sea-otter, ornithorhynchus,
and echidna.
It exists in animals with the utmost diversity of form and
habits.
_ It is sometimes present in one animal, e.g. the chimpanzee,
and absent in another very closely related to it, e.g. the ourang-
outang’.
Now is it possible to discern the conditions under which it
is present or absent ?
When the cavity of the acetabulum looks downward and the
head of the femur upward, in other words, when the direction
of the hip-joint is nearly vertical, and the weight of the body
falls through the centre of the joint, then the Ligamentum Teres
is absent, e.g. elephant.
When the acetabulum looks outward and the head of the
femur is inclined inward, in other words, when the hip-joint
is placed obliquely, so that there would otherwise be undue
1 There is great difference in the degree to which the Ligamentum Teres is
developed in Birds. In some it hardly appears, while in many it is very strong.
The great depth of the groove in the head of the femur of the Ostrich shews the
size it occasionally attains. In several birds in which I have dissected this
ligament I have always found its pelvic attachment to be, not to the border of
the acetabulum, but to the lower margin of the large foramen or foramina which
exist at the bottom; this, so far as its action is concerned, comes to the same
thing.
294 PROFESSOR SAVORY.
pressure at a particular part, then the Ligamentum Teres is
present, e.g. horse.
The exceptions to this occur in those animals in whom, al-
though it is an instrument of progression, the posterior ex-
tremity does but little in supporting the weight of the body,
e.g. seals, and the ourang-outang. :
These facts, that while the Ligamentum Teres is found in
the chimpanzee and other monkeys, it is almost or entirely
wanting in the ourang-outang, at first sight apparently so
capricious, are very suggestive. It is easy, I think, to under-
stand why it is generally present in monkeys, inasmuch as
in them the hip is placed obliquely, and the posterior extremity
can support the trunk. But the hip-joint is oblique also in the
ourang-outang. The conformation of the foot, however, is the
key to the explanation of its absence here. It is clear that in
the ourang-outang the posterior extremity cannot be such an
instrument of support to the trunk raised upon it, as in the
chimpanzee, and consequently the Ligamentum Teres is not
needed to counteract undue pressure at a particular point.
Again, it may be said that when an animal stands, in pro-
portion as the long axis of the head of the femur approaches
to a vertical line, so does the Ligamentum Teres become weak
until it disappears. On the contrary, it is strongest where the
head of the femur has a direction farthest from the vertical,
and has to support the greatest weight.
In conclusion, I should like to call attention, without at-
tempting to lay too much stress on it, to a specimen (2. 43) in
the pathological series of the museum of St Bartholomew’s
Hospital, which is thus described in the catalogue.
“Two hip-joints from the same person. In each joint the
Ligamentum Teres is completely wanting. The capsule of each
is perfect and exhibited no appearance of disease. In the usual
situation of the attachment of the Ligamentum Teres there is a
deep depression in the head of the femur, and just above this
the cartilage of each femur is slightly absorbed.”
It may be observed that the cartilaginous shell on the head’
of the femur is naturally thickest on the upper and inner
aspect.
USE OF THE LIGAMENTUM TERES OF THE HIP-JOINT. 295
The above Paper was read at a Meeting of the Cambridge Philo-
sophical Society, in April, 1874.
In discussions which followed Professor Humphry, after express-
ing his obligation to Mr Savory for affording the opportunity of
discussing the subject with him, observed that the suggestions made
with reference to the function of the ligament by Mr Savory rested
entirely upon the view that the ligament is tight in the erect posture.
Professor Humphry was one of those who had challenged this view,
of the accuracy of which Mr Savory had expressed himself to be
satisfied. He referred to his work On the Human Skeleton including
the Joints, in which he had stated as the result of careful observation
that the ligament is not tense and cannot be rendered tense in the
erect posture. He had lately reconsidered the question and re-ex-
amined the specimens, or some of them, upon which his statement
had been based, as well as other recent specimens made for the
purpose, and he was convinced that it was correct. In the first
place, the dimple in the head of the femur for the ligament is more or
less oblong or pear-shaped, and is directed from above downwards and
backwards with such obliquity that the ligament can lie in it, as it
must do when it is in astate of tension, only in the semiflexed position
of the hip, the thigh being inclined from the vertical to an angle of
about 45°. This can be seen in the dry bone, and still better in recent
specimens in which the direction of the insertion of the fibres of the
ligament are seen to correspond with this view. Secondly, the tre-
phine hole through the bottom of the acetabulum shews clearly that
it is at about this angle only, and when the thigh is adducted, that
the ligament is really tense. In the erect posture, and by the erect
posture he meant when the thigh descends vertically from the pelvis
and the capsular ligament, more particularly the anterior part of it,
is tight, neither adduction, nor rotation, nor any other movement
will throw it into a state of full tension. If this is so, which the
several specimens examined by the Professor proved to be the case,
then it is quite certain that the body cannot hang upon the Ligamen-
tum Teres when the person is erect, and the inferences based upon such
a view fall to the ground.
When resting upon one leg the body is tilted a little over to that
side so as to throw the line of gravity more directly over that limb,
the opposite side of the pelvis is slightly raised, the movement being
equivalent to that of abduction of the limb wpon which the weight is
borne, and the Ligamentum Teres is not stretched, but is still more
relaxed than in the erect posture. Even in the position of ‘stand at
ease,’ when the weight is borne upon one limb and the opposite side
of the pelvis is lowered, the other limb being placed upon the ground
slightly flexed, the movement now being equivalent to that of ad-
duction of the weight-bearing limb, though the Ligamentum Teres is
less relaxed than in the former position, and is also less relaxed than
in the erect posture, still it is not tight; and the body is slung, not
296 USE OF THE LIGAMENTUM TERES OF THE HIP-JOINT.
upon the Ligamentum Teres, but upon the thick and strongly resisting
upper portion of the anterior ligament of the hip. The use of the
ligament the Professor believed to be, as he had stated in his work, to
assist in bearing weight when the limb is placed upon the ground
partially flexed and adducted, when the capsule of the hip is compa-
ratively relaxed, and when, if the body be overweighted, dislocation
is most likely to oceur. In estimating, however, its value, even in
this position, it must not be forgotten that several instances have
occurred, some of which are noted in Meckel’s Archiv, vi. 341, in
which the ligament was wanting without its being known that any
inconvenience had resulted from its absence. In dislocation, too, it
must be severed, and it is highly improbable that it ever unites. It
has been found indeed ununited. Still the loss of it does not appear
to be much felt.
With regard to other mammals the ligament as stated in the paper
is commonly absent when the lower limbs do not bear much weight,
and also when they descend vertically from the pelvis. The Pro-
fessor had, however, pointed out in the Journal of Anatomy, Vol. 11.
p. 312, that it is present in the Bats. In most mammals in which it
exists the dimple or furrow or angular depression which it occupies
in the head of the femur is oblique, as in Man, indicating its tension
in them, as in him, to occur in the semiflexed position of the joint.
There were other points to which the Professor took exception,
but the important one was this of the position of the jot in which
the tension of the ligament takes place.
Mr Savory, in reply, remarked that he quite agreed with Pro-
fessor Humphry that, if he were wrong as to the assumption of the
tension of the ligament in the erect posture, his view fell to the ground,
but he could not agree with the Professor as to what really is the
erect posture. The skeletons In museums are commonly articu-
lated wrong, and give too much inclination to the pelvis, and he
thought Professor Humphry was in error on this point, and that if
the ligament be examined in the strictly erect posture it will be
found tight, or more nearly so than the Professor admitted. He
added that by applying his view he had generally been able to judge
from the direction of the limbs in well-articulated specimens of
animals whether the ligament had been present during life or not.
Still there were some exceptions, among the most notable of which
was the difference between the ostrich and the emeu. In the former
it is large, whereas in the emeu it is absent. Yet, though he had
visited the latter animal in the Zoological Gardens, and examined its
posture and movements with reference to this question, he had been
unable to make out why it should thus differ from the ostrich.
FURTHER EXAMPLES OF VARIATIONS IN THE
ARRANGEMENT OF THE NERVES OF THE
HUMAN BODY. By Pror. TURNER.
SINCE my last paper in this Journal on variations in the
arrangement of the nerves (November, 1871) some additional
examples have come under my observation, and that of some
of my pupils.
Fourth cranial nerve-——In an adult male, Mr W. J. Dodds
observed in the right orbit a well-marked branch to arise from
this nerve close to the superior oblique muscle, which branch
ran forward parallel to that muscle as far as the upper border of
the orbit, where it branched, the branches entering the orbicu-
laris palpebrarum. ‘This muscle was then dissected under
spirit with needles, and the finer subdivisions of the nerve were
traced into the fasciculi of the muscle, at the inner part of the
upper eyelid. In the opposite orbit the fourth nerve was
normal.
In an adult female subject, Mr H. S. Stone dissected in the
left orbit a small branch arising from the fourth nerve imme-
diately after it had passed through the sphenoidal fissure. It
ran forwards, clos¢ to the inner side of the superior oblique,
-and about three-fourths of an inch from the upper orbital
border formed a plexus with the infra-trochlear branch of the
nasal: from this plexus branches arose, those most in line with
the branch of the fourth passed to the orbicularis palpebrarum
external to the trochlea, and immediately beneath the orbital
arch, those most in line with the infra-trochlear were traced to
the mucous membrane of the upper eye-lid. The nerve on the
opposite side was normal. Murray, as quoted by Henle, had
also seen one case in which a branch of the fourth communi-
cated with the infra-trochlear. In another adult female,
Mr Stone traced in the left orbit a branch from the fourth
nerve, which ran forward parallel to the outer side of the su-
perior oblique, and near the upper orbital border broke up into
filaments which entered the periosteum ef the anterior part of
VOL. VIII. 20
.
798): PROFESSOR TURNER.
the roof of the orbit immediately to the outer side of the
trochlea.
Cervical Plecus—In my former paper in this Journal, I
recorded a case in which the middle of the three supra-clavi-
cular nerves passed through a hole in the left clavicle in its
course to the integument covering the pectoralis major. I have
since then seen two other cases, one on the left, the other on
the right side, the hole in each case being about the middle
of the clavicle. ‘This variation is obviously more common than
is usually supposed. Henle mentions that Bock, Gruber,
Luschka, Clason and Cruveilhier, have all recorded cases. The
formation of bone around this nerve as it crosses the clavicle
is due to the shaft of the clavicle being developed out of
membrane, and in all probability to the extension of the ossifie
process into the fibrous tissue, which surrounds this descending
branch of the cervical plexus.
Brachial Plerus.—Myr Stone has especially examined, during
the past winter, the arrangement of the nerve to the subclavius.
Commonly it gave a branch to the phrenic; most frequently,
a small branch ran almost transversely to join the phrenic
before that nerve entered the thorax. In one case, the nerve
to the subclavius gave origin to a long branch which entered
the cavity of the thorax a little in front of the phrenic and
nearer the mesial plane ; it descended in front of the arch of the |
aorta and root of the left lung, along the side of the pericardium,
and joined the phrenic immediately above the diaphragm. In
another case, two branches arose from the nerve to the subclavius,
bath ef which entered the thorax, and joined the phrenic before
it crossed the arch of the aorta. These cases examined by
Mr Stone, conjoined with those previously described by my-
self and by Mr Cunningham’, sufficiently establish that ac-
cessory roots to the phrenic nerve are by no means uncommon.
In another case the nerve to the subclavius gave off a branch
which joined the external anterior thoracic, and a second branch
which proceeded to the clavicular part of the sterno-mastoid
muscle.
1 This Journal, November 1871 and Noyember 1872.
ARRANGEMENT OF THE NERVES OF THE HUMAN BODY. 299
In the left upper arm of a subject, used in illustration of
my lectures on the nervous system, I saw the ulnar nerve give
origin to a slender branch about one inch below the tendon of
the latissimus dorsi, which after a course downwards of about
two inches joined a branch of the internal cutaneous nerve, and
with it was distributed to the skin of the inner side of the
upper arm, immediately above the elbow. In another subject,
the internal cutaneous nerve for the back of the left fore-arm
arose from the ulnar in the lower part of the axilla: about
one inch above the elbow it received a very slender branch of
communication from the internal cutaneous nerve. In the
right arm of another subject, the digital nerve for the ulnar
side of the ring-finger arose about the middle of the fore-arm
from the ulnar nerve and passed as a long slender branch in
front of the annular ligament to its distribution.
Sacral Plexus. In the left leg of a male subject the mus-
culo-cutaneous nerve gave off in the substance of the peroneus
longus a long slender branch which descended in the substance
of that muscle to about the lower third of the leg, then pierced
the muscle, but continued under the fascia as far as the external
malleolus, where it became superficial, and joined the external
saphenous nerve.
20—
bo
THE SYNTHESIS OF MOTION, VITAL AND PHY-
SICAL. By C. B. Rapcuirre, M.D.
INTRODUCTORY REMARKS.
More than five and twenty years ago my faith in all that
I had been taught to believe about vital motion received a
rude shock in this way. I had been watching the phenomena
of fatal tetanus in a rabbit poisoned by strychnia: I saw the
animal die: I expected to see its muscles relax at this time:
I was unable to find the slightest evidence of such relaxation
until the end of the fourth day after death, when putrefaction
had evidently set in. Over and over again, by feeling them
with my fingers, did I satisfy myself that the muscles remained
rigid all this while; and I had also additional evidence to the
same effect, in the fact that in dying the animal did not fall
down from a position in which it had been held up by the
spasms during life, and from which it must have fallen, as it
fell finally when the muscles were unstrung by putrefaction, if
death had been attended by muscular relaxation. Indeed, it
was this latter fact which arrested my attention in the first
instance, and made me use my fingers as I have said. For
until the muscular texture gave way in actual decomposition, it
so happened that the animal remained in the position in which
it had been kept by the spasm during life, that is, half-standing
on its hind legs, and half-leaning against the side of a box,
with its forepaws pointing directly upwards, and with its neck
and body bent backwards until the head almost rested upon
the scut—a position which could not have been preserved for a
single moment if the body had not been propped up by the
box as well as rigid. And how was this? Could it be that
the spasmodic rigidity which existed before death had passed
without any interval of relaxation into the cadaveric ngidity
which always comes on, sooner or later, after death, and which
is only relaxed by the actual decomposition of the muscular
tissue? Could it be that spasm had passed directly into rigor
mortis? All my prejudices were against such a notion, and
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL, 301
yet I could not bring my mind to think otherwise. Indeed, the
more I tried to do so, the more I felt constrained to assent:
and, in short, almost before I knew what I was about, I
came to believe that a radical change was necessary in the
doctrine of vital motion—that the interpretation of spasm was
to be sought, not on the side of life, but on that of death, even
in rigor mortis—that spasm and rigor mortis were to be
regarded, not as signs of vital action in certain vital proper-
ties of contractility, but as physical phenomena akin to, if not
identical with, the return of an elastic body from a previous
state of extension—that muscular contraction in all its forms
might be the simple consequence of the operation of the natural
attractive force or forces inherent in the physical constitution of
the muscular molecules—that life is concerned in antagonizing
contraction rather than in causing it—that this antagonizing
influence itself might have a physical basis—that, in short, a
true doctrine of vital motion involved an actual synthesis of
vital and physical motion.
And yet more did this conviction grow in strength on the
food supplied by two other facts to which my attention was
called not long afterwards.
Of these two facts the first was brought to light in an
epileptic patient in whom it had been thought expedient to
try and cut short a succession of most violent convulsions by
taking blood from the temporal artery. The artery was divided
when the fit was at its height, and the blood escaped by jets in
the usual way, but not of the usual colour. Instead of being
red, the blood was black: that is to say, instead of being
arterial, it was venous. The state during the convulsion was
evidently that of suffocation: and on this account, black, unaé-
rated blood had found its way into the arteries, and was
being driven through them at the time. The case was intelligible
enough as regards the suffocation, for in this state the simple
fact is, that black blood does for a time penetrate into and pass
along the arteries; but it was not intelligible as regards
convulsion, if convulsion was, as it is assumed to be, a sign of
exalted vital action. I could connect such exaltation with
increased supply of red blood to certain nerve-centres, but not
with the utterly contrary state of things involved in the actual
302 DR RADCLIFFE.
circulation of black blood; and, do what I would, I could see no
other conclusion open to me than that which had been already
forced upon me by the history of the poisoned rabbit, namely
this, that the convulsion pointed to a state of things which was
the very reverse of vital stimulation, even devitalization—that,
in short, the state of muscular contraction was due, not to the
black blood having acted as a stimulus, but to the simple with-
drawal of an inhibitory action which had served to keep up the
state of muscular relaxation so long as the system was duly
supplied with red blood.
And so likewise with the second of the two facts to which I
have alluded. I had the good fortune to be present on one
occasion when Matteucci was experimenting with strychnia
upon the electric ray of the Mediterranean, and to hear some of
the reasons advanced by this excellent physiologist for suppos-
ing that there was an intimate connection between the action of
the electric organ and muscular action: and, being very much
struck with the fact, that the discharge of the electric organ
caused by the poison was always accompanied by spasm, I
could not help but wonder whether muscular contraction might
not agree with the action of the electric organ m being accom-
panied by discharge—whether muscular relaxation was not
accompanied and produced by charge—whether muscular con-
traction did not hold the same relation to discharge, the dis-
charge acting, not as a stimulus to a vital property of irrita-
bility in nerve or muscle, but simply by allowing the play of
the attractive force inherent in the physical constitution of the
muscular molecules—a play which had been previously coun-
teracted by the presence of the charge. I could, mdeed, bring
myself to adopt no other conclusion than this: and thus it
was that this experiment upon the Torpedo proved to be the
means of adding not a little strength and definitiveness to the
conviction at which I had already arrived respecting vital
motion.
The question at issue, however, was not to be disposed of so
summarily; and, as I can now see plainly enough, I had long
to wait before I had evidence upon which my own reason or
that of any other person was entitled to rest with any feeling
of real satisfaction. Indeed, it is only of late that I have had
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 303
such evidence in my possession, and that I could safely venture
to challenge attention to it—that I could do what I propose to
do so soon as I have prepared the way a little by glancing at
certain historical points, which, as it seems to me, will make it
somewhat more easy to abandon the view of vital motion which
is at present in favour, and to adopt that which I would ven-
ture to substitute for it.
Very misty notions about vital motion prevailed in olden
times, and if the subject be looked into historically it will be
seen that much of this mistiness still clings about the notion at
present in favour.
In the beginning, as it would seem, all motion was looked
upon as essentially vital.
Thales talked about the world as being animated by a soul
and actuated by demons, and looked upon motion as being
brought about in one or other of these ways.
Hippocrates believed in the universal presence of a living,
intelligent, active principle, to which he gave the name of
nature (pvows), and to him, as to many in the present day, it
was enough to refer motion to nature—to regard it as natural.
The power of motion, indeed, was one of the faculties (Suvapecs)
with which the principle of nature was endowed.
Plato says little to the point. With him all philosophy
merged in theology: to him vital motion, and motion generally,
resolved itself into a display of divine power.
Aristotle, the great contemporary of Plato, recognized, not
a Divine Being as Plato did, but a First Moving Cause, a
primum mobile, one in essence, eternal, immaterial, immove-
able, and yet the spring of all motion. According to him, this
First Moving Cause worked in the living body (féov) through
the instrumentality of a principle which was distinctive of this
body, and to which he gave the name of soul (Wuyn)—a prin-
ciple possessing various energies or faculties of its own, distinct
from the organs in which it was manifested, and yet requiring
these organs for its manifestations. To this soul, when most
developed, belonged several faculties (Suvayers)—the faculty of
receiving nourishment (dUvaputs Opertixy), the faculty of sensa-
tion (6. aicOmrixn), the faculty of motion in place (8. kwyri«y),
the faculty of impulse or desire (6. épeti«n), the faculty of intel-
304 DR RADCLIFFE.
ligence (8. Ssavoutixn). Vegetables even, by having the lowest
of these faculties, the threptic, were supposed to have souls.
And—chiefly by chancing to witness the action of the intercostal
muscles under the transparent pleura in a living chameleon,
which he had cut open—Aristotle was also able to connect the
movement of animal bodies with the action of the muscles, and
to indicate, not only the difference in function between the
muscles and the nerves, but to define more clearly—a discovery
hinted at by Praxagoras two hundred years before his time—
the distinction between nerves of motion and nerves of sensation,
After this time, for a thousand years and more, when any-
thing was done in this direction it was little beyond a servile
copying of what had been seen said by Hippocrates and Aris-
totle. Even Galen made little progress in the matter of
originality ; nor yet the schoolmen of the middle ages, with
whom for the most part the notions chiefly in the ascendant
were those of alchemy and magic and astrology. At the revival
of letters, indeed, the only light of importance was that derived
from the old Greek fathers in science; and at the end of this
epoch no new light had arisen to dissipate the darkness. No
new light, for instance, was shed by the doctrine of occult
causes which found most favour in these times, for this doctrine
was no more than a copy of the doctrines of Hippocrates or
Aristotle, that various bodies had various powers (duvapess) by
which they were able to act in the various ways natural to
them. And still less did new light proceed from the notion of
Paracelsus and his followers, which in too many instances was
associated more or less closely with that of occult causes—the
notion, that is to say, that there were elementary spirits, inter-
mediate between material and immaterial beings, with special
names, in the four elements of air, water, fire, and earth—syl-
vans or fairies in the air; nymphs and undines in the water; sala-
manders in the fire; gnomes, trolls, pigmies, spirits of the mine,
little folks, little people, cobolds in the earth ;—that the human
body had its double or demon, called Arehzeus, whose primary
function was to superintend the work of the stomach, and who
managed the various functions of the body, that of motion
included, through the instrumentality of a legion of underling
deputies undignified by any distinctive names,
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 305
Indeed it was not until Von Helmont, Stahl and Hoffmann
appeared on the scene that the notions handed down from the
ancients began to be materially modified, and to take the
shapes belonging to modern times.
With Paracelsus, Von Helmont held that the Archzeus and
its underlings were the agents in all vital manifestations, and
he also thought for himself a little, for to him belongs the
credit, if credit it be, of being the first to maintain that the
living body had powers of a specific character altogether different
from those belonging to inanimate nature.
Accepting the doctrine that there was one law for animate
and another for inanimate nature, Stahl went further, and
maintained that matter is essentially and necessarily passive
and inert, and that all its active properties or powers are
derived from a specific and immaterial animating principle
imparted to it—a principle to which he gave the name of
anima. ‘The body, he held, as body, has no power to move
itself. All vital motion is a spiritual act. The physical powers
of matter, which have only free play after death, are in every
way opposed to, and counteracted by, the anima, of which he
further says, as the followers of Hippocrates said of nature,
that “it does without teaching what it ought to do,” and does
it “without consideration ;’ a remark which makes it evident
that the anima of Stahl is not to be confounded with the con-
scious personal Archeus of Paracelsus and Von Helmont.
What Stahl explained in this way, Hoffmann, who took
the next noticeable step in advance, explained on the hypo-
thesis of nervous influence or nerve-fluid, whatever that may
mean. By this influence or fluid, according to him, the moving
fibres have a certain power of action, or tone, which may be in-
creased or diminished. If increased unduly, spasm is the result:
if unduly decreased, atony.
Next in order have to be named Glisson, Haller, and the
Brown, known as the author of the Brunonian system of medi-
cine, men whose speculations form the basis of the doctrine of
vital movement now in favour.
Glisson, an eminent professor at Cambridge in his day, was
the first to advance the present doctrine of muscular irritability.
He asserted that there was in muscle a specific vital property,
306 DR RADCLIFFE.
to which he gave this name, and that contraction was the act of
this property.
Haller expanded this idea, and drew for the first time a line
of distinction between the special vital property of muscle and
the special vital property of nerve. He retained the name of
irritability for this property in muscle: he gave the name of
sensibility to this property in nerve. Each property was essen-
tially vital, something departing at death, and therefore in no
wise akin to any power in inanimate nature. The property was
a life of which muscular contraction or nervation were acts.
Brown, starting from this point, added another idea—that
of stimulation. Everything acting upon the vital property of
irritability or sensibility (to which he gave the common name of
excitability), according to him, acted as an excitant or stimulus.
Action is caused by a process of stirring-up, as it were, the ca-
pacity for action being asleep or at rest until it is so stirred-up.
The idea would seem to be none other than that all vital move-
ment in its nature is identical with that which is produced by
teasing a sleeping man until’he wakes up and strikes about
him in anger.
And this doctrine of vital motion, which thus took form in
the speculations of Glisson, Haller, and Brown, is, with little
change, the doctrine at present in favour. |
In point of fact, the position taken at present has but little
shifted since the days of the schoolmen, when occult qualities of
one kind or another were thought to be a sufficient explana-
tion for everything—when, for example, terreity, aqueity, and
sulphureity, the occult qualities of the three elements, earth,
water, and sulphur, of which, in varying proportions, according
to Paracelsus, all bodies were composed, were supposed to ac-
count for all that was general in these bodies,—when Petreity
was thought to bea sufficient explanation of the peculiarities
distinguishing Peter from men with other names,—when the
answer of Argan’ to the question, “quare op'um facit dormire,”
in the mock examination for the diploma of physician, would
have been listened to without a smile if it had been given in
sober earnest before the examiners of a real faculty of medi-
cine :—
1 Molitre, La Malade Imaginaire: 3itme interméde.
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 307
Mihi a docto doctore
Demandatur causam et rationem quare
Opium facit dormire.
Et ego respondeo
Quia est in eo
Virtus dormitiva
Cujus est natura
Sensus assoupire.
For in referring vital motion to a property of irritability, what
more is done than to say, that the moving body moves because
it is actuated by an occult quality which is suspiciously akin
to terreity, aqueity, or sulphureity, or to Petreity, or to the
“virtus dormitiva” of opium in the comedy? ‘To tell us,”
as Newton said, “that every species of thing is endowed with
an occult specific quality, is to tell us nothing.” Even to say
that the phenomenon is vwital, is, as Whewell remarks, “very
prejudicial to the progress of knowledge by stopping enquiry
by a mere word.’ Moreover, the very assumption upon which
the doctrine in question is based—that vital motion is altogether
distinct from physical motion—is itself not altogether satis-
factory, “At the best,” as Coleridge says’, “it can only be
regarded as a hasty deduction from the first superficial notions
of the objects that surround us, sufficient, perhaps, for the
purpose of ordinary discrimination, but far too indeterminate
and diffluent to be taken unexamined by the philosophic
enquirer. * * * * By a comprisal of the petitio principiw
with the argumentum in circulo—in plain English, by an
easy logic which begins by begging the question, and then,
moving in a circle, comes round to the point where it begins—
each of the two divisions has been made to define the other by
a mere re-assertion of their assumed contrariety. The Physio-
-logist has luminously explained y+x by informing us that it
was a somewhat that is the antithesis of y—x, and if we ask
what then is y—x, the answer is, the antithesis of y+x;—a
reciprocation that may remind us of the twin sisters in the fable
of the Lamiz, with one eye between them both, which each
1 Tints towards the Formation of a more Comprehensive Theory of Life.. By
8. T. Coleridge. Ed. by Dr Seth B. Watson. Churchill, 1848.
308 DR RADCLIFFE.
borrowed from the other as either happened to want it, but
with this additional disadvantage, that in the present case-it is,
after all, but an eye of glass.”
But this glance at the history of vital motion is not yet
ended. Up to the time of Von Helmont the idea of a well-
defined gulph between animate and inanimate nature was not
clearly defined: nor yet after this time did this idea gain uni-
versal acceptance.
At the time of Paracelsus the facts of chemistry began to
occupy a large share of the attention of philosophers, and soon
afterwards a school, called the Iatro-chemical school, propounded
various physiological doctrines founded upon chemistry. The
opposition of acid and alkali, and the workings of ferments of
one kind or another, were supposed to supply the solution of
many problems in vitality. Then came the hope, kindled na-
turally by the splendid discoveries of Galileo and Newton in
physical science, that the mechanical principles of the macro-
cosm would supply the key to all requirimg interpretation m
the microcosm—a hope which called into existence the so-called
Jatro-mathematical or mechanical Physiologists. The question
was of the cohesion, the attraction, the resistance, the gra-
vity, which operate in inert matter, and of mechanical impulse
and elasticity, not of powers of a higher order: it was believed
that all the various bodily functions were problems to be solved,
as so many hydraulic or hydrostatic problems chiefly, partly by
gravitation and the laws of motion, and parily by chemistry,
which itself, as far as its theory was concerned, was but a
branch of mechanics, working exclusively by imaginary wedges,
angles and spheres. The restoration of ancient geometry, aided
by the modern invention of algebra, had placed the science of
mechanism on the philosophical throne. It was thus, for ex-
ample, that Borelli dealt with the problem of muscular motion,
and after him Bellini.
As far back also as the time of the great Bacon, Gilbert had
struck out a new path in the same direction, the following out
of which has led to more satisfactory results than any of those
arrived at by the Iatro-mathematical School in their own par-
ticular lines of enquiry. He had investigated the phenomena
of magnetism with great success, and by continual poring
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 309
over this subject had come to believe that magnetism supplied
the key to vital movement, and to vital and physical problems
in general; but his speculations bore little or no fruit, and are
chiefly of interest as being the first step in an enquiry which
did not begin until two hundred years later, when an event
occurred in a house in Bologna which marks the birth of a new
epoch in the philosophy of vital motion, and on which it may
be well to say a word or two. The house is in the Via Ugo
Bassi, gia Strada Felice. The event is commemorated on a
marble slab let into the front in these words:—
LuIGI GALVANI
in questa casa
di sua temporaria dimora
al primi di settembre
dell’ anno MDCCLXXXVI
scoperse dalla morte rane
LA ELeTTRicITA ANIMALE
Fonte di maravighe
a tutti secolt.
The actual event was this. Experimenting with an ordinary
electrical machine at a short distance from a dish on which
lay a number of frogs’ legs prepared for cooking, and noticing
that these legs jumped whenever he drew a spark from
the prime conductor of the machine, it occurred to Galvani
that they might serve as very delicate electroscopes in some
experiments on atmospheric electricity in which he was then
engaged. Thereupon, he and his nephew Camillo Galvani, who
was with him at the time, each of them taking with him a
handful of the contents of the dish, mounted to a belvedere on
the top of the house which served the purpose of an electrical
observatory, and at once proceeded to put the idea in practice.
It was expected that these limbs might jump in obedience to
discharges of atmospheric electricity as they had been seen to
jump in obedience to discharges of Franklinic electricity ; and
in order to see whether they would do so or not, they were
suspended, by means of small hooks of iron wire bent suitably
upon certain iron bars or stays which stretched across the upper
part of the arched openings with which three sides of the belve-
310 DR RADCLIFFE.
dere were pierced. The time was a clear and calm evening,
without a gleam of either lightning or aurora; and yet the
limbs were found to jump whenever the iron hooks by which
they were suspended were pressed upon by the finger, and
not unfrequently when they were left untoucied. Describing
what happened, Galvani says, “Ranas itaque consueto more
paratas uncino ferreo earum spinali medulla perforata atque
appensa, septembris initio (1786) die vesperascente supra
parapetto horizontaliter collocavimus. Uncinus ferream laminam
tangebat: en motus in rana spontanei, varii, haud infrequentes.
Si digito uncinulum adversus ferream superficiem premeretur,
quiescentes excitabantur, et toties ferme quoties hujusmodi pres-
sio adhiberetur’.” The house, the ricketty wooden flight of steps
leading from the principal staircase to the belvedere, unmended,
unpainted, almost unswept, the belvedere itself, the iron bars
upon which the limbs were suspended, are still there, or were
there the other day when I made a pilgrimage to the spot;
and even the presence of Galvani himself may be recalled by
the help of a portrait which hangs or hung in the open landing
facing the locked door at the foot of the stairs leading to the
belvedere. In this place, and in this way, was the discovery
made which is commemorated on the slab on the front of the
house as the well-spring of wonders for all ages “fonte di
maraviglie a tutti secoli,” and of which a short time before
the close of the last century the illustrious author of Cosmos
wrote “le nom de Galvani ne perira point; les siécles futurs
profiteront de sa découverte, et, comme le dit Brandes, ils
reconnaitront que la physiologie doit & Galvani et & Harvey
ses deux bases principales*.” At this time, then, and in this
place, Galvani saw the contractions he describes, and dis-
covered, or rather divined, in them the existence of animal
electricity. How, he asked himself, were these contractions
to be accounted for? They could not be due to discharges
of atmospheric electricity, for the sky at the time presented
no indications of electric disturbance: they could not be due
to the discharges which gave rise to them within the house,
1 De Viribus Electricitatis in motu musculari Commentarius, 1791.
2 Expériences sur le galvanisme, et en général sur Virritation des fibres muscu-
laires et nerveuses. F, A. Humboldt. Traduit par J. F. N. Jadelot. 8vo. Paris,
1799, p. 361.
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 311
for the electric machine, which was also left behind, was not in
action: they could not be due, that is to say, to discharges of
either of the two kinds of electricity then known; and having
arrived at this point, he jumped from it to the conclusion, that
the limbs themselves must have an electricity of their own, and
that the contractions were brought about by discharges of this
electricity. It never occurred to him to doubt that electricity
was the agent at work in causing those contractions: and, in
short, he did not hesitate to conclude, not only that the contrac-
tions were in themselves abundant proof of the existence of
animal electricity, but also that the muscular fibres are charged
during rest as Leyden jars are charged, that muscular contrac-
tion is the sign and effect of the discharge of this charge, the
discharge, in one way or another, being brought about by an
electrical action of the nerves upon the muscles.
From this time until the day of his death, Galvani went on
performing experiment after experiment, sacrificing hecatombs
of fregs, and never wavering in his belief in the existence of
animal electricity, or in the conclusion he had come to respect-
ing the action of this electricity in vital motion: but during
his lifetime he was destined to be foiled in his hopes to bring
others to the same mind with himself, and that too by a weapon
which lay hid in one of his own experiments. The experiment
in question was one in which a galvanoscopic frog was thrown
into a state of momentary contraction by placing a conducting
are, of which one half was silver and the other half copper,
between the lumbar nerves and the crural muscles’. Galvani,
as was his wont, explained these contractions by supposing that
the conducting are had served to discharge animal electricity,
and that the contractions were the result of the discharge.
Volta, on the other hand, was of opinion that the electricity
producing these contractions originated in certain reactions
between the silver and copper portions of the conducting arc;
and he was not shaken in this view by what he did afterwards,
for, wishing to confirm it, he began a series of investigations
which ended in the discovery of the voltaic pile and battery—
1 The galvanoscopic frog was prepared from the hinder half of the animal, by
stripping off the skin, and cutting away all the parts between the thighs and the
fragment of the spine, except the principal nerves,
312 DR RADCLIFFE.
a discovery which filled all minds with wonder, and for a long
time afterwards diverted attention altogether from the con-
sideration of the claims of animal electricity. In the mean-
time, however, while Volta was demonstrating the existence of
that electricity which originates in the reaction of heteroge-
neous bodies, and which is now known as voltaic electricity,
Galvani continued his search after animal electricity, and made
many important discoveries as he went along. He discovered,
among other things, that a galvanoscopic frog would contract
without the help of a conducting are composed of heterogeneous
metals. He discovered not only that these contractions would
happen when this arc was composed of a single metal, but also
that an arc composed of muscle or nerve would answer the
same purpose as the metallic are. He also discovered that the
limb of a galvanoscopic frog, of which the nerve had been
divided high up in the loins, would contract at the moment
when the end of the nerve below the line of division was
brought down and made to touch a part of the trunk of the
same nerve. At last, indeed, he hit upon an experiment in
which he seemed to have to do with an electricity other than
that arising frem the reaction of heterogeneous bodies—an
electricity which must belong to the animal tissues themselves.
He did much, but he did not do enough to win the battle in
which he was engaged, for Volta still kept his position, denying
the existence of animal electricity, and maintaining that the
electricity which produced the contractions in the galvanoscopic
frogs was always due to electricity arising in the reaction of
heterogeneous bodies of one kind or other—silver and copper,
metal and organic tissue, muscle and nerve, nerve in one state
with nerve in another, as the case might be’.
In 1799, Humboldt took up the question at issue between
Galvani and Volta, and published a work’ in which he shews
by many new and curious experiments that there was error on
both sides—that Volta was wrong in ignoring altogether the
influence of animal electricity in Galvani’s experiments, and
that Galvani was not less wrong in recognising nothing but
this influence. He, himself, as is proved in the extract already
given, was a firm believer in animal electricity; but he failed to
1 Ann, de Chim., T. xxut. p. 276 and 301. aTOp err
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 313
supply reasons for this belief which can be regarded as thoroughly
satisfactory at the present day. Still, he did something in this
direction by making out—first, that the agent assumed to exist,
and to be animal electricity, has this in common with elec-
tricity, that its action is permitted by conductors and prevented
by nonconductors; and, secondly, that it is not to be con-
founded with voltaic electricity, because the action, which is
permitted by conductors, is possible across a gap in the circuit
which would allow the passage of frictional electricity, but
which would altogether prevent that of voltaic electricity—
that is to say, allow electricity of high tension to pass, but not
electricity of low tension. What Humbolt did, in fact, was to
increase the probabilities of the existence of animal electricity
not a little, and at the same time to make it appear that this
electricity would prove to be of higher tension than voltaic
electricity under ordinary circumstances.
In 1803, Aldini, Galvani’s nephew’, published an account of
certain experiments which furnish further evidence in favour of
the existence of animal electricity, by shewing that living
animal tissues are capable of giving rise to attractions and
repulsions which seem to be no other than electrical attractions
and repulsions. “TI held,” he says, “the muscles of a prepared
frog in one of my hands, moistened with salt and water, and
brought a finger of the other hand, well moistened in the same
way, near to the crural nerves. When the frog possessed a
great deal of vitality, the crural nerves gradually approached
my hand, and strong contractions took place at the moment of
contact.” And again :—“ Being desirous to render this pheno-
menon more evident, I formed the are by applying one of my
hands to the spinal marrow of a warm-blooded animal, while
I held the frog in such a manner that its crural nerves were
brought very near to the abdominal muscle. By this arrange-
ment the attraction of the nerves of the frog became very
evident.”
About this time, however, the discovery of the voltaic
battery had given the victory to the opinions of Volta—a vic-
1 “ Account of the late Improvements in Galvanism, with a series ‘of curious
and interesting experiments performed before the Commissioners of the French
National Institute, and repeated in the Anatomical Theatres of London, &c.”’
4to. London, 1803.
VOL. VIII. Dil
314 DR RADCLIFFE.
tory so complete that nothing more was heard about animal
electricity for the next thirty years.
In 1827, Nobili* brought back the subject of animal elec-
tricity to the thoughts of physiologists by discovering an
electric current in the frog. He made this discovery by means
of the very sensitive galvanometer which he himself had
invented a short time previously—an instrument which, as
perfected by M. Du Bois-Reymond and others, by Sir Wilham
Thomson more especially, ought to be as prominent an object
as the microscope in the laboratory of every physiologist. Im-
mersing each end of the coil of the instrument in a vessel con-
taining either simple water or brine, and completing the circuit
between the two vessels with a galvanoscopic frog—the fragment
of the spine being immersed in one vessel, and the paws in the
other—he found that there was a current in the frog from the
feet upwards, which current would cause a considerable perma-
nent deflection of the needle—to 30° or more if brine were
used, to 10°, or thereabouts, if water were substituted for brine.
Nobili supposed that this current was peculiar to the frog, and
in this he erred; but he did, nevertheless, a great thing, for, by
this experiment, he furnished, perhaps, the first unequivocal
proof of the real existence of animal electricity.
Twelve or thirteen years later, Matteucci published an
essay” which, as M. De la Rive says’, “ restored to animal elec-
tricity the place which it ought to occupy in electrical and
physiological phenomena.” This essay, moreover, had a great
indirect influence upon the fortunes of animal electricity, for
M. Du Bois-Reymond, as he himself tells us, was led to under-
take the investigations which have made his name famous in
this department of physiology by the inspiration arising from
its perusal. :
The joint labours of MM. Matteucci and Du Bois-Reymond
have left no room for entertaining any doubt as to the reality
of animal electricity. This will appear sufficiently in the
sequel, when many of the experiments which furnish the
demonstration will have to be referred to particularly. In the
' Bibl. Univ., 1828, T. xxxvit., p. 10.
2 Traité des Phénoménes Electro-physielogiques des Animaux. Paris. 1844.
3 4 Treatise on Electricity, in theory and practice. Translated by C. V.
Walker. 8yo. Longman. 1853-1858.
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 315
meantime, it may be said that Matteucci has demonstrated in
the most unequivocal manner that animal electricity is capable
of decomposing iodide of potassium, and of giving “signes de
tension avec un condensateur délicat’,’ as well as of producing
movement in the needle of the galvanometer; and not only so,
but also—a fact, the discovery of which will always give Mat-
teucci a place m the very foremost rank of physiological dis-
coverers—that muscular contraction is accompanied by an
electrical discharge analogous to that of the Torpedo. And as
for M. Du Bois-Reymond? it may be said that he has demon-
strated most conclusively that there are electrical currents in
nerve—in brain, spinal cord, and other great nerve-centres, in
sensory, motor, and mixed nerves, in the minutest fragment as
well as in masses of considerable size,—that the electrical current
of muscle, which had been already discovered by Matteucci, may
be traced from the entire muscle to the single primitive fasci-
eulus,—that Nobili’s “ frog-current,” instead of being peculiar
to the frog, is nothing more than the outflowing of the currents
from the muscles and nerves,—that the law of the current of
the muscle im the frog is the same as that of the current of
muscle in man, rabbits, guinea-pigs and mice, in pigeons and
sparrows, in tortoises, lizards, adders, glow-worms, toads, tad-
poles, and salamanders, in tench, in freshwater crabs, in earth-
worms—in creatures belonging to every department of the
animal kingdom,—that the law of the current in muscle agrees
in every particular with the law of the current in nerve, and
also with that of the feeble currents which are met with in
tendon and other living tissues,—and that there are sundry
changes in the current of muscle and nerve under certain
circumstances, as during muscular contraction, during nervous
action, under the influence of continuous and interrupted gal-
vanic currents, and so on, which changes, as I shall hope to
shew in the sequel, are of fundamental importance in clearing
up much that would otherwise be impenetrable darkness in the
physiology of muscular action and sensation.
Before the discovery of the galvanometer the attention of
those who cared to meddle in these matters was directed
1 Cours @Electro-Physiologie. Paris. 1858.
2 Untersuchungen iiber thierische Electricitiét. Berlin. 1849, 1853.
21—2
316 DR RADCLIFFE.
exclusively to the static phenomena of animal electricity. Then
the only definite electrical ideas were, charge on the one hand,
and discharge on the other. After the discovery of the galvano-
meter, the original point of view was abandoned altogether, or
almost so, and the attention diverted from the static to the
current phenomena of electricity. And herein, as I believe,
was an unmixed misfortune. As I take it, indeed, it is neces-
sary to go back to the standpoint occupied by Galvani and
Humboldt, and to work with the electrometer rather than with
the galvanometer ; and this conviction has now so much gained
upon me, that I am disposed to regard the New Quadrant
Electrometer of Sir Wiliam Thomson—the instrument which
for the first time makes it possible to arrive at an accurate
knowledge of the statical aspects of animal electricity—as an
instrument which is, to say the least, quite as indispensable as
the galvanometer itself to those who would do the work in
question. Already, indeed, as it will appear in due time, this
instrument has supplied proof of the existence of a definite
charge of electricity in nerve and muscle during rest, and of the
discharge of this charge when this state of rest is changed for
that of action, as well as of other facts without which, as I
believe, it would be impossible to gain any real insight into the
mystery of vital motion.
And thus, by the fact of the existence of animal lacie
being now established beyond question, the way is more pre-
pared than it was in the days of Galvani for the adoption of
any view of vital motion in which animal electricity has to
serve as the basis.
At the commencement of these introductory remarks I have
told how I came to doubt the truth of the view of vital motion
at present in favour, and to adopt in its stead a view which is
substantially that of Galvani. I did not then know that Galvani
had anticipated me; neither did I know that others were in the
same predicament, about whose labours it still remains for me
to say a few words.
The name to be mentioned first in order here is that of Dr
West, of Alford, in Lincolnshire. As early as 1832’, in some
1 “On the Influence of the Nerves over Muscular Contractility,” London
Medical and eeeroeat Journal, edited by Michael Ryan, M.D. Vol. 1. 1832.
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 317
remarks upon the influence of the nerves upon muscular con-
tractility, this writer maintains, “that the nervous influence
which is present in relaxed muscular fibre is the only influence
which the nerves of volition possess over that tissue; that its
office there is to restrain or control the tendency to contract
which is inherent in the muscle; and that contraction can only
take place when by an act of the will this influence is sus-
pended, the muscle being then left to act according to its own
innate properties ;” . . . and again, “that nervous influence is
imparted to muscular fibre for the purpose of restraining its
‘contraction, and that the action of the will, and of all other
disposers to contraction, is simply to withdraw for a while this
influence, so as to allow this peculiar property of muscular fibre
to shew itself.’ The co-existence of spasmodic action with
nervous debility, the efficacy of stimulants as antispasmodics,
and the postponement of rigor mortis until all traces of nervous
action have disappeared, are the principal facts which are
adduced in support of the probability of this theory.
Very shortly after the publication of these remarks, a similar
idea appears to have been hinted at by Sir Charles Bell in a
lecture at the Royal College of Surgeons in London, for, after
premising that the question could never be settled, the lecturer
said, “that relaration might be the act, and not contraction,
and that physiologists, in studying the subject, had too much
neglected the consideration of the mode by which relaxation is
effected.” This remark is preserved by Dr West in the essay
to which reference has just been made.
Six years later, in a chapter of his classical work on eom-
parative anatomy’, Professor Dugés, of Montpellier, argues with
much clearness that all organic tissues are the seat of two oppo-
site movements—expansion and contraction, and that contrac-
tion, which is in no sense peculiar to muscle, is nothing more
than the cessation of expansion ”—“ la contraction musculaire ne
consiste que dans l’annihilation de l'expansion.” The muscle is
supposed to contract in virtue of its elasticity, just as a piece of
caoutchoue might contract when set free from a previous state
of extension; and an analogy is hinted at between the expanded
1 Traité de Physiologie comparée de V Homme et des Animauz. 8yo. Mont-
pellier and Paris. 1838.
318 DR RADCLIFFE.
state of the muscle and the fluid state of the fibrine of the blood,
and between rigor mortis and the coagulated state of this
fibrine. Analogous in its effects to electricity, the vital agent is
supposed to accumulate in the muscles, and to produce expan-
sion by causing the muscular molecules to repel each other ;
and contraction is supposed to be brought about either by the
sudden discharge (as in ordinary contraction) or by the gradual
dying out (as in rigor mortis) of the vital agent. And, further,
it is supposed that the rhythmical movements of muscle are
caused by successive discharges of the vital agent, which dis-
charges are brought about whenever this agent acquires a cer-
tain degree of tension; and that the cramps of cholera, or the
spasms of tetanus or hysteria, are consequent upon the develop-
ment of the vital agent being for the time suspended.
More recently still, namely in 1847, Professor Matteucci
communicated a paper to the Parisian Academy of Sciences*
upon the influence of the nervous fluid in muscular action, in
which he writes :—‘“Ce fluide développé principalement dans
les muscles, s’y répand, et, doué d’une force répulsive entre ses
parties, comme le fluide électrique, il tient les éléments de la
fibre musculaire dans un état de répulsion analogue a celui
présenté par les corps électrisés. Quand ce fluide nerveux cesse
d’étre libre dans le muscle, les éléments de la fibre musculaire
sattirent entre eux, comme on le yoit arriver dans la roideur
cadavérique. . . . . Suivant la quantité de ce fluide qui
cesse d’étre libre dans le muscle, la contraction est plus ou moins
forte.’ Professor Matteucci appears ta have framed this hypo-
thesis, partly, in consequence of certain considerations which
seemed to shew that the phenomenon of “induced contrae-
tion” was owing to the discharge of electricity in the
muscle in which the “inducing contraction” was mani-
fested—an idea originating with M. Becquerel—and, partly,
in consequence of the analogy which he himself had found
to exist between the law of contraction in muscle and the
law of the discharge in electrical fishes; but he does not appear
to have attached much importance to the hypothesis. Indeed,
his own comment at the time is—“j’ai presque honte d’avoir eu
la hardiesse de communiquer 4 | Académie des idées si vagues,
1 Comptes Rendus. March 17th, 1847.
THE SYNTHESIS OF MOTION, VITAL AND PHYSICAL. 319
et apparemment si peu fondées, et contre lesquelles on pourrait
faire bien des objections, mais je pense que, parmi les théories
physiques les mieux fondées aujourd’hui, il en existe qui ont
débuté de cette manicre, et il est certain que des hypotheses,
aussl peu fondées que celles-ci, ont quelquefois pu produire
ensuite des découvertes remarquables.”
Next in order, and almost contemporaneously with the date
of my own first publication on the subject, Professor Engel, of
Vienna, wrote’:—‘‘So hat der Nerve die Aufgabe, nicht die
Zusammenziehungen des Muskels zu veranlassen, sondern den
Zusammenziehungen bis auf einen geringen Grad entgegenzu-
wirken. Im lebenden Organismus, in welchem Ruhe etwas
unmogliches ist, ist auch ein ruhender Muskel eben so wohl
wie ein ruhender Nerv undenkbar, der Muskel in seinem bestiin-
digen Streben, sich zusammenzuziehen, wird vom Nerven daran
verhindert, im Nerven macht sich das fortwaihrende Streben
kund, die Zusammenziehung des Muskels auf ein gerechtes
Mass zuriickzufiihren ; das Ergebniss dieser zwei einander ent-
gegengesetzten Higenschaften des Nervens und des Muskels ist
das, was man gemeinhin Zustand der Ruhe, Zustand des
Gleichgewichtes, oder an Muskeln auch Tonicitat nennt. Das
Verlassen dieses Gleichgewichts ist die Bewegung einerseits,
die Lahmung andererseits. Die Bewegung wird aber erzeugt,
indem entweder der Einfluss des Nervens auf den Muskel
herabgesetzt wird, oder indem die Contractionskraft des Muskels
unmittelbar gesteigert wird. Lahmung des Muskels findet sich
gleichfalls entweder durch unmittelbare Vernichtung der Con-
tractionskraft des Muskels oder durch eine iibermiissig gestei-
gerte Einwirkung des motorischen Nervens auf den Muskel.
Sollen daher abwechselnde Muskelcontractionen zu Stande
kommen, so ist die Gegenwart des lebendigen Nervens im
Muskel unerlisslich, und auch bei unmittelbaren Muskelreizen
kénnen abwechselnde Zusammenziehungen nur erfolgen, so
lange noch die Nerven lebensfihig sind ; hort letzteres auf, so
zieht sich der Muskel ohne Hinderniss zusammen. Diesen
Zustand nennen wir die Todtenstarre.” The chief grounds for
this opinion are, first, certain original experiments, some of
1 “Ueber Muskelreizbarkeit,” Zeitschrift der Kais. Kén. Gesellsch. der Aertze
zu Wien, Exster Band, pp. 205—-219, and pp. 252—270, 1849.
320 DR RADCLIFFE. THE SYNTHESIS OF MOTION.
them very remarkable, which afford additional proof that the
muscles of frogs are more prone to contract when they are cut
off from the influence of the great nervous centres, secondly,
the frequent spontaneous occurrence of cramps and other forms
of excessive spasmodic contraction in paralysed parts, and,
thirdly, the supervention of the permanent contraction of rigor
mortis when all signs of nervous irritability are completely
extinguished.
And, last of all, I find Professor Stannius, of Rostock’, arriving
at the conclusion:—“ dass es eine wesentliche Aufgabe der soge-
nannten motorischen oder Muskelnerven sei, die natiirliche Elasti-
citiitsgrdsse der Muskelfasern herabzusetzen und ihre Elasticitat
vollkommener zu machen; dass anscheinende Ruhe des Muskels,
zum Beispiele, wihrend des Schlafes, das Stadium solchen regen,
den Muskel zu seinen Aufgaben wieder befahigenden Ner-
veneinflusses anzeige; dass active Muskelzusammenziehung
einen geregelten und begrenzten momentanen Nachlass des
Nerveneinflusses auf den Muskel bezeichne; dass endlich die
Nachweisung einer Muskelreizbarkeit, in der iiblichen Auffas-
sungsweise, ein durchaus vergebliches Bemiihen sei.” M. Stannius
was led to this conclusion by certain original experiments, in
which he found blood to have the power of relaxing rigor mortis
and restoring muscular irritability, and these experiments are
advanced im evidence. Reference is also made to arguments,
to be brought forward on another occasion, which will prove—
“dass diese Anschauungsweise, so paradox sie immer auf den
ersten Anblick sich anlassen mag, mit unserem thatsichlichen
Wissen iiber Nerven- und Muskelthitigkeit keineswegs im
Widerspruch steht.” The essay from which these quotations
are taken was published towards the end of 1852—about two
years after the date of my own first publication on the subject.
I do not stand alone, then, in thinking that a great change
is necessary in the theory of muscular motion—a change
amounting to no less than a complete revolution; and I am glad
that it is so, for thus supported, I am the more bold to challenge
attention to the facts and arguments which will be advanced on
a future occasion.
1 “Untersuchungen iiber Leistungsfihigkeit der Muskeln und Todtenstarre,”
Vierordt’s Archiv fiir Physiol. Heilkunde. Stuttgart, 1 Heft, p. 22, 1852.
DISSECTION OF A LAMB WITH FISSURE OF THE
STERNUM AND TRANSPOSITION OF THE ORIGIN
OF THE RIGHT SUBCLAVIAN ARTERY. By LESLIE
OaILviE and CHARLES W. CatTHoart, Students of Medi-
_ cine, University of Edinburgh.
WE owe to Professor Turner the opportunity of dissecting a
ewe lamb, in which a median ventral fissure was conjoined with
other malformations. The specimen was presented by Dr Mac
Watt of Dunse.
On the ventral surface of the body there existed a distinct
mesial cleft, which extended from the root of the neck to
the place of exit of the umbilical cord two and a half imches
in front of the nipples. This cleft penetrated not only
through the skin and immediately subjacent structures, but
also through the sternum, which was divided into two parts,
to each of which were attached the appertaining rib-cartilages.
The linea alba in front of the place of exit of the cord was also
divided, but behind the cord the anterior wall of the abdomen
was complete. From this deficiency in the thoracic and abdo-
minal walls, the heart enclosed in the pericardium, the liver,
the stomach, the small intestine the ccecum and greater part of
large intestine, were exposed. The apex of the heart, the
greater part of the liver, the abomasum and psalterium, the small
intestine, the czecum and greater part of the large intestine, pro-
truded through the fissure. Owing to the longitudinal division
of the sternum, and the increased space between the ribs of the
two sides, a deep fissure, at the bottom of which could be seen
the posterior vena cava, existed in the muscular and tendinous
substance of the diaphragm.
Tracing the peritoneal membrane forwards from the anterior
surface of the liver, it was found that the falciform ligament
passed to the free border of the left half of the diaphragm,
where it split into two layers, one of which was prolonged over
the posterior surface of that half, whilst the other passed to the
posterior surface of the right half, covering in its course the
322 MR OGILVIE AND MR CATHCART.
posterior vena cava, and bridging across the fissure or interval
between the two parts of the muscle. In bridging across this
interval, just after leaving the margin of the left half of the
diaphragm, the peritoneal membrane came in relation to the
posterior part of the left pleura for about a quarter of an inch,
but did not unite with it. It next united with the posterior
surface of the pericardium, where this membrane would have
covered the anterior surface of the diaphragm, had that muscle
been complete. It then united with the posterior part of the
right pleura, after which it passed to the posterior surface of
the right half of the diaphragm, the pleura, as usual, covering
its anterior surface. After lining what existed of the dia-
phragm on each side, the peritoneum passed over the abdomi-
nal walls, and ended in a defined margin at the border of the
great cleft.
The pericardium was torn on its ventral surface, so as to
expose part of the heart, but it probably had formed a complete
sac. Its posterior surface was blended with the inter-dia-
phragmatic portion of the peritoneum which was not in relation
to the pleure. Its sides were in apposition with the pleural
membranes, but the ventral surface was free. The bag of the
pericardium was very thin and transparent, smooth and serous
on its inner surface, and was reflected on to the exterior of the
heart in the usual manner.
Each pleura existed as a shut sac attached below to the
corresponding half of the sternum. Owing, however, to the
fissure, the inter-pleural space was abnormally large. Upon
opening into the pleural cavities the left lung was found col-
lapsed, while the right was partially inflated. The posterior
part of the right pleura was in relation to a greater extent of
surface of the peritoneum than the left, for although the origin
of the diaphragm on both sides reached as high as the eighth
rib, on the left side it also had an attachment to the xiphoid
cartilage. Thus the gap left in the posterior thoracic wall by
the fissure in the diaphragm was occupied by a thin twofold
membrane composed of peritoneum and pleura on each side,
and of peritoneum and pericardium in the middle.
Blood-vessels.—The aorta divided in the normal manner into
anterior and posterior aorte, the latter of which arched over the
ee
DISSECTION OF A LAMB. 323
root of the left lung. The anterior aorta passed forwards for
about half an inch on the ventral aspect of the trachea and to its
left side, and divided into two branches. One of these, which
formed the common stem for the two carotids, was prolonged in
the same direction for about half an inch before bifurcating.
The other passed forwards and to the left, to be continued as
the axillary artery, having first given off the vertebral, the pos-
terior cervical and dorsal branches. The artery which corre-
sponded to the right subclavian, instead of being a branch of
the anterior aorta, arose from the posterior aorta to the left of
the cesophagus, and half an inch in front of the root of the left
lung. It then passed to the right side between the cesophagus
and second dorsal vertebra, from which it was separated by the
origins of the prevertebral muscles. It passed forwards out of
the apex of the thoracic cavity, and was continued to the right
anterior extremity in the normal manner.
It will be seen that the arrangement of these vessels in this
lamb differs from the normal arrangement in ruminants :—
(1) in the right division of the anterior aorta not being con-
tinued as the right axillary, but passing forwards to form a
common stem for the two carotids; (2) in the right subclavian
—or what corresponds to it—arising from the posterior aorta.
An explanation of such deviations in the origins of the vessels
is obtained by reference to the developmental changes which
take place in the vascular arches of the embryo, as was de-
scribed in a detailed manner, and with ample reference to
illustrative cases, by Prof. Turner, in a memoir published
in the Medico-Chirurgical Review for 1862. In this paper
numerous instances are mentioned where, in the human
subject, the right subclavian arose from the arch of the aorta
to the left of the origins of the right carotid, left carotid and
left subclavian, and passed, as in this lamb, behind the trachea
and cesophagus to the right side. The explanation given is
that the fourth right vascular arch becoming atrophied does
not form the right subclavian, and consequently that the right
aortic root, instead of disappearing, as is the case in the normal
development of the vessels, remains pervious, so as to convey
the blood to the right upper limb. The part of the artery,
therefore, which, in this lamb, lay behind the cesophagus, was
324 MR OGILVIE AND MR CATHCART.
really the persistent pervious right aortic root, and its point of
junction with the left aortic root, opposite the second dorsal
vertebra, was, without doubt, the point at which the two aortic
roots had united.
Bones—The cervical vertebre displayed no peculiarity
except that the. axis and third vertebrae were fused together.
The fusion was more complete at the articular processes, and,
on the ventral surface of the bodies, distinct traces of separation
were seen between the neural arches, and above and below the
articular processes. The two fused bones were displaced shghtly
to the left side, and the right halves of the bodies were more
closely pressed together than the left halves. The articular
processes also on the right side were very feebly developed,
while on the left they seemed normal.
The dorsal vertebrae, thirteen m number, were the most
irregular of the true vertebre. A well-marked lateral curva-
ture of the spinal column to the right existed between the first
and eleventh vertebre, and was greatest between the fourth
and seventh. Moreover, this part of the spinal column was
twisted on itself so as to depress the left sides of the bodies and
raise the right. The centres of the inferior aspect of the bodies
were displaced to the right side of the space between the ribs.
The second, third, and fourth dorsal vertebre were fused
together even more closely than the second and third cervical.
Their spines were blended into one deep blade, the top of
which preserved the same dorsal curvature as the other spines.
The relative position of the three fused bones made it appear
as if they had been fused while the spinal column was straight,
and that after their union their posterior part had been dis-
placed to the right, which had caused the vertebre behind to
share in the curvature. The lumbar vertebree were seven in
number and quite regular.
The sacrum, somewhat more developed on the right side
than on the left, was twisted at the coccygeal end so as slightly
to raise the right border.
There were thirteen pairs of ribs, of which eight were true
and five false on each side. From the third to the eleventh
they were more or less irregular on both sides, but more
especially from the fourth to the seventh. The shafts of these
DISSECTION OF A LAMB. 325
ribs on the left side were straight instead of being curved out-
wards, and this caused the thoracic cavity to be narrowed from
side to side. The right ribs were all inclined backwards at a
more acute angle to the axis of the spinal column than were the
left. The latter however all projected to a lower level than the
former owing to the twisting of the spine already referred to.
The lateral curvature of the spine, the concavity of which
was to the left and greatest between the fourth and seventh
vertebrae, caused the heads of the corresponding ribs to be
ap voximated, and the shafts to be thrust together so closely as
to\ vent growth. Accordingly they were much thinner in
their central parts.
On the right side the articular processes of the vertebre,
being placed on the circumference of part of a circle, were
thrown further apart than usual. Hence a line drawn from
the articular facet on any transverse process to the facet on the
side of the vertebra next anterior was made to slope more
acutely backwards than it would otherwise have done. From
this arrangement three results followed. 1. The ribs of this
side were brought closer to the spinal column than usual.
2. The inclination of the shafts downwards was more acutely
backwards than on the other side. 3. The broad surfaces of
the ribs looked anteriorly and posteriorly as well as outwards
and inwards.
The two halves of the sternum were not absolutely sym-
metrical. Owing to the difference in the obliquity of the ribs of
the two sides, the right half was projected further forward than
the left, and would have been on a lower plane, had it not
been that the cartilages of the true ribs on the left side passed
horizontally forwards and inwards, whilst those on the right
passed forwards, downwards, and only very slightly inwards.
The centres of ossification in each half of the sternum were
the following :—
On the left side six, all well developed. The first between
the cartilages of the first and second ribs was normal in shape.
The second was between the second and third costal cartilages.
In common with the remaining four, it was of the shape of
a quarter of a sphere with the two flat surfaces looking respec-
tively inwards and downwards. The third was between the
326 MR OGILVIE AND MR CATHCART. DISSECTION OF A LAMB,
third and fourth costal cartilages. The fourth was between the
fourth costal cartilage and a point above the level of the inser-
tion of the fifth. The fifth, from the insertion of the fifth costal
cartilage to above that of the sixth. The sixth from the inser-
tion of the sixth costal cartilage to above that of the seventh
and eighth conjoined.
On the right side were five centres, all well developed except
the first. The first between the first and second costal carti-
lages was a mere speck of bony matter about the size of a pin’s
head. The second was between the second and third costal
cartilages, and with the three following was of the quarter
sphere shape. The third was between the third and fourth
costal cartilages. The fourth was between the fourth and fifth
costal cartilages. The fifth, larger than the others, was placed
below the fifth costal cartilage, opposite to the sixth, and
anterior to the conjoined seventh and eighth.
The halves of the xiphoid cartilage were well developed, and
continuous with each half of the sternum. The left part was
larger than the right, and had an elongated well-developed
centre of ossification close to the posterior part of the sternum.
According to Geoffrey St Hilaire* median fissures generally
are to be accounted for by a want of development, t.e. by two
halves, which should have been prolonged to meet each other,
having been arrested in their growth so as to fall short of their
line of junction, and the only reasonable explanation of the
cleft in this lamb’s sternum that we can give is that at a
very early period in its foetal life the ventral lamin had failed
to unite along the mesial line. Why in this case a stoppage of
the ventral union should have occurred we cannot say. That
an abdominal cleft should be connected with the opening for
the exit of the umbilical cord is only natural, but this gives us
no clue to the persistence of the fissure anterior to the um-
bilicus. It is possible that the curvature of the spine and
consequent distortion of the ribs may have followed from the
ventral lamine not having united; for the non-union of the
ribs in a common sternum would render these bones less able
to resist any distorting influences which may have been brought
to bear upon them.
1 Histoire des Anomalies, &c. 1832. Vol. 1, p. 596.
ON THE SENSE OF ROTATION AND THE ANATOMY
AND PHYSIOLOGY OF THE SEMICIRCULAR CA-
NALS OF THE INTERNAL EAR. By Pror. A. Crum
Brown, M.D. University of Edinburgh.
For some time I have been convinced that we possess a sense
of Rotation quite distinct from all our other senses. By means
of this sense we are able to determine—Ist, the axis about
which rotation of the head takes place; 2nd, the direction of
the rotation; and 38rd, its rate.
In ordinary circumstances we do not wholly depend upon
this sense for such information. Sight, hearing, touch, and the
muscular sense assist us in determining the direction and
amount of our motions of rotation, as well as of those of trans-
lation; but if we purposely deprive ourselves of such aids we
find that we can still determine with considerable accuracy the
axis, the direction, and the rate of rotation. The experiments
that I have made with the view of determining this point were
conducted as follows: a stool was placed on the centre of a
table capable of rotating smoothly about a vertical axis; upon
this the experimenter sat, his eyes being closed and bandaged ;
an assistant then turned the table as smoothly as possible
through an angle of the sense and extent of which the experi-
menter had not been informed. It was found that, with mode-
rate speed, and when not more than one or two complete turns
were made at once, the experimenter could form a tolerably
accurate judgment of the angle through which he had been
turned. By placing the head in various positions it was pos-
sible to make the vertical axis coincide with any straight line
in the head. It was found that the accuracy of the sense was
not the same for each position of the axis in the head, and fur-
ther, that the minimum perceptible angular rate of rotation
varied also with the position of the axis. It was also found
that considerable differences of accuracy exist in different indi-
viduals.
328 PROFESSOR CRUM BROWN.
The sense of rotation is, like other senses, subject to illu- .
sions, rotation being perceived where none takes place. Vertigo
or giddiness 1 is a phenomenon of this kind.
When, in the experiments just mentioned, rotation at a uni-
form angular rate is kept up for some time, the rate appears to
the experimenter to be gradually diminishing, and to cease
altogether after a time, varying with the position of the head,
and different with different individuals; if the rotation be
then stopped, he experiences the sensation of rotation about
the same axis in the opposite direction. If the position of the
head be changed after the prolonged rotation has been made,
the position of the axis of the apparent rotation is changed, re-
taining always the same position relatively to the head as was
occupied by the axis of the real rotation. The readiness with
which this complementary apparent rotation is produced is not
the same for each axis. In such experiments, as long as the
eyes are shut, and the axis of rotation kept vertical, a sensa-
tion of giddiness is not experienced. That sensation appears
to be caused by the discordance between the testimony of the
sense of rotation, and that of some other sense. Thus if I ex-
perience a sensation of rotation, it makes no difference to my
mind whether that sensation corresponds to a real rotation or
not, as long as I have no means of ascertaining independently
the existence or non-existence of the real rotation. I am in
that position as long as my head is fixed and my eyes shut.
But if, while the complementary apparent rotation is felt, I
open my eyes, I still feel that I am being turned round, but
at the same time I see that external bodies retain their po-
sition relatively to me—if I am turning round so are they—
and this produces at once a feeling of insecurity or giddiness.
Similarly this giddy or insecure feeling is produced, if, while
the complementary apparent rotation is felt, the head be moved
so that the axis of this rotation is no longer vertical. _
The sense of Rotation, being a special sense, must neces-
sarily have a special peripheral organ physically constituted so
as to be affected by rotation, a sensory nerve, and a central
organ. The structure of the semicircular canals of the internal
ear is such as to fit them to act as such a peripheral organ,
and the experiments of Flourens and of Goltz support this
THE SEMICIRCULAR CANALS. 329
view. The bony canals are filled with liquid, in which float
loose connective tissue and the membranous canals with the
contained endolymph. Rotation of the head about an axis at
right angles to the plane of a canal will then produce, on ac-
count of the inertia of the liquid, &c., motion of the contents
relatively to the walls of the canal, and this may be expected
to irritate the terminations of the nerves in the ampulla. If the
rotation be continued at a uniform rate, fluid friction, friction
of the endolymph against the membranous canal, and of the
perilymph against the membranous canal and the periosteum,
will gradually diminish this relative motion, which will at last
cease. We should therefore expect, as we have seen to be the
case, that continued uniform rotation should be perceived less
and less strongly, and that the sensation should at last die away
altogether. The time required for this equalisation of the mo-
tion of the canal and its contents will depend upon the rate of
rotation and upon the dimensions of the canal and the amount
of attachment of the membranous canal to the periosteum.
These latter conditions are not the same in the three canals,
and therefore we ought to find, as we do, that the rate at
which the sense of rotation dies away is not the same for dif-
ferent positions of the head. Again, if the uniform rotation is
stopped, the contents of the canal will continue to move on,
thus causing an apparent rotation in a direction the reverse of
that of the original rotation, and this also will die away owing
to friction.
As the three canals are in planes nearly at right angles to
one another, rotation about any axis can be resolved into rota-
tions, each of which will produce the effect described above
upon one of the canals, and thus any rotation will have its
appropriate sensation.
So far then this view of the function of the semicircular
eanals seems to explain the phenomena of the sense of rotation,
and I find that an explanation almost identical with this was
given by Professor Mach, of Prague, and by Dr Breuer, of’
Vienna, shortly before I communicated the substance of this
paper, as a preliminary note, to the Royal Society of Edinburgh*.
1 Proceedings, 19th January, 1874.
VOL. VIII. 22
330 PROFESSOR CRUM BROWN. ©
But this explanation is not sufficient. As far as we know, a
nerve current can vary only in intensity and not in kind, so
that, if irritated at all, whether by rnght-handed or by left-
handed rotation, the nerve would convey the same message to
the central organ. The solution of this difficulty which I pro-
posed is as follows:—Each canal has an ampulla at one end
only, and there is thus a physical difference between rotation
with the ampulla first, and rotation with the ampulla last, and
we can easily suppose the action to be such that only one of
these rotations (say that with the ampulla first, in which case,
of course, there is a flow from the ampulla into the eanal) will
affect the nerve terminations at all’. One canal can therefore,
on this supposition, be affected by, and transmit the sensation
of rotation about one axis in one direction only, and for com-
plete perception of rotation in any direction about any axis six
semicircular canals are required, in three pairs, each pair having
its two canals parallel (or in the same plane) and with their
ampulle turned opposite ways. Each pair would thus be sensi-
tive to any rotation about a line at right angles to its plane or
planes, the one canal being influenced by rotation in the one
direction, the other by rotation in the opposite direction.
Now we have six semicircular canals, three in the one ear
and three in the other, and I find im all the animals that I have
examined that the exterior canal of one ear is very nearly in
the same plane as that of the other; while the superior canal of
one ear is nearly parallel to the posterior canal of the other.
The three axes are therefore—Ist, a vertical’ axis at right
angles to the plane of the exterior canals; 2nd, an axis which
may be roughly defined as passing (in the human subject)
through the left eye and the right mastoid process at right
angles to the planes of the right superior and the left posterior
canal, and 8rd, an axis passing through the right eye and the
left mastoid process at right angles to the right posterior and
left superior canals.
In different animals there are great differences in relative
1 In the preliminary note above referred to, I described a way in which this
might be supposed to take place.
2 In the human subject this axis is not quite vertical when the head is held
jin its usual position; it becomes se when the face is inclined slightly down-
avards,
THE SEMICIRCULAR CANALS. sel
size and position of the canals, but the relation just mentioned
appears to exist in all cases. This relation may be most simply
stated thus. In each ear there is one canal (the exterior) ina
plane at right angles to the mesial plane, and two other canals
(the superior and the posterior) in planes equally inclined to
the mesial plane. In no other way is it possible to harmonize
the bilateral symmetry of the two ears with the condition that
each of the three axes shall have two oppositely turned canals
in planes at right angles to it.
EFFECT OF WARMTH IN PREVENTING DEATH
FROM CHLORAL. By T. Lauper Brunton, M.D.,
Sc.D. Edin. Casualty Physician and Lecturer on Materia
Medica at St Bartholomew's Hospital.
SINCE chloral was first brought into notice, and its action inves-
tigated by Liebreich, it has been made the subject of numerous
experiments, and has not only proved a most useful medicine,
but a valuable aid to physiological research. During the stay
made in this country by Professor Stricker four years ago he
used chloral frequently as an anesthetic while making some
experiments with Dr Burdon Sanderson on the circulation in
mammals. At his suggestion I made the following experi-
ments as well as many others which it is quite unneces-
sary to give at length, as they simply confirm the observations
of Liebreich and others. The general results were that the
subcutaneous injection of a solution of chloral induced sleep,
which was light and readily broken if the dose were small, but
passed into coma if the dose were large. In dogs considerable
restlessness was observed before sleep came on. The power of
muscular co-ordination was affected in dogs before sopor was
induced, so that they staggered and fell when attempting to
walk either of their own accord or in obedience to a call. A
similar loss of co-ordination was observed in rabbits, but in
general they and guinea-pigs sat quietly after the administra-
tion of the chloral, and thus the moter affection was less percep-
tible in them than in dogs. In dogs the respiration occasion-
ally became very rapid immediately after the subcutaneous
injection of chloral, but it became slow after the animal began
to exhibit symptoms of drowsiness. In rabbits and guinea-pigs
the number of respirations was also diminished, but a pre-
liminary acceleration was not observed in them. The pulse
was not affected to the same extent as the respiration, and the
heart always continued to beat after the respiratory move-
ments had ceased. One of the most important phenomena,
EFFECT OF WARMTH IN PREVENTING DEATH FROM CHLORAL. 333
and the one to which I wish to call particular attention at
present, is the diminution of temperature which chloral in-
duces, and the extraordinary effects of warmth in hastening
recovery from its action, and preventing death from an over-
dose. The fall of temperature has been noticed by Liebreich
and most other writers, but the effect of warmth applied to
the animal’s body has not, I think, received sufficient attention,
although Dr Richardson has pointed out its usefulness in pre-
venting death. The diminution of animal heat is partly due
in all probability to greater loss from the surface caused by
the vessels of the skin becoming much dilated under the in-
fluence of the drug, and allowing the blood to be cooled more
readily by a low external temperature. It is partly due also
to the diminished production of heat which cessation of mus-
cular action always causes, whether it be induced by simply
tying down an animal so as to prevent motion, or by the ad-
mninistration of curare or narcotics.
Professor Stricker having noticed that the animals on which
he experimented often required a second dose of chloral to
maintain anesthesia, when they were wrapped in cotton wool
so as to prevent loss of heat, and still more when they were
laid in a warm place, I made the following experiments at his
suggestion. They shew clearly that an animal wrapped in
cotton wool may recover perfectly from a dose of chloral which
is sufficient to kill it when exposed to the cooling action of the
air (which in the laboratory was about 20° C.), and that recovery
from the narcotic action is much quicker when the temperature
is maintained in this way, and still more rapid when the animal
is placed in a warm bath. If the temperature of the bath is
too high the animal may die from excessive heat, as I have
shewn in a former paper’.
The bearing of these experiments on the treatment of
persons suffering from an overdose of chloral is so obvious as
hardly to require any observations from me. The patient
should be put to bed, and the temperature of the body main-
tained by warm blankets and hot-water bottles to various parts
of the body, and especially the cardiac region, Warmth over the
1 “On the Effect of Temperature on the Mammalian Heart and on the
Action of the Vagus.” St Bartholomew’s Hospital Reports, Vol. vit, 1871.
oot - DR BRUNTON.
heart is an excellent stimulant to the circulation, which, like
the respiration, is enfeebled by chloral, the heart according to
Rajewsky being more or less paralyzed by the drug. If respi-
ration threatens to fail it should be maintained artificially so as
to allow time for the chloral to be excreted and the normal
functions to be restored.
Expt. I. Into two guinea-pigs of nearly equal size °6
cubic centimetre of a 50 per cent. solution of chloral (equal to
about 5 grains or ‘8 gramme of chloral) were injected sub-
cutaneously.
Temp. of animal.
No. 1.
103°8°F.| 104:6°F.
No. 1 was almost motionless, and was
rolled in cotton-wadding.
No. 2 was left uncovered.
Before injection
After injection
5 min.
27 min.
Lhr. 20 min.
1 hr. 40 min.
2hr. 33 min.
95°2 Sale
94:3 about 80° | The graduation of the thermometer
did not extend so low as 80°, and
the temperature was calculated ap-
proximately, or guessed, by the
height of the mercury.
2 hr. 57 min 93°8 The legs of No. 2 are shivering.
4 hr. 87°3
4 hr. 42 min. 88:4 No. 1 was now deprived of its cover-
ing. It shivered and grunted but
lay in whatever position it was put,
and was still completely narcotised.
5 hr. 27 min. 88'8
5 hr. 42 min. No. 1 awakened.
No. 2 remained as before, motionless,
except for the shivering.
95°8 It did not recover, and died some
little time afterwards.
5 hr. 52 min.
EFFECT OF WARMTH IN PREVENTING DEATH FROM CHLORAL, 335
Expt. II. Into two guinea-pigs, No. 1 weighing 655
grammes, and No. 2 weighing 670 grammes, 1‘1 cub. cent. of
50 per cent. solution of chloral hydrate was injected.
Femp.| Resp. | Temp.} Resp.
102°4°
After injection
Tmin.|101:8 | 89
101° Both animals nearly narco-
tized.
No. 1 put into a warm-air
bath,
No. 2 allowed to lie without
cover,
91°8 24
Kicks vigorously when pinched.
abt.87; 12 | Does not kick when pinched.
13. | No. 1 is now awake.
Temperature of No. 2 cannot
be taken or estimated, as it
is so low that the mercury
does not rise from the bulb
of the thermometer. No re-
spiration is visible, but occa-
sionally the animal opens
its mouth convulsively. No
reflex. It was put in the
warm bath, but respiration
did not become re-establish-
ed, and all signs of life
shortly disappeared.
336 DR BRUNTON.
Expt. III. Into a guinea-pig (No. 1) weighing 272 grammes
was injected °65 cub. cent. of a 50 per cent. solution of hydrate
of chloral. Into another (No. 2) weighing 330 grammes the
same quantity of the solution was also injected. |
No. 1. | No. 2.
At injection 100°9
After injection
7 min. No. 1 fast asleep. It was put
into a warm-air bath. No. 2 |
not quite asleep.
12 min. No. 2 nearly quite asleep. Lies
asitis placed. Reflex move- |
ments are slight.
26 min.
50 min. | 100°7 62
ihr. 26 min.
Shivers very much with ex-
piration, so that the respira-
tions are difficult to count.
Grunts slightly when the
thermometer is introduced
into the rectum.
about
895° | 30 | Both animals are chewing.
88:5 No. Lis awake. Though still
somewhat sleepy it will no
longer lie on its back.
No. 2 eries when pinched. It
was put into the warm bath.
No. 2 is now awake.
2hr. 4 min.
2hr. 38 min.
102° 104
101°8
In this Exp. (No. HT.) the dose was small and guinea-pig
No. 2 recovered, although it was not kept warm, but not till an
hour and a half after No. 1, although the latter was the smaller
animal, and the dose it received was therefore much greater in
proportion to its size.
EFFECT OF WARMTH IN PREVENTING DEATH FROM CHLORAL. 337
Into each of three guinea-pigs 1°I cubic centi-
Expt. IV.
metre of 50 per cent. solution of hydrate of chloral wa
ted
Ss injec
No. 1 weighed 640 grammes, No. 2, 670
rammes, and No. 3, 717 grammes.
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subcutaneously.
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338 DR BRUNTON. -
Expt. V. Into the flank of a guinea-pig (No. 1) weighing
392 grammes was injected ‘75 cc. of a 50 per cent. solution of
chloral, and into the axilla of another (No. 2) weighing 335
grammes ‘9 cc. of the same solution.
Temp. of animal.
No.1, No; 2:
After injection
3-min. No. 1 lies quite quiet. No. 2 put into
a warm-air bath at 98°6°.
5 min. 98-2 98
12 min. 98°3 No. 1is dead. The heart beat after
the respiration stopped.
The respirations of No. 2 became very
rapid and deep after it was put in
the air-bath.
2hrs. 26 min, 111-4 No. 2 was heard to give a grunt, and
on taking it out almost immediately
after it was found to be dead.
The dose was here either too large or the temperature of
the bath rose too high,
EFFECT OF WARMTH IN PREVENTING DEATH FROM CHLORAL. 339
_ Expt. VI. Into each of three guinea-pigs, No. 1 weighing
490 grammes, No. 2 weighing 425 grammes, and No. 3 weigh-
ing 415 grammes, 1'1 cubic centimetres of a 50 per cent. solution
of chloral hydrate were injected subcutaneously.
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This experiment shews the effect of warmth in preventing
death, and the rapidity of recovery when the animal is put in a
warm bath, compared with that which it makes when encased
in cotton-wool.
All these experiments were made in August, 1870, in the
laboratory of my friend Dr Burdon Sanderson, and I gladly
take this opportunity of returning him my most hearty thanks
for the facilities and aid he afforded, and the kindness which.
he then and ever has shewn me.
THE SEPTUM ATRIORUM OF THE FROG AND THE
RABBIT. By F. CHampneys.
For convenience sake, the heart, both of the frog and rabbit, is
imagined with its long axis vertically placed, not in the obligue
position which it has tn situ; and the relations of the various
parts are given accordingly.
Septum AtTrRIoRUM OF FROG.
In order to learn thoroughly the structure of the septum, it was
necessary to have recourse to various reagents. Of these three were
found to furnish together all the necessary data, viz, gold chloride.
solution, silver nitrate solution, and alcohol. In all cases injection |
by means of a barometer-tube was found most effective. Some ex-
perience was necessary to accomplish this satisfactorily; the two
difficulties being, Ist, to avoid coagulation of blood on the septum,
2nd, to avoid tearing the very delicate auricles by attempting too
much preparation, or by trying to tie too many vessels. The heart
should be injected while still beating if possible, at any rate as fresh’
as possible. It should never be removed from the body till after in-
jection, partly to avoid the risk of tearing the auricles, partly to keep
the pulmonary veins, which are much too small to be tied, intact,
so as to be able to maintain the pressure.
Ligatures were placed on the large veins, and all were secured
except the cava inferior, into which a canula was tied. The right
aortic root was tied close to its origin, and a ligature was placed
round the left aortic root as far distally as possible, but not tied.
Silver solution.—Silver nitrate solution of 1 per cent. was simply
injected for fifteen minutes, the left aortic root being cut beyond the
ligature so as to rapidly clear the heart of blood. When the solution
had run through for some time this vessel was tied, and the pressure
maintained by adding fresh solution. The heart was removed quickly
and the septum at once dissected out under water, exposed to the
light, and mounted in glycerim. This gave a beautiful picture of the
endothelium and muscular fibres.
Gold solution.—The heart was injected in the same manner with
water acidulated with acetic acid for about four minutes, the left’
aortic root being left open; then with 3 per cent. solution of gold
chloride for ten minutes, the left aortic root being tied soon after the
beginning of the gold injection, but not at first, in order to remove
the acidulated water. After ten minutes’ injection of gold solution
acidulated water was again injected, the left aortic root was cut to
allow the gold to be removed, and then tied again. The heart was
then removed from the body, and allowed to hang in acidulated
water exposed to the light till it had acquired the proper colour, the
pressure of acidulated water being maintained. This gave a beautiful
picture of the muscular and nervous structures.
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 341
Alcohol was simply injected as above, except that the heart was first
washed out with } per cent. Na Cl solution to clear it of blood, which
otherwise would have coagulated on the septum. The Na Cl solution
was removed in the way above described, the aortic root not being
closed till the entrance of the alcohol was seen through the transparent
wall of the auricle, to cause no disturbance by its mixture with the
Na Cl solution, the latter being in fact removed. The pressure
with alcohol was maintained for some hours, the heart being allowed
to hang in a vessel of alcohol, and subsequently prepared under
alcohol. This gave a beautiful picture of all the constituents of the
septum except. the endothelium, but especially of the connective
tissue and muscular striz.
In dissecting out the septum It was always found convenient to
cut off the apex of the ventricle and to blow a few air-bubbles into
the auricles, especially the left. By this means the somewhat difficult
relations of the very delicate septum are made plain. It should be
prepared from the left auricle.
General structure-—The septum atriorum of the frog is a
complete partition’, as is that of the embryo of the bird and
mammal. With regard to the description of Sindes and Roki-
tansky, that the septum atriorum in an early embryonal stage
is quite compiete, and begins then to form a network, one hole
of which remains as foramen ovale—the question now is,
whether the septum atriorum of the frog corresponds with the
early stage, before the network (in which case the network
never exists in the frog), or whether the network first exists
and then disappears.
There is no trace of a foramen ovale. Sabatier®?, in a work
published in the last two or three months, takes the following
position. The venous division of the heart, he says (pp. 187—
190), is composed of two divisions, the sinus venosus and the
auricle proper; these are quite separate in fish, and really so
in Batrachia, but in the higher orders they generally become
blended. The septum atriorum of Batrachia is the “septum
auriculorum,” and not the “septum sinus venosi,” and the
foramen ovale (foramen Botalli) is found in Batrachia in the
sinus venosus. I have not been able, having so lately gained
access to his book, to examine the sinus venosus’*.
1 Ueber Defect der Scheidewand der Vorhéfe. Fragment yon Carl Roki-
tansky, Wiener Med. Jahrbuch, 1871, p. 113.
2 Etudes sur le Ceur dans la Série des Vertébrés. 1873.
3 He says (p. 190), ‘‘Chez les Batraciens, dont les oreillettes sont uniquement
constitués par les auricules, la cloison antérieure (septum auriculorum) forme
342 MR CHAMPNEYS.
Thus the septum atriorum of the frog corresponds only to
the very restricted anterior portion of the septum atriorum of
Fig. 1.
Septum atriorum of the Frog (diagrammatic), :
Anterior muscle,
Posterior muscle
1. Lower horizontal portion seen from the right.
2. - Upper portion radiating vertically seen from the left.
C. Place of insertion of muscle A.
D. Free lower margin of the septum.
E and F. Nerves.
G. Cavity of the ventricle.
pp
entitrement la cloison des oreillettes. C’est naturellement au point de reneontre
de ces deux cloisons qu’a du se trouver Vorifice de communication des deux
systémes veineux, c’est & dire le trou de Botal (foramen ovale). Cetorifice setrouve
done, chez le crapaud, la grenouille, la salamandre, et peut-étre chez tous les
Batraciens, dans la partie antérieure et supérieure de la cavité du sinus, et ap-
partient chez tous au sinus, et non aux auricules.” Each of the two divisions
has its septum (p. 187). ‘A chacune des deux parties des oreillettes (sinus et
auricule) correspond, chez les Vertébrés qui ont deux oreillettes, une cloison qui
lui est propre; l’une antérieure, placée entre les auricules, ‘cloison des auri-
cules,’ et autre postérieure, ‘cloison du sinus,’ partageant le sinus en deux
compartiments plus ou moins inégaux; le sinus des veins caves, et le sinus
des veines pulmonaires. La rencontre et la soudure de ces deux demi-cloisons,
qui s’4vancent lune vers l’autre, donnent lieu & la séparation complete des deux
oreillettes. L’intervalle plus ou moins grand qui existe entre ces deux demi-
cloisons, avant lépoque de leur rencontre et de leur soudure, constitue précise-
ment l’ouverture & laquelle chez les Vertébrés supérieures on a donné le nom
“Trou de Botal.’”
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 343
Mammals’, the whole of the rest being represented in the frog
by the “septum sinus venosi.”
The septum atriorum of the frog hangs with a thin free
edge, Fig. 1 D, looking into the ventricular cavity. It shuts
off a small room (the left auricle), since it lies much to the left
of the middle line. It is very delicate and transparent; pos-
sessing only im the boundary line between its upper and middle
third any considerable amount of muscular fibres, though these
spread from hence over the whole septum. It is formed of a
ground substance, of muscular fibres, nerve-fibres, and nerve-
cells, and of two layers of endothelium.
The ground substance is immensely rich in connective tissue-
cells, lying in very beautifully contoured wavy fibres; the
whole having an aspect like that of the embryonal connective
tissue of Mammalia. Sometimes the connective tissue-cells are
granulated, at other times they are oblong, stellate or irregu-
larly shaped. Sometimes there are wavy bundles, and between
them fibres arranged like those in a teazed specimen. The
connective tissue fills the interstices in the network of mus-
eular fibres.
The muscular fibres are most marked in the upper part of
the septum. They are derived almost exclusively from two
sources, ‘he greater part come originally from a well-marked
muscular band, which runs in the horizontal furrow seen on
the anterior aspect of the right auricle, and which curving to
the left and then backwards, becomes united with the right
aspect of the septum in a line at the junction of the upper
with the middle third, and at some distance from its anterior
edge, Fig.1,4,c. The triangular space, bounded in front by the
anterior wall of the auricle, on the left by the anterior part of
the septum, between its anterior edge and the insertion of this
It is difficult to decide whether these two “ demi-cloisons”’ of Sabatier corre-
spond with the “zwei Leisten” of Rokitansky (loc. cit. p, 112), for Sabatier
makes no mention of the early embryonal Gitterwerk. The accounts of Roki-
tansky and Sabatier as to the formation of the foramen ovale obviously differ in
this way:—Sabatier considers it as a hole between the septum sinus venosi
and septum auriculorum, Rokitansky as one persistent hole, remaining from the
innumerable holes in the primitive Gitterwerk.
The ‘‘cloison du sinus’ or posterior demi-cloison is the ‘‘valyule du trou
ovale” (p. 189), and all that lies anterior to this in the mammalian heart corre-
sponds to the ‘‘cloison des auricules” or anterior demi-cloison, the same pro-
bably as the “eigentliches septum atriorum” of Henle. Gefésslehre, Fig. 6.
1 Sabatier, Pl. xv. Fig, 6, 1.
344 MR CHAMPNEYS.
muscular bundle, A, c, and on the right by the muscular
bundle itself, is filled with a very delicate membranous net-
work, almost destitute of muscular fibres. Thus the muscular
band forms the strong edge of an otherwise very delicate
frenum. The second principal source of the muscular fibres
ef the septum is a posterior band, also joining the septum
at the junction of the upper with the middle third, and derived
from the posterior wall of the auricle, Fig. 1,B. This, how-
ever, joins the posterior edge of the septum at once, and forms
no frenum. With it enter the nerves of the septum from
behind. The disposition of these bundles is very similar.
Some fibres run horizontally across from each to the bundle of
the opposite side, some curve upwards, others downwards. The
posterior bundle further requires remark, since the fibres, enter-
ing in a flat bundle, are folded on themselves in the same
way as the fibres of the tendon of the pectoralis major muscle
of man. Viewed from the right side, the fibres.at first in-
ferior, Fig. 1, B1, run superficially and quite horizontally and
join those of the anterior bundle; those originally next above
them cross under (to the left of) them and run forwards and
downwards, while those originally most superior hie deepest
(most to the left) of all. and have a nearly vertical course,
Fig. 1, B2. This arrangement seems to denote a somewhat
complicated development. It will be seen that the line of
junction of the upper with the middle third is by far the most
muscular part of the septum’.
The arrangement of the muscular fibres resembles that in
the frog’s urinary bladder, and im the best gold specimens one
would believe that they are truly smooth fibres, since they are
very thin, and have oblong nuclei like those of smooth mus-
cular fibres. Sometimes the oblong nucleus represents the
1 Sabatier, loc. cit., p. 149, says of the septum of Batrachia: “Une particula-
rité digne de remarque de la cloison inter-auriculaire, c’est son indépendance &
peu plus complete des faisceaux musculaires qui tapissent les parois. En effet,
elle passe au dessus d’eux en y adhérant légérement, mais sans interrompre leurs
parcours et sans influer en rien ‘sur leur direction et leur disposition de telle
sorte que, la cloison enlevée, il ne reste, sur les parois auriculaires, pour ainsi
dire, aucun temoin de sa presence.”
I must differ from the latter part of this statement. The bands which I have
described, always thick and well marked, especially the anterior, prevent this
description from being correct; but, as I have said, the greatest number of
muscular fibres in the septum are derived from these two bands, and apart from
them the septum is almost independent of the auricular walls.
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 345
thickest part in the fibres. But when we prepare a septum
in alcohol, as above described, all the fibres appear striated.
It is to be noticed that naturally isolated fibres of the thinnest
possible diameter are striated like the fibrille in the muscles
of a frog killed in an alcohol bath. The network of muscles is
in some places so thick as to cover the whole microscopical
field, and sometimes they have interstices nearly as extended
as the field of vision in Hartnack, Oc. 3, obj. 8, 25 cent. The
whole septum of the frog prepared in alcohol offers us one of
the most beautiful combinations of the tissues we have men-
tioned.
Nerve-fibres and -cells.
Through the complicated network of muscular fibres and
connective tissue run nervous fibres, the course of which is
marked by ganglion-cells. The physiologists recognise these
ganglion-cells as the centra motoria of the frog’s heart ; compare
especially the experiments of Stannius. Ludwig, Miiller’s
Arch. 1848, p. 140, has described these nerves, as has also
Bidder, Ib. 1852, p. 163°.
To the observations of Bidder I have to add that there are
to be found single medullated nerve-fibres, running a long
solitary course, and furnished at intervals with single ganglion-
cells, often set on alternate sides, like the buds on the twig of a
tree. I have seen one of the two principal nerve-trunks divide
l The latter says, ‘‘ Die beiden gleichstarken Herziste des Vagus an der Vor-
hofswand zusammentreffend sich zu einem Knotenoder Plexus vereinigen, aus
welchem zwei Fiiden von verschiedener Dicke hervorgehen, die divirgirend am
vorderen und hinteren Rande der Scheidewand verlaufen. Der vordere ist
diinner und linger, der hintere kiirzer und stirker, beide aber bilden, sobald
sie mit dem Septum an dem Rande der Kammer gelangt sind, jedenfails eine
starke sehr kenntliche Ausschwellung...... Auf ihrem Verlauf in der Scheidewand
zeigen diese beiden Nerven wiederum reichliche Ganglien-formation, die aber in
Einzelnen grosse Verschiedenheiten darbietet, indem die Kugeln entweder
vereinzelt ligen und auf der ganzen Strecke ohne erhebliche Unterbrechung
sich folgen, oder in gréssere und kleinere Gruppen mit dazwischen liegenden
gangliosen Partheien gesammelt sind. Immer aber, welches auch vorher
das Verhiltniss der Kugeln zu den Fasern gewesen sein mag, erscheint an
beiden Nerven nahe yor ihrem Uebergang’in den Ventrikel ein microscopisches
Ganglion yon welchem aus dieselben in einer kurzen Strecke ganglienlos zu dem
ringformigen klappenartigen Wulst sich fortsetzen, der den Uebergang in den
Ventrikel bezeichnet. Au fdiesem Wege geben beide Scheidewandnervstiamme
mehrere feine, oft nur aus zwei Primitivfasern bestehende Aestchen ab, die ge-
wohnlich bei ihrem Abgange am Hauptstamme, nicht aber in weiterem Verlaufe
mit Nervenzellen versehen sind. Diese Aestchen endigen iibrigens nicht an dem
Septum selbst, sondern setzen sich in die aiissere Wand der Atrien fort.”
VOL. VIII., 23
346 MR CHAMPNEYS.
and then unite again. The longer (anterior) and somewhat
thinner nerye-trunk (F) runs at first horizontally along the
band of muscular fibres formed by the horizontally running
fibres of the two principal muscular bundles, but near the
insertion of the anterior bundle (c) it suddenly curves down
and runs almost perpendicularly to the ventricle. With the
exception of the first part of the course of this nerve, the
nerves run generally independently of the muscular bundles.
The posterior and somewhat thicker nerve-stem (E) runs almost
immediately perpendicularly straight to the ventricle. I must
differ from Bidder in the last sentence of his which I have
quoted; in no case have I found even one single nervous
fibre unconnected with nerve-cells in any part of its course.
The best preparations for studying the nerves are certainly
those with gold choride, and Bidder used no reagents.
The nerve-cells are in some cases imbedded in the substance
of the larger nerves, but often they are laterally placed, though
in close apposition to the nerve. They are mostly pyriform and
unipolar, with a maximum diameter about three times that
of a red blood-corpuscle. Their arrangement and appearance
on the larger trunks correspond with those figured by Mayer’
in “the Sympathetic of the Frog.”
Endothelium.—The Endothelium has the usual character of
large plates, varying in size and shape in different parts of the
septum, and on directly opposite sides. Their shape is irre-
gularly square or long and diamond-shaped, and between these
all intermediate forms are found. The contours are tolerably
straight or very wavy, and silver shews round dark balls in
many places between adjacent cells. Sometimes we find among
them pigment-cells. The endothelium covers both sides of the
septum.
Blood-vessels.—There are no blood-vessels in the septum of
the frog as there are in that of the rabbit. Hyrtl has shewn
that the whole of the frog’s heart is destitute of blood-
vessels.
1 Beobachtungen und Reflexionen tiber den Bau und die Verrichtungen des
Sympathischen Nervensystemes. Fig. 10.
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 347
Septum ATRIORUM OF THE RABBIT.
Methods of Preparation.
Injection of the heart with various reagents was tried, but fur-
nished no good results. The method at last adopted with some suc-
cess was to carefully cut out the septum from the heart, if possible
still beating, to stretch it out on wax with glass pins, raising it a
little from the wax so that the septum was nowhere in contact with
the wax, then to plunge the whole into acidulated water for a few
minutes, then to remove the superfluous water from beneath the
septum by shaking it thoroughly ; then either to plunge the whole
into 3 per cent. gold chloride solution, or to keep it constantly wet
with this solution, by adding the solution continually in small quanti-
ties from a drawn-out glass tube. It was exposed to this fluid for a
time varying from five to ten minutes, according to the thickness of
the septum, then left in acidulated water till it acquired the proper
colour. After this it was removed and laid on a glass slide with
glycerin and covered with a cover-glass. But only in thin septa did
the glycerin make it transparent enough for examination ; in grown-
up animals it was too thick. The best results were furnished by re-
moving the septum, after it was coloured, from the acidulated water,
laying it for some time in alcohol, pressing it gently with a glass rod
to help the removal of the water; then a few minutes in oleum caryo-
phyllorum, and mounting it in damar. This shewed the course of the
nerves and the position of the cells better than any other method,
but the specimens soon became too dark for use.
To learn the course of the muscular fibres the fresh septum in
glycerin, stretched out on the edges of a wax-cell mounted on a glass
slide, furnished the best results ; hyperosmic acid furnished no good
results.
For sections the whole septum was treated as above with gold
chloride, but it was exposed to the solution for fifteen minutes to
penetrate the thicker parts, and only exposed to the light in acidu-
lated water for a short time in order that the surface might not be too
dark. It was then imbedded in a mixture of wax and oil and cut
with free hand in the usual manner.
Nerves.—There was often a considerable, though microsco-
pic, nerve-trunk, which ran in general vertically downwards
across the fossa-ovalis, and was apparently intended for distri-
bution in the septum-ventriculorum, which destination its
direction, its considerable size throughout, and the fact that it
rarely gave off a single branch, seemed to denote. This nerve
was sometimes accompanied by an artery of equal size. Some-
times this nerve ran from the anterior and superior part of the
limbus, near the foramen-ovale, curving backwards; oftener it
23—2
348 MR CHAMPNEYS.
was found at the posterior part of the fossa-ovalis, sometimes
even in the muscular bundle forming the posterior boundary of
that fossa. In two cases I found two or three very minute
ganglion-cells imbedded in this nerve as it crossed the fossa-
ovalis. In addition to this there were sometimes one or two
minute nerves having a similar direction, but not extending
far, and ending in the septum.
In the limbus and muscular boundaries of the fossa-ovalis
there were sometimes many nerves to be seen, having in
general the same direction as the above, and in certain cases a
plexus, which was almost invariably in connection with the
ganglion-mass, to be shortly described, and therefore in the
posterior and superior part of the limbus, above and behind
the fossa-ovalis. In one case where the ganglion was situated
lower down than usual, viz. behind and below the fossa-ovalis,
the plexus was there also, but in one case where no ganglion
could be found in the usual place, the plexus was still there’.
Nerve-cells—The nerve-cells in the septum atriorum of the
rabbit are of very different sizes and kinds. I distinguish three
varieties. (1) The first sort are very small, having a maximum
diameter generally half that of a red blood-corpuscle. These
I have found embedded deeply in the substance of the larger
nerves, and are the only nerve-cells which I have found any
distance removed from the muscular parts; I have for instance
found two or three of these as above described, embedded in a
large nerve as it crossed the fossa-ovalis. They form no gan-
ghon-mass. (2) The second have a maximum diameter about
twice that of a red blood-corpuscle, and are by far the most
numerous. These constitute the principal ganglion masses.
(3) The third are rare. They have a maximum diameter
about five times as great as that of a red blood-corpuscle, and
are found not thrust into the substance of nerves or forming
considerable ganglion-masses, but I have found them connected
with a single medullated fibre, as in the frog. These are
pyriform and unipolar, and resemble those found in the septum
of the frog, except that they are larger’.
1 Scarpa in his work on the nerves of the heart mentions (Tabule Neurologi-
c@, 1794, p. 2) nerves in the auricle, and figures them (Tab. vit. Fig. 2), but only
on the exterior.
2 Lee in his treatise on the nerves and ganglia of the heart (Phil. Trans. of
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 349
With regard to the position of the ganglion-cells I can
affirm that the first variety (1) are the only nerve-cells which
I have found removed from muscular tissue. I have found
these embedded in the substance of the principal nerve which
I have described as crossing the septum, and also in other
large nerves. JI have never found more than six of these
together, and these not in close apposition.
The second variety (2) form the principal ganglion masses.
With regard to the position of these, I can affirm, that, in spite
of some irregularities, I have found them in by far the greatest
number of instances forming a well-marked mass, occupying
a constant position, viz. in the muscular boundary of the fossa-
ovalis at its posterior and superior part, between the openings
of the right vena superior and cava inferior (Fig. 2c). Sec-
tions shew that it les not embedded in the muscular substance,
but on the right side of the septum and superficial to the mus-
cular fibres. It is often in connection with a considerable
plexus of fine nerves, as above stated. The number of cells
varied with the age of the animal, as will be mentioned.
As irregularities I have found this mass lower down than
usual, viz. behind and below the fossa-ovalis, and in one soli-
tary instance, where the place usually occupied by this ganglion
was devoid of cells and of the usual plexus of nerve-fibres,
I found a large mass of several hundred of these cells below
and a little behind the foramen-ovale, but this mass lay on the
left side of the septum. This was in an adult rabbit. I have
also in one large rather old individual found nerve-cells in
nearly the whole of the limbus, but as this was a lamellated
preparation, I cannot give their exact position; I never subse-
Royal Soc. of London, 1849, p. 47), treats, neither in the text nor figures, of the
nerves or ganglia in the auricle, still less of those in the septum. His descrip-
tions are quite general, as the following extract shews) (p. 47): ‘It can be
clearly demonstrated that every artery distributed throughout the walls of
the uterus and heart, and every muscular fasciculus of these organs, is supplied
with nerves upon which ganglia are formed.” If this isso the innervation of the
mammal’s heart must be essentially different from that of the Frog, according
to Bidder’s investigations, for only in the neighbourhood of the septum auricu-
lorum could he find ganglia or nerves.
Neither Searpa nor Bidder mention the nerves or ganglia of the septum
auriculorum in man or in any of the mammals which they have examined. :
Krause, in his work on the anatomy of the rabbit, also says no word on this
subject.
350 MR CHAMPNEYS.
quently found them so dispersed. These were, however, excep-
tions to the vast majority of instances,
Fig, 2.
Diagram of a longitudinal section from the upper half of the septum
atriorum of the rabbit.
A. Muscles cut longitudinally,
B. Muscles cut transversely.
C. Ganglion.
Differences in relation to age.—Two great differences may be
noticed between the septum of young and adult rabbits; age
increases the richness, first of muscular fibres, then of nerves
THE SEPTUM ATRIORUM OF FROG AND RABBIT. 351
and nerve-cells. In the young rabbit the floor of the fossa-
ovalis is entirely composed of connective tissue, and has no
muscular fibres. Sabatier makes the same remark (p. 189) of
Birds as well as of Mammals, and he says that the addition of
muscular fibres is most marked in the larger animals. I have
not had time to study the genesis of the muscular fibres among
the connective tissue of the septum, but as this takes place long
after birth, and as the floor of the fossa-ovalis is very thin in
young animals, there could hardly be found a better place for
studying this most interesting process. I may state, however,
that I have seen no signs of the process being one of gradual
encroachment from the periphery, and imagine the genesis
takes place immediately in situ and independently of the sur-
rounding muscular masses, though these also grow. There is
often a marked growth of muscular fibres, constituting the
vertical band which sometimes divides the fossa-ovalis into two
parts. This absence of all muscular fibres in the young animal
in the floor of the fossa-ovalis is one of which any one may
convince himself, the thinness of the septum favouring the
examination. The muscular and nervous constituents seem to
grow in dependence on one another, for both of these are far
less plentiful in the young than in the old individual; and,
moreover, with the single exception of the nerve which I have
described, which seems destined for the septum ventriculorum,
and the two or three nerve-cells which I have twice found
embedded in it, nerve-fibres were, however small, hardly ever,
nerve-cells never found, except in the close neighbourhood of
muscular fibres.
Anomalies.—The following anomalies have incidentally struck me
in my work :—
(1) In one young rabbit I found a small free band of fibres
running from the anterior border of the septum, i.e. from the anterior
part of the limbus, across the fossa ovalis, and joining the septum
again at its posterior border. Henle (p. 8) says, ‘‘ Zuweilen spannt
sich ein Gitterwerk feiner Fiden iiber die Oeffnung:” this of course
refers to man.
(2) I found, in both a young and an old rabbit, the foramen ovale
entirely shut, but the “valvula foraminis ovalis” with a perfectly free
anterior border. This apparently contradicts the usual account which
says that the foramen ovale is closed by the forward growth of the
352 MR CHAMPNEYS. SEPTUM ATRIORUM OF FROG, &c.
left of the two falces (“ halbmondférmige Saiime,” Henle), or at any
rate shews that this is not always the method. Indeed, from the
observations of Rokitansky the septum is developed as a perfect parti-
tion which afterwards, becomes perforated with innumerable holes
forming “ Gitterwerk,” and the falces are a later growth’.
(3) I have found two openings in the septum, which was very
thin, in an adult rabbit. Neither of these occupied the position of
the norma! foramen ovale, but were more posteriorly placed ; they were
not surrounded by any rim of fibres, nor provided with any valve,
but were simply formed by the divergence of the scanty fibres, in this
case formed simply from the left falx or valvula foraminis ovalis.
These were no doubt persistent holes in the original “‘ Gitterwerk” of
Rokitansky *.
In conclusion I may add, that in the course of my work
I have many times coloured with gold and carefully examined
the semilunar and atrio-ventricular valves for nerve-fibres and
cells, as well as the papillee from both ventricles, but with the
exception of one doubtful case, in which a mitral valve seemed
to have a few cells which looked like nerve-cells close to its
attached border, there was never any trace of either to be seen.
The papille, however, must sometimes, or in some animals,
possess nerve-cells, for Stricker has once seen the freshly
excised papilla from one of the ventricles of a dog, independently
and rhythmically contract.
1 He says (loc. cit. p. 113), “Indem diese Courtine sich weiter herablasst,
erreicht ihr unterer freier Rand endlich die oben beschriebene quere auriculo-
ventricular-Spalte, iiberbriickt dieselbe, sich mit ihr rechtwinklig kreuzend,
und verschmilzt mit deren Lippen, welche unter einem auch mit einander ver-
wachsen. Hs ist der Vorhofsack hiemit in zwei Ratime gesondert,” and (p. 114),
‘Der das urspriinglich edurchbrochene haiitige Septum—das Gitter—umfassende
dickere Rahmen wiachst ringsum.”’
2 He says (p. 115, 3), “‘Dabei kommen seine Licher durch Wachsthum der sie
umfassenden Balken in grosser Anzahl zum Verschlusse, wahrend andere
bleiben und sich vergréssern.”’
PRELIMINARY NOTE ON AN EPITHELIAL ARRANGE-
MENT IN FRONT OF THE RETINA AND ON THE
EXTERNAL SURFACE OF THE CAPSULE OF THE
LENS.—By J. C. Ewart, Student of Medicine, University
of Edinburgh.
Max ScHuLtzeE states that the membrana limitans interna of
the retina immediately invests the vitreous humour to which it
is often intimately connected, that it is partly formed by the
supporting fibres which traverse the layer of the optic nerve
fibres, these fibres being arranged serially and prolonged into
conical flattened enlargements, or after undergoing division,
like roots of a tree, are continuous with several such terminal
dilatations that ultimately coalesce to form the smooth mem-
brana limitans interna.
Kolliker described this membrane as intimately connected
with the retina, but differing from it in consistence; hence he
ranked it amongst the vitreous membranes. Henle also con-
sidered it an independent membrane, to the outer surface of
which the radial-supporting fibres of the retina, with their
expanded extremities, are applied, calling it the limitans hya-
loidea, to shew its identity with the special membrane of the
vitreous humour; but Max Schultze, though fully recognising
the importance of the separability of the m. limitans, and
the difference in resistance between it and the spongy subja-
cent layer, considers it identical in structure with the supporting
spongy connective tissue. That the internal limiting mem-
brane is not composed of spongy connective tissue, or that,
besides the limiting membrane already described, there is
another epithelial membrane lying on the anterior surface of
the retina, may be demonstrated by the following method.
Take a fresh ox’s eye, divide it transversely into an anterior
and posterior part. From the posterior half evacuate the vitre-
ous humour, and pour in some half per cent. solution of nitrate
of silver; in ten minutes wash this out with distilled water, and
expose for a short time in a good light. On examining the
surface of a retina, thus prepared, next the vitreous humour
354 MR EWART.
with a No. vil. objective and No. 4 eyepiece (Hartnack) a
beautiful mosaic of epithelial cells is seen. The cells, varying
from a faint yellow to a dark brown, are multiform, fitting
closely into each other, constituting a continuous layer, which
can be traced over the whole retina. Some of the cells (with
No. vil. and 4) are nearly half an inch long, and from one to
two lines in breadth. Filling up the small spaces between the
larger cells are many small ones, sometimes only half the size of
a blood-corpuscle, and more or less hexagonal, thus forming a
centre, from which larger cells are often seen radiating. The
edges of the cells are slightly irregular but not serrated, the
surface next the vitreous humour is smooth, and if the eye has
been quite fresh, even after the addition of nitrate of silver,
well-marked nuclei may be seen, either as small round spots, or
as irregular crescent-like bodies. When treated with hema-
toxylin these. nuclei are brought well out, and have often the
same irregular appearance. If after staining with silver the
preparation is left some time in strong acetic acid the whole is
rendered more transparent, and the nuclei of the cells, both of
the epithelial layer and those of the vascular sheath, become
visible. The retina being very vascular, at almost any part, on
altering this focus, you see one or more of the vascular sheaths
discovered by His. The sheaths are easily distinguished from
the fine epithelium on the surface by the different arrangement
of the cells, which are generally larger and often run at right
angles to those of the epithelial layer. The epithelium cells of
the epithelial layer over the smaller vessels and capillaries are
long and narrow, bridging over the space like so many small
arches. Over the large vessels sometimes there are two, or
three, lengths of elongated narrow cells; sometimes they have
their ordinary form, but with many small cells between the
large ones, as is well seen where the vessel has ruptured, leaving
the epithelium above entire. The sheaths of the small vessels
and capillaries are chiefly composed of elongated cells similar
to, but better marked than, those figured by Dr Thin in his
paper on the “Lymphatic System of the Cornea’ In each
cell an oval nucleus can be seen when hematoxylin and acetic
acid are added to a silver preparation. Besides these long
1 Laneet, Feb. 14, 1874.
EPITHELIAL ARRANGEMENT IN FRONT OF RETINA, &c. 355
pointed cells in the larger vessels numerous irregular cells run
across the tube, seemingly on a deeper plane; these are best
seen when the silver has entered the lumen of the tube through
its probably staining the cells of the intima. The sheaths
envelope all the vessels, and not only the small and medium-
sized ones as formerly described.
The epithelial layer above described extends over all the
surface of the retina, but I have not been able to trace it be-
yond the ora serrata on to the ciliary processes. Immediately
under it the vessels and connective tissue can be seen, and
between the bundles of connective tissue spaces are left which
in the fresh state are filled with nerve-fibres.
Lying on this layer of epithelium is the vitreous humour.
The vitreous humour is described as having no distinct capsule,
and the posterior three-fourths of it can easily be removed, but
at and near the ora serrata some of the fibres of the outer layers
of the vitreous humour dip into the substance of the retina
through narrow spaces left in the epithelial lining, and thus the
two firmly adhere. The vitreous humour is also adherent by
some of its external and connective tissue-like fibres to the
ciliary processes, but has no attachment to the posterior surface
of the lens.
In the sheep the epithelial layer is easily demonstrated, it
only differs from that of the ox in that the majority of the epi-
thelial cells are smaller. At some parts, especially over the
vessels, there are groups of large cells; these however are not
elongated, as in the ox, but are generally circular, and, when
treated with acetic acid, present a distinct round nucleus with
a nucleolus. The lymphatic sheaths are composed of long
pointed cells with oval nuclei; at a deeper plane numerous
nuclei run across the tube, corresponding to the cells already
described running across the-vessel. On the surface of the retina
next the choroid there is exactly the same appearance, as that
figured by Professor Henle in the epithelial layer of the ol-
factory region of the horse (Fig. 642, p. 834, vol. IL).
In the cat the epithelial cells are nearly all of the same size,
they are smaller but more regular than those of the ox. The
sheaths round the capillaries are especially well marked, and as
in the sheep, numerous nuclei are seen on some of the sheaths.
356 MR EWART.
In the common fowl the epithelial cells are exceedingly small
—few being as large as a human blood-corpuscle. The greater
number are nearly circular, others are four, five, or six-sided,
all uniting to form what appears with a No. vi. a very fine
network. Under this thin layer fine parallel bundles of connec-
tive tissue are seen, which have a striated appearance like a
muscular fibrilla. This striated appearance is due to clear
spaces which, in the fresh state, are filled with fine nerve-fibres.
On the surface next the choroid the inner segments of the cones
—each ending in a bright red or yellow lenticular body—are
seen passing through the round spaces of the external limiting
membrane.
LENS.
On taking the anterior half of an ox’s eye, removing the
vitreous humour from the posterior surface of the lens, and
treating with silver, as before, a layer of epithelium is found-on
the posterior surface of the capsule.
Babuchin says that several authors have found epithelial
cells on the posterior surface of the capsule of the lens, which
probably resulted from the circumstance, that the inner surface
of the anterior capsule has been described as covered with epi-
thelial cells. He thinks it would be more natural to say that
the epithelium which forms the anterior layer, and the direct.
continuation of the posterior, as well as this last itself, is covered
by the capsule. He thinks that the posterior extremities of the
fibres of the lens, which directly abut against the capsule, or
the spherical bodies which arise from the breaking up of these
ends, have been taken for the epithelial cells.
Whatever kind of epithelial cells may have been described,
those seen by treating the capsule as above could not be any of
the structures Professor Babuchin mentions. In the ox the cells
are large, sometimes measuring half an inch each way with
No. vu. Hartnack: they are polygonal: the nucleus is of a
bright yellow colour, and sometimes under that objective seems
as large as a small pea. The edges are irregular, and processes
from one cell dovetail into those immediately in contact with it.
The surface is smooth, and under the cells the outer fibres of the
capsule of the lens are seen. At some parts spaces are left
EPITHELIAL ARRANGEMENT IN FRONT OF RETINA, &. 357
between the fibres which communicate with each other, thus
corresponding to the spaces in the cornea which contain the
branching cells. At a lower level the characteristic serrated
fibres of the lens come into view.
In the domestic fowl all the cells but a few round the
margin are hexagonal—the sides of each being twice as long as
the ends—there is a nucleus about half the size of the one in
the ox, and, as in the ox, there is a nucleolus. These cells are
arranged in parallel rows, and under each row is a large bundle
of connective tissue ; between each bundle is a small lymphatic
space, similar to those between the bundles of connective tissue
of tendons; these spaces as well as the bundles of fibres are
covered over by the epithelial cells—each row of cells fitting
exactly into those on each side of it.
Near the margin of the capsule the cells are nearly square,
and not arranged in parallel rows as in the centre. Immediately
behind this epithelial layer is the vitreous humour, which is seen
to be in contact with epithelial cells nearly all round, but only
adherent at the ora serrata and the ciliary processes.
In a puppy-dog four days old well marked lymphatic sheaths
are seen, similar to those of the retina, immediately under the
posterior epithelium. The cells of the posterior epithelium are,
except at the margins, hexagonal, and the edges are distinctly
serrated. At the margin they are often irregular in shape, and
sometimes not serrated. On the anterior surface of the capsule
of the lens next the aqueous humour the epithelial layer of cells
is similar to those on the posterior surface of the cornea.
NOTE ON AN INTERESTING CASE OF MALFOR-
MATION. By Joun W. Octe, M.D., Physician to St
George's Hospital. (Pl. VIII. figs. 1 and 2.)
THE following description of the case of a man who was born
in the year 1845 without lower extremities was sent to me by
a physician in Brazil, viz. Dr J. A. Alves Ribeiro of Ceara, a
most able and intelligent physician. He is by trade a saddler,
and has always had good health: his parents were quite healthy,
and none of his relations were in any way malformed, and when
he was of full age he had attained the height, as regards his
trunk, of a fully grown person. The trunk and the upper ex-
tremities present nothing whatever remarkable; and on exami-
nation the bones of the pelvis appear to be well formed, and the
penis, the scrotum, the testicles, the nates and anus are per-
fectly natural and well formed: but on each side of the nates,
where the legs ought, as it were, to be attached, masses or
cushions of soft flesh exist without any appearance of a rudi-
mentary limb: -on the left side this mass is simply rounded and
rather small, whilst on the right side it is larger and peduncu-
lated. The whole circumference of the surface corresponding to
the pelvis terminating in a horizontal line surrounding the
body at about two inches above the root of the penis is covered
by hair. The man is of very active and energetic habits, and
though he cannot read or write, has a clear intellect and quick
perception. Huis powers of locomotion are peculiar. He can
ride well on horseback, mounting and dismounting entirely
without assistance. He progresses (or walks) quickly, lifting
the trunk in a remarkable way upon his arms and propelling it
forward: and he runs fast, resting first on one arm and then on
the other, and throwing the trunk forward with a swing from
right to left.
Of the illustrations appended, No. I. shows the man sitting
upright upon the nates; and No. II. shows him recumbent,
thus exhibiting the rounded masses on each side of the nates,
and the surface extensively covered with hair as above
described.
AN ILLUSTRATION OF THE RELATIONS OF THE
CONVOLUTIONS OF THE HUMAN CEREBRUM
TO THE OUTER SURFACE OF THE SKULL.
By Proressor TURNER.
In the last number of this Journal I gave a short account
of the relations of the convolutions of the human cerebrum
to the outer surface of the skull and head. At that time
I had not had prepared a drawing in illustration of these rela-
tions. During the early part of the winter I made a special
dissection of the head of an adult man, and requested Mr C.
Berjeau to prepare a careful drawing of the dissection, which
drawing he has now reproduced on wood. M. Broca and Prof.
Bischoff, the only authors who had, previously to the publi-
cation of my paper, attempted the accurate localization of the
cerebral lobes, determined their relations by drilling holes in
the skull, inserting wooden pegs through them into the brain,
and then, after removing the skull cap, ascertaining the part
of the surface of the hemisphere into which the pegs had
penetrated. This plan:of drilling holes through the skull and
inserting pegs into the brain is one, which though it may
conveniently be employed, when the object is merely to obtain
an idea of the extent of the lobes of the cerebrum in relation
to the surface of the head, requires the expenditure of too much
time and labour, when the position of the individual convolu-
tions is to be established. I adopted therefore another method
in this dissection. With a fine saw I cut out definite pieces of
bone along the sutures and other lines, which marked the
boundaries of the different areas, into which, as I described in
my previous communication, the surface of the skull may con-
veniently be subdivided. As each piece was raised from its
position, the particular portion of the surface of the brain
which was covered by it was drawn, and in this manner a
projection of the convolutions on the outer surface of the skull
was obtained.
Before describing the figure it may be convenient to restate
the names of the regions into which I subdivided each lateral half
360 PROFESSOR TURNER.
of the skull. Taking the sutures as the guides to the primary
divisions, I spoke of an occipital, or post-lambdoidal ; a frontal,
or pre-coronal; a parietal, subdivided into an antero-parietal
or post-coronal, and a postero-parietal or pre-lambdoidal, by a
line drawn vertically through the parietal eminence from the
squamous to the sagittal suture; and asquamoso-sphenoid, The
frontal region was then sub-divided into a supero-frontal, a
mid-frontal and an infero-frontal area: the antero-parietal into
a supero- and an infero-antero-parietal: the postero-parietal into
a supero- and an infero-postero-parietal: the squamoso-sphe-
noid into a squamoso-temporal and an ali-sphenoid region.
The occipital area was undivided.
R. Fissure of Rolando, which separates the frontal from the parietal lobe.
P.O. Parieto-occipital fissure between the parietal and occipital lobes.
S.S. Fissure of Sylvius which separates the temporo-sphenoidal from the
frontal and parietal lobes.
SF, MF. IF. The supero, mid, and infero-frontal sub-divisions of the frontal
area of the skull: the letters are placed on the superior, middle and inferior
frontal convolutions.
RELATIONS OF CEREBRAL CONVOLUTIONS TO THE SKULL. 361
SAP the supero-antero-parietal area of the skull: S is placed on the
ascending parietal convolution, AP on the ascending frontal convolution.
TAP the infero-antero-parietal area of the skull: I is placed on the ascend-
ing parietal, AP on the ascending frontal convolution.
SPP the supero-postero-parietal area of the skull, the letters are placed on
the angular convolution. ;
IPP the infero-postero-parietal area of the skull, the letters are placed on
‘the mid-temporo-sphenoidal convolution.
X the convolution of the parietal eminence, or supra-marginal gyrus.
O the occipital area of the skull, the letter is placed on the mid-occipital
convolution,
Sq the squamoso-temporal region of the skull, the letters are placed on the
mid-temporo-sphenoidal convolution.
AS the ali-sphenoid region of the skull, the letters are placed on the tip of
the supero-temporo-sphenoidal convolution,
VOL. VIII. 24
ON THE PLACENTATION OF THE SLOTHS.
By Proressor TuRNER.
In this Journal, June, 1873, a short abstract was given of a memoir
communicated to the Royal Society of Edinburgh on the Placenta-
tion of the Sloths. This memoir has now been printed in eatenso
and illustrated with four quarto plates, in Vol. xxvii. of the Trans-
actions of that Society. The memoir opens with a short historical
introduction, in which the meagre observations of Rudolphi and
G. C. Carus on the placenta in Sradypus are referred to, and the
comments thereon by Von Baer, H. Milne-Edwards, Owen, Huxley,
and Rolleston. A detailed description of the gravid uterus’ of
Cholopus Hofimanni and of the naked-eye and microscopic cha-
racters of its placenta is then given. The characters of the fetus
and of the remarkable envelope named by Welcker the Epitrichium
are then described. The memoir concludes with a chapter entitled
“General Observations on the Placentation of the Edentata ;” and as
in this chapter the results arrived at by the dissection are summarised,
it is reproduced here—
General Observations on the Placentation of the Edentata.
From the description given of the form and structure of the
placenta in this specimen of Cholopus, and from the accurate inter-
pretation which I am able to offer of Carus’s figure of the placenta
in Bradypus, it is evident that in the Sloths the placenta is not
cotyledonary, 7.e. if we use the term cotyledon, in the sense in which
it is usually employed by zoologists, to express a particular form of
non-deciduate placenta, subdivided into distinct and scattered masses
as in the Ruminants. It can only be called cotyledonary, if the
term be employed, as is sometimes done in speaking of the deciduate
human placenta, as equivalent to lobes. To avoid confusion in the
use of terms, it may be well to speak of the sloth’s placenta as a
dome-like,- multilobate, aggregate placenta, the lobes of which are
discoidal. It is a deciduate placenta in the fullest sense of the
word; for not only is a decidua reflexa shed along with the feetal
membranes, but if the plane at which I separated the placenta from
the uterus be, as I think there can be no doubt it is, the natural
plane of separation of these parts during parturition, then the feetal
membranes carry away with them the deciduous serotina, the curling
arteries, the utero-placental veins, and the intra-placental maternal
vessels. I have been able, therefore, to put on the basis of an actual
demonstration the deciduate structure of the placenta in the Sloth,
which Rolleston, Owen, and Milne-Edwards, from the study of Carus’s
drawing, had regarded as not improbable.
The character of the placentation in the Sloths having now been
determined, it will be interesting in the next place to compare it
THE PLACENTATION OF THE SLOTHS. 363
with what has been recorded respecting the placentation of the other
mammals included in the order Edentata. Unfortunately, however,
we are not provided with such detailed information relative to the
placental characters of these animals as to enable one to make so
complete and exact a comparison as could be desired. Any remarks,
therefore, which may be based on this comparison must be regarded
as provisional merely, and to be subject to revision when more precise
knowledge is obtained.
Of the placenta of the Armadillos nothing further has apparently
been recorded than is contained in the brief statement made by Prof.
Owen’, that it is a single, thin, oblong disc, with which the maternal
deciduous substance is interblended.
‘ Our information as to the placenta of Orycteropus is also equally
brief, and is limited to the observation recorded by Prof. Huxley in
his “Tatroduction to Classification’,” based on the examination of a
Specimen in the stores of the Royal College of Surgeons, London,
that in this genus the placenta is discoidal and deciduate.
Of the Hairy Anteaters, A. F. J. C. Mayer states* that in
Myrmecophaga (Cyclothurus) didactyla a single orificium uteri exists,
and in the gravid state the uterus and vagina are blended into a
common cavity, and form a round membranous sac. The foetus in
the specimen he examined was well developed and strongly haired,
with the head presenting. The placenta was a thick roundish cake
(Kuchen), and lay to the right in a special pouch of the uterus. The
chorion and amnion were distinct, but the erythrois and allantois could
not be distinguished in the torn membranes. Prof. Welcker, in his
Memoir*, incidentally mentions that in J. didactyla the amnion and
chorion cireumscribe in the usual way the border of a fungiform (pilz-
férmig) placenta. The elder Milne-Edwards°® states that he has found
in IV. didactyla a discoid placenta, but composed at its borders of small
branched tufts: it did not appear to be united to the uterus by
a decidua. His son, M. Alphonse Milne-Edwards, deseribed only
last year® a placenta of Tamandua tetradactyla, which had been
hardened in spirit for some years before it came into his possession.
It occupied the larger part of the surface of the ovum, and was not
composed of simple villosities like the placenta of the Pachydermata,
but of vascular very compact vegetations. Its central part was thick
and spongy; its borders well defined, with a smooth chorion beyond
them, The vascular vegetations, he says, do not apparently resemble
the reticulated folds and alveoli seen by Dr Sharpey in Manis. Some
débris of uterine tissue indicated the presence of a decidua, but he
could not speak positively on the subject. The placenta is unilobed,
dome-like in the mode in which it is set on the chorion, and is named
by Milne-Edwards placenta discoidal envahissant. The surface of
1 Anatomy of Vertebrates, 111. p. 731. 1868.
2 P. 104. London, 1869.
3 Analecten fiir Vergleich. Anatomie. Zweite Sammlung, p. 54. Bonn, 1839.
4 Abhand. der Natur. Gesell. zu Halle, Bd. 1x. 1864.
» Lecons sur la Physiologie, rx. part 2, p. 563, note. Paris, 1870.
5 Annales des Sciences Naturelles, xv. 1872.
24—2
364 PROFESSOR TURNER.
the placenta next the embryo did not possess the projections seen by
Carus in Bradypus, or by myself in Cholopus, but was smooth. No
trace of an allantois was found.
From the above descriptions it is clear that in the hairy ant-eaters
the placenta is not diffused over the whole surface of the ovum, but
is localised in a particular area. From Milne-Edwards’ figure the
proportion of chorion occupied by the placenta is about equal to what
T have seen in Cholopus, but the organ is not, as in the latter animal
and in Bradypus, subdivided into lobes. From the expressions Kuchen
employed by Mayer, pilz-formig by Welcker, discoid and spongy by
Milne-Edwards, pere et jils, it is also clear that the organ possessed
considerable thickness; and although the younger Milne-Edwards
could not, from the condition of his specimen, speak positively of the
presence of a decidua, the thick spongy character of the organ with
the compact arrangement of the villi points rather to a deciduate
than a non-deciduate structure.
In the Tardigrada, the Dasypodide, and the Orycteropodidee, we
have evidence then that the placenta is deciduate, and composed of
one or more dise-shaped lobes. In the Myrmecophagide the evidence
is not so complete, though I think it inclines in favour of the
deciduate nature of the placenta.
But when we turn to the Scaly Anteaters we find a placenta of
a very different character. Some years ago Dr Sharpey examined
the gravid uterus of a Manis, which, from the size of the foetus, was
presumably near the full time. He observed several most important
features in the arrangement and structure of the placenta, which he
communicated to Professor Huxley, who incorporated them in his
“‘ Klements of Comparative Anatomy'.” ‘The surface of the chorion
is covered with fine reticulating ridges, interrupted here and there
by round bald spots, giving it an alveolar aspect, something like the
inside of the human gall-bladder, but finer. The inner surface of
the uterus exhibits fine low ridges or villi vot reticulating quite so
much.. The chorion presents a band free from villi, running longi-
tudinally along its concavity, and there is a corresponding bald space
on the surface of the uterus. The ridges of the chorion start from
the margins of the bald stripe, and run round the ovum. The
umbilical vesicle is fusiform.”
In a letter to me, in reply to a request for further information
about this specimen, Dr Sharpey states that it had been in spirit be-
fore coming into his possession, and that the substance of the uterus
and the tissues of the embryo were brown and fragile. An injection,
both of the uterus and membranes, was attempted, but, from the
condition of the parts, was unsuccessful. The elevations on the
chorion corresponded to the finely-corrugated inner surface of the
uterus. “I turned off the chorion like an everted stocking, and got
the arrangement of the allantois, fusiform umbilical vesicle, omphalo-
mesenteric and umbilical vessels. The ramifications of the umbilical
vessels extended generally over the inner surface of the chorion, and
1 Pp, 112, London, 1864.
a)
THE PLACENTATION OF THE SLOTHS. 865
were lifted off with it from the receptacular part of the allantois,
which was very extensive, and passed into the diverticula of the
chorion. The uterine glands were abundant and easily seen, but IE
could never distinctly trace their orifices ; it seemed to me as if the
ducts opened not abruptly, but gradually and funnel-like among the
placental rugee. I think you found this condition in the whale.”
1 have not only to express my thanks to Dr Sharpey for this
additional information, but to state that with great liberality he has
allowed me to examine the original drawings of the chorion and
uterine glands, and the specimen itself as it had been dissected by
him, and to supplement his description by the following particulars.
Although the preparation had now been for many years in spirit of
wine, yet I had no difficulty in recognising the diffused arrangement
of the fine villi of the chorion and the elongated band free from villi,
such as Dr Sharpey had described. In its general aspect the villous
surface resembled the appearance which I had seen and described in
Dalenoptera and Orca’, though the ridges, folds, and villi of the
chorion were finer than those seen in the Cetacea. The condition
and age of the specimen were such as to render it impossible to make
a satisfactory microscopic examination of the structure of the villi.
The two extremities of the elongated chorion were unequal in ca-
pacity, and the non-villous band extended in closer proximity to the
more dilated than to the narrower pdle of the chorion. In the more
dilated end part of the foetus had, in all likelihood, been lodged ; and
it is probable that the poles of the chorion had been contained in two
pouch-like recesses about the size of walnuts, one situated at each
lateral extremity of the transversely elongated uterus. Each com-
municated freely with the general cavity of the uterus by an orifice
somewhat less in diameter than that of the pouch itself, and was
lined by a prolongation of the uterine mucosa. Into the outer end
of each pouch the very fine orifice of the Fallopian tube opened.
T examined carefully the poles of the chorion to ascertain if a spot
bare of villi, similar to what I had seen in Orca and in the mare,
existed. At the narrower end the chorion was torn, so that the
examination was not satisfactory, but the more dilated pole was
entire, and in it no bare non-villous spot was recognised, so that
if, as I suppose, the chorion enters the pouch-like recesses of the
uterus, the villi investing it would have a relation to the mucosa
lining each pouch as the villi covering the body of the chorion have
to the mucosa lining the general cavity of the uterus. I saw no
stellate bare spot on the chorion corresponding to the orificium uteri
similar to what I have described in Orca and in the mare. Branched,
eylindriform utricular glands, as figured in one of Dr Sharpey’s
drawings, closely resembling those T have figured in Orca, and con-
taining plenty of epithelial cells, the form ‘of which was not very
distinct, though apparently columnar, could be seen without difficulty
in the uterine mucous membrane, but I could not precisely ascertain
1 As described and figured in my memoirs in the Transactions of this Society,
Vol. xxyi. pp. 207 and 467, 1871.
366 : PROFESSOR TURNER.
the mode in which they opened on the surface of the mucosa, which
was thickly studded with minute pits or fosse ; it is probable, how-
ever, that they opened obliquely into the bottom of these pits. The
free surfaces, both of the chorion and uterine mucosa, presented,
without doubt, the appearance which one recognises as characteristic
of a diffused non-deciduate placenta.
The placenta, therefore, in Manis, differs in several most im-
portant particulars, both in arrangement and structure, from what
I have described in Cholopus. Not only is it diffused and non-deci-
duate,.but both the allantois and umbilical vesicle remain as distinct
sacs, and the utricular glands persist throughout the gravid uterine
mucosa.
The demonstration, therefore, of the diffused, non-deciduate cha-
racter of the placenta in Manis by Dr Sharpey, and of the multilo-
bate, discoid, deciduate placenta in Cholopus by myself, will render it
necessary for the systematic zoologist to reconsider either the value
of the placenta as the basis of a system of classification, or the pro-
priety of retaining the Sloths and the Scaly Anteaters in the same
order. If the characters of the placenta are to be regarded as of
More importance in classification than those furnished by any other
organ or combination of organs, then it is clear that the non-decidu-
ate Manis can no longer be placed in the same order as the deciduate
Tardigrade, But if the charaeters of the other organic systems in
the animals belonging to the order Edentata, as at present accepted,
exhibit a series of affinities, which to the mind of the zoologist may
seem to outweigh the differences in placental structure, not only
between Manis on the one hand, and Cholopus, Dasypus, and Oryc-
teropus on the other, but, if my inference as to the deciduate nature
of the placenta in Myrmecophaga be correct, between Manis and
Myrmecophaga also—aftinities which would render it advisable that
they should be retained in the same order,—then the placental system
of classification is obviously not universally applicable, and will have
to be abandoned. It would be out of place in this communication to
enter into the consideration of the anatomy of the other organic
systems in Cholopus, and to discuss how far its various organs resem-
ble, or differ from, those of the genera with which it is usually
associated ; but I hope, from the materials in my possession, to sup-
plement this memoir, and draw up, in the course of the next few
months, a detailed account of the structure of this animal, and to
compare it, as far as the materials at my disposal will allow, with the
other animals usually grouped with it in the order Edentata.
Before bringing this memoir to a conclusion, it may not be with-
out interest briefly to compare the placentation of the Sloths with
that of the other orders of Deciduate mammals.
It may help to make this comparison more complete if I introduce
here some observations which I have recently made on the structure
of the unimpregnated uterus of the Sloth. Through the kind permis-
sion of Dr Sclater I received from A. H. Garrod, Esq., Prosector to
the Zoological Society, the fresh carcase of a female Hoffmann’s Sloth,
which had died in the Gardens early in the month of November.
i
THE PLACENTATION OF THE SLOTHS. 367
An injection of coloured gelatine was passed into the arterial system
from the abdominal aorta, and the uterus was then removed from the
abdomen. The uterus was 31 inches long by ;ths inch broad.
The Fallopian tubes were slender, the ovaries the size of peas, and
lodged in peritoneal pouches, The cavity of the uterus was remark-
ably large for a non-gravid organ, and through its somewhat con-
stricted orifice which opened into the vestibule, the finger could be
introduced into the cavity, which shewed no subdivision into cervix
and body.
The mucous membrane both on the anterior and posterior walls
was elevated into longitudinal ridges such as I have already described
in the pregnant uterus, but these ridges terminated about $ths inca
from the orificium uteri, leaving a smooth surface of mucosa, The
mucous membrane was very vascular; the small arteries both in it
and in the submucous coat presented a corkscrew-like twist, the coils
of which were close together, so that there can be no doubt that the
curling arteries of the placenta pre-exist in the mucosa, and merely
grow larger during pregnancy contemporancously with the develop-
ment of the placenta. The veins were larger than the arteries, and
serpentine in their course. The surface of the mucous membrane of
the fundus uteri, and both horizontal and vertical sections through
its thickness, were examined with reference to the presence of tubular
utricular glands. Distinct evidence of their existence was obtained,
though they were much more difficult to see and much less numerous
than in the pig, mare, cetacean or Manis; they were short tubes, and
did not appear to give off more than one or two branches, which ter-
minated in rounded closed ends. They did not lie perpendicular to
the plane of the surface, for in vertical sections they were irregularly
divided, and they were arranged in groups so that they were more
numerous in some than in other portions of the fundus. Their orifices
on the free surface of the mucosa, which were recognised with some
difficulty, were nearly circular in form, and a somewhat elongated
epithelium projected from the wall towards the centre, leaving, how-
ever, a small lumen; the polygonal ends of the epithelial cells were
seen through the walls of the glands as they lay horizontally in the
mucosa. Capillary loops surrounded the glands in the deeper part
of the mucosa. In the smooth part of the mucosa near the orificium,
although a careful search was made, no glands were seen, and the
arteries did not possess the corkscrew-like twist, so that from these
structural differences this part of the mucosa did not present the same
characters as that of the fundus uteri. The connective tissue of the
mucous membrane contained numerous well-marked corpuscles, and
its surface was covered by a layer of cells, only the ovoid nuclei of
which could be defined with precision.
The placenta of the Carnivora is at once distinguished from that
of the Sloth by several striking characters. By its zonary form; by
the presence not only in the unimpregnated uterus, but in the non-
placental area of the gravid uterine mucosa, and in the maternal part
of the fully developed placenta, of utricular glands; by the intra-pla-
368: 3 PROFESSOR TURNER.
cental maternal vessels retaining the form of a capillary net-work";
by the brevity of the umbilical cord and by the persistence both of the
umbilical vesicle and allantois.
In the Insectivora, Rodents, and Cheiroptera again, the placenta,
though with some slight modifications in its shape, forms a single
“discoid” organ, which in some cases at least shews a subdivision into
lobes; the allantois and umbilical vesicle, the latter of which is of
large size in many genera, persist as distinct sacs throughout intra-
uterine life, and no evidence has been advanced that the intra-pla-
cental maternal vessels are dilated into sinuses. Reichert has shewn*
that in these orders a decidua reflexa, more or less complete, exists.
The condition of the mucous membrane, as regards the utricular
glands, exhibits some variations in different genera. Leydig has ob-
served them in the mole*; Ercolani in the hedgehog*; Reichert in
the guinea-pig’; Ercolani in Jus musculus* ; in which animal he says
they are few in number, simple, and slightly sinuous. In the rabbit
there is some difference of opinion as to the nature of the deep de-
pressions which exist in its uterme mucosa, for though Reichert com-
pares the windings and folds of that membrane to the appearance
exhibited by the convolutions of the brain, and speaks (Miiller’s Archiv,
1848, p. 80) of the involutions of the mucous membrane as short and
wide, yet he evidently regarded them as essentially the same as the
tubular utricular glands in the pig, guinea-pig, bitch, &e. Ercolani
again states that the rabbit, instead of possessing utricular glands, has
numerous very short glandular follicles, which are only inflexions of
the epithelial layer, and represent in this animal the uterine mucosa.
These follicles, he says (p. 146), develope largely during pregnancy,
are transformed into a glandular organ, and appear destined to re-
place, during pregnancy, the utricular glands which are wanting in
these animals. I have had the opportunity of examining sections
through not only the non-gravid uterine mucosa of the rabbit, but
through the non-placental portions of the mucosa in the gravid uterus,
which have been carefully prepared by Mr Stirling, and, from the
appearances presented by these sections, there is, I think, good reason
to regard these “glandular follicles” as only utricular glands some-
what modified in their shape. For though the part which lies next
the uterine cavity has not the cylindrical tubular form usually exhi-
1 T am aware that Eschricht, in his important memoir (De Organis, &e.,
p. 24), speaks of the vessels in the feline placenta as exhibiting dilatations, and
that Kolliker (Entwickelungsgeschichte, p. 163) states that in the biteh the
maternal blood-vessels are very strongly developed, and appear as very thin-
walled capillaries 2’ in breadth, but I have not seen in the placenta of this
animal sinuses at all comparable with the sinuses in the sloth, which possess a
transverse diameter of from ‘003 to 008 inch.
2 Beitriige zur Entwickelungsgeschichte des Meerschweinchens in Abhand. der
Konig. Akad. der Wissensch. zu Berlin, 1861.
3 Lehrbuch der Histologie, p. 517. 1867.
4 Sur les Glandes Utriculaires de V Uterus, p. 10. French translation. Al-
giers, 1869.
~ 9% Op. cit. plates i. ii., p. 117, and Miiller’s Archiv, p. 80. 1848.
. ® Sulla le Glandole Otricolari dell’ Utero. Mem. del Acad, dell Sc. di
Bologna, p. 26. 1873. /
THE PLACENTATION OF THE SLOTHS. 369
bited by the utricular glands, the deeper extremities of these follicles
lying next the submucosa present, on transverse section, a circular or
oval form, and on longitudinal section an elongated tubular form,
such as the utricular glands themselves exhibit. Moreover, both the
dilated and tubular portions of the so-called follicles are lined by a
columnar epithelium which projects into their cavity, but leaves a
central lumen. It is clear, therefore, that these follicles or glands
exist in the uterine mucosa prior to impregnation, and are not occa-
sioned by the gravid condition.
The very important observations recently made by M. Alphonse
Milne-Edwards’, on the placentation of the Lemurs, furnish some
material for comparing this group of animals with the sloths. M.
Edwards has examined specimens of the genera Propithecus, Lepi-
lemur, Hapalemur, and Chirogaleus. He found the chorion almost
entirely covered by dense compact villosities constituting a sort of
vascular cushion, the result of the confluence of a multitude of irregu-
lar cotyledons. The placenta had the appearance of a large sac which
almost completely enclosed as in a hood the amnion. This form of
placenta, he calls, bell-like (placenta en cloche), for the villi are most
numerous at the upper and middle parts of the chorion, but almost
entirely disappear towards the cephalic pole. The uterine mucosa
corresponding to the villosities exhibited numerous irregular anfrac-
tuosities, and had developed a caducous layer. An enormous sac-like
allantois was situated between the chorion and amnion. In its general
mode of disposition on the chorion, the placenta of the lemur is not
unlike that of the sloth, but in the latter animal the lobes or coty-
ledons are apparently less intimately blended than in the former,
which has in addition a highly developed allantois; but as no infor-
mation is given on the arrangement of the utricular glands or maternal
blood-vessels, [ can make no comparison between the disposition of
these important structures in the lemurs and sloths’.
In the new-world Monkeys the placenta consists of a single dise-
shaped organ, which is probably also the case in the anthropomor-
phous apes, though in the tailed apes of the old world, as Hunter
to)
and Breschet’s observations have sufficiently shewn*, the placenta is
1 Annales des Sciences Naturelles, xv., 1871.
2 Since this Memoir has been put in type my attention has been directed by
Prof. R. O. Cunningham, to a paper ‘‘On the Lemurs,” by Mr St George Mivart,
in Proc. Zool. Soc. London, May 20th, 1873. Mr Mivart states that from a
private communication made to him by M. Alphonse Milne-Edwards, that
naturalist is now of opinion that the Lemurs have no decidua, and that the
placenta is diffuse. In does not appear from Mr Mivart’s paper whether M.
Milne-Edwards had received additional specimens since the publication of his
Memoir quoted in the text, in which, when describing the uterine mucosa of
Propithecus, he says, “Et la surface en est hypertrophée fagon & former un
couche caduque, trés-analogue a celle qui, dans une tresfaible étendue, adhére
au placenta discoide des Singes, des Insectivores et des Rongeurs.” From this
extract it is clear that when his Memoir was written, M. Milne-Edwards had no
doubt of the presence of a decidua.
3 Figures of the placenta in the Quadrumana, or descriptions of its naked-
eye characters, will be found in John Hunter’s Collected Works, tv. 71; Plates
XXXY., XXXvi. and fig. 2, xxxiv.; in Rudolphi’s Memoir, ‘‘ Ueber den Embryo der
370 PROFESSOR TURNER.
subdivided into two large lobes by a greater or less interval, John
Hunter has pomted out that each of these large lobes is made up of
smaller ones, united closely at their edges—a feature which Breschet
also has confirmed (op. cit. p. 445). The subdivision of the placenta
in these monkeys into two parts is interesting in connection with the
arrangement seen in the sloth’s placenta, where a partial separation
into a right and left lateral half was found, each of which in its turn
consisted of smaller lobes. But though the form of the placenta, the
existence of a decidua, the absence of the sac of the allantois, the
absence, or at least the rudimentary condition, of the umbilical vesicle’,
the arrangement of the amnion, and the comparative length of the
wmbilical cord, have all been determined in the quadrumana, and cor-
respond in most particulars with what I have described in the sloth,
yet there is, unfortunately, a want of precise information on the
arrangement of the maternal blood-vessels, and the condition of the
utricular glands in the fermer group of animals. John Hunter, in-
-deed, says, veins or sinuses were placed in the fissures between the
lobes which received the blood laterally from the lobes, and that the
substance of the placenta seemed to be ‘cellular,’ as in the human
subject." Dr Rolleston, from the examination of a spirit-preserved
placenta of Macacus nemestrinus (op. cit. p. 301), obviously inclines
to the view that in it intraplacental maternal sinuses existed, though
he points out that, from the age of the specimen, the examination was
not so satisfactory as he could have desired. Prof. Ercolani states’,
from an examination of a specimen preserved in spirit, of the placenta
of Cercopithecus sabaeus, that in this monkey the placenta in its micro-
scopic characters, as well external as internal, does not present any
difference from that of woman, so much alike are they, that as he had
just described the human placenta, he did not think it necessary to
go into the details of structure in the ape. He does mention, how-
ever, that the intraplacental lacune in Cercopithecus, which contain
the maternal blood, are smaller than in woman; and that on the
uterine face of the placenta are manifest traces of serotina, which is
continued on to the fetal villi forming the external membrane, or the
walls of Ercolani’s glandular organ. As he had stated on p. 36 that
he had never seen utricular glands in the human placenta, we must
assume trom the similarity in structure that he had not observed
them in this monkey.
As I could find no record of observations on the utricular glands,
even in the unimpregnated uterus in the quadrumana, I examined
the mucosa of a non-gravid spider-monkey, apparently the Ateles gri-
cescens, preserved in spirit. I found numerous short glands, dilated
Affen,”’ already quoted; in Breschet’s important Essay in Memoires de U Institut,
1845, xix.; and in Owen’s Comparative Anatomy of Vertebrates, 11. 747, im
which volume it is mentioned that the pregnant monkey, dissected by Hunter,
the name of which that anatomist had omitted to give, was a specimen of
Macacus rhesus.
- 1 Compare Breschet’s description of Simia Sabaea with that of Simia nasua,
pp. 444, 470.
. * Mem. del Acad, di Bologna, 1870, p. 53.
THE PLACENTATION OF THE SLOTHS. 371
into pouch-like recesses, and containing an abundance of epithelium,
opening into the uterme cavity by constricted mouths. Some few
which possessed the form of short tubes were interspersed amidst
these pouch-like glands. In this ape, as in the human female, the
mucosa was in close relation to the subjacent muscular coat, and was
not attached to it, as in the sloth and mammals generally, by a lax
coat of submucous connective tissue.
I owe to Dr Rolleston the opportunity of examining a slice of the
placenta of the Macacus nemestrinus in the Oxford University
Museum. Both to the naked eye and with a simple microscope, a
section through the organ shewed a spongy character similar to that
exhibited by the human placenta. The arborescent arrangement of
the villi, and the processes of decidua, prolonged from the serotina
into the interior of the placenta, corresponded to Dr Rolleston’s
description. J examined the bud-like offshoots of the villi micro-
scopally; the capillaries filled with a yellow injection were arranged,
not in loops, but in networks. Between these vessels and the peri-.
phery of the villus, a relatively thick layer of tissue intervened,
which seemed to consist of two strata; one next the capillaries, which
consisted of cells such as Ercolani regards as the glandular organ,
continuous with the serotina; the other, and more external, was
apparently composed of flattened cells which, on the supposition of an
intra-placental circulation of maternal blood, and an involution of the
utero-placental vessels, probably represents the wall of the maternal
blood-vessels reflected on to the villus. A microscopic examination
of the serotina displayed numerous large fusiform cells with ovoid
nuclei, mingled with which were circular or spherical cells possessing
granular contents and nuclei often of a globular form. The processes
of the decidua contained fusiform cells smaller than those just de-
scribed, and a nucleated protoplasm in which a differentiation into
definite cell-forms was not clearly demonstrable.
The great attention which has been paid by various eminent
anatomists to the structure of the Human placenta, enables one to
institute a closer comparison between it and that of the sloth than
could be done with regard to the quadrumana. In man the lobes
are more closely fused together into a single organ, though the
original subdivision into separate lobes is not unfrequently shewn by
deep fissures extending from the uterine surface deeply into its sub-
stance, and in a case described by M. Breschet’ the outer face of the
chorion is said to have exhibited, not a regular placenta, but a number
of distinct cotyledons. But, further, cases have been seen® in which
the human placenta was divided into two not quite equal parts, a
condition which is normal in the tailed monkeys of the old world, and
an approximation to which, as I have already stated, is seen in the
placenta of the sloth. Some recent observations by Reichert’, on a
young human embryo, at the twelfth or thirteenth day, have shewn
1 Répertoire Général, p. 3, Paris, 1826.
2 See Hecker, as quoted by Dr J. Matthews Duncan, in Edinburgh Medical
Journal, Novy. 1873.
3 Reichert und du Bois Reymond’s Archiv, p. 127. 1873.
372 PROFESSOR TURNER.
that, at this very early period, distinct, elevated islets or cotyledons
had formed in the uterine mucosa, and a median cleft separated these
islets into two halves, so as to give a bilaterally symmetrical arrange-
ment. If, as is not improbable, these islets are the rudiments of the
lobes of the future placenta, the human placenta approximates in its
bilateral arrangement at this stage of development to what I have
seen in the sloth.
Both in the human and sloth’s placenta, curling arteries and utero-
placental veins are present, though in the former their subdivision in
the substace of the placenta into branches cannot be followed out as
in the latter, in which animal, moreover, the dilatation of the veins
into large sinuses within the muscular wall of the uterus and the
serotina does not exist as in the human female. In both, intra-
placental maternal sinuses, communicating on the one hand with the
curling arteries, on the other with the utero-placental veins, are met
with. In the sloth, however, these sinuses retain their individuality,
their walls can be isolated from the adjacent villi, they have a tubu-
lar form, and their arrangement as an anastomosing network is pre-
served’. The mode in which they wind in and out between the
villi bears a strong resemblance to the description and figures given?
by Dr Priestley of the arrangement of the maternal vessels in a
human embryo at the second month; only in his case the vessels,
though described as “capacious capillaries,” were not dilated into
large sinuses as in the sloth. In the fully formed human placenta,
on the other hand, though the walls of the curling arteries and utero-
placental veins are distinct structures, yet the intra-placental maternal
blood-sinuses are not tubular, but consist of a system of irregularly
formed and freely communicating spaces. Whether they possess
delicate walls separating them from the tissue of the feetal villi, or
whether the villi float naked in the mother’s blood, are questions
which have been much debated amongst anatomists. Every one who
has examined the villi of the human chorion is familiar with the
layer of nucleated cells which invests the vill like a cap. It has
been repeatedly figured, and is seen to lie immediately outside the
single or double capillary loop, which the bud-like offshoots of the
human villi contain. Opinions are divided whether this layer of cells
belongs to the villus, or is a layer of cells derived from the decidua
ser otina, which has become blended with the tissue of the villus ; but
whether we regard these cells as proper to the villus, or as belonging
to the serotina, no corresponding layer was seen in the sloth, In the
course of observations made during the past two years on the minute
structure of the human chorionic villi, I have more than once seen
an appearance which led me to believe that outside this layer of well-
defined cells, z.e. nearer, or rather next to the maternal blood, was a
layer of squamous cells, which would represent, therefore, the endo-
thelium of the maternal blood-vessels, blended with the villus owing
1 The arrangement in the sloth is, indeed, not unlike what E. H. Weber
conceived to be the arrangement in the human placenta.
2 Lectures on the Development of the Gravid Uterus, p. 62, figs. 19, 20-
London, 1860.
THE PLACENTATION OF THE SLOTHS. 373
to the great expansion of these vessels into the irregularly-shaped,
freely-communicating blood-sinuses. In the sloth, again, as the intra-
placental sinuses retain the individuality of their walls, their endothe-
lum remains in its proper position as a layer of cells lining the
maternal blood-tubes. In the sloth, therefore, the capillaries of the
chorionic villi lie closer to the periphery of the villus than is the case
in the human placenta; but, further, in the sloth these capillaries are
arranged as a distinct network, whilst in man they form single or
double loops. The branches which arise from the stems of the villi
are move elongated, and have more of a laminated arrangement than
is the case in the human placenta,
The presence of a system of maternal sinuses within the placenta
of the sloth, though, as I have just pointed out, it differs in several
points from the corresponding arrangement in man, is of great ana-
tomical interest. For it presents a transitional form between the
simple maternal capillary plexus met with in a diffused, cotyledonary,
or zonary placenta, and the irregularly formed blood-spaces which
exist in the placenta of the human subject. Several obstetrical
writers have, from time to time, denied the existence of intra-pla-
cental maternal blood-sinuses in the human female, and the objection
has been advanced that the presence of such sinuses receives no
support from comparative anatomy. In a communication, read to
this Society, May 20, 1872’, I adduced a number of facts, derived
from the study of the human placenta, in support of the Hunterian
doctrine of the intra-placental circulation of maternal blood, and
I may now add to these facts the confirmatory evidence afforded by
the structure of the placenta in the sloth.
The condition of the utricular glands in the mucosa of the human
and sloth’s uterus is also a feature of much interest. In both, in the
quiescent, non-gravid state, the glands are short, comparatively simple
in form, and are not demonstrable with the same readiness as in the
uteri of animals, which, in the gravid state, possess a diffused, a
cotyledonary or a zonary placenta. In the human uterus, at the
commencement of gestation, the glands are well marked, but as
pregnancy advances to its middle period they disappear, so that no
traces can be seen of them, either in the decidua or in the fully
formed placenta*. Of the condition of the glands, in the early period
of gestation in the sloth, we have no information, but that they are
absent in the later period I have little doubt, for the careful exami-
nation to which I subjected both the placenta and the non-placental
area of the mucosa would have disclosed them had they been present.
Both in the sloth and in the human subject the decidua reflexa is
well marked. The serotina is not, however, so strongly developed in
the sloth, more especially is there a deficiency in the granular colossal
1 Abstract in Proceedings of that date, and more fully in Journal of Anatomy
and Physiology, November, 1872.
* Dr Priestley states (p. 27), that he could see them distinctly in the parietal
decidua near the seat of the placenta in the third month, but they were under-
going granular degeneration, and, instead of being lined with epithelium, were
filled with granules and molecules.
374 PROFESSOR TURNER.
cells, which in the human placenta not only lie on its uterine aspect,
but pass into its substance in the decidual dissepiments.
The great importance in feetal nutrition, which has been ascribed
by Ercolani, in his several valuable and most instructive memoirs on
placental structure, to a system of gland-follicles forming the maternal
part of the placenta, which he believed to be new formed during
pregnancy in all placental mammals, and in which the feetal villi are
lodged, naturally led me to examine the sloth’s placenta with care, to
ascertain if in it such gland-follicles could be recognised. I failed,
however, to see any indications of follicular structure either in
the defined form described by him to exist in the diffused, cotyle-
donary or zonary placenta, or as a layer of cells, belonging to the
decidua serotina, investing the fcetal villi as in the human female.
I have already, in my memoir on the Placentation of the Cetacea’,
criticised and advanced some objections to the general applicability of
Ercolani’s theory, and from the study of the placenta in the sloth
there appear to be additional reasons for doubting that anatomical
unity in placental structure which he advocates. The presence of a
gland-secretion as an osmotic medium in feetal nutrition, whether we
regard it as produced by new-formed gland-follicles, as Ercolani sup-
posed, or by the utricular glands themselves, as Eschricht argued,
does not, I believe, necessarily occur in all forms of placente. That
a white fluid, subsequently termed uterine milk, which serves as
aliment for the feetus, is present in the cotyledons of the ruminants,
was known to Harvey and the older school of physiologists, and it is
very probable that a similar fluid is produced in all placente where
uterine glands or follicles continue to secrete during the whole period
of placental formation. But in those placente, as the sloth, the apes,
and the human female, where an unusual development of the ma-
ternal blood-vessels into large sinuses takes place, a modification in
the anatomical structure is introduced, which seems to render the
presence of such a secretion unnecessary; the utricular glands seen
in the non-gravid uterus disappear, no new-formed follicles are pro-
duced, and the nutritive changes, in all probability, take place directly
between the fcetal and maternal blood.
The amnion in the sloth is related to the chorion, placenta, and
umbilical cord, as in the human female. The sac of the allantois
and the urachus have disappeared, and I could see no trace of an
umbilical vesicle. Further, I may state that the uterus is simple
and uniparous, and that the mammez are two in number and pectoral
in position.
In classifying the Sloths and the other members of the Order
Edentata it has been customary for zoologists to rank them with
the lower orders of mammals. Professor Owen’, for example, directs
attention to the supernumerary cervical vertebre supporting false
ribs, and the convolution of the windpipe in the thorax of Brady-
pus, as manifesting its affinity to the oviparous vertebrata, and to
1 Trans. Roy. Soc., Edinburgh, Vol. xxvi. p. 467.
2 Reade Lecture On the Classijication of the Mammalia, p. 31, 1859.
THE PLACENTATION OF THE SLOTHS, . Bye)
the unusual length of the dorsal and short lumbar spine in Cholopus
as recalling a lacertine structure; whilst the abdominal testes,
single cloacal outlet, low cerebral development, absence of medullary
canals in the long bones in the sloths, and long-enduriug irritability
of the muscular fibre in both the Sloths and Anteaters shew the
same tendency to an inferior type. In his system of classification,
based on the cerebral characters, he places them in the group
Lissencephala, along with the Rodentia, Insectivora, and Cheiroptera.
Professor H. Milne-Edwards, in his most recent defence of his
placental system of classification’, whilst admitting the insufficiency
of the information on the mode of development of the Edentata,
considers that, from the structure of the teeth and the absence of
incisors, these animals have affinities with the Cetacea more than
with other mammals, though they appear to have some relations
with the Monotremata, and he does not hesitate to form a separate
phalanx for their reception. Professor Haeckel, whilst ranking them
amongst the Indeciduata?, admits that the genealogy of the Edentata
is very difficult. Perhaps, he says, they are nothing but a peculiarly
developed offshoot of the Ungulata, but perhaps their root may lie in
-a very different direction.
The comparison which I have just made between the placenta
of the sloth and that of the other deciduate mammals reveals a
correspondence in important features, both of arrangement and
structure, between the placenta of the sloth, that of the human
female, and of the monkeys, greater than exists between it and
the same organ in auy of the other orders of the Deciduata, so
far as has yet been described. This correspondence in placental
form and structure between mammals, which on general zoological
grounds are so widely separated, affords room for much speculation
and thought, and throws a new light, not only on the position of
the sloths in the order Edentata, but on their relations generally to
the placental mammals.
Professor H. Milne-Edwards, in the Memoir above referred to,
argues that similarity in the form of the placenta and in the
arrangement of the membranes is associated with resemblances in
other important structural characters, so that the classification of
mammals founded upon the placenta rests on a natural basis. Thus
Man, the Quadrumana, Cheiroptera, Insectivora, and Rodents, are
grouped together by him in the Micrallantoid legion of the phalanx
Hématogénetes, as they possess in common a discoid placenta, a small
allantois, and a caduca uterina. But further, they are all markedly
unguiculated, their teeth are provided with a covering of-enamel, and
the dental series is continued around the front of the jaws.
As regards their placental characters the Sloths would fall into
this Micrallantoid legion, with which also they would be associated
1 Considérations sur les affinités naturelles et la classification methodique des
Mammiferes, being the first chapter in the Recherches pour servir @ Vhistoire
naturelle des Mammiféres, now in course of publication by himself and his son
M. Alphonse. Preface, dated 27th April, 1868. Paris.
2 Natiirliche Schipfungsgeschichte, Berlin, 1868, p. 480.
376 PROF. TURNER. THE PLACENTATION OF THE SLOTHS.
by their long claws; but in the structural characters of their teeth
and the absence of incisors, they are at once markedly distinguished
from them, so that in these respects the correspondence between
placental form and structure, and these other well pronounced natural
characters, breaks down.
Between Cholopus and Homo the divergence in most of thats
organic systems is so great that it is difficult to find evidence of any
affinity except in their placental characters. With the Prosimii
and Apes, however, affinities may be found. De Blainville had
indeed many years ago’ indicated correspondences in the skeleton
of the Sloths and the Apes, more especially the Gibbons; I may
here refer to the very remarkable vascular plexuses which exist in
the limbs both of the Sloths and Lemurs; and now that I have
called attention to the evidences of aflinity with these higher
mammals it is not improbable that other features of resemblance
may in time be recognised. From the point of view of those who
hold the descent-hypothesis it is possible that between the Sloths
and the Lemurs genealogical relations may exist.
In conclusion, I may state that the study of the placenta in the
Sloth has shewn how difficult it is to predicate, from the arrangement
and structure of the other organic systems, what the character of
the placenta may be, and how necessary it is, before a proper estimate
can be formed of the nature of the placentation, not only that the
form of the organ and the arrangement of the membranes in the
different orders of mammals should be worked out, but the modi-
fications in its minute structure should also be determined. Moreover,
it would seem that affinities in placental form and structure may
exist between mammals which in many other respects are widely
separated, so that the placenta is not in itself sufficient to deter-
mine the position of an animal in the mammalian series, and the
use of this organ as a basis of classification, though in many
instances it may be relied on, yet, from the complex cross-rela-
tions which exist between the several organic systems in the pla-
cental mammals, is not universally applicable,
1 Ostéographie des Mammiféres. Paresseux, p. L.
NOTES OF SOME MUSCULAR IRREGULARITIES. By
Joun Curnow, M.D., London; Professor of Anatomy in King’s
College, London.
Since the publication of my paper in the June number of the Journal
for last year, 42 bodies have been dissected at King’s College, and in
every one, without an exception, some deviation from the normal ar-
rangement of the muscles has been found. Although they were most
frequent in the more muscular bodies, yet important abnormalities
were observed in many very thin subjects, whilst in an excessively
muscular male, they were quite insignificant. I am therefore inclined
to attach less importance to the mere muscularity of the individual
than many anatomists seem to do. I have selected from amongst my
notes some of the rarer forms of irregularity in the muscles of the
neck and trunk, reserving those of the limbs for a future opportunity.
1. Cletdo-occipital. Besides several instances of the usual form
of this muscle, it was twice. made up of a sternal as well as of a clavi-
cular factor. Both were muscular throughout, and on the left side
only, while the usual single variety was present on the right side. In
the first case, the inner head arose from the anterior surface of the
manubrium sterni, just internal to the first costo-sternal articulation,
distinct from, and external to the sternal portion of the sterno-cleido-
mastoid, and the outer head arose from its usual position on the
clavicle a little external to the clavicular portion of that muscle. The
inner fibres crossed the sterno-mastoid superficially, and joined the
outer about the middle of the neck, forming a large muscle quite two
inches in width. The sterno-cleido-mastoid was also very irregular,
for its two parts were almost entirely separate, a few fibres only of
the clavicular portion uniting with the sternal within an inch of the
mastoid process. Its sternal origin was split into three divisions, for,
in addition to the normal attachment, two slips passed down from its
inner border over the manubrium—one to take origin from the lower
half of that bone, and the other to become continuous with a well-
marked rectus sternalis which was also present.
In the second case, the sternal origin of the cleido-occipital was
exactly similar, but its clavicular head was divided into two parts by
an areolar interval one inch wide, and extending upwards fortwo inches.
The sterno-cleido-mastoid in the middle third of the neck gave off a
cross slip from its inner (sternal) portion to the cleido-occipital.
Two doubled-headed specimens of this muscle are described by
Prof. Wood (Phil. Trans. 1870), and its relations to similar forms in
several animals is pointed out, while a third example is recorded by
Mr Bradley in Vol. vi. of this Jowrnal.
2. Rectus Sternalis, (Rectus thoracicus superficialis.) This muscle
was seen in three subjects ; in one on both sides, and in the others
on the right side only. The body in which it was double is the same
as that in which the first cleido-occipital occurred. On both sides
VOL. VIII. DAR)
378 PROFESSOR CURNOW.
the muscles were well developed, one inch wide, crossed the thin
sternal fibres of the pectoralis major superficially,—the left extending
from the above described tendinous prolongation of the mesial tendon
of the sterno-mastoid, and from the adjoining surface of bone for a
slight extent outwards, to the level of the seventh costal cartilage,
where it became attached to the anterior lamina of the sheath of the
‘rectus- abdominis. The other specimens were less complete, for
neither of them passed upwards to the sterno-mastoid, and one only
extended downwards as far as the upper margin of the fifth rib-carti-
lage. There were no tendinous intersections.
3. Supra-costalis. (Rectus thoracicus profundus.) A very
singular form of this muscle was found on the leéft side of a male
subject. It was attached to the upper border of the third rib, just
internal to the origin of the serratus magnus, and passed upwards
behind the pectorals and clavicle, over the upper two ribs, lying
partly on the inner margins of the digitations of the serratus. It
continued along the outer border of the scalenus medius, slightly
overlapping it, and was crossed superficially by the subclavian vessels
and the cords of the brachial plexus, while the external respiratory
nerve of Bell appeared just behind it, It then crossed inwards over
the scalenus medius, giving off a very few fibres thereto, and ended
in a strong but thin tendon, which joined the scalenus anticus at its
origin from the anterior tubercle of the fourth cervical vertebra. The
scaleni were quite normal in every other respect. In most instances
the supra-costalis either stops at the first rib, or is lost in the fascia
over the lower part of the scaleni; but two cases in which it was
blended with the middle scalenus are on record ; viz. one by Lawson
Tait in this Jowrnal (Vol. Iv. p. 236), and another by Pye-Smith in
Guy’s Hospital Reports for 1871. I have found no notice of such an
intimate connexion with the anterior scalenus as is above described.
4, Omo-hyoid. Although this muscle is so very variable, yet
cases of complete duplicity are very infrequent. It occurred on the
left side only of a thin, aged female. Both muscles were digastric,
with an intervening tendon. The more anterior muscle was attached
to the upper costa of the scapula, from half an inch behind the notch
to the angle, and followed the usual course of the normal omo-hyoid
to the hyoid bone. The other muscle was attached to the ligament
over the supra-scapular notch and reached backwards a little behind
the origin of the former. It passed forwards along the posterior
border of, and then a little above the clavicle (to which the tendons
of both were bound down by the cervical fascia), and the anterior
belly joined the sterno-hyoid about its middle. The anterior bellies
of both were of much the same size, and were larger than the
posterior ; the difference being especially marked in the lower or
supernumerary muscle.
5. Sterno-thyroid. An additional sterno-thyroid is more common
than the preceding, and I noticed it once also. It was half an inch
wide, and lay internal to the usual muscle. .
6. Crico-hyoid. On the left side of a male larynx, a bundle of
‘muscular fibres, one-third of an inch in width, extended from the
NOTES ON MUSCULAR IRREGULARITIES. 379
upper border of the cricoid cartilage to the lower border of the body
of the hyoid bone, just internal to the greater cornu. It arose
internally to the crico-thyroid and ran up near the median line, being
separated by a very distinct interval from the inner border of the
thyro-hyoid, with which it was parallel. The only reference to this
muscle is by Zagorsky, in 1806, who gave it the above name. In the
case described by him it occurred on both sides,
7. Digastric. From the junction of the posterior belly with
the tendon, a muscular slip, two inches long and half an inch wide,
passed downwards and inwards over the styloid muscles and the
middle constrictor of the pharynx, and was blended with the upper
edge of the inferior constrictor, a little external to the thyroid carti-
lage. ‘The styloid muscles were quite normal.
In another subject a thin digastric muscle (which I looked on as
an abortive Occipito-hyoid), was attached behind to the fascia on the
superficial surface of the splenius capitis near its insertion. It
crossed the sterno-mastoid lying beneath the platysma and was lost
on the deep cervical fascia between the carotid sheath and the hyoid
bone. It was present on the right side only. Fasciculi somewhat
similar to these are described by Perrin in Vol. v., and by West in
the last number of this Journal.
8. An additional small muscle, triangular in shape, was once
seen in the sub-occipital triangle. It arose from the posterior tubercle
of the atlas, close to the rectus capitis posticus minor, and crossing
between the recti muscles was inserted into the occipital bone
external to the greater rectus and under cover of the superior oblique.
9. Rectus abdominis. On both sides of a moderately muscular
male, the posterior division of the sheath of this muscle ceased
abruptly about an inch below the umbilicus, and from this point to
near the pubis, a large quantity of wnstriped muscular fibres lay on,
and was intermixed with the fascia transversalis, which was less
developed than usual. The fibres were principally transverse, but
some crossed superficially in every direction, and their appearance
forcibly reminded one of the bladder after its peritoneal investment
has been stripped off. They became gradually indistinct externally,
internally, and below, and did not reach the linea semilunaris,
the linea alba, or Poupart’s ligament, but above a few fibres were
continued behind the sheath for a very short distance. The deep
epigastric artery was immediately superficial to the muscular layer,
having pierced the fascia transversalis very low. The character of
the fibres was determined microscopically. I can find no description
of such an arrangement, and Prof. Wood tells me that he has never
seen unstriped muscle in this situation.
bo
Or
ho
NOTES OF A DISSECTION OF AN EXCISED ELBOW.
By G. J. Matcotm Situ, M.B., Demonstrator of Anatomy in
the Unwwersity of Edinburgh.
Ear.y in the present session a male subject, apparently about 65
years of age, was brought into the Dissecting Room of the University.
On examination, before dissection was commenced, it was found that
the right elbow-joint had at some former period been excised by the
H method; and the result evidently was a flail joint, and an almost
useless arm, proved by the atrophy of the limb, and the conditions on
dissection. 7
As one seldom has a chance of examining the result of excision of
this joint, I gladly availed myself of the opportunity afforded, and
append the notes which I made of the dissection.
Exanination before dissection.—The whole limb was atrophied,
and presented a marked contrast to that of the opposite side. The
cicatrix on the back of the elbow was in the form of the letter H,
and apparently of considerable age. The forearm was shortened, the
superior extremities of the radius and ulna resting on the front of the
lower extremity of the humerus: it could not be flexed more than a
degree or two past a right angle, but could be completely extended.
There was very slight lateral motion. On rotating the forearm with
one hand, with the fingers of the other hand manipulating the elbow-
joint, it was found that both radius and ulna moved on the humerus,
in fact that there was little or no independent motion of the head of
the radius. Careful examination of the relative position and shape
of the new joint was easy, on account of the atrophy and thinness of
the limb, and of the absence of any new bone round the articulation.
The humerus ended in two sharp projections, the outer extending
one and a half inches lower down than the inner. The inner of these
projections was posterior, inferior, and external to the upper end of the
ulna. The breadth of the humerus at its lower extremity was one
inch and a half. The upper end of the radius lay in front of the
inner projection of the humerus, and behind the ulna. The ulna ter-
minated superiorly in a process which ended in a sharp projection,
and which lay in front of the outer projection of the humerus, sepa-
rated from it by about half an inch of soft parts. The forearm was
shortened and the upper ends of its bones were one inch and three
quarters above the lower end of the humerus.
On reflecting the skin and fascia of the limb the biceps was found
much atrophied (as were all the other muscles) ; its surfaces looked
outwards and inwards, its inner border being turned forwards. The
long tendons of insertion curved round the upper extremity of the
ulna to be inserted into the upper end of the radius. The fibres
of the brachialis anticus passed downwards and forwards, arising
MR SMITH. DISSECTION OF AN EXCISED ELBOW. 381
from the whole of the external half of the anterior and lower aspect
of the humerus, to be inserted into the apex of the superior projec-
tion of the ulna,—this projection then was without doubt the coro-
noid process. The triceps over the lower extremity of the humerus
was very thin, and was inserted by fibrous bands into the ulna, two
inches from its upper margin, and also into the fascia of the forearm.
These fibres of the triceps formed the posterior ligament of the new
joint. The anterior muscles of the forearm arose from the internal
projection of the humerus; those of the posterior aspect arose from
the external projection. These projections were evidently the re-
mains of the condyloid eminences and ridges. With the exception of
the biceps, triceps and brachialis anticus, the other muscles presented
no peculiarity or deviation from normal arrangement worthy of de-
scription.
Arteries.—Brachial artery, tortuous on account of the shortening
of the forearm; lay on coraco-brachialis, brachialis anticus, passed
over the ulnar projection, then curving outwards lay on the radius
and gave off the radial and ulnar arteries one inch below the upper
border of the extremity of the ulna. The ulnar artery was twice
the size of the radial, and after arising from the brachial, where it
lay on the radius, pursued its usual course. The radial artery re-
quires no description. There was a free anastomosis round the joint
between the profunda, anastomotica, and recurrent vessels, which
were all markedly enlarged.
The ulnar nerve, opposite the lower extremity of the humerus,
was expanded and flattened out: the length of this expansion was
three quarters of an inch, the breadth half an inch, This expansion
was attached to the subjacent muscles and fibrous covering of the
joint, and from it two nervous processes arose ; one internal, and of
the normal size of the ulnar nerve in this situation, was one inch in
length, and was the upper cut end of the ulnar nerve; the other
external, at first very thin and connected very slightly to the expan-
sion, after about an inch became thicker and was continued down
he forearm without any peculiarity. Round the extremities of the
three bones was an imperfect fibrous capsule, with a small synovial
membrane, resembling that of a bursa rather than of a joint. There
was almost no cavity, the two synovial surfaces lying in contact
opposite the lower extremity of the humerus, and upper and posterior
surfaces of the radius and ulna. There was no separate or special
articulation between the radius: and ulna, and no sort of orbicular
ligament. The radius, as before stated, inclined behind the ulna,
the tendon of the biceps hooking round the latter bone. The fibres
of the capsule were attached to the summits of the radius and ulna
and to the humerus along the anterior aspect of the lower end of the
bone: posteriorly, the fibres were mainly those of the triceps muscle.
The bones require but short notice, since they corresponded with
what has been described as felt on external manipulation. The
upper end of the radius lying between the other two bones was
smooth, and the biceps tendon was attached to its anterior aspect
382 MR SMITH. DISSECTION OF AN EXCISED ELBOW.
half an inch from the upper border: the sharp projection of the ulna
had exactly the appearance of the coronoid process, and had the
tendon of the brachialis anticus attached to it. The point of greatest
interest was the entire absence of Any new bone: the three bones
had exactly the appearance of normal and healthy bones excised in a
dead subject, save that the sections were everywhere rounded off, and
the eancellated texture was covered in. The bones showed marked
atrophy on their section a few inches above and below the joint.
NOTE ON AN UNUSUALLY LARGE RENAL CALOU-
LUS. By J. A. Russett, M.A. and M.B., Demonstrator of
Anatomy, University of Edinburgh.
Durine the dissection of a female subject apparently above mid-
dle age, last winter, in the Anatomical Rooms of this University,
the left kidney was felt to contain a calculus of large size; and on
removing the kidney and making a more careful examination it was
found that the pelvis of the organ was distended by the stone, which
could also be readily felt in the calices. The sinus contained much
fat surrounding the vessels, and the vein and wall of the pelvis were
adherent where they came in contact. The kidney, which measured
only 35 inches by 12 by 1 inch, was longitudinally bisected from
hilus to outer border so as to display the calculus in situ. The
calculus was now seen to form a cast of the interior of the pelvis,
infundibula and calices of the organ, and its pointed lower end
extended along the ureter as far down as on a line with the lower
end of the kidney. The extreme length of the calculus was three
inches, and it divided into three main branches, one of which was
situated in each infundibulum. These branches gradually enlarged
from the point of division, and attained their greatest size at the free
extremity. That for the upper end of the kidney was longest and
thickest, and that for the middle part smallest.
The calculus was slightly tuberculated and stained brown over
the greater part of the surface, but over the remainder and internally
it was white and presented a crystalline appearance. All over the
surface were seen bright points, due to crystals of ammonio-mag-
nesian phosphate, of which the chemical tests shewed the calculus
to be mainly composed. The structure was dense, tolerably hard,
and though brittle it was not easily crushed. As many of the
papillary apices of the pyramids of the kidney presented a natural
appearance, and the atrophy of the cortical substance, though very
considerable, was not extreme, it is probable that a certain amount
of urine had been secreted by this kidney up to the time of death.
The right kidney of the same subject was normal in size, and had
a large cyst upon its surface.
SINGULAR MALFORMATION OF WRIST AND HAND.
By Epwarp Betiamy, Esq. F.R.C.S.
THE rarity of fusion of the bones of the carpus has led me to place
the following specimen on record. The body from which it was
taken was that of a powerful middle-aged labourer, of . great
muscular development, and with no other noticeable irregularity.
The os scaphoides was nearly normal as far as its surface articulating
with the radius was concerned. The os lunare, cuneiforme, trapezoides,
magnum and unciforme, were fused together into a quadrilateral
mass articulating superiorly with the radius and the ulna, radially,
with a tolerably normal scaphoid and trapezium, and on the portion
of the mass corresponding to the os cuneiforme, with the os pisiforme.
The unciform process was wanting in the mass, the articulating sur-
faces of which were very broad and continued far on to its dorsal and
palmar aspects. The representative of the carpo-metacarpal articula-
tion existed between the anterior surface of the aforenamed mass, the
anterior surface of the representative of the trapezium, and three
peculiarly shaped digits. The trapezium carried a fairly normal
pollex, the phalanges of which were of great length and massiveness.
The anterior articular surface of the fused mass carried two metacar-
pal bones, the inferior articular extremities of which corresponded
almost entirely to the inferior articulating extremities of (1) the
index and middle fingers, (2) the ring and little fingers. The
phalanges were again enormously large. There was a congenital
dislocation of the elbow-joint, which frequently occurs associated with
a malformation of the hand, flexion and extension of which, how-
ever, appear to have been tolerably complete, the acquired articulat-
ing surfaces being very large. The fore-arm was capable of limited
pronation, but not of supination. I regret very much that I am
unable to give any account of the musculature of the hand, as the
specimen came into my possession too mutilated to examine it effectu-
ally. The arrangement of the grooves over the posterior aspect
of the radius leads me to imagine that the movements of this
abnormal thumb were very limited, and an accurate description of
the muscles and their tendons would have been of considerable
myological interest,
NOTICES OF BOOKS.
Catalogue of the Preparations of Comparative Anatomy in the
Museum of Guy's Hospital. By P. H. Pye-Smiru, B.A., M.D.
London, 1874.
Unver the unassuming title of a Catalogue of Preparations, Dr. Pye-
Smith has compiled not only a useful descriptive catalogue of the
comparative anatomical specimens in the Guy’s Hospital Museum,
but a work which may serve as an introduction to the science of com-
parative anatomy. He prefaces the catalogue with an introduction,
in which he points out the leading characters and the principles of
classification of the animal kingdom, and the distribution of animals
both in space and time. The vertebrate specimens are arranged in
physiological series, and an abstract of the leading characters of each
class and order is prefixed to the description of the corresponding
specimens. The invertebrate specimens again are arranged in zoolo-
gical order, and are in a like manner prefaced by a short but succinct
statement of the characters of the different classes and orders. With
this book as his guide, the Guy’s student is in a position to obtain
not only a knowledge of the specimens in the museum, but an intelli-
gent conception of the principles of anatomical science.
Animal Physiology: the Structure and Functions of the Human
Body. By Joun Cienanp, M.D., F.R.S. London and Glasgow,
1874.
The Students’ Guide to Zoology, a Manual of the Principles of
Zoological Science. By ANDREW Witson. London, 1874.
Were anything needed to prove how important a part the study of
the biological sciences is beginning to play in the general education
of the country, the publication in various quarters of so many manuals,
which have for their object to popularize these sciences, would be a
sufficient testimony. Dr Cleland’s little manual is produced by the
firm of Collins and Sons, as one of their Advanced Science Series. In
preparing the work, the author tells us that he has kept constantly in
view the desire of the publishers to supply the information required
for the advanced course of the directory of the Science and Art
Department. As might be expected from so able an anatomist and
experienced a teacher as Dr Cleland, this book furnishes not only a
lucid exposition of the current facts and opinions on human anatomy
and physiology, but not unfrequently groups these facts together in a
new and more instructive manner than had previously been done,
and draws fresh and original conclusions from them. The chapters on
the Nervous System and on Reproduction and Development may
especially be referred to in illustration of the author’s independence
NOTICES OF BOOKS. 385
of mind, when matters of opinion are under consideration. We cor-
dially recommend this book to the notice not only of the general
student, but to students of medicine, as an excellent and useful com-
pendium of physiological science.
Mr Wilson’s book is published by the firm of Churchill, London.
Though called a guide to zoology, it is not so much a guide to the
principles of zoology, in the restricted sense in which it is now cus-
tomary to use that term, asa guide to the general Principles of Biology,
and it would have afforded a better conception of the character of the
book if the author had given it that name. The book does not pre-
tend to discuss the various biological questions of which it treats
from an original point of view, but is intended to give in plain and
intelligible language an account of the opinions which are entertained
on the leading biological questions of the day. The author has suc-
ceeded in writing a readable book, one in which the student may
find much from which he may derive interest and instruction.
A Monograph of the British Annelids. Part 1. The Nemerteans.
By W. C. M‘Intoso, M.D. Printed for the Ray Society,
1873—1874. London.
Amonecst the various important monographs by British naturalists
which have been published by the Ray Society, none exceeds, either
in the beauty of its illustrations or the careful preparation of the
text, that on the British Nemerteans, by Dr M‘Intosh, which the
society has presented to its subscribers for the years 1873 and 1874.
The monograph contains chapters on the habits, food, anatomy and
physiology of this group of Annelids: a chapter on the reproduction
of lost parts, and another on the parasites which infest them. Their
classification, homologies and general distribution are also described,
and an account of the several genera and species is given. Every
page of the book exhibits the care which the author has taken in its
preparation, and the desire he has shown to make it worthy to hold
its place in the series of important monographs produced under the
auspices of the Ray Society.
Grundriss der Vergleichenden Anatomie. Von Cari GEGENBAUR.
Leipzig, 1874.
Proressor GEGENBAUR’S large work on Comparative Anatomy, the
Grundziige, is doubtless well known to most of our readers. As it
treats the subject with an amount of detail greater than is needed by
those who desire only to obtain a general acquaintance with the science,
the author has been induced to prepare a shorter and more elementary
work which he names the Grundriss. This book is not, however,
a mere abstract of the larger work, but exhibits many modifications
in the mode of treating the subject. As a general exposition of the
principles of comparative anatomy, it is worthy of the reputation of
its distinguished author.
REPORT ON THE PROGRESS OF ANATOMY.’
By Proressor TURNER.
OssEous SysteM.—Wenzel Gruber gives in Wém. de ? Acad. Imp.
de St Pétersb. 1875 a minute description, with illustrative figures, of
BonEs SITUATED IN THE FRONTAL FoNTANELLE. He has seen 43 cases
in the 10,000 human crania which, in the course of 25 years, have
been macerated in the Institute for Practical Anatomy. He also
gives an example in a hydro-cephalic skull, and refers to specimens
seen in the skulls of mammals. In the MJemovrs of the same Acad.
1874, Gruber describes and figures a series of crania, both in man
and other mammals, in which the SQUAMOUS-TEMPORAL ARTICULATED
WITH THE FrontTau. He relates two modes in which the articulation
may take place in the human skull: a, by direct union of the two
bones which he has seen in only two crania; 6, by the prolongation
of a process from the temporal to the frontal. He has seen this in 58
human crania. In those mammals in which these bones articulate,
the direct union occurs much more frequently than through the
intermediation of a process. In Reichert u. du Bois Reymond’s
Archiv, 1873, p. 337, SUPERNUMERARY BoNEs IN THE ZYGOMATIC ARCH
are described by Gruber, and on p. 348 modifications in the DenTaL
ForamEn and Myto-nyoip Groove in the lower jaw. Th. Simon in
Virchow’s Archiv, tvi11. 572, makes some observations on PERSISTENCE
OF THE FrontTaL Suture. He found this suture 76 times in 809
cases, 7. e. nearly 10 per cent.; the skulls were from persons between
10 and 100 years old. Of the 809 skulls 452 were males, 357
females, the percentage of cases of persistent suture being 8-4 in the
men and 10°1 in the women. As a general rule the other sutures
were well marked. In some cases the frontal suture was in part
obliterated. As a rule the frontal is not continuous with the sagittal
suture, but begins either to its right or left. 8. M. Bradley makes
observations on the NarionaAL CHARACTERISTICS OF SKULLS in Mem.
Int. & Phil. Soc. Manchester, y. 213. He argues against Retzius’s
mode of classifying crania, He states that the skull-forms of (pro-
bably) every living nation range from extreme types of dolicho-cepha-
lism to extreme types of brachy-cephalism, and constantly present
examples of every intermediate form. His results were obtained by
measuring the outline of the living head in some hundreds of persons
with an apparatus used by hatters. He reproduces many of the out-
line figures to illustrate the paper. T. Zaaijer describes (Neder.
Tijdschr. v. Geneesk, 1874) a ScApHocePHALIC CRANIUM and the
head of a scaphocephalic man. Fr. Merkel gives an account of the
1 To assist in preparing the Report Professor Turner will be glad to receive
separate copies of original memoirs and other contributions to Anatomy.
REPORT ON THE PROGRESS OF ANATOMY. 387
Femur (Virchow’s Archiv, u1x. 237). He describes its position in the
body; its external form; the direction in which weight presses on it,
and its internal structure. A, Kolliker continues (Verh. d. Phys.
Med. Gesellsch. Wiirzburg, 1873) his observations on ABSORPTION OF
Bone and Inrerstir1aL Growth oF Bone (feports May and No-
vember, 1872). It principally consists in an adverse criticism of the
observations of Strelzoff on the same subject, who, in the course of
his researches into the histo-genesis of bone, had come to conclusions
opposed to those of Kolliker. J. v. Rustizky in Virchow’s Archiv,
LIx. 202, relates his observations on the ABSORPTION OF BoNnE and on
giant-cells. He acknowledges the great importance of Kolliker’s
observations on the same subject referred to in Report May, 1872,
but he does not attach the same importance to the giant-cells, as he
considers that absorption of bone may go on without the interme-
diation of these colossal cells. C. Robin describes in his Journal,
1874, 35, some comparative observations on the Marrow or Bongs.
Tretu.—C. Legros and E. Magitot consider, in Robin's Journal,
1873, 449, the Origin AND ForMATION OF THE DENTAL FOLLICLE IN
THE Mammautia. Their conclusions are: 1. The first indication of
the dental follicle is a cordon arising from the epithelial layer of the
mucous membrane of the gum. 2. The cordon which gives rise to
the milk-follicles springs directly from a prolongation of the buccal
epithelium, whilst that for the permanent teeth is a diverticulum
from the primitive cordon. When the permanent teeth are not preceded
by temporary teeth, they arise sometimes from the mucosa, sometimes
from the cordon of the preceding molar. 3. The cordon is invariably
epithelial; its peripheral part consists of the prismatic elements of
the epithelial layer, its centre of polyhedral cells. 4. The extremity of
the cordon constitutes the enamel-organ. 5. The dentary bulb ap-
pears spontaneously in the midst of the embryonic tissue close to the
enamel-organs, 6. The enamel-organ is moulded like a hood on the
dentary bulb. 7. The wall of the follicle is directly produced from
the elements of the bulb, from the base of which it rises over the
sides and summit of the follicle to constitute the sac of the follicle.
8. When the follicle closes the epithelial cordon is ruptured, and the
follicle then loses its connection with the mucosa. 9. The phenomena
of evolution of the follicle, either in the milk or permanent dentition,
are the same. 10. The genesis of a tooth-follicle and a hair-follicle
are identical. C. 8. Tomes describes (Quart. Journ. Mic, Sc. Jan.
1874) the ExisTENCE oF AN ENAMEL-ORGAN IN AN ARMADILLO. In
Tatusia Peba, a mammal which has no enamel on its teeth, the first
histological structure recognisable in the tooth-germ is a well-de-
veloped enamel-germ, and the enamel-organ in its teeth has an
arrangement similar to that attained to by the enamel-organs in the
teeth of mammals which possess a distinct cap of enamel. The pre-
sence of an enamel-organ in the early stage of development of teeth
on which a cap of enamel does not form, had however been previously
described by W. Turner in the Narwhal, in a paper published in this
Journal, November, 1872.
388 PROFESSOR TURNER.
REsPrrATORY System.—E. W. Collins laid before the Roy. Irish
Acad. April, 1874, a specimen illustrating an Accessory LoBE oF THE
Rieut Lune, takenfrom the body of a male subject, aged about 50 years,
in the dissecting-room of the University of Dublin. The specimen pre-
sented a three-fold morphological peculiarity as regarded lung, pleura,
and azygos vein. An accessory lobe, measuring four inches in length
by two and a half in breadth at its widest portion, sprang from the angle
between the root and the upper portion of the right lung, immediately
above the bronchus. It was somewhat pyriform in shape, and rested
upon the right side and front of the bodies of the five upper dorsal
vertebra, in an accessory pleural pouch. The pouch was formed by
a duplicature, which depended from the cone of the pleura. It was
continuous with the costal pleura externally along a line correspond-
ing to the heads of the five upper ribs, internally along the mesial
line of the five upper dorsal vertebree. Between these points it arched
over the accessory lobe, and isolated it from the remainder of the
lung. It extended underneath the trachea, and invested the right
side of the esophagus. The azygos vein, instead of arching over the
bronchus, arched over the peduncle of the accessory lobe, lying in
the lower free margin of the pleural duplicature. This was the only
specimen of the kind that had been noticed in Dublin, though seven
similar cases had been recorded elsewhere. Stress was laid upon the
suggestion of Prof. Cleland, made in this Journal (May, 1870), that an
abnormal course of the azygos vein during the process of its develop-
ment, whereby it drew down around it a pleural fold, and thus isolated
a probably adherent portion of the lung, offered a satisfactory solution
of the mode of formation of this accessory lobe. The author pointed
out that a remarkable confirmation of this theory was to be found in
an unique case recorded by Wrisberg of a similarly situated accessory
lobe of the left lung, where the left azygos or superior intercostal vein
preserved its foetal condition by opening into the left vena innominata.
The author regarded such a case as that described by Pozzi (Report,
vit. 174), and a similar one on the left side by Rektorzik, as merely
higher developments of pulmonary notches. The paper concluded
with an allusion to accessory bronchi in their connection with this
subject.
Bioop-vascuLaRk System.—C, Giacomini described to Acead. di
Med. di Torino, Nov. 1873, a case in which the Porta anp RicurT
Iu1ac Verns freely communicated with each other——W. Macdonald
has published a pamphlet in which he explains his peculiar views on
the Source anpD CouRSE OF THE CIRCULATION in the embryos of
warm-blooded vertebrates (Edinburgh, 1874).——A. Sabatier records
his observations (Ann. des Sc. Nat. 1874) on the TRANSFORMATIONS
OF THE Aortic ARCHES in the vertebrata.
LympH-vascuLar System.—The researches of Recklinghausen,
of Ludwig, and his pupils have of late years added much to our
knowledge of the lymphatics, more especially in their relations to
the serous membranes. E. Klein has just published an important
REPORT ON THE PROGRESS OF ANATOMY. 389
monograph on THE SerRous Mempranes, London, 1873, in which
their structure and relations to the lymphatics are described with
great care and illustrated by numerous beautiful plates. In chapter
1. the endothelium of the free surface of these membranes is described
more especially in connection with the presence of individual polyhe-
dral, club-shaped, or even short columnar cells, with granular contents,
an ovoid or sometimes spherical distinct nucleus, with large shining
nucleolus: these cells he names germinating endothelium. In the
frog they showed amceboid movements. In chapter u. the cellular
elements of the ground-substance are described. In the omentum of
the rabbit two kinds of lymphangial structures are recognised ; a.
patches, the matrix of which consists of groups of ordinary more or
less flattened, more or less branched cells, which on the one hand
multiply by division, so that the patch increases in size, and from
which on the other hand grow up lymphoid cells. The branched
cells lie in the lymph-canalicular system together with the lymphoid
cells, At an early stage of development these patches do not contain
a special system of blood-vessels ; at a later they are especially rich
in capillary blood-vessels ; by growing in length these patches join so
as to form whole tracts; 6. patches and:tracts the matrix of which
consists of a reticulum the meshes of which contain a variable number
of lymphoid corpuscles : they are generally. provided with more or less
abundant blood-vessels. In the omentum of the guinea-pig, cat, dog
and monkey similar lymphangial structures are met with, but from
their greater thickness they are named nodes or nodules. Intimate
relations exist between these lymphangial. structures and the fatty
tissue of the omentum, the former indeed may and does become con-
verted into the latter, and with this conversion the lymphoid cells
diminish in number in the structure. He agrees with Fleming in
saying that fat-cells are transformed branched cells. In chapter 11.
he describes the lymphatics of the serous membranes ; some remark-
able relations of the lymph to the blood-vessels are pointed out, more
especially the invagination of the veins by lymphatics, and capillaries
hanging in a lymph-sac. A growth of a network of branched cells
may proceed from the endothelium of a lymph-sac, which may trans-
form the sac into a cavernous structure in which lymph-corpuscles
lie and fill up the meshes. Hence arise endo-lymphangial nodules,
and as the proliferation increases the nodule extends more and more
along the lymphatic vessel invaginating or accompanying a large
blood-vessel. The relations of the lymph-vessels to the surface
of the serous membrane are described in this chapter. Two kinds
of stomata are recognised, stomata vera and pseudo-stomata. The
stomata vera are of two kinds, a. the mouth of a vertical
lymphatic channel, which is lined by a special layer of endothelium,
and which channel leads into the lumen of a superficial lymph-vessel ;
b. a discontinuity between the endothelial cells of the surface, leading
into a simple lymphatic sinus near the surface, which represents a
cavity lined only on one side with an endothelium, Both kinds of
stomata vera are bordered by endothelial elements of a more or less
distinct germinating character, so that the mouths may he recognised
390 PROFESSOR TURNER.
by the endothelial cells surrounding them differing in appearance
from the ordinary endothelial cells. The lymphatic capillaries and
the blood-vessels are then described. The book concludes with an
important chapter on the pathological conditions of the serous mem-
branes. E. Klein makes a communication to the Roy. Soc. London
(Proceed. Jan. 29, 1874) on the Lympnatics oF THE Lunes. He
points out that the endothelium of the pleura pulmonum consists of
polyhedral or shortly columnar, granular cells with very marked
nuclei, while the pleura costarum has flattened, almost hyaline, endo-
thelial plates. Beneath the endothelium of the pleura pulmonum is a
very thin connective tissue membrane with numerous elastic fibres and
a layer of flattened connective tissue corpuscles. In the guinea-pig a
layer of non-striped muscular fibres lies beneath the proper pleural
membrane, in rats, rabbits, cats and dogs, bundles of unstriped muscle-
fibres occur sparingly. In chronic inflammation these bundles increasé
so as to form a continuous membrane. A system of inter-muscular
lymphatics lined by an endothelium lies in the meshes of the muscular
layer and communicates freely by stomata with the pleural cavity. A
network of anastomosing lymph-vessels lies in grooves which corre-
spond with the most superficial groups of alveoli of the lungs, which
communicates on the one hand with the inter-muscular lymphatics,
on the other with deeper inter-alveolar lymphatics. Lymphatic
capillaries arise in the alveolar septa from branched cells and lead into
lymph-vessels which accompany branches of the pulmonary artery
and vein; they run either in the adventitia of these vessels, or the
blood-vessel is entirely or only half invaginated in a perivascular
lymph-vessel. The branched cells send a process between the epi-
thelial cells into the cavities of the alveoli, which processes Klein
names pseudo-stomata. Other lymphatics are distributed in the
adventitia of the bronchi, thin capillaries originate in the branched
cells of the mucosa of the bronchi and penetrate through the tunica
muscularis. The branched cells penetrate between the epithelial
cells of the mucosa and project on its free surface. Vascular lymph-
follicles are in continuity with the endothelial wall of a lymph-vessel
so that they are surrounded by it, just as a lymph-follicle of a Peyer's
patch is surrounded by a lymph-sinus. The paper concludes with
observations on the pathological conditions of the several structures.
Fr. Tourneux communicates to Robin's Journal, 1874, 66, his
observations on the EpirHenium oF THE SErrRous Mempranes. He
regards the distinction recently drawn between epithelium and endo-
thelium as arbitrary. His observations were made on batrachians.
G. Thin describes and figures (Lancet, Feb. 14, 1874) a Lym-
PHATIC SYSTEM INTHE CorNEA. He has seen endothelial lined lymph-
vessels traversing its substance, the endothelial cells possessing
precisely the form of the endothelial lining of the lymphatics. He
regards these vessels as the corneal tubes of Bowman and believes
that the mercury in Bowman’s injection had passed along these
vessels. He describes lacunse in the cornea in which the cornea-
corpuscles lie, which lacunee communicate with each other and with
the lymphatic system. These lacune are lined by an endothelium
REPORT ON THE PROGRESS OF ANATOMY. 391
which is directly continuous with that lining the lymph-vessel. He
states that nerves lie in the lymph-vessels and nearly fill them, a
narrow space only being between the nerve and the wall of the lym-
phatic.—On THE LyMPHATICS OF THE NORMAL NON-PREGNANT UTERUS.
Dr G, Leopold has arrived at the following conclusions with regard
to the lymphatics of the normal non-pregnant uterus (Archiv fiir
Gyndcol. Vol. v1. Abst. in Lond. Med. Rec. 1. 35).
I. Mucous Membrane.—1. The mucous membrane consists of a
framework of the finest connective tissue, the bundles of which are
covered with endothelium, and whose interspaces form the lymph-
spaces (Lymphrdume). 2. In the deeper layers, the membrane of the
glands consists of a fine layer of delicate connective tissue bundles,
whose epithelium lies externally, but more superficially it is formed
only of a sheath composed of the cell-plates (Zellplatten, pldttchen-
Jormigen Zellen). 3. The blood-vessels, from the finest capillaries
onwards, have a number of fine endothelial sheaths increasing with
their size. 4. The framework of connective tissue stands by means
of fine twigs in direct connection with both sorts of sheaths. 5. The
glands and blood-vessels, therefore, pass directly through the lymph-
spaces, separated only from the latter by their sheaths, formed from
the framework of connective tissue. 6, At the limits of the muscular
layer the lymph-spaces reach a short distance into the filter-shaped
hollows between the muscular bundles, and become gradually nar-
rowed into the intermuscular lymph-vessels and spaces.
If. Muscular Coat.—1. The muscular layer contains in animals
and in the human subject lymph-vessels and lymph-spaces (Lymph-
spalten). The walls of each are formed of the fine intermuscular con-
nective tissue. The former are lined by fine endothelial lamelle,
which exhibit here and there openings and slits ; the latter are lined
by delicate cell-plates (Zellplatten). 2. In animals, the characteristic
net of lymphatics is arranged parallel to the long axis of the two
layers of fibres ; they therefore cross each other. Those of the inner
layer pass into the lymph-spaces of the mucous membrane, and those
of the outer into the subserous lymph-vessels. The large lymph-col-
lecting tubes, provided with valves, and spread in the form of a net
over the horns of the uterus, lie between the muscular layers, and re-
ceive the whole lymph-vessels from both sides: externally those of
the subserous and outer muscular layers, and internally those of the
inner layers and the mucous membrane. 3. In the uterus of the
human subject the lymph-vessels are much more complicated, on
account of the arrangement of the muscular fibres. They are most
richly developed in the outer layer, and in the other layers specially
in the neighbourhood of the large vessels, and are connected with the
subserous membrane as in animals, but with the mucous more by
lymph-spaces. They come together in the outer layer, especially at
the side of the uterus. 4. The lymph-spaces, in the human subject
and in animals, surround the smaller bundle of a large muscular
bundle, and pass into the lymphatics. In animals, they stand in in-
direct connection with the subserous and CAseD channels ; in the
human subject, however, in direct connection. 5. For the most part,
47
392 PROFESSOR TURNER.
large blood-vessels lie in the neighbourhood of the large collecting
tubes; the other lymph-vessels are partly accompanied by blood-
vessels for a certain distance, and the lymph-spaces are almost regu-
larly penetrated by small vessels.
Ill. Serous Coat.—1. Under the serous membrane only lymph-
vessels are found. They lie in the subserous connective tissue, and
form large characteristic nets. 2. They are much less numerous than
the subserous blood-vessels lying over them, but are from eight to ten
times stronger than the latter. 3. They have large ampulle, points
of union, constrictions, valves, and swellings, and send branches
towards the deeper parts, either in a vertical direction or at an angle.
4, In the pig, rabbit, and sheep, the net has mostly a direction cor-
responding to the long axis of the uterine horns. In the human sub-
ject, on the contrary, they cover the anterior and posterior wall, in
irregular large or small groups, and have, especially at the insertion
of the Fallopian tube, large ampulle, and then pass as an extended
net upon the tube.
Nervous Systew.—In Schultze’s Archiv, 1873, p. 208, Rudolf
Arndt gives an account of his researches into the GANGLIA OF THE
SympatHetic. All the ganglion-bodies (cells) of the sympathetic
are provided with several processes, indeed.all bipolar and multipolar
bodies correspond with complex cells, and are derived from complex
cells. All unipolar ganglion-bodies, on the other hand, correspond
to simple cells and proceed from them. All so-named apolar ganglion-
bodies, when they are large, represent anomalous forms of deve-
lopment of the original formative cells ; when they are small, they
are the formative cells themselves. In the same Archiv, p. 255,
H. Zuppinger relates a method of demonstrating the AXISs-CYLINDER
Processes of the ganglion-cells of the spinal.marrow. F. Darwin
contributes to Quart. Journ. Mic. Science, April, 1874, a paper on
the SyMPATHETIC GANGLIA OF THE BLappeER.in their relation to the
vascular system. The.course of the nerves corresponds in a general
way with that of the chief blood-vessels. Branches from the ganglia
could be traced :to the arteries; veins appear to be very scantily
supplied with nerves. The author distinctly saw delicate nerve-fibres
arise from the cells of a ganglion and supply the neighbouring capil-
laries, which in some cases formed part of the vascular plexus which
surrounds the ganglion. Moritz Benedikt communicates (Virchow’s
Archiv, tix. 395) some observations on the INNERVATION OF THE
INFERIOR CHOROID-PLEXUS. On p. 511 of the same Archiv, R.
Arndt enquires into the Paruonocican Anatomy of the CENTRAL
Oreans of the Nervous System.
ELEecTRIcAL OrcAns.—F. Boll in Schultze’s Archiv, x. p. 208,
gives an account of the structure of the electrical plates of Malapte-
rurus, and on p. 101 of Torpedo.
Kipyey.—R. Heidenhain in Schultze’s Archiv, x. p. 1, contri-
butes to the Anatomy and Physiology of the Kidney.
= REPORT ON THE PROGRESS OF ANATOMY. ~ 393.
Ovary.—W. Romiti makes some observations on the structure
and development of the ovary and Wolffian duct, Schultze’s Archiv,
x. p. 200. He confirms Waldeyer’s statement that the ovary posses-
ses a celiular investment different frem the endothelial celis of the
peritoneum.
Piacenta.—J. Mauthner communicates a short paper (Sitzd.
Akad. der Wiss., Vienna, 1873, p. 118) on the Maternal Circulation
in the Radbit's Placenta, with reference especially to its relations to
the human placenta. When a section is made through the placenta
whilst still attached to the wall of the uterus, a uniformly thick clear
layer, which consists of a very fine network with interspersed nuclei,
is seen next the uterus. Then follows the cellular part of the pla-
centa, commonly named the pl. uterina, but which Winkler has
named basal-platte (Report, vu. p. 163); this layer is much thicker
than in man, and consists of an indistinct, granular-looking conglo-
merate of cell-elements, which possess distinctness only where the
maternal vessels pierce the basal-plaite. Then succeeds the sonamed
feetal part of the placenta, which is subdivided into many small
lobules, separated from each other by furrows. Each lobule contains
a strong-walied maternal artery and comparatively thin-walled veins,
so that each such lobule fer itself alone corresponds to the entire
human placenta, because, in both, arteries and veins are separated
from the fcetal villi by intermediate blood-spaces: but these blood-
spaces differ fundamentally from the human, in that they are in the
rabbit fine capillaries, while in man they are widely expanded spaces..
In both, however, the maternal blood-channels have no proper walls,
but are bounded directly by the epithelium of the villus. The villi
are so adapted to each other that the epithelial investments of adja-
cent villous folds become blended with each other, and the maternal
blood-spaces lie between the two rows of investing epithelial cells.
M. X. Delore in Gaz. Médicale, 21 Feb. 1874, confirms the “ demon-
stration by Weber, Kolliker, Turner and Winkler of the circulation
of the maternal blood through the human placenta.” He finds an
epithelium in the circular sinus, but has not seen it on the placental
villi. G. B. Ercolani communicates (Mem. dell Accad. delle Scienze
di Bologna, tv. 1874) a memoir on the Structure of the Caduca Ute-
ring in two cases of extra-uterine pregnancy. He arrives at the
following conclusions: the whole extent of the uterine decidua has
the power of forming a placenta, bué its actwal place of formaticn is
determined by the position which the ovum takes up; there the
development of the placenta goes on, whilst elsewhere its develop-
ment is arrested. There is ro difference in the decidua in its first
stage of development between a case of normal pregnancy and an
extra-uterine pregnancy ; but while in the normal decidua develop-
ment ceases except in the position of the ovum, in the extra-uterine
it advances further, for a rich vascularity appears amongst the cellular
elements, just as in the serotina, and a partial ectasy of some ef the
vessels, indicating the beginning of the formation of lacuns, takes
place. Whilst the new formation in normal cases only invades the
VOL. VIIL 26
394 PROF. TURNER. REPORT ON THE PROGRESS OF ANATOMY, —
subjacent uterine tissue in the region of the ovum, in extra-uterine
the whole of the subjacent uterine tissue is invaded. He does not
believe in the existence of a mucosa in the human uterus, as he finds
only a fine membrane on which the epithelium rests. As the deci-
dual formation in these extra-uterine cases extended to the musculo-
glandular layer, it could not be formed exclusively of the uterine
epithelium, nor, owing to the absence of a true mucosa, from a sub-
epithelial connective tissue. From the multitudes of new cells, having
the appearance of white blood-corpuscles, he believes them to be
derived from the blood by migration, and he associates with this the
well-known fact, that white blood-corpuscles abound in the blood of
pregnant women. The utricular glands are altered in the decidua in
extra-uterine cases ; in one case at two months large spaces, lined by
layers of conical epithelium, were seen near the muscular layer, and
in the layer of decidua, superficial to these spaces, traces of com-
pressed gland-tubes were seen. The new-formed elements of the
decidua compress the gland-tubes, and the secretion, being prevented
from escaping, dilates the terminal branches, and thus the muscular
bundles are broken up or pushed aside so as to present a trabecular
arrangement. Amidst the muscular elements of the trabecule there
was an enormous infiltration of the white corpuscles, more especially
in the second case at the fifth month of pregnancy, and a similar
infiltration amidst the cells of the serotina. As from the want of the
ovum the feetal part of the placenta cannot be completed, the dilated
vessels of the decidua become stopped up by the formation of clots
in their interior.
REPORT ON PHYSIOLOGY. By Witttam Sriruine, D.Sc.,
M.B., C.M. (Edinb.), jormerly Demonstrator of Practical
Natural History in the University of Edinburgh’.
Nervous System.
Functions OF THE Brain.—Nothnagel (Virchow’s Archiv, tvut.
420) has continued his researches on this subject. He experimented
exclusively on the brain of the rabbit. On puncturing with a fine
microscopic needle a spot lying in the interior of the posterior end
of the cerebrum, the animal sprang from the table and exhibited
unusually violent spasmodic movements, which appeared either at the
time of puncture or a second or so thereafter (at the latest two
minutes), and lasted not longer than three minutes. No alteration
of the sensibility of any part of the body was to be observed. The
author for the present refrains from suggesting any hypothesis as to
the probable cause of these movements. As to the Cornu Ammonis,
on puncturing it with a fine microscopical needle no disturbance
in any direction was observed. Many of the animals so injured had
meningitis with dyspnea, but dyspnea and death without meningitis
was also observed. [These results are at variance with those of
Fournier obtained by the injection-method, Jowrn. of Anat. and Phys.
vit. 179.] More than forty experiments in different ways and
different directions were made on the thalamus opticus. Slight dis-
turbances of the superficial layers were without effect. In a few cases
the paralysis of the extensors of the finger observed by Schiff oc-
curred. If punctured more deeply and towards the middle line, the
limbs of the opposite side were directed towards the middle line.
This was specially and almost exclusively observed of the fore-limbs.
The deviation was the more pronounced, the more basal the direction
of the puncture. The deviation in all cases was only temporary,
disappearing sometimes after a few hours, in most cases after twenty-
four hours. In other cases, immediately after puncture, the head
was turned to the opposite side, the fore-limbs strongly divergent,
the one directed outwards, the other (opposite side from injury)
towards the middle line. No disturbance of sensibility. The phe-
nomena persisted, but with decreasing intensity. After death the
seat of injury was found in the posterior half of the thalamus,
unusually deep towards its base, and near to the region of the
pedunculus cerebri. Horizontal-section of the thalamus from above
downwards was followed by the phenomena already described by
Schiff (Lehrbuch der Muskel- und Nervenphysiologie). No change of
sensibility was observed. Sometimes it appeared that just after
the operation a slight hyperalgesia specially corresponding to the
wounded side occurred, but it was not very convincing. This much
1 To assist in rendering this report more complete, authors are invited to
send copies of their papers to Dr Stirling, Edinburgh University.
26—2
396 DR STIRLING.
is certain, that on division of the thalamus, no anesthesia of the
anterior extremities was produced. “ Extirpation of both nuclei
lenticulares.” Nothnagel, Centralblatt, No. 56, 1873. “ EXAMEN
DE QUELQUES PorINTs DE LA PuHysIoLoGiI—E pU CERVEAU,” par le
Dr Eugéne Dupuy, Paris, Ad. Delahaye, 1873. Dupuy has ar-
rived at results different from those of Ferrier. The conclusions he
draws from his experiments are: (1) It is possible, by irritating
certain limited points of the cortical layer of the brain, to produce
contractions, sometimes of an entire limb. (2) Generally it is the
fore-limb, and of the opposite side to that of the point of irritation,
which is the seat of contraction. (3) The electrical current must
be propagated to the base of the brain in order to excite it; either
to the nerves which arise on the base itself, or medulla. (4) If
the dura mater be excited by electricity, we also obtain contrac-
tions in one of the fore-limbs, generally in a crossed manner. (5) The
fact that a galvanoscopic frog has been thrown into a state of con-
traction when its nerve touched a part of the cerebral mass, far
from the point of excitation, confirms the idea that the electric
current is propagated. (6) Contrary to the results obtained by
Ferrier, we have never succeeded in obtaining the effects upon the
tongue, whether of projection or of retraction. (7) The whole of
the cortical layer of the brain is probably a centre of reflexion of a
certain kind of sensibility, capable of acting in a reflex manner on
motor or sensitive centres, but its integrity is not indispensable to
the manifestation of voluntary and even intelligent actions. (8) In
the case of the animals (dogs) on which he experimented, it was
possible to excite contractions of the muscles of the entire limbs on
the opposite side of the body, even after the removal of the opto-
striated bodies of the opposite side. (Also Lond. Med. Ree. Vol.
u. No. 56.) Carville and Duret, in a communication read before
the Soc. de Biologie, enter into a criticism on the results of the
experiments of Fritsch, Hitzig and Ferrier, and add a series of
experiments of their own to determine whether secondary currents
are caused in the brain by the application of induced electricity
when used as a means of stimulating that organ. They praise very
highly the injection of chloral into the veins for producing sleep in
the animals to be operated on, and rank it high above chioroform
for this purpose. (This method is that of M. Oré, for the results
of whose experiments with this drug see the Lond. Med. Rec. Vol.
u. No. 64.) These authors have arrived at the following conclusions:
(1) Even feeble induced currents diffuse on the surface of the brain
from one point to another. (2) This diffusion on the surface is
caused by the fluids and by the cerebral pulp. (3) Even feeble in-
duced currents cannot be localised to the depth of the gray cere-
bral matter; they diffuse more or less deeply into the subjacent
white strata. It is probable that in this case they follow a certain
determinate direction, perhaps that of the principal white bundles
which lead to the corpora striata or the peduncles. In the Gaz.
Méd. de Paris, No. 4, a continuation of these experiments on the
excitability of the hemispheres is published. They show that the
REPORT ON PHYSIOLOGY. 397
phenomena described by Ferrier, and which these authors have also
obtained, are not due to stimulation of centres placed in the cortical
substance of the brain, and ascribe them to a stimulation transmitted
from the surface to the cerebral ganglia and the peduncles. They
find a difference in the results when the animal (dog) is only incom-
pletely or completely narcotised. When incompletely narcotised,
the movements obtained by stimulating the surface of the hemi-
spheres with Faradic electricity are of two orders: (1) Movements
‘due to the diffusion of the current by the liquids, movements of ro-
tation of the head to the opposite side, quivering of the jaw, and
movements of the eyelids. (2) Movements due to the diffusion of
the current into the depth of the cerebral mass, and to the exci-
tation due to the corpora striata, movements of the limbs, and of the
trunk of the opposite side. On the other hand, when the anesthesia
was complete (tested by stimulating with the induced current the
central end of the divided sciatic nerve) the experimenters did not
obtain any of the effects of stimulation of the convolutions, whatever
the intensity of the current. Neither the centres described by Ferrier,
as regulators of the combined movements of: the anterior extremity,
‘nor those of the posterior extremity, nor those of the lips and eyelids,
became manifest by the electric stimulation. The authors conclude
from their experiments that the peripheral layer of the hemispheres
is inexcitable, it is insensible and does not contain any special
‘motor centres. The effects obtained by Faradisation, which pene-
trates to the corpora striata and to the peduncles, are those of
direct excitation of these organs. These effects cannot be attributed
to any reflex action (as maintained by Schiff and Dupuy). Complete
anesthesia, which prevents these effects, does not change the con-
ditions of the gray matter of the hemispheres; it only acts by
diminishing the excitability, in a more or less pronounced degree,
of those parts of the brain already known to be excitable.
E. Hitzig (Reich. und Du Bois Rey. Arch. 1873, 397) has also con-
tinued his researches on the physiology of the brain. He confirms
the results of his previous experiments with Fritsch that with in-
creasing intensity of the current on applying both electrodes to the
surface of the brain of dogs, the first contraction is produced by
changing the direction of the currents, when thereby the anode comes
upon the proper centre. The anode is throughout more effective
than the cathode. With regard to the influence of narcotism pro-
duced by ether or morphia, the author found that during the deepest
zetherisation, one or other centre did not react for a short time, whilst
in others irritability remained. All reaction could only be abolished
for a minimum time. With morphia narcotism the reaction of the
centres remained undiminished. The same is true during the con-
dition of apnea. The author has found a centre for the single move-
ments of the eyes, which coincides with a part of the centre of the
facialis.
Insury To THE Brain witH Putmonary H#MoRRHAGE.—H.
Nothnagel (Centralblatt, No. 14, 1874) finds on injiring with a
398 DR STIRLING.
needle a certain spot on the surface of the brain of the rabbit that
peculiar disturbances occur, above all, hemorrhage in the lungs and
in the tissue of the same, often so pronounced that almost the whole
lung is traversed by the hemorrhage. Brown-Séquard, as is known,
has also observed this, not however from injury to the surface of the
brain, but of its basilar portion. Secondly, in the same way, menin-
gitis can be regularly produced, chiefly bilateral, very seldom on the
injured side, sometimes only on the half opposite to the injured side.
This meningitis, the author thinks, is not a mere accidental cireum-
stance.
INFLUENCE OF THE BRAIN ON THE TEMPERATURE OF THE Bopy.—
J. Schreiber (Pfliig. Archiv, vin. 576) has operated on a large
number of rabbits, with a view to determine the effect of injuries to
the brain on the temperature of the body. The injury was made by
means of a lancet-shaped needle introduced through the skull. The
temp. was measured in the rectum. From about 70 experiments the
author concludes that after injury of the pons in all parts, of the
pedunculi cerebri, of the cerebellum and cerebrum, increase of the
body temperature occurred when the animals were protected arti-
ficially from losing warmth; that the same results followed uncon-
ditionally and constantly on injury to the limit between the medulla
and the pons. For the present the author leaves the question as to
whether these results are due to the presence of a moderating excito-
caloric centre, as T'scheschichin supposed, or are pure vaso-motor
phenomena.
Functions oF THE Lincuat Nurve.—Prévost (Arch. de Phys. v.
253 and 375) has arrived at the following conclusions: 1. Ablation
of both spheno-palatine ganglia does not affect, in dogs and cats, the
sense of taste, in parts supplied by the linguals. 2. After section of
the chorda tympani, in dogs and cats with cut glosso-pharyngei, the
taste was little modified in some cases, notably diminished in others,
and completely abolished in one. Our results do not permit us to
specify the réle the chorda plays in relation to the function of taste,
but we are inclined to accord to it only an accessory part. 3. Con-
trary to the old views of Vulpian, and coinciding with his recent
researches, the author finds that the chorda carries fibres to the ter-
minal branches of the lingual as well as to the sub-maxillary gland.
After section of the chorda, in cats, dogs, rabbits and guinea-pigs, de-
generated nerve-fibres were found in the terminal branches of the
lingual, as well as in the mucous layer of the tongue and sub-maxillary
gland. 4. The chorda has not a trophic centre in the papille of the
tongue, and if the sub-maxillary gland acts as a trophic centre on it,
this influence ought at least to be very limited. After section of the
chorda in the ear, the central end of this uerve (on the side of its
emergence facially) remains healthy.
INFLUENCE OF EXTIRPATION OF THE SUPERIOR CERVICAL GANGLION
oy THE MoveMENTs oF THE Irnis.—Vulpian (Arch. de, Phys. vt. 177)
removed completely this ganglion in dogs, on the left side, At the end
REPORT ON PHYSIOLOGY. 399
of from 10 to 15 days the animals were curarised and artificial respira-
tion kept up. The skin and subjacent tissues of different parts of the
body were stimulated by strong induced currents. Each time, the
pupil on the left side dilated a little from one quarter to a third of
its radius. It is therefore certain that all the sympathetic nerve-
fibres which act on the iris do not traverse the superior cervical
ganglion, nor do all of them pass through the part of the sympathetic
above this ganglion, for it was excised at the same time as the
ganglion.
“Vulpian on Section of the Chorda Tympani.”—Abstract in
Lond. Med. Rec. No. 46. “Experiments relative to the physiology
of the vaso-dilating nerves.” Vulpian, Arch. de Phys. vi. 175.
INFLUENCE OF CHANGES OF TEMPERATURE ON THE CENTRAL ENDS
OF THE CarpiAc Nerves.—Tarachanoff, under Cyon’s direction
(Pjliiger’s Archiv, vit. 347), finds, in opposition to Fick, who says that
the central ends of the cardiac nerves are not influenced by changes
of temperature, that, in passing defibrinated blood under normal pres-
sure through the vessels of the brain, after ligature of the carotid and
vertebral arteries, and of the veins, sudden increase of temperature of
about 18° C. acts as a stimulant in the most powerful manner, on
the central ends of the vagus, and quite in accordance with the action
on the peripheral ends of this nerve.
INFLUENCE OF THE PosteRIOR NervE-Roots oN THE SENSI-
BILITY OF THE ANTERIOR.—Cyon (Pfliiger’s Archiv, vin. 340), in
opposition to the negative results of G. Heidenhain, cites the follow-
ing experiment from Steinmann’s paper, as demonstrating Brondgeest’s
tonus. The gastrocnemius of a frog is weighted with twenty or thirty
grammes, and the muscle is allowed, while at rest, to write upon a
rotating cylinder. The posterior roots are then cut carefully with
sharp scissors, and the muscle is allowed to write its length further.
The weighted muscle increases in length in a marked degree, either
immediately or in the course of a minute.
Action or Strycunra on Sensory Nerves.—Busch (Berl. Klin.
Wochensch. No. 37) finds that sensibility is so much impaired in frogs
poisoned by strychnia that pinching the animals’ toes or burning the
central end of the divided sciatic nerve may be performed without being
followed by a reflex action. “The Physiology of Man. Vol. Iv.
The Nervous System.” By Austin Flint, jun., M.D. 8vo. pp. 470.
New York: D. Appleton & Co. 1872. ‘Physiological Studies on
the Motor Nerve of the Heart; Relation of Motor and Sensory Nerves
to their centres of Nutrition; Relation cf the Roots of the Spinal
Nerves to the Sympathetic ; Excitability of the Spinal Cord, etc.”
Abstract of these papers from the Italian by Boll, in Centralblatt
No. 52, 1873. “Experimental Investigation of the simplest Psy-
chical Processes.” Sig. Exner, Pjliig. Arch. vir, 601 (Abstract in
Lond. Med. Recd. 11. No. 55).——“ Paralysis of the Vagus in Man.”
P. Guttmann, Virch. Arch. u1x. 51. “ New formation of Brain Sub-
stance in the form of a Tumour on the surface of the Convolutions.”
400 DR STIRLING.
T. Simon, Virch. Arch. tv. 310. “Effects of a rise of Tempera-
ture on Reflex action in the Frog.” M. Foster, Journ. of Anat. and
Phys. vi. 45; also Nature, Dec. 4th, 1873. ‘“‘Sensation in the
Spinal Cord,” by G. H. Lewes, and Reply in Nature, Dec. 11th, by
M. Foster. “Double Nerve Stimulation.” Dew-Smith. bid.
(Abstract of both papers in Lond. Med. Recd. Vol. 1. No. 59.)
GENERAL PHysioLoGy oF THE NerveEs.—<Electrotonus.” 4.
Hermann, Pjliigz. Arch. vu. 301, 323, and 497, Abstract in Central-
blatt, No. 43, 1873. “Transverse conduction in the Nerves of the
Frog.” E. Hitzig, Pjliig. Arch. vu. 263. “Form of the Curve in
the so-called transverse conduction in the Nerves of the Frog.”
W. Filehne, 7bid. vit. 71. * Action of the electrical current in
different directions of the nerves and muscles.” Bernheim, 60, Jbid.
— “Nerve Degeneration and Nerve Regeneration.” H. Eichhorst,
Virch. Arch. wrx. I. “On the union end to end of Sensory with
Motor Fibres.” Vulpian in Gaz. Méd. de Paris, No. 7, 1874.
Eys.—“ Influence of Spectacles on the acuteness of Vision.”
Donders, in v. Grdfe’s Arch. xvi. 2. 245 (Abstr. in Centralblatt, No.
47, 1873). ‘‘New apparatus for measuring the field of vision.”
Scherk, Zehend. Klin. Monatsbl. f. Augenheilk. x. 1873, 151 (Ab-
stract in Centralblatt, No. 40, 1873). “Development of Traumatic
Keratitis.” A. Bottcher, Dorpat. Medic. Zeitschr. 1873, tv. 66 ( Ab-
‘stract in Centralblait, No. 52, 1873). ““Micrometry of the posterior
part of the Eye.” Laqueur, Centralblatt, No. 59, 1873.-_—“ Lymph
Sheaths of the Choroidal Vessels.” Morano, Centralblatt, No. 1, 1874.
“Inflammation of the Cornea.” C. J. Eberth, Centralblatt, No.
6, 1874. “Change of Fluids in the Eye.” Th. Leber, Arch. f-
Ophthal. x1x. 87 (Abst. in Centralblatt, Nos. 9 & 10, 1874). “On
the accommodation movement of the Choroid in the Eye of Man, the
Ape and the Cat.” Hensen and Vélkers, Arch. f, Ophthal. xrx. 156
(Abst. in Centrelblatt, No. 11, 1874). “Estimation of the Refrac-
tive Index of the Fluid Media of the Human Eye.” J. Hirschberg,
Centralblatt, No. 13, 1874. “Action of the Galvanic Current on
the Human Eye.” H. Schliephake, Pjliig. Arch. vin. 565.
Ear.—‘The Mechanism of the Ossicles of the Ear and Membrana
Tympani.” By H. Helmholtz, translated from the German, by Albert
H. Buck and N. Smith. New York: William Wood & Co, 1873.
DP. 69. “Mechanism of the Ear.” J. G. M*Kendrick, £dinr,
Med. Jowr. coxxut 577. “Mechanism of Opening and Closing
the Eustachian Tube.” Yule in Journ. of Anat. and Physiol. vit.
127. “Effect of the Galvanic Current upon the Acoustic Nerve.”
C. J. Blake, Arch. of Scient. and Prac. Med. New York, 1873. No.
4. 326.——“The External Ear a Synthetic Resonator.” C. H.
‘Burnett, Philad. Med. Times, 1873, No. 101.
FUNCTIONS OF THE SEMICIRCULAR CANALS.—Solucha has made
‘experiments on this subject under Cyon’s directions (Pfliiger’s Archiv,
Vol. yur.). Phe results obtained by Flourens are already well known.
eh te
—
ee ee ee
REPORT ON PHYSIOLOGY. 401
Goltz ascribes the disturbance of movement produced by section of
these organs, to the loss of the feeling of equilibrium; while Liwen-
berg regards the collective disturbance of movement as only con-
sequences of a reflex stimulation of the nerves, which run in the
membranous canals. The first point investigated by Solucha was,
how far is an abnormal position of the head able to disturb the feeling
of equilibrium of the animal, and so to produce the abnormal move-
ments? The author confirms the experiment of Longet that mere
section of the recti capitis posticl majores et minores in the dog
renders the movements of the animal uncertain and insecure; the
dog was unsteady on its feet, moved from side to side, kept the fore
feet widely apart from each other, running was rendered difficult, de.
After five or six days the head generally assumed the normal position,
and at the same time the walking became normal. In a second series
of experiments the author sought to give pigeons a pecuhar position
of the head, without wounding important parts——a position such as
occurs on section of the canals, with the beak directed upwards, and
the occiput towards the ground. On fixing the head to the breast in
this position with a thread, the animals conducted themselves partly
like those in which the horizontal as well as the vertical semicircular
canals were destroyed. They could not retain their equilibrium, but
moved to and fro on their legs, and always sought for a third point of
support, made movements de manége, &c. As soon as the head was
freed, the whole disturbance of movement disappeared, and locomo-
tion became normal. This experiment also shows the importance of
a normal position of the head, so that the animal may be able to pre-
serve its position of equilibrium, as well as to execute co-ordinated
movements. In what way is the feeling of equilibrium dependent on
a normal position of the head? The answer to this is, for the greatest
part, to be sought in the changes which our conceptions of the posi-
tion and distance of cuter objects bear in relation to our own body.
In pigeons, provided with spectacles with prismatic glasses, which
produced artificial strabismus, disturbance of movement distinctly
analogous to the higher degrees of that which occurs after section of
the semicircular canals was observed. In some cases even ‘pendulum
movements,’ corresponding to those seen on section of the horizontal
canals, were noted. On section of one horizontal canal, the animal
made several lateral movements of the head, beginning from the in-
jured side, which soon ceased. On section of the corresponding canal
on the opposite side ‘pendulum movements’ of the head occurred,
and persisted very long. The violence of the movements increased
from the beginning onwards, until they reached a maximum, when
the animal lost its equilibrium, fell over, executed movements de
manége, &c. In a few cases the animals recovered completely, but
generally after four or five days the animal was found in a corner
with the peculiar position of the head above described, and quite
quiet, but when disturbed it resumed the pendulum movements, &e.
Most of the animals died in from ten to twenty days. On section,
the neighbouring parts of the skull were bloody and infiltrated, and
the cerebellum was softened on the posterior surface and of a yellow-
402 DR STIRLING.
ish-green colour. The results of section of the smaller vertical canals
were in some respects similar to the above, but differed in some
important points. In section of the horizontal canals, the head moved
in a horizontal plane from right to left, and back again; but in animals
with cut vertical canals, the pendulum movements of the head occurred
from above downwards and back again—then in a vertical plane. The
subsequent movements of the trunk were also different, in that the
whole trunk tumbled round on its transverse axis, and that mostly
from before backwards. On section of all four canals, violent move-
ment of the head, resembling a screw motion, occurred immediately,
accompanied by general swinging movement of the whole body. That
disturbance of equilibrium is a direct consequence of section of the
semicircular canals is certain. 1. These disturbances occur ummedi-
ately after the operation, and this when it is free from all other com-
plications, as section of muscles, bleeding, injury to the cerebellum, &c.
2. The two sorts of movement, as well of the head as of the trunk,
scarcely admit of a doubt, that the semicircular canals stand in rela-
tion to certain conceptions of space, and sensation. By means of the
nerves which end in the membranous canals, a series of unconscious
impressions are continually communicated, which lead to unconscious
conclusions as to the position of the head in space. The semicireular
canals contribute only indirectly to the retention of the equilibrium
of our body, in that they direct the position of the head in space.
The chief results of these experiments are the following: 1. For re-
tention of equilibrium, it is necessary that the animal has correct
conceptions as to the position of its head. 2. The semicircular canals
possess the functions of informing the animal, by a series of uncon-
scious (auditory?) impressions, as to the position of his head in space,
and each semicircular canal has an exact relation to a dimension of
space. 3. The movements which occur after section of the semi-
circular canals are of three sorts: a Disturbance in equilibrium, as
the direct consequence of the injury; 6 Swinging movements, as
consequences of stimulations arising from abnormal auditory sensa-
tions ; c. Consecutive phenomena, produced by inflammation of the
cerebellum, occurring several days after the injury (Lond. Med. Ree.
Vol. 1. No. 58). “On the Sense of Rotation and the Function of
the Semicircular Canals.” A. Crum Brown, Proceeds. Roy. Soc. Edin.
1873—74, and this Journal, 1874.
Sxivy.—“ Capillary Circulation in the Skin,” Bloch, Arch. de Phys.
v. 681. “Ueber den Raumsinn der Haut des Unterschenkels.”
Ad. Riecker, Zeitsch. f. Biol. 1x. 99. “Eifect produced on the
Skin by various injuries.” Bloch, Arch. de Phys. v1. 158.
Circulatory System.
Bouillaud “on normal and abnormal pulse.” Abstract in Lond.
Med. Reed. 1. No. 42. “Action and Sounds of the Heart,” G. Paton,
Edin. Med.. Journ. ccxxi. 407 (Abstract in Lond. Med. Recd. 1.
No. 48). ‘Blood-pressure in the heart and arteries,” Fick, Verhandl.
d. physik. medic. Gesellsch. zu Wiirzburg, tv. 223 (Abstract in Lond.
REPORT ON PHYSIOLOGY. 408
Mted. Reed. No. 49).——“ Quinine and Blood.” Binz, Arch. f. Exper.
Pathol. 1873, 1. 18. “Action of Atropin and Physostigma on the
pupil and heart.” Rossbach and Frohlich, Wiirzb. phys. med. Verh. 1.
(Abstract in Centralblatt, Nos. 58 and 59, 1873). “Origin of
Hemorrhages after ligature of the vessels.” Virch. Arch. ivi. 436
(Abstract in Centralblatt, No. 59, 1873). “The Capacity of blood
for Oxygen at different barometric pressures.” P. Bert in Soc. de
Biologie (Gaz. Méd. de Paris, No. 2, 1874). ‘Demonstration of the
Pulse by means of the flame.” R. Klemenziewiz, Untersuch. aus d.
Institut. in Graz, Heft 11. “On Ecchymosis and other effusions
of blood, caused by a nervous influence.” Brown-Séquard, Arch. of
Scien. and Pract. Med. 1873, 148. “The red blood-corpuscles.”
C. Faber, Archiv der Heilkunde, xiv. 481 (Abs. in Centralblatt, No. 6,
1874). “On Diapedesis.” J. Arnold, Virch. Arch. uvut. 203 (Abs.
in Centralblatt, No. 8, 1874). “Action of Calabar Bean on the
Heart.” H. Kohler, Arch. f. exp. Pathol. 1873, 1. 277 (Abs. in
Centralblatt, No. 8, 1874). “On the migration of white corpuscles
into the lymphatics of the frog’s tongue.” KR. Thoma (Z'ageblatt
d. 46 Versam. deutsch. Naturf. und Aerzte in Wiesbaden, 1873 (Abs.
in Lond. Med. Recd. 1. No. 51). “Kin Beitrag zur Mechanik der
Herzcontractionen,” Inaugural Dissertation, by E. A. Lutze, Cothen,
1874.——“Causes of the secondary waves in the Pulse,” Galabin,
Journ. of Anat. and Phys. vii. 1. “The Law which regulates the
frequency of the Pulse.” A. H. Garrod, Jbid. p, 54. ‘““Vaso-motor
and uterine Nerve-centres.” Schlesinger, Al/gem. Wiener Med. Zeitung,
Nos. 42 and 43 (Abstract in Lond. Med. Reed. 11. No. 55). “The
role of Oxygen in the formation of Pus.” C. Binz, Vireh. Arch. urx.
293. “The proportion of the white to the red Corpuscles after
Suppuration.” Apolant, /bid. tix. 299. “Doctrine of the reflex
stimulation of the vaso-motor nerves.” Cyon, Pjliig. Arch. vit. 327.
“The resistance of the walls of the vessels in the normal condition
and during Inflammation.” F. v. Winiwarter, Sitz. d. k. Akad. d.
Wissensch. txviu. 3 Abth. 1873. “ Direct electrical stimulation of
the Mammalian Heart.” 8S. Mayer, Jbed.
“OBSERVATIONS ON WELCKER’S METHOD OF ESTIMATING THE
Quantity or Bioop.”—R. Gscheilden, Pjliig. Arch. vu. 530, disposes
of objections which had been urged by Ranke against the method of
estimating the quantity of blood in the body, employed by Heiden-
hain and himself, for which see the original.
I. Brozeit suggested that the quantity of blood would be found
to be smaller when the animal had been previously subjected to an
important injury. The author, to determine this, experimented upon
rabbits. From the animal first a small quantity of blood was with-
drawn and its amount of solid constituents determined. It was then
subjected to various operations, and then a second quantity of blood
withdrawn and its solid constituents likewise determined. The
operations that the rabbits were subjected to were, throwing the whole
animal into a state of tetanus by the application of induction shocks
to the spinal cord ; poisoning with strychniaj; section of the spinal
404 DR STIRLING.
cord and artificial respiration ; suffocation, by the mechanical cessation
of the respiration, by CO, and common gas. In all cases the blood
showed a slight diminution in solid constituents, but this was also
found in the second blood-test when the animal was left uninjured,
and was the greater, the smaller the animal employed.
It. The quantity of blood in rabbits, guinea-pigs, and dogs,
Ranke gives great variations in rabbits, from 1 : 12,25 to 1 : 34,4,
In two new experiments the author found it in comparison to the
nett weight of the body (weight of animal minus intestines) as 1; 19,2
and 1: 18,2. His former result was 1 : 19,5.
III. Colouring matter of the muscles of different animals. The
author shows that Brozeit’s view as to the colour of muscle being due
to diffusion of hemoglobin is quite untenable. Whether the quantity
of the colouring matter in muscle depends upon the activity of the
muscle the author’s experiments do not definitely determine.
PERIODICITY OF THE ACTION OF THE HEART.—Regarding the expe-
riments of Metschnikoff and Setschenow (Journ. of Anat. and Phys.
vit. 193), A. B. Meyer adds a note in Centralblatt, No. 59, 1873,
to say that he has observed the phenomena described by these authors
and announced it under the term “Intermittenz,” in his paper “ Ueber
das Hemmingsnervensystem des Herzens.”
CHEMICAL COMPOSITION OF THE BLOoD-coRPUSCLES.—J. Paquelin
and Jolly, in a memoir presented to the Soc. de Biologie (Gaz. méd.
de Paris, No. 7, 1874), sum up the results of their researches on this
subject thus : 1. We do not know any process which yields hema-
tosine ina state of purity. 2. This substance is always more or less
mixed with albuminous matter. 3. The albuminous matter varies in
quantity according to the degree of alkalinity of the solvents employed.
4, Other substances, such as iron, may be mixed with hematosine
according to the mode of preparation. 5. The differences of opinion
regarding the constitution of heematosine are a consequence of the diffi-
culties inherent to its preparation. 6. Heematosine contains no iron.
7. The ferruginous principle of the corpuscles is combined with an
albuminous body scluble in heat, alcoholic fluids, but strongly in
alkaline ones. 8. Ineineration changes the constitution of the
mineral principles of the corpuscles, 9, Carbonization does not alter
these principles. 10. Iron exists in the corpuscles in the state of
phosphate of the protoxide. 11. This phosphate of iron is tribasic.
12. All the phosphates of the organism may be represented by the
formula 3 (MO), PO,. 13. The basicity of these phosphates is less
stable, 14. 100 grms. of dried corpuscles yield
Phosphoric acid 0,415
Lime 0,015
Protoxide of Iron 0,600
Potass 0,031
REPORT ON PHYSIOLOGY, 405
which corresponds to
Phosphate of Potass 0,046
4 of Lime 0,027
re of Tren 0,994
1,067
The method of (1) extracting the corpuscles, (2) separating the
heematosine, (3) analysing the mineral substances, is detailed in the
original. The experiments were made on the blood of the ox.
Oricin or Mecnanican (ipema.—A. Hehn (Cenéraldlatt, No. 40,
1873) has repeated Ranvier’s experiments on this subject and arrived
at the following conclusions. The author experimented on dogs. 1.
Extraperitoneal ligature of the vena cava inferior below the renal
veins was follewed, in a large number of animals, by neither cedema
of the lower extremities nor collection of transudation in the peri-
toneal cavity. 2. Ligature of the vena jugularis on both sides
yielded aiso negative results. 3. Section of one sciatic nerve with
sunultaneous ligature of the vena cava inferior produces edema in
the corresponding limb only when the nerve is divided at a certain
height after its exit from the ischiadic foramen.
Action oF Zinc on Buoop-Sotution.—H. Struve (Journ. of Pract.
Chem. N. F. vu. 1873, 346). Schdnbein has shown that when Zn is
shaken with water a small quantity of hydric peroxide is formed.
The author finds that simple contact of Zn with water is sufficient
for this. Place Zn in a solution of blood in a similar way and allow
it to stand, gradually a brownish red precipitate is formed and the
fluid becomes less and less coloured, and finally becomes as clear as
water. The precipitate contains the whole colouring matter of the
blood, and the filtrate is free from albumen.
TRANSFUSION OF Bioop.—L. Landois (Centralblatt, Nos, 56 and
57, 1873) injected into dogs the blood of the sheep, cat, guinea-pig,
rabbit, man, pig, calf, and pigeon. The blood of the hare, sheep, calf,
and man was transfused into the rabbit. In a special series of ex-
periments the frog was employed, and into it blood from the frog,
rabbit, sheep, calf, guinea-pig, pigeon, pike, and man, was injected.
With regard to the experiments upon the mammalia, the author
remarks—A. That the blood-serum of many mammals dissolves the
blood-corpuscles of other mammals. The most active serum (as yet
examined) is that of the dog, the least active is rabbit-serum. Bb. The
blood-corpuscles of mammals possess quite another capability of
resistance in the serum of other animals, e.g. the blood-corpuscles of
the rabbit are exceedingly easily dissolved, whilst those of the dog and
cat manifest great resistance. ‘The manner of solution can be observed
under the microscope. At blood-heat the solution takes place quicker
than at a lower temperature. Amongst others the author draws the
following conclusions from his experiments. The dissolved constitu-
ents of the blood-corpuscles are excreted principally by the urine, less
406 DR STIRLING.
richly and not so constantly by the intestine, uterus, bronchial tubes,
and into the serous cavities. A certain quantity of the dissolved
materials can be used for the construction of the body of the recipient.
The beginning and the end of the excretion of blood by the urine
varies. Already after 12, 24 hours after the injection hemoglobin
and albumen were found in the urine. The same condition existed
even twelve hours thereafter and also later. The quantity and kind
of blood transfused and the condition of the vascular system is in
this respect of influence. When foreign blood is transfused into an
animal, some of the animal’s own blood-corpuscles may disintegrate
and dissolve. ‘This is the case when the blood-corpuscles of the
recipient are soluble in the serum of the animal which yields the
blood. In animals with easily soluble blood-corpuscles, e.g. the rabbit,
the injection of serum of different sorts, e.g. of the dog, man, pig,
sheep, cat, produces, according to the quantity, very menacing symp-
toms; increase of the frequency of respiration, dyspneea, convulsions,
even death or asphyxia: whilst animals with resisting blood-cor-
puscles, e.g. the dog, can bear the injection of other kinds of serum,
e.g. of the sheep, ox, pig, without any of these phenomena. After
too great a transfusion with solution of the blood-corpuscles the for-
mation of clots of fibrin sometimes occurs, causing the death of the
aninal,
AcTION oF BitTER SUBSTANCES ON THE CIRCULATION AND Buioop-
PRESSURE.—Kohler (Vageblutt d. 46 Versam. deutsch. Naturf. und
’ Aerate in Wiesbaden, 1873) finds that Cetrarin, Columbin, and pro-
bably the other bitter substances, when injected into the veins pro-
duced a diminution of the arterial blood-pressure (8—20 Millim. Hg.)
followed by a gradual increase of 12—18 Millim. Hg. above the
original tension. ‘The cause of the diminution of pressure, which oc-
curred even after section of spinal cord and vagus, is to be sought
in the heart itself; whilst the cause of the increase, which did not
take place after section of the spinal cord, must be sought for out-
side the heart. ‘The increase of the blood-pressure after injection
of cetrarin and columbin is to be ascribed to the excitation of the
vaso-motor centre. The initial diminution was also observed after
paralysis of the termination of the vagus in the heart. The frequency
of contractions of the heart, even when poisonous doses of the above
substances had been given, remained unchanged till shortly before
death.
A PECULIARITY OF CAPILLARY-BLOOD.—F. Falk (Virch. Arch. urx.
26) asks why it is that blood is found fluid in the capillaries post-
mortem, and why even after removal from the body it remains in the
same condition. He set himself to determine which of the three
substances necessary for the coagulation of the blood [(fibrino-plastic
substance, fibrinogen, and the coagulating ferment (Schmidt) ] is absent,
The blood was obtained by puncture from the human lungs or from
those of the horse. The presence of fibrino-plastic substance was
proved by diluting the blood with twenty parts of water and cuare-
REPORT ON PHYSIOLOGY, 407
fully adding acetic acid, or leading through it a strong stream of
CO., when a precipitate showing the properties of paraglobulin was
obtained. This precipitate placed in the fluid of a hydrocele or in
pericardial fluid yielded fibrin. On treating the blood after the
manner recommended by Schmidt for obtaining the fibrin-ferment, a
fluid was obtained which accelerated the coagulation, but not to an
extent equal to that obtained from normal blood or serum. No
coagulation occurred however on adding to the capillary post-mortem
blood, defibrinated blood (which contains fibriu-plastic substance).
This points to the absence of the fibrinogen. In fact the author could
not obtain this substance by any of the means recommended for its
separation. The non-coagulability therefore of capillary blood from
the dead body is due to the absence or considerable diminution of the
fibrinogen.
Inriuencr or Nutrition on Carprac Muscies.—L. Perl (Virch.
Arch, ux. 39, Abst. in Centralblatt, No. 15, 1874), in his first series
of experiments (on dogs), performed seldom and large bleedings (every
5—7 days, 3—3% per cent. of the body weight at each time), in a
second series more frequent and smaller bleedings (every 3—4 days,
1—14 per cent.) were practised. ‘The animals endured the operative
procedures well. The wounds healed well without fever, and only
in one case did embolus of the lungs vccur. Whilst the animals of
the second series, on which ten as the minimum, and seventeen as
the maximum, of bleedings were practised, remained quite cheerful
and well, and when killed, from the 36—39th day, showed no signs
of change in the muscles of the heart, the seven dogs of the first
series, on the contrary, on which 5—7 bleedings were practised,
became lean, lost appetite, became sad, had partial edema of the
extremities, and died (6) with the phenomena of marasmus within
eleven weeks. With a single exception, all the animals dying after
four weeks showed a very flabby heart, with a yellowish colour, and
under the microscope a large number of the muscular fibres were
found to have undergone extensive fatty degeneration. These fibres
lay irregularly scattered amongst normal ones, for the most part in
the papillary muscles, especially of the left half.
Reriex INNERVATION oF VeEssELs.—E. Pick (Jnaug. Dissert.
Berlin 1873, 8", 30 p., and Reichert und du Bois’ Archiv, 1873, 1.)
experimented on the web of the second interdigital space of Rana
temporaria. ‘The main artery and vein run close to the bone, and
give off a series of branches which divide dichotomously until they
end in the capillaries. Dilute acetic acid or the induced current
were used as the stimulating agents. The greater the stimulation,
and the smaller the vessel, the quicker and more intensive is the
reflex contraction. Different portions of skin require different in-
tensities of the stimulant to produce the same effect. The most
sensitive part is the skin of the back, the least that of the face.
Intense irritation of the larger arteries is followed by dilatation,
The vaso-motor channels run exclusively in the ischiadicus.
408 ; DR STIRLING.
NumBer of Rep anp Ware CorruscLes IN THE Broop.—
Melassez (De la Numération des Globules Rouges du Sang, Paris,
1873, and Arch. de Phys. v1. 32) describes his method of estimating
the number of corpuscles, red and white, in the Llood. Amongst
others he has arrived at the following conclusions.
Arterial blood seems to have the same richness in corpuscles in
the great trunks; in an arteriole he has found the corpuscles increased
in number. Venous blood is very different according to its source and
to the state of activity of function of the organs from which it comes.
In the skin, the richness in corpuscles (“la richesse globulaire”) in-
creases. This augmentation is less marked on suppression of the
cutaneous evaporation and when congestion is produced either by
paralysis or by stimulation of the sympathetic ; ; 1t is, on the contrary,
more considerable when the cutaneous evaporation is increased, or
when congestion is produced by an obstacle to the circulation. A
similar augmentation is observed in muscles. It is least when the
muscle is at rest or paralysed, and is much increased when the muscle
contracts. In glands, the increase is mere considerable during the
state of repose, than when the gland is in action. The number of red
corpuscles 3 is increased in the spleen, and most so during the period
of digestion. In the liver, their number appeared to the author to
be diminished. ‘Then follows an estimation of the number of the
red blood-corpuscles in the mammalia Nea Rend. Dec. 1872
and Abs. in Lond. Med. Rec. No. 1, Vol. 1. 1873). The author
also estimates the richness of the blood in white corpuscles in cases of
erysipelas and in some cases of suppuration. The author finds by
his method that in a cubic millimeter of the arterial blood of the
dog there are from 3,410,000 to 3,780,000 red corpuscles; in the
arterial blood of the rabbit 4,700,000, and in the venous 4,900,000;
in the arterial blood of the grinea-pig 3,900,000, and in the venous
4,300,000.
ActTIon or GASES IN THE COAGULATION OF ALBUMEN.—H. Mathieu
and V. Urbain (Comptes Rendus, 1873, rxxvut. 706, Abs. in Central-
blatt, No. 58, 1873) have found that blood-serum, when deprived of
its gases, does not coagulate at 100°, and the same is true of the
albumen of eggs. The gas obtained from the albumen of the egg is
for the most part CO,. If the solution of albumen is again saturated
with CO,, then heat will cone it. When the albuminous coagu-
lum is treated with an acid, e. g. tartaric, CO, can be obtained, about
60—80 ccm. frem 100 ccm. of albumen. ‘The authors regard the
coagulation occurring under heat as a combination of the albu-
men with the CO, present in the fluid. In an albuminous solution
freed from CO. ,, alcohol, acids and the metallic salts cause coagulation.
If the pumping out of the gases is continued long enough, ‘not only
the dissolved gases are obtained, but also traces of ‘carbonate of am=
monia, as well as sulphide of ammonium, and then the solution
exhibits the properties of globulin, which, according to the authors,
can be precipitated in the cold by CO, The air-pump is not neces-
sary to cause this change of albumen i into globulin! it is sufficient to
REPORT ON PHYSIOLOGY. 409.
dilute freely the solution of albumen and place it under a bell-jar with
sulphuric acid and caustic potash. The addition of a small quantity
of phosphate of soda gives the globulin solution the properties of
cesin. Coagulated albumen, and the different albuminous substances
dissolved in NH,, and evaporated, yield a solution, exhibiting the
properties of globulin, and can be.compared with Mulder’s Protein.
Respiratory System.
* Action of protoxide of N. on germination and respiration ; on
the properties of chloropyll.” Jolyet and Blanche, Gaz. Hebdom.
1873, p. 389.—‘ The respiratory centre,’ Gierke, Pfliig. Arch. vil.
583 (Abstr. in Centralblatt, No. 59, 1873).—“ The respiratory move-
ments” (A. physiological and pathological study), F. Riegel, Wiirz-
burg, 1873, pp. 176, 12 plates. Abs. in Cenétralblatt, No. 5, 1874.
INFLUENCE OF ARTIFICIAL RESPIRATION OVER POISONING WITH
Srrycunia. Jochelsohn (Verhandl. d. Physikal. Medicin. Gesellsch.
zu Wiirzburg, New Series, v. 107) finds in support of the experiments
of Rossbach (Jour. of Anat. and Phys. vit. 203), and in opposition
to those of Leube and Brown-Séquard, that artificial respiration has
no distinet effect upon the duration and intensity of the convulsions ;
and these occurred even when the animal had been rendered apneic
‘before the administration of the drug. The author does not believe
that the strychnia is excreted or destroyed in the lungs, and thinks
that artificial respiration prolongs the life of the animal poisoned by
strychnia in the same way as that of animals whose spinal cord has
been divided high up in the neck (fuller report in Lond. Med. Kee.
No. 52).
On APNGHA AND THE ACTION oF AN ENERGETIC STREAM OF CO,
Gas oN THE Mucous MrewBRANE OF THE RESPIRATORY APPARATUS,
&c. The results of Brown-Séquard as to the effects of a strong
stream of CO, in cutting short an epileptic attack in guinea-pigs whose
sciatic nerve or lateral half of the spinal cord has been divided, have
been already noticed (Jowrn. Anat. and Physiol. vit. 343), W. Filehne
(Reich. und Du Bois Reym. Arch. 1873, 361) has repeated the ex-
periments of Brown-Séquard, and has always arrived at only negative
results. He finds further that powerful stimulation of the soles of
the feet with induced current is without effect on the epileptic attacks
in guinea-pigs.
Alimentation.
“On the Separation of Digestive Ferments.” V. Paschutin, Pech.
und Du Bois Reym. Arch. 1873, 382. “ Amylotic Ferment of
the Pancreas.” <A. Liversidge, Journ. of Anat. and Phys. vu. 23.
“Chemical Properties and Physiological Action of Fat.” J. Day,
Australian Med. Journ. (abstract in Lond. Med. Journ. Vol. it No.
‘62. “Contributions to the Question of Nutrition.” J. Forster,
4eitsch, f. Biol. 1x. Heft 3. “On the Butyrie Acid Fermentation.”
VOL. VIII. vag
410 DR STIRLING.
V. Paschutin, Pfliig. Arch. vit. 352 (abstract in Centralblatt, No. 9,
1847). “Tntervention of Electro-Capillary Forces in the Pro-
duction of the Phenomena of Animal and Vegetable Life.” Becquerel,
Robin's Journ. de 1 Anat. Jan. 1874.——“ Physiology of Defecation,”
Gobrecht in the Clinic of March, 1873 (abst. in Lond. Med. Ree.
No. 49). “ Physiological Action of Alcohol.” Sée, Translation
in the Lond. Med. Rec. No. 50. “ Absorption of Fat in the
Small Intestine.” S. Thauhoffer, Pest. Medizin. Chirurg. Presse,
1873, No. 22 (abstract in Centralblatt, No. 44, 1873, and Pfliig.
Arch, Viil.). “ Withdrawal of Alkali from the Living Body.” E.
Salkowski, Virch. Arch. 1873, tv111. 1 (abstract in Centralblatt, No.
49, 1873). “Mineral Inanition and Influence of Phosphate of
Lime on the Transformation of Albuminous Substances.” Brothers
Dusart in Société de Biologie (Gaz. Méd. de Paris, No. 5, 1874).
“On the Cataphoric Changes of Moist Porous Bodies.” H.
Munk in Reich. und Du Bois Rey. Arch. 1873, 241.——“ Prepara-
tion of Solutions of Albumen free from Salts by means of Diffusion.”
R. Aronstein, and observations on the same by A. Schmidt. Pjliig.
Arch. vit. 75 and 93 (abstract in Centralblatt, No. 1, 1874).
“On the Origin of Sulphuric Acid and the Condition of Taurin in
the Animal Organism.” E. Salkowski, Vireh. Arch. Lyiu., and
“Synthesis of Taurocarbaminacid,” Berich. d. Deutsch. Chem. Ges. zu
Berlin, vi. 1191. “The History of Uramid Acid.” H. Huppert,
Ibid. (abstract of these three papers in Centralblatt, No. 11, 1874).
“On the Albuminous Bodies.” O. Nasse, Pjliig. Arch. yin. 381
(abstract in Centralblatt, No. 13, 1874). “ Estimation of N. in
Albuminous Bodies by means of Soda Lime.” Ritthausen, Journ. f.
Pract. Chem. N. F. vu. 10.
SECRETION OF THE Panoreas.—-L. Landau (Znaug. Dissert.
Breslau, 1873, pp. 34, abstract in Centralblatt, No. 56, 1873). A
graduated canula was introduced into one of the pancreatic ducts,
whilst the other was ligatured along with the ductus choledochus.
Heidenhain’s method of using a poison to paralyse the secretory
nerves was employed. The dogs were curarised and artificial respi-
ration kept up. The mean quantity of secretion, in seven experi-
ments, was 0,2 ccm. in 60 mins. (The results of Bernstein’s ex-
periments gave about thrice as much.) Injection of Atropin, Calabar
bean, Nicotin, had no constant effect on the secretion, nor had increase
or diminution of the blood-pressure any material influence upon it.
Stimulation of sensory nerves, as well as electrical stimulation of the
lingual, yielded only negative results. In opposition to Bernstein,
who employed a permanent fistula, the author found only seldom a
diminution of the secretion, after stimulation of the vagus. Often
section of the vagus was without any effect, sometimes however this
(along with general clonic spasms of the body) as well as its stimula-
tion was followed by increase of the secretion. Positive results were
first obtained on stimulating the medulla, either directly or by
suspending the respiration. Stimulation of the medulla with in-
duced electricity increased the pancreatic secretion, and this the more,
REPORT ON PHYSIOLOGY. 411
the oftener the irritation. Often the rapidity of outflow appeared
first after cessation of the irritation.
Estimation oF THE N. ry Atspumen.—M. Maerker (after the
researches of O. Abesser), Pjliig. Arch. vi. 195. A large num-
ber of analyses. were made to determine whether the mean per-
centage of N. in flesh is to be trusted. It is found that the Will-
Varrentrapp method gives results very nearly accurate. The different
results of Nowak and Seegen are to be referred to differences in the
conduction of the analyses.
SIGNIFICANCE OF THE ASH-CONSTITUENTS IN Foop.—J. Forster
(Zeitschrift fiir Biologie, Vol. 1x.) has selected pigeons and dogs for
his experiments on this subject. These animals were fed with food
containing as few salts as possible, ‘ with albumen, in the form of the
residue of flesh after the preparation of the “ extract of meat.”’ This
residue, as is known, is not quite free from salts, but these were
extracted from it, as far as possible, by repeated boiling and washing
in distilled water. By this process a powder, containing in the dried
state 14°445 per cent. of nitrogen was obtained ; 100 grammes of this
dried substance contained—
Phosphoric acid anhydride - 0:548 gramme.
Chalk (calcium oxide) a wOO7SSes
Tron . < 2 . . 0:023
Potassium E ° . Ont b: XS 5,
0-800
Magnesia and chlorine were found in too small quantities to be
weighed. Casein from ordinary milk, boiled in distilled water, was
also employed. The starch used was treated several times with a
0-08 per cent. solution of hydrochloric acid, and washed in a filter
with distilled water until the filtrate yielded no precipitate on the
addition of nitric acid. A mixture of one part of casein and seven
parts of starch, as used in the experiments on the pigeons, contained
0-279 grammes of phosphoric acid in 100 grammes of the dried
mixture. Fat, in the form of the best butter, formed part of the
food, and in addition distilled water was freely given.
A prepared mixture of the above was given to pigeons and dogs,
and when it was rejected they were fed artificially. The author con-
cludes from his experiments that the addition of certain salts is neces-
sary for the retention of the balance of materials (Stoff/gleichgewicht)
in the animal economy. When this supply sinks below a certain
limit or is completely withdrawn, the body excretes salts, and the
animal dies.
The following table shows the quantity of nitrogen taken in in
the food, and that excreted, in a dog. The duration of the experiment
is divided into three periods of eight days each.
This table clearly shows that the removal of salts from the food
has no effect on the transformation of albumen, but that this chiefly
27—2
412 F DR STIRLING.
depends upon the quantity and kind of supply of the combustible ali-
mentary materials.
NITROGEN NrrroGen
TAXEN IN
EXCRETED DIFFERENCE
Flesh Fat Starch Nitrogen Urine | Feces | Total
met ie |
I. 1,433] 1,200 | 300 | 207-0 || 197:5| 7:5 | 205-2 + 18
PE 3st 650) 9 189-3 || 188-2} 15-0 | 203-2 — 13-9
TIT. 1,249} 689) 663 | 180-4 || 182-1 | 16-0 -| 198-1 -17-7
Apart from the contents in salts, the solid as well as the fluid
products excreted, during deprivation from salts, are the same as in
normal nutrition. At the beginning of the experiments, the digestive
juices secreted had the normal constitution, but they gradually
changed. A time then arrived when they became inactive, or no
longer had the normal composition.
“In all animals fed with food from which the salts had as far as
‘possible been extracted, a condition of weakness of the muscles and
tremblings occurred, best characterised as general exhaustion. Thé
weakness, of the posterior limbs of the dogs assumed in the second
week of the experiments 9 paralytic character. The activity of the
brain was disturbed, as shown by .the stupid appearance of the
animals, ete. Phenomena of increased sensibility showed themselves
later. By the greatest possible removal of the mineral constituents
from the food of adult animals, the process of the changes of materials
and decomposition m the body proceed in the same way, till the
death of the animal, as by a diet which, in addition to the above
“necessary constituents, also contains the ash constituents. Latterly,
however, disturbances in the functions of the organs occur, which
hinder on the one side the transformation of the nutritive material
into modifications capable of being absorbed, and thereby prevent the
reparation of the decomposed material of the body, and on the other,
by suppression of processes necessary to life, bring about the destruc-
tion of the organism, before the impossibility of a continued reception
of food is followed by decline and death.
With regard to the salts excreted, when these are removed from
the food, the tables show that the excretion of phosphoric acid is
never interrupted. In animals deprived of salts the exeretion of this
substance by the urine is largely diminished. The less the quantity
of food poor in salts introduced, the greater is the loss in phosphoric
acid which the body suffers. The smallest quantity exereted corre-
‘sponded to the time when the greatest quantity of combustible material
was introduced.
Although the food was quite free from chlorine, and the urine in
the latter ‘period of the experiment contained only a trace of this
substance, still the vomitings on the thirty-fourth day showed that a
large quantity of chlorine was mixed in the stomach with the food in-
‘troduced. 246 grammes of food vomited contained 1:63 grammes of
REPORT ON PHYSIOLOGY. 413
chlorine, while the urine excreted on the same day contained only
0-04 grammes, Chlorine was continually excreted in the stomach
and absorbed again, for the feces contained no chlorine.
The author thinks that all these conditions are to be explained by
the condition of the salts in the body. He divides these into two
classes. By far the greatest part is in combination with the com-
bustible substances, chiefly with the albuminous bodies in fixed or
loose combinations. A small fraction, previously in combination,
but freed by decomposition and oxidation, is present in the blood,
with the products of the metamorphosis of the tissues. The latter
are excreted on the blood passing through the kidneys. Food free
from salts, introduced into blood from the digestive canal, unites in
the blood with the free salts arising from the chemical decompositions,
The quantity of salts excreted must increase with the quantity of free
salts in the blood. The salts excreted are increased during hunger,
and this because the salts in combination with the body-substance
are set free to enter the blood. An increased excretion was also
observed when a surplus of salts was added to the food, as was the
ease ut the end of one experiment.
The author is of opinion that the supply of nutritive salts, or of
these salts in the food which can prevent a loss of salts from the
body, is less than till now has been supposed.
RESEARCHES ON DIGESTION AND ABSORPTION IN THE HUMAN LARGE
InTEsTINE.—V. Czerney and J. Latschenberger, Virch. Arch. tix. 161.
{Translation in full in Lond. Med. Ree. Vol. u. Nos. 62 and 63.)
The authors give a full account of the literature of the subject, and
then cite the history of the patient upon whom they experimented.
The researches were conducted on a man with preternatural anus (of
the sigmoid flexure) in the left inguinal region. The case was dis-
tinguished from the others which have been employed for physio-
logical investigation, in the circumstance that the rectum was com-
pletely exposed by the prolapsed loop of intestine ; it could be filled
from above with the articles of food on which the experiments were
made, and, at any desired time, emptied per anum. As the rectum
could be washed out from above like a retort, the discharge gave at
once the amount of material absorbed. While Voit and Bauer
deduced from the excreta, by an indirect process, the amount of water
absorbed, we were able to directly determine the quantity of unab-
sorbed residue.
The method of experimenting is then described. The authors have
arrived at the following conclusions :—
The human rectum and its secretion has no digestive action
either on coagulated albumen, or soluble albumen, or on fat.
In the normal condition, soluble albumen (dissolved in water) is
absorbed from the human rectum as such, not being changed by the
intestine; and a greater percentage is taken up, the longer it remains
in the bowel. Any irritative condition in the bowel impedes absorption,
or completely arrests it. Chloride of sodium also impedes absorption,
_but is itself taken up notwithstanding that the intestine is in a state
414 . DR STIRLING.
of irritation and that absorption is suspended. In the hen’s egg
albumen is contained in a form unfavourable for absorption. .
Fat in emulsion is absorbed by the human rectum. The quantity
absolutely absorbed is in proportion to the degree of concentration ;
and the amount per cent. absorbed is in proportion to the time during
which the fluid has been in contact with the absorbing surface.
Starch in the hydrated form is absorbed by the intestine ; whether
as such, or after being changed into sugar, is not decided by these
experiments.
The greatest quantity of albumen absorbed in 29 hours was about
13 grammes. As the large intestine is on an average about four
times as long as the portion which served for experiment, this indi-
cates an absorbent power, in the whole large intestine, of 6 grammes
of soluble albumen from a solution containing 4} per cent. This
quantity is far from being sufficient for nutrition, as about 120
grammes are necessary for a healthy man (Voit and Bauer). The
quantity of albumen absorbed is evidently increased when more con-
centrated solutions are used.
General Result.—The human rectum absorbs soluble albumen
unchanged, as such; it also takes up emulsion of fat; starch also is
absorbed, but it remains undecided whether it be absorbed as starch,
or whether it must first be changed into sugar. Chloride of sodium
impedes or completely arrests absorption.
Tables are appended showing the results in each of the experi-
ments.
THE Move or Secretion or Gastric J uice.—H. Braun (Eekhardt’s
Beitrage zur Anatomie und Physiol. vu. 1.27. From abst. in Cen-
tralblatt, No. 15, 1874). The general view held regarding the secre-
tion of the gastric juice is that it is caused by stimulants (chiefly of a
mechanical and chemical nature). In opposition to this view, the
author renders it probable that, just like the urine, the gastric juice is
continually being secreted. He experimented on dogs with gastric
fistula. Through these fistule the stimulating substances, small
pieces of sponge, portions of feathers, sand, alkalies, and pieces of
flesh, were introduced, and this without the quantity of secretion,
which was obtained by means of small pieces of sponge, shewing any
increase. Quite as inactive was the alkaline saliva of dogs and man.
According to the author’s experiments no near relation exists between
the conditions of the secretion in the mouth and stomach, for neither
had stimulation of the former any pronounced influence on the secre-
tion of the gastric juice, the secretion of the gastric juice had as little
effect on the secretion of the saliva. Schiff’s hypothesis that pepsin
is first extracted from peculiar so-called peptogenous bodies in the
stomach is rejected by the author, for completely starved animals,
whose salivary ducts were cut, showed a not unimportant secretion of
active gastric juice. Much more are pepsin and acids, as Spallanzani
first hinted, continually secreted. The mucous membrane of the
empty stomach is only seldom covered with tough mucus, often with
a fluid, which has an acid reaction. Just the same as the urine, is
REPORT ON PHYSIOLOGY. : 4AL5
the gastric juice secreted in larger quantities after injection of large
quantities of water into the vena femoralis, This is not a simple
transudation, for the secretion has an acid reaction and digests discs
of albumen normally. Often, to give the secretion digestive power,
the addition of the HCl was necessary ; a fact which is brought by
the author in connection with the results of Manassein, that in the
gastric juice of acutely anemic animals the acids are absent. The
injecting fluids employed were a 1—2 per cent. solution of urea, and
1 per cent sol. of NaCl. Whether section of one or both splanchnici
increases the secretion of the gastric juice, the author, in spite of
several positive results, leaves for the present undecided, for not
seldom the secretion in the course of the experiment increased with-
out appreciable cause.
FEEDING WITH FLESH AND CARBOHYDRATES AND CARBOHYDRATES
ALONE.—M. Pettenkoffer and C. Voit (Zeitschr. f. Biol. 1873, 1x. 435.
From Salkowski’s report in Centralblatt, No. 18, 1874). This paper
is a continuation of their previous one on feeding with flesh and fat
alone (Journ. of Anat. and Phys. vut. 207). The experiments were
made upon the same dog and after the same manner as their previous
ones, with the aid of their respiration apparatus. The gain of fat in
the body was estimated by the authors from the deficit of carbon in
the excreta as opposed to the quantity taken in, and this in refer-
ence to the excretion of N, which represented the measure for decom- -
position of the albuminous bodies. The results are the following:
A great quantity of starch can be changed into grape-sugar and ab-
sorbed from the intestinal canal of the dog. As a minimum one
kilo dog digests 15 grms. starch in 24 hours, whilst one kilo fattened
ox, with copious feeding, did not absorb more than 12,7 grms. The
assumption therefore that the dog, as a flesh-eater, is not suited for
experiments with feeding on carbohydrates is not correct. Even
when great quantities of starch were given, the quantity of feces
excreted was inconsiderable, and consisted, for the most part, of
the residue of the intestinal juice, and not till the quantity of
starch exceeded 379 grms, in the 24 hours did unchanged starch
appear in the feces. The sugar absorbed split up completely into
CO, and water. The quantity of fat formed is not proportional to
the quantity of carbohydrates given. On the contrary, the quantity
of fat stored up depends upon the quantity of flesh present; then
with 379 grms. starch and 1800 grms. flesh it was 112 grms., with
800 grms. flesh and 379 grms. starch only 55 grm. From the N.
excreted during exclusive feeding with starch, it can be calculated how
much flesh the animal has given up from its body. Leave this aside,
and calculate how much fat must be stored up when flesh and starch
were given, and a tolerably exact agreement with the number obtained
by the experiment is got. This shows that the fat arises from the albu-
men of the flesh. Of course a certain proportion must always exist be-
tween the quantity of carbohydrates introduced, and of the fat formed,
in that the starch protects from consumption the fat resulting from
the decomposition of the albumen. ‘This is the only action of the
: aa
416. 3 DR STIRLING.
carbohydrates; a formation of fat from them was not found through-
out, From their former experiments the authors calculated the sepa-
vation of 11,22 grms. fat from 100 grms. flesh. If large doses of starch
are added to the flesh, so in fact almost 11 per cent. of flesh as fat
will be added.
RECHERCHES EXPERIMENTALES SUR LA QUESTION DE SAVOIR, SI
CERTAINES GELLULES DES GLANDES (DITES A PEPSINE) DE L’ HsToMAC
PRESENTENT UNE REACTION ActpE.—R, Lepine (Gazet. Med. No. 51,
18738) placed thin sections of the mucous membrane of the stomach
im a mixture of ferrocyanide of potassium.and sulphate of iron, which
yas treated with so much caustic potash, that the precipitated Prussian
blue was again dissolved with a yellow colour. Such a fluid produces,
with a trace of free acid, the separation of Prussian blue. In these
experiments with microscopic investigation nowhere in the glands of
the stomach was the separation of Prussian blue detected. The ex-
periment was varied in another way. Pieces of the mucous mem-
brane several Cm. long were carefully removed and used as a small
dialyser,, On the one side was placed lactate or sulphate of iron in
watery alcoholic solution, and on the other side ferrocyanide. of potas-
sium. The free surface of the mucous membrane was directed at one
time upwards, at another downwards. The result was in all eases the
same. The formation of Prussian blue occurred, but exclusively on
the free surface. In the glands of the stomach themselves no separa-
tion of the Prussian blue was to be detected, and the author concludes
from his experiments that the acid of the gastric juice is not. formed
in the cells of the glands.
MOVEMENTS OF THE GisopHacus.—Mosso (Movimenti dell’ Esofago,
Torino, 1873, pp. 44) employed the following simple methods for
studying the movements. of this tube. The esophagus was exposed
in the neck and into it a small wooden ball of the size and shape of
an olive was introduced. A: thin flexible wire was attached to the
ball, so that it could be easily extracted from the stomach, The
experiments were performed on-cats and dogs. A: transverse section
of the cesophagus does not hinder the propagation of the peristaltic:
movements when the upper half of the tube is stimulated. Excision
of a part of the cesophagus, e.g. from under the larynx to the archi:
of the aorta, does not arrest the peristaltic movements from above
downwards, under similar conditions. Even when the csophagus
was ligatured two or more times over a cylinder of wood introduced
into it (the ligatures were separated about 1 cm. from each other),
the mov oes were propagated from above downwards.
In a second series of experiments, on stimulating the vagi by the
opening shocks of an ordinary induction apparatus, the contractions
of the cesophagus formed a gradually ascending ‘Treppe,’ such as Bow-
ditch has described for the heart. The excitability of the vagi- con-
tinued exceedingly long after death. A dog showed, four and a half
hours after death, these movements when the recurrent nerves were
stimulated. The movements of the lower part of the cesophagus,
REPORT ON PHYSIOLOGY. 417
described by Magendie and Schiff, have not been observed by the
author. He finds that the lower part of the cesophagus is always at
rest, except when the upper portion is stimulated and caused to con-
tract.
Movements oF THE Intestines.—V. Basch (Die Hemmung der
Darmbewegung durch den N. Splanchnicus, Sitzungsb. der k. Akad.
der Wissensch. txvint. June, 1873). Basch and Oser have previously
shown that nicotine, at a certain stage, produces regular. peristaltic
movements of the intestines. The author employed the movements
.so produced for studying the inhibitory action of the splanchnicus on
the movements. of the intestine. The experiments were performed
upon dogs. Not only the phenomena exhibited by the intestines
were observed, but the blood-pressure was measured at the same time
in the carotid (this Journal, vur. 226).
Stimulation of the splanchnicus only inhibits the intestinal move-
ments, when it, at the same time, produces a considerable increase in
the arterial blood-pressure. The time when the intestines come to a
standstill always corresponds to the time when the blood-pressure has
reached its highest value, and this equally whether the stimulation acts
for a longer or shorter time. The intestinal movements could be
inhibited when the medulla eblongata was stimulated electrically,
after previous section of both splanchnici. Also in these experiments
-the inhibitory action occurred only when at the same time the arterial
blood-pressure was considerably increased—i.e.. in animals with
divided splanchnics, where not all the nervous connections between
the vaso-motor centres and the vessels of the abdomen were divided.
Asphyxiated blood or nicotin, after previous section of the splanch-
nicus, also inhibited the intestinal movements. It is therefore certain
that the splanchnicus is not the only inhibitory nerve for the intesti-
nal movements, but that also this inhibition is always accompanied
by the contraction of the vessels of the intestines, and that the in-
hibitory action is to be considered as a function of the vaso-motor
properties of the splanchnicus. Van Braam Honckgeest (Plt tig.
Arch. vit. 163), in opposition to V. Basch, concludes from his experl-
ments, that the inhibitory function of the splanchnicus is specific
and distinct from its vaso-motor functions, and as supporting this
view he finds that, 1. After section of the splanchnicus the spontaneous
movements become more lively than before. 2. Stimulation of the
vagus after section of the splanchnicus produces constant move-
ments. 3. On exposing the intestines freely to the air, hyperemia,
‘in consequence of paralysis of these vessels, i.e. paralysis of the peri-
.pheric expansions of the splanchnicus, is produced; then the spon-
taneous movements are neither so lively, nor is stimulation of the
vagus followed by movement, or, if so, only by very weak and irregu-
lar movements in some loops of the small intestines. 4, During the
inhibition from stimulation of the splanchnicus, the intestines do not
- become pale, but remain hyperemic as before. Blood-pressure not
measured.
418 DR STIRLING.
Inver.
“Tectures on Diabetes and the Glycogenic Function of the
Liver.” M. Claude Bernard. Lond. Med. Rec. Nos. 40, 41, 42, 43,
44, 45, 46, 47, translated from the Revue des Cours Scientifique.—
« Lussana and others on the Portal Circulation and the Biliary Se-
cretion.” Abstracts by Dr Brunton, in Lond. Med. Rec. 1. No, 56.—
“The formation of Glycogen in the Liver.” B. Luchsinger, Pjliig.
Arch. vit. 289. (Abstr. in Centralblatt, No. 10, 1874.)
Bittary Fistuta. H. Westphalen, Deutsch. Arch. f. klin. Med.
1873, xt. 588 (Abstr. in Centralblatt, No. 49, 1873). The bile was
collected from a case of empyema of the right side, in which paracen-
tesis had heen performed. At first the pus was mixed with bile and
had a bad odour, but fourteen days after the operation, whilst the
feces were devoid of colour (this lasted three weeks), only pure bile
was obtained. The bile was collected by a special apparatus. The
bile was quite fresh, of a bright gold yellow colour, neutral or scarcely
alkaline, and of a thinly fluid consistence. The smallest and most
concentrated quantity was evacuated between 2—4 a.m. In 24
hours a mean of 498,85. grms. was evacuated. Weight of patient,
67,5—64 kilos. S.G. 1008—1012,5. Percentage in solid residue
2,24—2,28 per cent.
The dried bile yielded with ether, cholesterin 2,49 per cent., un-
saponifiable fats with some oleate of soda 0,44 per cent. Lecithin
(reckoned from the P. per cent) 0,21 per cent. The substances in-
soluble in ether and alcohol (coloured mucin) made up 10 °/, of the
solid residue.
The following table gives the results of the chemical analysis of
the ash of the bile.
In 100 parts In 100 parts In 100 parts
Ash, | dried Bile. fiuid Bile.
K Cl 3,39 1,276 0,029
Na Cl 65,16 24,508 0,557
CO, Na, |- iti lik 4,108 0,095
PO, Nag 15,90 5,984 0,136
2(PO,) Cag | 4,44 1,672 0,038
| 100,00 "37,620 | 0,855
The alcoholic extract of the dried bile contained, glycocholate of
soda 4,48 per cent., palmitinate and stearinate of soda 6,4 per cent.
In the decomposition of these with baryta, only glycin and no trace
of taurin was obtained.
Urea, sugar, or other substances capable of reducing an alkaline
solution of Cu, were not found in the bile; only several times traces
of albumen and leucin (no tyrosin), of bile-pigments, bilirubin and
biliverdin, and in the ash in three tests each time some Cu, quinine or
calomel taken did not pass into the bile. The latter had no influence
oe
REPORT ON PHYSIOLOGY. 419
whatever on the quantity of bile. The same was the case with water,
which however doubled the quantity of urine and diminished its
S.G. from 10,21 to 10,11.
On THE ALBUMINOUS SUBSTANCES OF THE Hepatic CELLS,
P. Plész (Pjliig. Arch. vit. 371), under Ktihne’s direction, has in-
vestigated this subject. It is well known that the liver-cells undergo
marked changes soon after the removal of the organ from the body.
Sections of the fresh liver, even when free from blood, always have
an alkaline reaction ; but if kept for a short time at ordinary tempe-
rature, and still more quickly when at blood-temperature, they be-
come acid. The tissue of the liver as a whole becomes also more
resistant and rigid (Kiihne). The change in character of those cells,
which have become altered by post-mor tem changes, or, as the author
terms them, cells in a state of rigor mortis, are described. To ex-
amine these, all blood, bile, and lymph must be removed from the
liver by injections of weak solution of NaCl (0,75 °/,). The liver
after being washed for an hour or two appears pale, and neither
hemoglobin nor glycogen are to be detected in it. Some, not incon-
siderable quantity of soluble albuminous compound is also diffused
out and washed away. The liver is then finely minced, and passed
through fine linen, The pap is treated with a 0°75 °/, solution of
NaCl, and set aside for the cells to precipitate. The supernatant
fluid can be drawn off in a few hours, and is found to be opalescent,
partly owing to the presence of a little glycogen, and partly owing
to extremely fine granules which proceed from broken up cells.
When these have settled, or after filtration, the clear fluid is found to
contain—1l. An albuminous compound coagulating at 45° C., and not
redissolving at a higher temperature, soluble in solutions of NaCl,
sulphate and carbonate of soda, HCl, and acetic acid. It closely
resembles the albuminous substance found by Kiihne in muscular
fibre. 2. An albuminous nuclein combination, which coagulates at
70° C., and only differs from the preceding in this point. It agrees
in all its characters with the albuminous compound found by Mie-
scher in the nuclei of pus-cells. If the cells exhausted with the 0-75°/,
solution of NaCl, be now treated with a 10°/, solution, a considerable
quantity of an albuminous substance is obtained, coagulating at 75° C.,
-which is precipitated by the addition of much water, and by an excess
of a concentrated solution of NaCl, and presents other features showing
that it belongs to the globulin-like albuminous bodies, and closely
agrees with myosin. The liver-cells from which the above albuminous
substances have been extracted yield nuclein (Lancet, Oct. 18, 1873).
Formation OF GiycoGEN IN THE Liver. G. Salomon (Central-
blatt, No. 12, 1874) employed rabbits for his experiments. They
were fed on different substances, and the glycogen in the liver esti-
mated by Briicke’s process. He confirms Luchsinger’s statement as
to the difficulty of extracting glycogen and sugar completely from the
liver. The author confirms Hoppe-Seyler’s view, that on feeding
with gelatine, glycogen is formed in the liver. This “ gelatine-glyco-
420 DR STIRLING.
gen” had the same reaction as ordinary glycogen. Under feeding
with neutral fat (olive oil) and dilute glycerine, in each case glycogen
was found in the liver. Milk-sugar and fruit-sugar yielded similar
results.
_- ON THE HamaTocEeNous FoRMATION OF THE CoLOURING MATTER
oF THE Bite.—Steiner (Leichert und Du Bois’ Archiv, 1873, p. 160),
after referring to what is already known upon this subject, cites the
results of his own experiments upon the injection of water (at the
temperature of the body) into the carotid artery and external jugular
of rabbits. He finds that an injection of 20 cc. of distilled water into
the common carotid is not followed, either by a solution of the red
blood-corpuscles in the urine, or by the presence of the colouring
matter of the bile in the same. From 17 cther experiments the author
finds, after the injection into the right or left external jugular, of
from 30 to 50 cc, of distilled water, at the temperature of the body,
that in twelve cases there was a solution of the blood-corpuscles in
the blood-vessels, ¢. e. the appearance of free haemoglobin in the urine.
After precipitation of the albumen and examination of the filtrate, in
no case was the colouring matter of the bile proved in the latter.
The method of precipitating the albumen by heat, and examining the
filtrate for the presence of bile-pigment, is quite reliable. These re-
sults are completely negative, and stand in direct opposition to the
positive results of M. Hermann, who, however, operated upon dogs,
and collected the urine by canule introduced into the ureters.
Propuction oF Icrerus.—Audigné (Gaz. Méd. de Paris, No. 1,
1874) read a communication on mechanical icterus. He ligatured
the bile-duct in a dog. The animal lived 19 days, and though it
continued to have a voracious appetite, it emaciated visibly. The
colouring matter of the bile was found four hours after the operation
in large quantity in the urine, and on the second day the feces were
quite discoloured. The icteric tint in the skin and conjunctiva ap-
‘peared on the eighth day. The appearance of the bile-pigment in the
‘urine is at variance with the results of Frerichs, who observed this
only after twenty-four hours. A. histological examination was made of
the liver.
| CHANGES IN THE LIVER WHICH FOLLOW LIGATURE OF THE BILE-
Ductrs.—J. W. Legg (St Barthol. Hosp. Rep. 1x. 1873) operated
upon cats. The animals survived the operation for varying times up
to 20 days, and peritonitis when present remained local. This author
also, in opposition to Frerichs, observed the icteric tint of the con-
junctiva only on the 10th, nay even on the 14th day, after the
-operation. Hein. Mayer also obtained a similar result upon operating
‘on cats. The cats became emaciated and died without convulsive
- phenomena, and only became comatose shortly before death. With
‘regard to the cause of death, the author lays stress upon the decided
‘diminution or absence of glycogen of the liver (tested with iodine
‘solution), Sugar was not found in three cases where a watery ex-
-tract of the liver was made; still in one case, on the sixth day after
REPORT ON PHYSIOLOGY. 421
the ligature of the bile-ducts, diabetes was produced by puncturing the
floor of the fourth ventricle. If the animals lived after the fourth
day, a decided increase in the connective tissue of the liver was found
on histological examination. The cells of the liver were atrophied or
infiltrated with fat, but not more strongly pigmented than normal.
In the kidney, as a rule, the epithelial cells were cloudy or had
undergone fatty degeneration.
Gentro-UrINARY SysteM.—“Ascertained Facts regarding Diabetes
and Hydruria.” Résumé by Dr Brunton in Lond. Med. ec. 1. No.
42. “Erection in Birds.” C. Eckhard, Centralblatt, No. 53, 1873.
“The Carbonic Acid in the Urine in Fever.” C. A. Ewald,
Reich. und du Bois Reym. Arch. 1873. ‘Elimination of Urea.”
Rabuteau, 1’ Union Médicale, 1873, No. 107. “Hstimation of the
quantity of Albumen.” Esbach in Société de Biologie (Gaz. Méd.de Paris,
No. 5, 1874). “Condition of the Circulation in the Kidney.” A.
Hogyes, Arch. f; Hap. Pathol. 1873, 1. 289. Abstract in Centralblatt,
No. 1, 1874). “Influence of the activity of the Skin on the Secre-
tion of Urine.” K. Miiller, Arch. f. exp. Pathol. 1873, 1. 429. Ab-
stract in Centralblatt, No. 3, 1874.
PHYSIOLOGY OF THE COLLECTION OF URINE IN THE BLADDER.—G.
Edlefsen (Pjliig. Archiv, vir. 499) took a sufficient quantity of water be-
fore going to bed, and evacuated the contents of his bladder at stated
periods (7 times), and then took the sp. gr. of each portion. The first
portion evacuated in the morning showed the highest sp. gr. The
sp. gr. of those portions following became gradually less. The layers
of urine in the bladder became mixed during the day, owing to the
movements of the body ete. If the author evacuated his bladder
while on his hands and knees, the first portion in the morning had
the lowest sp. gr., and the last portion the highest. The gentle motion
of the body had not changed the position of the contents of the blad-
der. The author’s experiments show that the walls of the bladder
move round its contents during the movements of the body.
To Propuce GrycosuriA.—Ewald (Centralblatt, No. 52, 1873)
injected subcutaneously from 0,5 to 2,0 grms. of nitro-benzol (sp. gr.
1,078) into a rabbit. On pressing on the bladder to collect the urine dur-
ing the first three hours after the operation, a substance having all the
properties of sugar was found in the urine. This substance can also
be obtained plentifully till 20 hours after the injection, after that it
gradually disappears, and after 24—36 hours it can no longer be
detected, The author could not produce mellituria in dogs on inject-
ing nitro-benzol subcutaneously ; but when the drug was given by the
mouth large quantities of sugar (in one case 2°5 per cent.) were found.
CoMPARATIVE OBSERVATIONS ON THE CONSUMPTION OF SUGAR IN
Diapetic AND Non-piabpetic ANimALs.—L. Seelig, Jnaug. Dissert.
Konigsberg, 1873. (From Abs. in Centralblatt, No. 59, 1873.) The
author experimented on diabetic and non-diabetic rabbits, both of
429 DR STIRLING.
which had been allowed to hunger. Diabetes was produced by
Eckhard’s method. The author then convinced himself that in the
starving animals after the diabetic sugar had disappeared from the
urine or occurred only in traces, corresponding to the results of Dock
(the hunger period lasted 3—5 days, the collected urine was evacuated
by pressure, after it had collected for six hours), that only once by
tibiation with Fehling’s solution did he find 0-4 grms, of sugar. A
solution of sugar (generally 20 ccm. of a 10 per cent. sol. = 2 grms.
sugar) was then injected into the jugular vein, in the one case into
the starved animals, and in the other into the starved diabetic ones.
In the former case only traces of sugar appeared in the urine, when
the animals had starved for 5, 6 and 7 days, somewhat more, when
the hunger-period was shorter. As a mean of 16 experiments, 0-2 grms.
of sugar were found in the urine, while in the diabetic animals the quan-
tity of sugar excreted was much greater ; the maximum 0-925 grms.,
the minimum 0-27 grms. (in this case only 0-91 grms. were injected),
mean 0-6 grms. The mean quantity of urine in non-diabetic animals
was 25 ccm., in the diabetic 41 ccm. The diabetic animal, there-
fore, is distinguished from the non-diabetic one by its incapability of
using the sugar for the nutrition of its body. For the theoretical de-
ductions we must refer to the original.
REACTION OF URINE AND Sweat.—A. Moriggia (Moleschoti’s Un-
tersuch. 1873, 1x. 129), from experiments upon himself and animals,
found that the sweat of plant-eating animals is generally alkaline, that
of flesh-eaters acid. In animals and man the urine becomes acid during
fasting or under flesh diet, under vegetable food alkaline; but the
sweat has a reaction peculiar to the individual. To change the re-
action of the urine, it is only necessary to have a long and continued
change of the diet.
ANATOMY AND PuysioLocy oF THE KipNney.—R. Heidenhain,
Arch. f. Mikroscop. Anat. x. 1.—This paper is chiefly devoted to the
histology of the secretory apparatus of the kidney. The author de-
scribes peculiar rod-shaped bodies (Stdébchen) in the contorted tubules
and in the broad part of the loops of Henle. Experimenting with
indigo-sulphate of soda (that bought in the shops is generally impure),
the author concludes that the kidneys are the specific secreting organs
for this substance, in the same sense as they are for urea. In this
process the Malpighian capsules do not act a part, the excretion is
accomplished by the tubuli contorti. The single tubuli contorti can
act independent of each other, so that in one a living secretion may
take place, whilst in those in its immediate neighbourhood no secre-
tion is to be observed. The straight urinary tubules do not excrete
the indigo-sulphate, but serve only for conducting the formed secre-
tion. Not all the constituents which occur in the urine are secreted
in the Malpighian capsules, the author agreeing with Bowman that
the capsules secrete only water, and perhaps salts of small atomic
weight; whilst he thinks that the secretion of urea devolves upon the
tubuli contorti. What is true of the contorted tubules, is also true of
REPORT ON PHYSIOLOGY. 423
the broad part of the loops of Henle. Rod-shaped bodies were also
found in the fine ducts of the nasal, parotid, and sub-maxillary glands,
but not in those of the sub-lingual. In animals which excreted blue
urine on stimulating these glands electrically, no blue secretion was
obtained. [See fuller abstract in Lond. Med. Reed. 1. 823.]
Uterus.
On THE INNERVATION OF THE UTERUS.—Scherschewsky, under
Cyon’s direction, has arrived at the following results (Pfliiger’s
Archiv, vit. 349). 1. The uterine plexus contains the most im-
portant, if not the only motor nerves, which can produce actual
movement of the uterus, on stimaulation of their peripheral ends
(stimulation of the central ends produces only violent vomiting).
2. Stimulation of the central ends of the first two sacral nerves pro-
duces reflexly violent uterine movements, which disappear after pre-
vious section of the uterine plexus (stimulation of the peripheral ends
produce only violent contraction of the urinary bladder and rectum).
Stimulation of the brachial, crural, median, sciatic, &c., nerves, does
not produce peristaltic movements of the uterus, but only a slight
rigidity and paleness of the organ. 4. The consequences of stimula-
tion of these nerves disappear when the aorta is previously tied.
Stimulation of the central ends of the sacral nerves is still accom-
panied by peristaltic movements of the uterus, after ligature of the
aorta.
Muscle.
“Shortening of Muscle and Tendons.” Hermann, Pjliig. Arch.
vu. 417 (Journ. of Anat. and Phys. vu. 214), M. W. Englemann,
in Pfliig. Arch. vitt. 77, and Reply by Hermann at 275,——“ Differ-
ence of the Physiologica] Action of Induced Currents, according to
the Nature of the Wire forming the Secondary Spiral.” Onimus,
Acad. d. Scien. in Gaz. Méd. de Paris, No. 1, 1874. “ Das Myo-
physische Gesetz.” W. Preyer, Jena, 1874, pp. 144; critical and ex-
perimental observations on the same. B. Luchsinger, Pfliig. Arch.
vit. 538. “On two Electro-Physiological Disputed Points,” A,
Griinhagen, Pjliig. Arch. vut. 519.
Rep anp Waite Muscies oF THE RABBIT AND Ray,—Ranvier,
Comptes Rendus, txxvit. 1105. It is well known that certain animals
have two kinds of voluntary muscles, the red and the pale. Thus the
semi-tendinosus in the rabbit is red, whilst the vastus internus in
which it is lodged is pale. In the rays and torpedoes there are
muscles formed of the two kinds of fibres. The difference of colour
does not depend upon the quality of blood in the capillaries. On
stimulating the semi-tendinosus of a rabbit with an interrupted
current, it contracts gradually and progressively, when tetanised it
remains contracted as long as the irritation is continued, and returns
gradually to its original length on cessation of the irritation. A pale
424. ‘ DR STIRLING.
muscle (adductor magnus) of the rabbit stimulated by the same:
current contracts quickly and abruptly, and on cessation of the stimu-
lation returns as abruptly to its original length. The red muscles of
the rays and the few pale fibres found under the skin of the back,
exhibit similar physiological properties. These two kinds of muscles
exhibit slight differences in histological structure. A more complete
description of the physiological properties and histological structure
of these two kinds of voluntary muscle is given by Ranvier in the
Arch. de Phys. v1. 1. The time which elapses between the beginning
of the stimulation and the contraction of the pale muscle is jy of a
second, whilst for the red muscles with the same apy and in-
tensity of the stimulating current the time occupied is 75 of a second,
or about four times as long a period of latent contraction or irritation
as for the pale muscle.
As a corollary to the above the important histological fact may be
added, that Ranvier (Soc. de Biologie, Gaz. Méd. de Paris, No. 4, 1874)
has found that the transverse branches connecting the parallel
capillaries in the red muscles are provided with small dilatation-
ampulle, or little aneurysmal-like sacs. The importance of this
arrangement upon the circulation in muscle is pointed out by the
author.
DEVELOPMENT OF Muscunar FIBRES IN THE FRoG.—Petrowsky
(Centralblatt, No. 49, 1873) has arrived at the following results :
1. In the larval stage the muscle consists of fusiform fibres with
a series of oval nuclei in the middle. Neither sarcolemma nor peri-
pheral nuclei are present.
2. On the periphery of several fibres, at the end of the larval
stage, and in most of the fibres of the frog when 10 mm. long, the
nuclei and sarcolemma appear at the same time together. "These
nuclei appear for the most part as rods, but are oval nuclei, when
seen in profile.
3. The axial series of nuclei disappear, the greatest part of the
peripheral nuclei separate from the sarcolemma, increase by division,
and thus form rows of parallel peripheral nuclei. Increase of the
fibres goes hand in hand with increase of the rows.
4. In the increase of the muscles the formation of new fibres
participates, but the formation of new fibres does not take place
through division of the older fibres.
Retative Provorrion of Nerve to Muscunar Fisres.—P.
Tergast (Arch. f. Mikroskop. Anat. Bd. 1x. 36) gives the following as
the result of his investigations upon the relative proportion of nerve-
‘fibres to muscular fibres in the ocular muscles of the sheep. The pro-
portionate number of primitive nerve-fibres to muscular fibres varied
considerably in the several muscles. In the obliquus inferior there
are three or four muscular fibres for each nerve-fibre, in the obliquus
‘superior six or seven, in the rectus inferior seven or eight, and in
the externus ten. In the ocular muscles of man there appear to be
three primitive nerve-fibres to every seven muscular fibres. In the
REPORT ON PHYSIOLOGY. 425
general muscles of the body the proportion of nerve-fibres is much
less. In the biceps of a young dog the author found only one primi-
tive nerve-fibre to eighty-three muscular fibres, whilst in the sartorius
they were in the ratio of one to forty or sixty muscular fibres.
Action oF Nitrite or Amyt on Musciz.—R. Pick (Centralblatt,
No. 55, 1873) shows that this substance produces a rapid and direct
paralysis of muscular fibre.
Bone.
“Qn the Marrow of Bone.” Ch. Robin, in Journ. de [ Anat.
Jan. and Feb. 1874, 33. “Bone Absorption by means of Giant-
cells.” A. Morison, Adin. Med. Journ. ccxx. 305.“ Intercel-
lular Growth of Bone.” Schachowa, Cenétralblatit, No. 57, 1878.
“Resorption of Bone and Giant-cells.” Rustizky, Virch. Arch.
ix. 202. “Composition of Bone under~Diet poor in Lime or
Phosphoric Acid.” H. Weiske and E. Wildt, Zeitsch. f. Diol. 1x. 541).
(Abst. in Centralblatt, No. 14, 1874.) |
FEEDING with Mapprer.—Centralblatt, No. 47, 1873.—Strelzoff
in his experiments with this substance has arrived at the following re-
sults: 1. The bones of old as well as of young pigeons are coloured by
madder. The coloration occurs more quickly in young than in old
pigeons. 2. The bones of very old pigeons are either not at all or but
slightly coloured on feeding with madder, 3. Not only the osseous
tissue formed during the feeding with madder, but all that formed
previously is coloured by the madder, 4. A juice-canal system which
is in connection with the processes of the bone-corpuscles, is inter-
calated between the blood-vessels and the osseous tissue, and may be
regarded as a lymphatic system. 5. The bones are coloured in the
direction of their juice-canals by the madder.
ARTIFICIAL ARREST OF THE GRowTH OF Lone Bongs By IRrI-
TATION OF THE EpipHyses.—A. Bidder (Archiv f. exper. Pathol. und
Pharmacol. 1873, 1. 248. Plates vit. and vu.) operated on the
superior epiphysial cartilage of young rabbits. The cartilage was
either exposed and transfixed with needles or destroyed by section.
Growth of the bone was arrested either on one side only or over the
whole extent of the terminal surface, according to the part irritated,
and this effect was marked throughout the whole length of the bone as
far as the distal epiphysis. Destruction of the cartilage on the fibular
side was followed by growth of the opposite side, causing curvature of
the bone with the convexity inwards.
ConTRIBUTIONS TO THE CHEMISTRY OF Bonr.—R. Maly and
J. Donath (Site. d. Wien. Acad. d. Wissensch. uxvut. ii, Abth. Juni-
Heft, 1873) made experiments upon the solubility of bones and phos-
phate of lime in different fluids. They examined three preparations:
1. Phosphate of lime precipitated from lime-water, by the addition of
phosphoric acid and preserved under water. 2. Phosphate of lime
VOL. VIII. 28
426 DR STIRLING.
from chloride of calctum, ammonia, and phosphate of soda, dried and
ignited. 3. Powdered bones purified with alcohol and ether. 100,000
parts of water dissolved with shaking and long standing, of the first
preparation 2°36, of the second 2°56, and of the third 3:00. Under
certain circumstances salts in the water increased its power of solu-
bility: 100,000 parts of a 1 per cent. solution of chloride of ammo-
nium dissolved 16°8 of bone-powder. Pieces of the femur of an ox,
after lying several days in 2 per cent. solutions of different substances,
lost most in weight in water rich in Co,, then in sal-ammoniac, bile,
common salt, simple water. Some substances diminish the power of
water for dissolving phosphate of lime, sugars, gelatine, glycerine,
lactate of soda, &c. Co, dissolves phosphate of lime easily. Liberal
potations of water increase the quantity of phosphoric acid excreted
per urethram—a fact already well known. No increase, however, of
this acid was noted under the use of water impregnated with Co,,.
The authors then discuss the question whether the calcareous salts in
bone are really in chemical combination with the ground substance of
the bone or not. ‘They cite the facts that are favourable to such a
view, and then give experiments of their own. The authors tried to
impregnate bone-cartilage with phosphate of lime (after extracting
the bones with HCl), but in vain. They added to solutions of gela-
tine of different concentrations ammoniacal solution of phosphate of
soda and chloride of calcium solution, and each time so much, that
therefrom 1:96 grms. dry phosphate of lime Ca, (Po,), must arise. The
precipitates always contained gelatine, but the quantity varied with
the concentration of the gelatine solution; from 1-96 grms. of phos-
phate of lime 0°37, 0:47, to 0°67 grms. of gelatine were obtained, pro-
portions which distinctly speak against a chemical combination of the
gelatine with the phosphate of lime. Other gelatine-containing pre-
cipitates (oxide of iron, silicic acic, zinc oxide) produced in gelatinous
solutions, also contained gelatine, often in a high degree; thus the
iron oxide contained 51°8 of gelatine, and the zine oxide 47°8 per
cent.; other substances had also this property, as albumen of egg,
whose precipitate contained 32:4 of organic substance, that of gum
27:7 per cent. The phenomenon is therefore a purely mechanical
one, but the mixture is a very intimate one, so that even treat-
ment for days with hot water does not serve to extract all the
gelatine. In complete unison with the above is the fact that phos-
phate of lime in bones conducts itself with regard to solvents in
exactly the same way as precipitated phosphate of lime. As sup-
porting the mechanical nature of the union of phosphate of lime in
bone, see ‘‘Relation of Bone-Cartilage to Phosphate of Lime.” By
C. Aeby, in Centralblatt, No. 44, 1873.
Propuction or Rickets ArtiFic1aLLy.— L. Tripier (Arch. de Phys.
vi. 108) has tried to produce rickets artificially in young animals.
Guérin has answered this question in the affirmative. He fed young
dogs on raw flesh, instead of the milk of their mother. Tripier has made
similar experiments on young cats and dogs, and fowls, giving them
insufficient nutriment, keeping them both in the town and country to
REPORT ON PHYSIOLOGY. 427
test the effect of hygienic conditions. In no case was any affection of
the skeleton observed. Careful analyses were made of the bones of
the animals experimented on.
Milk.
PERCENTAGE OF Far ry Human Mitx.—Brunner (Abs. in Journ.
of Anat. and Phys. viti. 210) gave as the mean percentage 1°73. A.
Schukowsky (Zettsch. f. Biologie, 1x. 432), from numerous analyses
which he has made in Moscow, is convinced that this figure,is too
low. He finds that the normal milk of a healthy woman, suckling
her child, contains very seldom Jess than 3 per cent. of fat. Brunner
estimated the fat by Trommex’s method.
Excision oF THE Mamma purine Lacration.—Sinéty read a
communication before the Société de Biologie (Gaz. Méd. de Paris,
No. 3, 1874) upon this subject. Rabbits and dogs did not survive
extirpation of all the glands. Guinea-pigs were therefore employed.
The guinea-pigs operated on in September are still alive. The re-
moval of these glands is followed by the occurrence of sugar in the
urine; under the microscope globules of fat are to be detected. In a
further communication (Gaz. M/éd. No. 8, 1874) the author finds that
the absence of the mamme has no effect on the triple function of
fecundation, gestation, and parturition. Further, that in ‘female
guinea-pigs, deprived of these organs, glycosuria is not produced
during gestation or after parturition. Gestation is without influence
on the appearance of sugar in the urine, this phenomenon depending
entirely on lactation. “Reaction of Milk to Litmus.” Vogel,
Jour. f. Pract. Chem. N. F. vint. 137.
Temperature.
“Increase of Temperature by Pyrogenic Substances.” 8.‘ Dobez-
canski and B. Nannyn, Arch. f. Exp. Pathol. u. Pharmac. 1873, 1.
Sh; “Effects of Alcohol on Warm-blooded Animals.” Read
before the British Association at Bradford. Abstract in Vatwre, 1x.
132. “Temperature in Health.” Ed. Laurie. Indian Med. Ga-
zette. Abstract in Lond. Med. Rec. No. 46. “Influence of Alcohol
on the Bodily Temperature.” TF. Riegel, Deutsch. Archiv f. Klin. Med.
1873, x11. 79.——*“ Temperature of the Heart and Lungs.” Albert
and Stricker, Wien. Med. Jahrbuch, 1873, 29. Abst. in Centralblatt,
No. 3, 1874.———“Influence of Nerve Lesions upon Temperature.”
W. Mitchell, Arch. of Scient. and Pract. Med. 1873, 351. “Calo-
rification in Asphyxia.” Claude Bernard, Lancet, Oct. 3, 1873.
“‘ Doctrine of the Regulation of the Temperature.” F. Riegel, Virch.
Arch. rx. 114.——“ Anesthesia from Cold.” Horvath, Centralblatt,
No. 14, 1873. “ Action of Alcohol on the Temperature and Pulse.”
Rabow, Abs. in Centralblatt, No. 21, and Daub in Ceniralblati, No.
29, 1873. “Effects of Exercise on the Temperature and Circula-
tion.” ©. H. Jones, Proceeds. of Roy. Soc. xxt. 374. “Effects of
Exercise on the Bodily Temperature.” Allbutt, Jowr. of Anat, and
28—2
498 DR STIRLING.
Phys. vu. 106. “On the Substances which Increase the Temp. of
the Animal Body.” P. Lewitzky, Centralblatt, No. 46, 1873.
INSENSIBLE Excretion IN Fever.—Fr. Neumann (Exp. Unter-
such. iiber das Verhalten der insensibelen Ausgabe im Fieber), Jnaug.
Dissert. Dorpat, 1873. From experiments upon dogs the author
arrives at the following results: 1. During hunger the insensible ex-
cretion sinks from day to day. Variations caused by the varying con-
ditions of the atmosphere occur. 2. During the fever the insensible
excretion was in general greater than without fever under the same
conditions. 5. A parallelism seems to be present between the
degree of temperature and the insensible excretion. 4. During and
after the decrease of the fever (“crisis and epicritical stage”) an in-
crease of the insensible excretion takes place.
ON THE FoRMATION AND REGULATION oF ANnimMAL Heat (Del
Potere regulatore della Temperatura Animale, Studio Critico-speri-
mentale, by A. Murri, pp. 79. Firenze, 1873). This is an exceed-
ingly able paper. We have only space to cite a few of the author’s
more important results. A fuller abstract will be found in the Lond.
Med. Rec. No. 47. 1. The increase in the excretion of Co, during a
cold bath is probably the consequence of more complete expiration,
and even if increased formation of the same was demonstrated, this
can be explained without increased production of heat, which is not
the case. 2. The increase of temperature in the axilla during the
cold bath is quite compatible with a considerable diminution of the
quantity of the heat in the body. There is actually a diminution
of heat. It is, therefore, not necessary to assume an increased
production. 3. The calorimetric measurements of Liebermeister
and Kernig are not exact, because the cooling which the body
experiences in the cold bath was deduced from erroneous caleu-
lations. The direct proof shows that no more heat is produced
in the cold bath than under ordinary circumstances. 4. The hypo-
thesis that there exists a nervous centre which fixes the tempera-
ture in health and disease is quite untenable. The heat-regulating
centres are neither proved nor rendered probable. 5. Fever is
produced by a series of unusual chemical changes (fever-process)
which thus increase the heat-production, so that the bodily tem-
perature rises. Even continued increase of temperature, which is
not based on a special unusual biochemical process, is not feverish
(e. g. hysterical, epileptic, and tetanic convulsions).
Action oF Sweat.—A. Rohrig, Jahr. f. Balneol. 1873, 1. 1, col-
lected sweat from his forehead, and injected 3:5 ce. of it, fresh and
filtered, into the jugular vein of a rabbit at mid-day. The temperature
of the animal was 37°2° C. before the injection. Towards evening
the temperature rose, and during the night reached 40:2°C. The
heart-beats rose from 122 per min. to 326, and the respiration from
82 to 105. Next morning the temperature was 40:2° C. heart-beats
315, and respiration 215 in the minute. In two days the animal
REPORT ON PHYSIOLOGY, 439
returned to the normal. Urine during the fever contained albumen,
which disappeared as the fever subsided. What elements in the sweat
produced these effects the author has left undetermined.
INFLUENCE OF VARIOUS. DEGREES OF TEMPERATURE ON THE [RIS
OF MAMMALS AND UPON THE STRIATED MuscuLar Frere oF THE
Froc.—Griinhagen (Wageblatt der 46. Versam. deutsch. Naturf. und
Aerzte in Wiesbaden) stated that certain temperatures between 0° C.
and blood-heat exert a considerable influence on the size of the pupils
in mammalia after extirpation. In the cat, if the eye after death is
exposed to the temperature of the blood, the pupils remain widely
dilated; whilst, if temp. is lowered to thé mean temp. of the room,
they contract strongly, and dilate again strongly when the temp,
is reduced to 0°C. The author believes that this is not due to
contraction and dilatation of the sphincter pupille, as Brown-
Séquard and H. Miiller are inclined to believe, but that the cause
of the phenomenon in question is due to the varying capacity of
the tissue of the iris for the imbibition of water at different tem-
peratures. He is of opmion that the tonus of the tissue of the
iris is lost after death by the absorption of water; whilst on ex-
posure to low temperature (0°C.) it is restored by the giving up
of water. That water is really given off in the latter case is rendered
probable by the reaction of the lens to cold, for this body becomes
cataractous at a freezing temp., owing to the formation of vacuoles in
its substance. The impaired irritability of the iris can be restored
even after two days by placing the iris in a chamber at blood-heat.
The author also showed that on cooling down the muscular tissue of
the frog to the freezing point, its irritability, upon the application. of
a mechanical stimulus, was greatly increased. In this we possess a
means of producing muscular contraction mechanically, without the
participation of nerves, The contractions so produced can produce
secondary contractions. “On the Substances which increase the
Temperature of the Animal Body.” P. Lewitzky in Centralblatt, No.
46, 1873. “Tnerease of Temperature in Fever.’ Dobezcanski and
B. Nannyn. Arch. f. exp. Path. and Pharmac. 1873, 1. 1. 181.
On THE HEAT PRODUCED IN THE Bopy, AND THE EFFECTS OF
Exposure To Cotp.—Draper (American Jour. of Scien. and Arts,
Dec. 1872) wished to determine the quantity of heat passing from
the surface of the body by finding how much it would elevate the
temperature of a known mass of cool water during a given period of
time. By lying quietly for one hour in a bath of 74° F., enough heat
was lost from the body to raise the temperature of the water 2° F. and
to lower that of the body 1° F. (temp. measured in mouth and axilla).
The volume of water in the bath was 7} cub. ft., and that of the body
3, and the author concludes that enough heat is evolved from the
body in one hour to warm the body itself 5°F. Thermometers in the
mouth and axilla indicate a steady fall of temperature during the
bath and for a short time after, leaving it accompanied by a diminished
rate of respiration and pulse. When the experimenter executed con-
430 DR STIRLING.
tinued muscular movements in the bath, the temperature of the water
around him was not raised any higher than when he remained perfectly
at rest.
BEHAVIOUR oF CoLD-BLOODED ANIMALS AT FREEZING TEMPER-
ATURE.—Doehnoff (Reich. und du Bois’ Arch. 1872, p. 724) states that
cold-blooded animals conduct themselves like plants in cold. The honey-
bee dies at —1°C., the spider —3°; the flesh-fly can endure a temper-
ature of — 6°, and the silkworm’s egg survives a cold of — 21°. Part of
the water contained in the leech can be frozen, and the silkworm can
be solid ice without this being prejudicial to their life.
On THE Capacity or Frocs FoR RESISTING High snp Low
TEMPERATURES.—Mueller (Reichert’s Arch. 1872, p. 754) placed both
Rana esculenta and temporaria in water which was then frozen. The
vessel was allowed to remain in the open air five hours (temp. of air
5° and 7°R.). Afterwards the vessel was brought into a moderately
warm room, and when the frog was freed from the ice, after about 13
hours, it breathed quite lively at the surface of the water. Leuciscus
rutilus treated in the same way died after a short time. When a frog
has been for a long time removed from the water, so that its skin is
dry, it becomes motionless, but the vital processes are not interrupted,
as is shown by the circulation. Freezing completely controls the cir-
culation at first; upon thawing, not the slightest trace of movement
is to be observed in the web of the foot. Later, but slowly, the vessels
begin to show signs of life. Frogs placed in water at a temperature
of 20° R. were lively, at 26° they seemed tired, and at 28° death en-
sued rapidly. It seems probable that the temperature of warm-blooded
animals produces death in the cold-blooded.
THe TEMPERATURE OF THE Bopy 1n HeAttH.—Juergensen, Leip-
zig, pp. 60, 1 plate, 1873. These papers have already appeared in the
Deutsch. Archiv f. klin. Medic. The temperature measured in the
rectum of a person at rest has a daily variation of about 1°C., the
highest temp. being in the afternoon, and the lowest between mid-
night and morning. Taking of food slightly increases the temp.
With regard to the influence of the extraction of heat on the tem-
perature, the person experimented on found that the subjective feel-
ing of cold during and after the bath at 9°—11°C., lasting 25’—30, is
so intense as scarcely to be endured. Bath of 30° C. and 25’ duration :
during the bath the temperature measured in the rectum showed an
increase. Bath of 9°—11°C. and 25’ duration: during the bath the
temp. measured in the rectum showed a greater or less fall. In
these cold baths, soon after the beginning of the experiment, the
development of heat is diminished; it then remains some time con-
stant, and then diminishes further. Thus, after 5 mins.’ immersion in
such a bath, the rectal temp. in different experiments on the same
person showed a variation of 0°5°, 0-7°, 0-2°C. less than the temp. at
the beginning of the experiment. Then followed a longer or shorter
period when the temp. remained constant. After 25 mins.’ immersion
REPORT ON PHYSIOLOGY. 431
the rectal temp. showed a diminution of 1°4°, 0:9°, 1-1°C. under the
same circumstances. The after-effects of immersion in such a bath.
The greatest cooling of the body takes place not in the bath, but some
time (hours) thereafter. After a bath of 10°—11°C. the minimum temp.
of the after-effect 33-1° (3-6° under the normal), and only after seven
hours was the normal reached. The duration of the bath also influ-
ences the after-effects. Period of the day or night also. During the
night the remote effects appear less than during the day. The after-
effect increases with the time, and is cumulative. Thus, after a
series of experiments, the after-effect was so pronounced that even
after 25 mins.’ immersion in 9°C. bath no diminution of the
temp. was observed. Then follow chapters on the influence of quinine,
muscular exercise, etc. on the temp., and also a chapter on the temp.
during the first week of life.
La CHaLeur AnIMALE.—Bernard (Revue Scientifique, 1. 133 et seq.)
in the above lectures discusses the theory of Koerner and Heidenhain
(Pjliig. Arch. tv. 558), that the difference between the temperature of
the blood in the right and left ventricles is due to the proximity of
the right ventricle to the abdominal organs (Journ. Anat. and Phys.
Vol. vil. p. 358). He rejects this theory, for Hering has found that
in a case of ectopia cordis the temperature of the right ventricle, even
in this malformation, was higher than that of the left.
Miscellaneous.
Nucteme.—Worm Miiller, as the results of his researches upon
this substance, has arrived at the following conclusions (Pfliig. Arch.
vu. 190). Nucleine, as at present prepared, is no single characteristic
chemical substance, and this is shown by the different results of
different investigators as to the quantity of P. contained in it. Nu-
cleine is probably only a mixture of organic phosphorous compounds
with albuminous or albuminous-like bodies. It cannot with certainty
be asserted that nucleine belongs entirely to the nuclei..
INDEX TO VOLUME VIII.
A.
Abnormalities, see ‘‘ Variations ”
Absinth, effects of, 220
Absorption, of bone by giant-cells, 425 ;
in large intestine, 413
Achscharumow, aconitia, 22
Acid, uramid, 410; taurocarbamin, syn-
thesis of, 410; carbonic, in urine
in fever, 421; nitric and sulphuric,
toxicology of, 222; origin of, in ani-
mal organism, 421; of bone, 425
Ackermann, action of digitalin, 227
Aconitia, action of, 223
Afonassiew, oxygen in blood, 196
Albert, temp. of heart and lungs, 427
Albumen, 204; destruction of, 206;
action of gases on, 408; preparation
of, 410; quantity of 410, 421; of he-
patic cells, 209, 419; nitrogen in,
410, 411
Alcohol, effects of, 220, 233, 427
Alimentation, see “ Digestive”
Alkali, withdrawal of, from living body,
410
Allbutt, bodily temp., 427
Amez-Droz, nitrite of amyl, 221
Amphibia, tissue-metabolism in, 122;
anat. report, 177
Amyl, nitrite, effects of, 221; on muscle,
425
Anesthesia, from cold, 247
Annelids, 385
Antagonism, 23I—232
Apolant, white and red corpuscles after
suppuration, 403
Apomorphia, emetie action ef, 223
Apneea, 204, 409
Araneina, embryology of the, 170
Arctictis binturong, anat. of, 176
Arloing, the vagus, 185
Armadillo, enamel-organ in, 387
Arndt, R., nerve-endings in striated
muscle, 161; chloral, 220; ganglia
of the sympathetic, 392; path. anat.
of central organs of nervous syst.,
392 :
Arnold, J., Diapedesis, 403)
Aroustein, R., preparation of albumen,
410
Arsenic, not poisonous, 218
Arteries, see ‘‘Circulatory ”
Arthropoda, tissue-metabolism in some,
121
Asphyxia, calorification in, 427
Atropia, action of, 225; antagonism
between physostigma and, 232, 403
Aubert, carbonic’ acid excreted by skin
in man, 188
Audigné, production of icterus, 420
B.
Bacteria, in sweat, 188, in blood, 198
Balandin, J., curvatures of spine, 159
Balbiani, M., embryology of the ara-
neina, 170
Balfour, F. M., develop: and growth of
blastoderm of hen’s egg, 168, of blood-
vessels, 169; primitive groove, a
temporary structure, 169
Basch, effects of nicotia, 226; move-
ments of intestines, 417
Baumstark, cholic acid, 209
Béchamp, organic particles in milk,
209; coagulation of caseine by ren-
net, 209 :
Becquerel, electro-capillary phenomena
of life, 410
Bellamy, E., malformation of wrist and
hand, 383
Beneden, P. J. Van, cetaceans, 176
Benedikt, M., innervation of infr. cho-
roid-plexus, 392
Bennett, A., action of thein, caffein,
guaranin, cocain, theobromin, 225
Benzine, toxicology of, 222
Bergeret, absorption, of bismuth, 218,
of mercury, 219, of lead and gold,
21
Ted R., Nudibranchs, 178
Berlin, section of optic nerve, 187
Bernard, M. Claude, diabetes: and the
glycogenic function of liver, 418}
calorification in asphyxia, 427; la
chaleur animale, 431
——
INDEX.
Bernheim, electricity in muscle and
nerve, 213, 400
Bernstein, electrotonus, 213
Bert, phenomena of life, 215; capacity
of blood for oxygen at various pres-
sures, 403
Bidder, A., bone-growth arrested by
irritating the epiphyses, 425
Binz, action, of quinine on blood, 198,
224, 402, of alcohol on warm-blooded
animals, 233, of oxygen in formation
of pus, 403
Birds, report, anat., 177; erection in,
421
Bismuth, detection of, in-tissues, 218
Blake, C. J., galvanic current upon
acoustic nerve, 400
Blake, J., introduction into blood of
inorganic substances, 243
Blanche, nitrous oxide in plants and
animals, 221, 409
Blasius, frog’s heart under varying
blood-pressure, 189
Bloch, capillary circulation in skin, 402 ;
injuries to skin, 402
Blood-vessels, see ‘‘ Circulatory ”
Bochefontaine, phys. of spleen, 185
Bohm, action, of arsenic, 218, of vera-
tria on muscular fibre, 213, of amor-
phous aconitia, 223, of digitalin, 227
Bottcher, movements after section of
the semi-circular canals, 187; trau-
matic keratitis, 400
Bogoslowsky, action of flesh-broth, &c.,
20
Boll, F., histology and histogenesis of
central organs of nervous syst., 172;
electric plates, of Malapterurus, 392,
of Torpedo, 392; phys. of the ner-
vous syst., 399
Botany, Madagascar poison, 102
Bouillaud, normal and abnormal pulse,
402 :
Bowditch, H. P., nerves of heart, 193
Bradley, S. M., natural characteristics
of skulls, 386
Brain, see ‘‘ Nervous syst.”
Braun, H., secretion of gastric juice,
I
Bromides, reflex excitability, 229
Brown-Séquard, ecehymosis by nervous
influence, 403
Brunner, J., comp. of human milk,
209, 427 ;
Brunton, J. Lauder, action, of drugs in
animal organism, 95, of digitalis on
temp. and circulation, 189; warmth
in preventing death from chloral,
3323; diabetes and hydruria, 421
Buchanan, G., Wilson’s Anatomist’s
Vade Mecum, 157
Buck, H., auditory ossicles, 187
435
Bunge, chloride of sodium and potassium
salts in man, 205
Burnett, auditory ossicles, 187; ex-
ternal ear, a synthetic resonator, 400
Bus, M. du, remains of the Delphinide,
176
Busch, strychnia on sensory nerves, 399
Butzke, V., the eerebral convolutions,
171
Caffein, 225, 229
Calabar-bean, action of, on heart, 403
Calcium-salts, action of, 218
Carbohydrates, 415
Carnivora report, anat. 17
Cartilages and synovial membranes of
joints, 127
Carville, cerebral currents, by induced
electricity, 396
Caseine, coagulation of, by rennet, 209
Cassowary, ‘‘moderator band” in heart
of, 173
Cat, action of Madagascar poison on,
107
Cathcart, C. W., lamb with fissured
sternum and transposed origin of
right subclavian artery, 321
Cetacea, anat. report, 176—177
Champneys, F., septum atriorum of
frog and rabbit, 340
Chatin, J., visceral anat. of Viverra
civetta, 176
Chauveau, Prof., Comparative Anatomy
of Domesticated Animals, 157
Chick, develop. of blood-vessels of, 169
Chloral, 220, 229, 332
Chtapowski, nerve-influence on arteries
of submax. glands, 184
Circulatory system ;—synthesis of mo-
tion, 300; reports, phys., 188—199, 402
—40g;—arteries, irregular, 155, 3215
nerve-influence on dilatation of, 183 ;—
blood-vessels, develop. of, 169 ; inner-
vation of, 182, 185 ;—blood; size of
corpuscles in the Salmonide, 178 ; in-
troduction of inorganic substarcesinto,
243; formation of colouring matter
of bile, 420 ;—-cizculation in capillaries,
402, 427; in kidney, 421 ;—heart, atro-
pia on, 226; temp. of, 427 ;—lympha-
tics, of lungs, 390, of cornea, 390, of
uterus, 391, of serous membranes,
388, in dura mater, 174, in retina and
vitreous body, 174 ;—lymph, secretion
of, in the fore-limbs of dog, 199;—
pulse, secondary waves, 1; frequency
of, 54, 189; action of alcohol on, 427;—
spleen, innervation of, and its rela-
tion to leucocythemia, 185; pbysio-
434
logy of, 185 ;—veins, communication
between umbilical and portal, 149;
innervation of, 392; portal circulation,
418
Clapham, W. C. §., weighings of the
brain, 173
Claus, veratria, 228
Cleland, Prof., double-bodied monsters
aud develop. of tongue, 250 ; accessory
lobes of lung, 388; Animal Physio-
logy, 384
Cocain, action of, 225
Collins, E. W., accessory lobe of right
lung, 387
Convulsants, action of, 2
Costa, intestinal glands, 205
Coxghtrey, M., tracheal pouch of Emeu,
177
Crab, anat. of American King, 178
Crampe, H., length of intestines, 175
Crum-Brown, <A., sense of rotation
and function of the semicircular ca-
nals, 327, 402
Crustacea, anat. report, 178
Curnow, Se irregularities, in ophthal-
mic and meningeal arteries, 155; in
some muscles, 377
Cutaneous syst., see ‘Integumentary ”
Cyon, influence of posterior nerve-roots
on sensibility of anterior, 399; reflex
stimulation of vaso-motor nerves, 403
Czerney, V., digestion and absorption
in human large intestine, 413
D.
Darwin, F., sympathetic ganglia of
bladder, 392
Daub, alcohol on temperature and pulse,
247
Davidson, A., Madagascar ordeal poison,
97
Day, J., fat, 409
Death, muscular’ irritability after sys-
temic, 213
Decidua, see ‘‘ Genito-urinary”
Defecation, 410
Delore, M. X., maternal circulation in
placenta, 393
Delphinide, remains of the, 176
Development, see ‘‘ Embryolo
Dewar, J., physiological action of light,
187
Dew-Smith, A. G., double nerve-stimu-
lation, 74, 400; sugar-forming sub-
stance in penicillium, 82
Diabetes, 209, 418, 421
Diapedesis, 403
Digestive system of Arctictis binturong,
176, of Viverra civetta, 176, of Su-
matran Rhinoceros, 176; epithelium
INDEX.
of cesophagus, 162; stomach and nerve-
centres of circulation, 182; gastric
juice, 274; blood-vessels of small in-
testine, 174;— liver, physiological re-
ports on, 209, 418—421 ;—pancreas,
23 ;—ectopia viscerum, 171 ;—tongue
of the genus Nestor, 177, development
of, in double-bodied monsters, 250 ;—
teeth ; bidental skull of Narwhal, 133 ;
supernum. molars in Orang, 140; den-
tition of rhinoceros, 176; anat. report,
378 ;—reports, anat., 175, phys. (on
alimentation), 204—209, 409—417
Digitaln, aie 231
Dobczcanski, , increase of temperature
by pers substances, 427, 429
Doehnoff, cold-blooded animals at freez-
ing temperature, 430
Dog, olfactory epithelium of, 42
Dolphins and whales of New Zealand
Seas, 176
Donath, J., chemistry of bone, 425
Donders, expulsion of nitric oxide from
blood, 197; influence of spectacles on
acuteness of vision, 400
D’Ornellas, A. E., action of certain
enietics, 230
Draper, heat of the body, 429
Dromaius Nove-Hollandiz, anat. of,
177
Deckert M. G., anat. of Dromaius
Nove-Hollandiz, 177
Dupuy, Dr E., la physiologie du cer-
veau, 396
Duquesnel, aconitia, 223
Duret, secondary cerebral currents, by
induced electricity, 396
Dusart, Brothers,albuminous substances,
410
Dwight, T., ischio-trochanteric lig., 134
Dybowski, B., skull of Phoca baica-
lensis, 176
Dyspnea, from warmth, 203
E.
Ear, innervation in rabbit, 184; semi-
circular canals, function of, 327, 400,
402
Eberth, keratitis after section of trige-
minus, 187; bacteria in sweat, 188;
inflammation of cornea, 400
Ebstein, formation of pepsin, 207
Ecker, Dr A., Convolutions of Human
Brain, 158
Eckhard, C., sympathetic on the eye,
187, on erection in birds, 421
Edlefsen, G., phys. of collection of
urine in bladder, 421
Egg, see “Ovum”
Egli, T., glands in pelvis of kidney, 175
INDEX.
Eichhorst, H., nerve degeneration and
nerve regeneration, 400
. Electricity, 423; on heart, 403; on life,
410
Electrotonus, 213
Elephant, Indian, 85
Embryology; develop., of frontal bone,
159, of cranium of Sus scrofa, 176, of
sternum in tortoises, 177, of nerve-
cells, 181, of tongue, 250, of muscular
fibre in frog, 424, of Wolffian duct,
393; of dental follicle, 387; imperfect
of limbs; anat. report, 168—170
Emetic, 223, 230
Emeu, tracheal pouch, 177
Englemann, irritation of nerves and
muscles, 186; shortening of tendons,
214, 443
Epithelium, olfactory, 39; stratified,
162; of serous layer of ovum of
Rabbit, 170; of serous membranes,
390
Breland G. B., placenta, 165, 393
Ergotin, action of, 228
Esbach, albumen, 421
Eustachian tube, opening and closing
of, 127
Ewald, C. A., apncea, 204; carbonic
acid in urine in fever, 421; glyco-
suria, 421
Ewart, J. C., epithelium of retina and
capsule of lens, 353
Exner, §., simplest psychical processes,
3
Eye, of lobster, 178; sup. cerv. gan-
glion and iris, 398; lymphatics of
retina and yitreous body, 174, of
cornea, 390; epithelium, of cornea,
162, of retina and of capsule of lens,
353; reports, anat., 173; phys., 187,
400
F,
Faber, C., red blood-corpuscles, 403
Falck, phys. of water, 205; chloride of
sodium as a poison, 219; action of
hydrocotarnin, 224; peculiarity of
capillary blood, 406
Fasciz and muscles about urethra, 160
Fat, 204, 207; absorption of, 205, 209;
assimilation of, 410, action of, 409;
in milk, 427
Feinberg, suppression of perspiration,
182
Ferment, in stomach, 207 ; pancreas, 23,
409
Ferrier, Dr, researches in cerebral phys.
and path., 152, 179
Fever, urine in, 421, 428
Fibrin, digestion of, 204
435
Fick, electrical irritation of nerves, 186;
blood-pressure, 189, 402; stomach-
ferment, 207; veratria, on muscular
fibre, 213
Filebne, W., carbonic acid in apnea,
409 ; curve in transyv. conduction of
frog’s nerves, 186, 400
Fishes, respiration of, 203; anat. report,
177—178
Fistula, biliary, 418
Flesch, M., malform. of thorax, 170
Flesh, feeding with, 204, 207, 415
Flint, A., nervous syst., 399
Fokker, alkaline blood, 199
Food, mechanical coefficient of, 204
Forster, J., nutrition, 409; ash-consti-
tuents in food, 411
Foster, M., temperature on reflex ac-
tions of frog, 45, 400; reply to Lewes’
‘*sensation in spinal cord,” 400
Fournié, cerebral phys. and path., 179
Fraser, T. R., report on pharmacology,
217—232
Frohlich, atropin and physostigma on
pupil and heart, 403; hemorrhage
after ligature, 403
Frog, transv. conduction in nerves of,
186; reflex actions, 45, 400; Auer-
bach’s plexus, 173; develop. of mus-
cular fibres, 424 ; action, of Madagas-
car poison, 105, 108, of temp. on
striated muscle of, 429; resistance to
high and low temp., 430; heart, 189,
191; septum atriorum, 340; lung,
188 ; migration of corpuscles, 174,
403; olfactory epithelium, 42
Fuchs, equilibrium for stimulated and
non-stimulated muscles, 213
G.
Galabin, A. L., causes of secondary
waves of pulse, 1, 112, 403
Galton, J. C., translation of Ecker’s
Convolutions of the Brain, 158
Galvanism, current of, on eye, 400,
on acoustic nerve, 400; action of in-
duced currents, 423
Garrod, A. H., law regulating frequency
of pulse, 54, 189, 403; placenta in
Hippopotamus, 167 ; anat. of Arctictis
binturong, 176; visceral anat. of Su-
matran Rhinoceros, 176; nasal bones
of some birds, 177; tongue of the
genus Nestor, 177
Gegenbaur, Carl, Grundriss der ver-
gleichenden Anatomie, 385
Genito-urinary system, of Arctictis bin-
turong,176; phys. report, 421—423;—
bladder, epithelium of, 162, extrover-
sion of, 171, sympathetic ganglia of,
we
436. INDEX.
392 ;—kidney, large calculus of, 382;
anat. reports, 175, 392; ovary, anat.
report, 393 ;—penis, muscles and fas-
ciz, 160 ;—placenta, 162, 164, 165, of
hippopotamus, 167, of sloths, 362 ;
anat. report, 393 ;—uterus, pregnancy
in rudimentary cornu of, 171, re-
flex movements of, 214, lymphatics
of, 391; decidua, 167, 393; nerves,
403
Genzmer, A., path. of lungs after section
of vagi, 185
Gerlach, L., nerve-endings in cortex
of cerebrum, 171; myenteric plexus
of Auerbach, 173; minerals of blood-
serum, 196
Gervais, P., monsters, 171
Giacomini, C., veins of leg, 174
Gierke, respiratory centre, 409
Glands ; Brunner’s, 205 ; intestinal, 205 ;
in. pelvis of kidney, 175; of head of
Ophidians, 177; effects of stimuli on,
206
Glycerine, nitro-, toxicology of, 222
Glycogen, source of, in liver, 209, 418,
419: diabetes, and function of, in
liver, 209, 418
Glycosuria, 421
Gobrecht, defecation, 410
Gotte, A., develop. of the Vertebrata,
170
Gold, in tissues, 219
Goldstein, dyspncea from warmth, 203
Goltz, the nerves of erection, £86
Gray, J. E., pig-skulls, 476; dentition
of rhinoceros, 176; skeletons of Kogia
Macleayii, 176; sternum in young
tortoises, 177
Gréhaut, absorption of oxygen by blood,
199; aconitia, 22
Gruber, W., os zygomaticum biparti-
tum, 159; supernumerary bones in
zygomatic arch, 386; variations, in
foramen mentale, 159, in dental fora-
men and mylo-hyoid groove, 386, in
cranial bones, 159; articulation of
temporal with frontal bone, 386 ; bones
in frontal fontanelle, 386
Griinhagen, A. V., anat. of iris, 173;
‘temp. on iris and on striated muscie
of frog, 429 ; irritation of nerves and
muscles, 186; contractions in muscle,
186; on two electro-physiological dis-
puted points, 423
Griitmer, nerve-influence on submax.
glands, 184; pepsin, 207; reaction of
muscle, 213
Gscheidlen, R., chemical reaction of
central nervous system, 181; quantity
of blood, 199, 403
Guaranin, action of, 225
Gubler, action of chloral, 220
Gulliver, G., blood-corpuscles of the
Salmonide, 178
Guttmann, P., paralysis of vagus, 399
Guyochin, absorption of quinia, 225
iH:
Hemoglobin, variations of, in the zoo-
logical series, 1g6
Handyside, P. D., new species of Polyo-
don, 178
Harting, P., the swimming bladder, 177
Hayem, G., changes in spinal cord after
excision of sciatic nerve in rabbit, 186
Hector, J., whales and dolphins of New
Zealand Seas, 176
Hehn, A., mechanical cedema, 405
Heidenhain, R., anat. and phys. of
kidney, 392, 422
Hein, R., ectopia viscerum and imper-
fect limbs, 171
Heinzmann, gradual thermal irritation
of nerves, 186
Heller, A., blood-vessels of small intes-
tines, 174
Helmholtz, H., ossicles and membrana
tympani, 400
Henle, Dr. J., Handbuch der systemati-
schen Anatomie des Menschen, 157
Hennig, C., human placenta, 164
Hensen, accomm. movement of choroid
in eye of man, ape and cat, 400
Hermann, L., galvanic currents on
muscles and nerves, 186; electroto-
nus, 213, 400; shortening of tendons,
214, 423
Hesse, O., hydrocotarnin, 224
Heubel, E., comp. of tobacco-smoke,
226
Hippopotamus, placenta in, 167
Hirschberg, J., refractive index of eye,
400
Histology, and phys. of nerves, 186
Hitzig, E., trans. conduction in frog’s
nerves, 186, 400; phys. of the brain,
397
Hees A., circulation in kidney, 421
Hoffmann, F. A., nitrite of amyl, 222
Hoffman, assimilation of fats, 205
Hoggan, blood-pressure in inspiration,
203
Hollis, W. A., tissue metabolism, 120
Holmes, ergotin, 228
Honckgeest, Van Braam; movements
of stomach and intestines, 209
Hoppe-Seyler, destruction of albumen,
206
Horvath, movements of intestines, 208 ;
anesthesia from cold, 427
Hiiter, circulation in frog’s lung, 188
Humphry, Prof., depressions in parietals
‘
tie iti a i i Nt i i ee he
INDEX. 437
of orang and man, supernum. molars
in orang, 136; use of ligamentum
teres of hip, 295
Huppert, H., uramid acid, 410
Hydrobilirubin, absorption spectrum of,
209
Hydorcotarnin, action of, 224
Hydruria, 421
Hyperoodon bidens, rudimentary <finger-
muscles in, 114
Icterus, 420
Incanity, cerebral histology of the insane,
173
Integumentary system in Ophidians,
177, in larva of Salamandra maculosa,
177; irritation of, 186; touch-cor-
puscles, 30; sweat, 422, 428; reports,
anat , 175, phys., 188, 402; see also
» “ Epithelium”
J.
Jacobson, pressure in the pericardium,
18
ieebeh: R., pregnancy in rudimentary
uterine cornu, 171
Jago, J., visible direction, 187
Jhering, H. V., develop. of frontal bone,
159
Jochelsohn, artificial respiration with
strychnia, 409
Jolyet, nitrous oxide, 221, 409
Jones, C. H., exercise on temp. and
circulation, 427
Juergensen, temp. of body, 430
K.
Keratitis, after section of trigeminus, 187
Kessel, accommodation of the ear, 187
Kiewiez, A., pressure in the pericardium,
189
Klein, E., Auerbach’s plexus in frogs
and toads, 173; relation of serous mem-
branes to lymphatics, 388 ; lympha-
tics of the lungs, 390
Klemenziewiz, R., demonstration of
pulse by flame, 403
Koehler, H., saponin and digitalin, 231;
Calabar bean on heart, 403; bitter sub-
stances on circulation and blood-pres-
sure, 406
eee A., absorption, &¢c., of bone,
3°7
Kogia Macleayii, skeletons of, 176
~
Korowin, secretion of pancreas and paro-
tid, 206
Krause, contraction of muscular fibre,
213
Kreatin, 204
Krolow, Brunner’s glands, 205
Kronecker, H., fatigue of striped muscle,
210
L.
Lemargus borealis, anat. of, 285
Landau, L., secretion of pancreas, 410
Landois, L., transfusion of blood, 405
Langerhans, P., stratified epithelium,
162; histology ef heart, 173; touch-
corpuscles and rete Malpighi, 175;
skin of larva of Salamandra maculosa,
177
Lankester, E. Ray, primitive cell-layers
of embryo, 170
Laquer, miecrometry of eye, 400
Latschenberger, J., digestion, &c., in
large intestine, 413
Laurie, E., temp. in health, 427
Lead, detection of, in tissues, 219
Leber, T., change of fluids in the eye, 400
Lee, R. J., sight in birds, 173
Leeches, tissue-metabolism in, 120
Legg, J. W., changes in liver after liga-
ture of bile-ducts, 420
Legros, C., dental follicle, 387
Lehmann, nitro-benzine, 223
Lemur, Madagascar poison, 106
Leopold, G., lymphatics of uterus, 391
Lepine, R., peptic gastric glands, 416
Lesshaft, P., muscles and fascixe about
urethra, 160
Leucocythemia, and
spleen, 185
Lewes, G. H., sensation in spinal cord,
400
Lewitzky, P., temp. of body, 428, 429
Leydig, F., glands of head in Ophidians;
177; skin of Ophidians, 177
Life, 215, 410; synthesis of motion, 300.
Ligaments; ischio-trochanteric, 134; use
of ligamentum teres of hip, 291
Light, phys. action of, 187
Litzmann, C.C., Th., extroversio vesice,
7
Liver, see “Digestive syst.”
Liversidge, A., amylolytic ferment of
pancreas, 23, 409
Lobster, eye of, 17
Lockenberg, E., movements of respira-
tion, 203
Lubimoff, embryonal develop. of nerve-
cells, 181
Luchsinger, B., glycogen in liver, 418;
Preyer’s “das myophysische Gesetz,”
423
innervation of
438 INDEX.
Luciani, L., periodic function of isolated
frog’s heart, 191
Lussana, portal circulation, &., 418
Lutze, E. A., Mechanik der Herzcon-
tractionen, 403
Lymph, see ‘‘Circulatory syst.”
M.
Macaques, obs. on, 175
Mach, accommodation of ear, 187
Madder, feeding with, 425
Maerker, M., estimation of N. in albu-
men, 411
Magitot, E., dental follicle, 387
Magnan, alcohol and absinth, 220
Major, H. C., cerebral histology of the
insane, 173
Malapterurus, electric plates of, 392
Malformation, of pelvic limbs, 358, of
wrist and hand, 383; anat. report,
I7O—I7I
Maly, R., chemistry of bone, 425
Mamme, excision of, during lactation,
Mammalia, placenta in, 165; dental fol-
licle in, 387; temp. on iris, 429
Marey, uniformity of heart’s action, 190
Marrow of bone, 387, 425
Martin, H. N., structure of olfactory
mucous membrane, 39
Mathieu, E., action of gases in coagula-
tion of albumen, 408
Mauthner, J., maternal circulation in
rabbit’s placenta, 393
Mayencon, absorption of, bismuth, 218,
mercury, 219, lead and gold, 219
Mayer, 8., relations of stomach to nerve-
centres of circulation, 182; electrical
stimulation of heart, 403
M‘Intosh, W. C., A Monograph of the
British Annelids. Part 1., 385
M°Kendrick, J. G., phys. action of light,
187; mechanism of ear, 400
Mechanism of Eustachian tube, 127, 187;
of auditory ossicles, 187
Mégevand, action of digitalis, 228
Meihwizeen, reflex excitability, 229
Melassez, number of red and white cor-
puscles, 408
Membranes;—serous, structure and rela-
tions of, to lymphatics, 388; lympha-
tics, 391; enithelium, 390
Mercury, detection of, in tissues, 219
Merkel, F., the femur, 386
Metabolism, 120
Metschnikoff, action of vagus on heart,
I 5
Meyer, A. B., action of digitalis on cir-
culation and temp., 189; periodicity of
action of heart, 404
Michell, J., blood and lymph-channels
of dura mater, 174
Michelsohn, post-mortem rigidity of
muscles, 213
Mihalkovics, V., pecten of birds, 177
Milk, organic particles in, 209; coagu-
lation by rennet, 209; comp. 209;
phys. report, 427
Mitchell, W., influence of nerye-lesions
on temp., 427
Mohlenfeld, peptones, 204
Moin, cubic space and volume of air,
203
Mollusca, tissue-metabolism, 121; re-
ports, anat., 178
Monsters, 171
Morano, lymph-sheaths of choroidal
vessels, 400
Moreau, innervation of ear of rabbit,
184; action of purgatives, 231
Moriggia, A., reaction of urine and
sweat, 422
Morison, A., bone-absorption by giant-
cells, 425
Morphia, reflex excitability, 229
Morphology, of skull, 62
Mosso, chem. irritation of nerves of
heart, 193 ; movements of cesophagus,
416
Miller, sensibility to sound, 187; activity
of skin, &c., 421; skin of frog, 430
Worm, nucleine, 431
Munk, H., of moist porous bodies, 410
J., excretion of bile, 209
Murie, J., the Macaques, 175; Upu-
pide, 177
Murri, A., animal heat, 428
Muscular system ;—in digits of toothed
whale, 114}; intercostal, 203; influ-
ence of temp., 429; irregularities in,
150, 377; galvanic currents on, 186;
secondary contractions in, 186; irri-
tation of nerves, 186; of larynx, 203;
nerve-lesions on, 186; respiratory,
161, 203 ; effect of section of semicircu-
lar canals on, 187; of intestines, 208 ;
of uterus, 214 ;—reports, anat., 160—
162, phys., 210—214, 423—425.
N.
Nannyn, B., increase of temp. by pyro-
genic substances, 427, 429
Narwhal, bidental skull of, 133; rete
mirabile of, 176; enamel-organ in,
8
Nae albumen, 204, 410
Nedsvetski, moving particles in blood,
198
N anaus system ;—nerve-centres; rela-
tions of human cerebrum to skull and
head, 142, 359; phys. and path,
152; blood and lymph-channels, 174 ;
INDEX.
vaso-motor and uterine, 403; convo-
lutions of the brain, 158 ;—nerves,
variations, 297; proportion of, to
muscular fibre, 424; functions of, in
larynx, 203; action of electricity on
muscle and, 213; influence of certain
substances upon, 45, 229; double
stimulation of, 74; chemical irritation
of, of heart, 193; of liver, 209; of
limbs, 402; of striated muscles, 161;
of olfactory mucous membrane, 39;
touch-corpuscles, 175; electrical plates,
392; reports, anat., 171—173, 392,
phys., 179—188, 395—402
Nestor, tongue of the genus, 177
Neumann, F., insensible excretion in
fever, 428
Newt, olfactory epithelium
tissue-metabolism in, 122
Newton, E. T., eye of lobster, 178
Nicotia, effects of, 226
Nitrous oxide, on plants and animals,
221
Norton, A. T., anat. of ciliary body,
of, 39;
7
Mot magal: H., cerebral phys. and
path., 179, 3953; extirpation of nuclei
lenticulares, 396; injury to brain with
pulmonary hemorrhage, 397
Nowak, nitrogen in albuminates, 204
Nucleine, 431
Nudibranchs, 178
Nussbaum, respiration, 203
O.
Obermeier, O., varicose axial cylinders,
173
Cidema, 405
Ogilvie, L., lamb malformed, 321
Ogle, J. W., absence of lower limbs, 358
Olivier, A., poisoning by hyperchloride
of mercury, 219
Onimus, intercostal muscles, 203; in-
duced currents, 423
Ophidians, see ‘‘ Reptilia ”
Orang, depressions in parietals of, 136;
supernum. molars in, 140
Oser, effects of nicotia, tobacco-smoke,
226
Osler, bacteria in blood, 198; atropia
and physostigma, 232
Osseous system ;—of Kogia Macleayii,
176 ;—cranium of Phoca baicalensis,
176; of Pig, 176; of Rhinoceros, 176;
relations of human cerebrum to, 142,
359; depression in parietals, 136;
jaws of the porpoise, 176; bidental,
of narwhal, 133; auditory ossicles,
187, 400; nasal bones of birds in
439
classification, 177 ;—malformation of
thorax, 170 ; sternum in tortoises, 177 ;
subdivision of scaphoid, 177 ;—reports
anat., 159—178, 386—394, phys.,
425—427
Ovum, 168, of trout, 170; serous epi-
thelium of, 170
Owen, R., anat. of American king crab
178
Owsjannikow, nerves of arteries, &c.,
183
12%
Pachydermata, anat. report, 176
Pancreas, see ‘* Digestive”’
Paquelin, J., comp. of blood-corpuscles,
404
Parker, W. K., morphology of skull,
62, 176
Paschutin, lymph in limbs of dog, 199;
digestive ferments, 409; butyric acid
fermentation, 409
Pathology, cerebral, 152, 179, 392; of
lungs, 185
Paton, G., action and sounds of heart,
402
Penicillium, sugar-forming substance,
82
Pepsin, digestive action of, on fibrin,
204; formation in stomach, 207, 414,
416
Peptones, 204
Perissodactyla, anat. report, 176
Perl, L., nutrition of cardiac muscles,
407
Petrowsky, composition of matter of
brain, 181; skin under slight mecha-
nical irritation, 186; develop. of mus-
cular fibre in frog, 42
Pettenkofer, feeding with flesh, &c.,
204, 207, 415
Pettigrew, circulation in plants, &c.,
18
Sears eee report on, 217—232
Phoea baicalensis, skull of, 176
Phosphorus, action of, 217
Physostigma, and atropia, 232, 403
Pick, E., innervation of blood-vessels,
185, 407
Pick, R., nitrite of amyl on muscle, 425
Pinnepedia, anat. report, 176
Placenta, see ‘‘Genito-urinary ”’
Plész, P., ferment of blood, 199; albu-
minous substances of hepatic cells,
1Q
Podolinski, expulsion of nitric oxide
from blood, 197
Polyodon, new species, 17
Porpoise, jaws, 176
Pozzi, S., lobus impar of right lung, 174
440
Prévost, lingual nerve, 185, 398; regene-
ration of nerves, 186
Pribram, stomach and circulation, 182
Pulse, secondary waves, I, 112; fre-
quency of, 54
Purgatives, 231
Pye-Smith, P. H., Catalogue of the
Museum of Guy's Hospital, 384
Q.
Quehl, emetic action of apomorphia,
223
Quinine, on blood, 198, 224, 403; on
reflex excitability, 230
Quinguand, variations of hemoglobin
in the zoological series, 196; respira-
tion of fishes, 203
R.
Rabatel, M., circulation in coronary
artery, 189
Rabbit, innervation of ear, 184; spinal
cord after excision of sciatic nerve,
186; retardation of pulse on closure
of nostrils, 193; circulation in pla-
centa, 393; septum atriorum, 340;
red and white muscles, 423
Rabow, alcohol on temp. and pulse, 427
Rabuteau, urea, 222, 421 ; calcium salts,
218
Radcliffe, C. B., synthesis of motion,
vital and physical, 300
Radziejewski, absorption of fat, 205;
action of purgatives, 231
Ransome, A., respiratory movements,
161
Ranvier, degeneration of nerves after
section, 186; histology and phys. of
nerves, 186; red and white muscles ~
of rabbit and ray, 423
Ray, red and white muscles of rabbit
and, 423
Reichert, C. B., early human decidua,
167
Reoch, J., acidity of gastric juice, 274
Reptilia, anat. report, 177
Respiratory system ;—lurgs, pathclogy
of, after section of both vagi, 185;
temp. of, 4275; lymphatics of, 390;
circulation and its disturbances in, of
frog, 188 ; injury to brain with he-
morrhage in, 397 ;—respiratory move-
ments in man, 161;—tracheal pouch of
Emeu ;—reports, anat., 174—175, 387
—388, phys., 203—204, 409
Reyher, C., cartilages and synovial
membranes of joints, 261
INDEX.
Rhinoceros, visceral anat. of Sumatran,
176
Rhus, venenata and toxicodendron,
action of, 22
Richardson, B. W., muscular irritability
after systemic death, 213
Rickets, 426
Riecker, A., skin of lower limb, 402
Riegel, F., respiratory movements, 409 ;
alcohol and temp., 427; regulation of
temp., 427
Rindfleisch, E., nerve-endings in cortex
of cerebrum, 171
Ritthausen, N., albumen, &e., 410
Robin, C., marrow of bones, 387, 425
Rochefontaine, parasiticide action of
quinia, 225
Rohrig, secretion of bile, 209; action
of sweat, 428
Rolleston, G., Harveian Oration, 1873,
158
Romiti, W., structure and develop. of
ovary and Wolffian duct, 393
Rosenthal, spinal cord, 182; galvanic
currents on muscles and nerves,” 186
Ross, alcohol.and absinth, 220
Rossbach, artificial resp. in strychnia
poisoning, 203; atropin and physo-
stigma, 403; hemorrhage after liga-
ture, 403
Roth, M., varicose hypertrophy of cere-
bral nerve-fibres, 173
Rouget, action-of silver-salts, 217
Riidinger, closure of the Eustachian
tube, 187; perception of high musical
notes, 187
Rumbold, functions of Eustachian tube,
187
Russell, J. A., communication between
umbilical and portal veins, 149; large
renal calculus, 382
Rustizky, J. V., absorption of bone and
on giant-cells, 387, 425
Rutherford, retardation of pulse on
closure of rabbit’s nostrils, 193
S.
Salamandra maculosa, skin of larva of,
--
77
Salkowski, E., withdrawal of alkali from
living body, 410; sulphuric acid and
taurin in the animal organism, 410;
synthesis-of taurocarbamin acid, 410
Salmonide, blood-corpuscles, 178
Salomon, G., glycogen in liver, 419°
Sanson, mechanical coefficient of food,
204
Saponin and digitalin, 231
Savory, W. S., ligamentum teres of hip,
2Q1
INDEX.
Schachowa, intercellular growth of bone,
425
Schiifer, bacteria in blood, 198; action
of arsenic, 218
Schech, laryngeal nerves and muscles,
203
Scherk, apparatus for measuring field of
vision, 400
Scherschewsky, innervation of uterus,
423
Schiff, electrical irritation of nerves, 186;
sensibility of heart lessened by atro-
pia, 226
Schlesinger, reflex movements of uterus,
214; uterine nerve-centres, 403
Schliephake, H., galvanic current on
eye, 400
Schmidt, A., preparation of albumen,
410
Schmuziger, F., migration of red and
white b. corpuscles in frog, 174
Schreiber, J., influence of brain on temp.
of body, 398
Schroetter, resp. movements, 203
Schultz, nerve-section on nutrition and
regeneration of tissues, 186
Schwalbe, lymph-channels of retina and
vitreous body, 174
Sée, phys. action of alcohol, 416
Seegen, nitrogen in albuminates, 204
Seeliy, L., consumption of sugar in dia-
betic and non-diabetic animals, 421
Shark, anat. of Greenland, 285
Siebert, emetic action of apomorphia
Silver-salts, 217
Simon, T., new formation of brain-sub-
stance, 173, 399 ; persistence of frontal
suture, 386
Sinéty, excision of mamme during lacta-
tion, 427
Skin, see ‘‘Integumentary”’
Skull, see ‘‘Osseous”
Slavjansky, K., ovum of rabbit, 170
Smee, A. H., coagulation of blood, 199
Smith, C. J. M., dissection of excised
elbow, 380
Sodium chloride, a poison, 219
Solucha, semicircular canals, 400
Sound, sensibility to, 187; perception of
high musical notes, 187
Sphygmoyraph, secondary waves of pulse
in tracings of, I, 112
Spina, A., structure of tendons, -162
Spleen, see ‘‘Circulatory”
Starkow, benzine, nitro-glycerine, nitric
and sulphuric acids, 222
Steinbe g, quantity of blood, 197
Steiner, bile in absorption of fat, 209;
colouring matter of bile, 209, 420;
diabetes, &c., 209
Stieda, L., nerve-coils at roots of hairs,
175
VOL. VIII.
441
Stirling, Dr, report on phys., 179—216,
395—431
are P. B., stimuli on parotid gland,
20
Strelzoff, feeding with madder, 425
Stricker, temp. of heart and lungs, 427
Struthers, J., subdivision of scaphoid,
113; rudimentary finger-muscles in
toothed whale, 114
Struve, H., colouring matters of blood,
198; action of zinc on blood-solution,
405
Strychnia, influence of on reflex excita-
bility, 229; on sensory nerves, 399;
influence of artificial respiration over
poisoning with, 409
Sugar, consumption of, in diabetic and
non-diabetic animals, 421
Sweat, see ‘‘ Integumentary ”
AM
Tanghinia venenifera, 97
Tappeiner, H., blood-stream after liga-
ture of vena porte, 189
Tarachanoff, innervation of spleen and
its relation to leucocythemia, 185 ;
temp. in the central ends of cardiac
nerves, 399
Taurin, 410
Teeth, see ‘‘ Digestive”
Temperature, influence of brain on, of
body, 398; changes of, on cardiac
nerves, 399; on sensibility of nerves,
186; preventing death from chloral,
3323 on circuation, 189; on reflex
actions in frog, 45; dyspncea, 203;
phys. report, 427—431
Tendons, structure of, 162; shortening
of, 214, 423
Tergast. S., proportion of nerve to mus-
cular fibres, 42
Thanhoffer, S., absorption of fat in small
intestines, 410
Thein, 225
Theobromin, 225
Thin, G., structure of tactile corpuscles,
30; lymphatics of cornea, 390
Thiry, purgatives, 231
Thoma, R., migration of white cor-
puscles into lymphatics, 403
Tievel, E., sugar-producing ferment of
blood, 199
Tissue, nietabolism of, 120; connective
t., 162; nutrition of, 186
Toad, Auerbach’s plexus in, 173
Tobacco-smoke, 226
Tomes, C. 8., enamel-organ in an arma-
dillo, 387
Torpedo, electrical plates of, 392
Tortoises, sternum in, 177
29
442,
Tourneux, F., epithelium of serous mem-
branes, 390
Pens L., rickets artificially produced,
42
Trissier, the vagus, 185
Tschiriew, nerve-influence on arteries,
&e., 183
Turner, Prof, bidental skull of Nar-
whal, 133; relations of convolutions
of brain, to outer surface of skull, 142,
359, to intelligence, 173; anat. of
Greenland Shark, 285; placentation
of Sloths, 362; enamel-organ in Nar-
whal, 387; var. in human nerves,
297; reports on anat., 159—178,
386—394
U.
Upupide, obs. on the, 177
Urbain, V., coagulation of albumen,
408
Urea, absorption of, 222; elimination of,
421
Urethra, see ‘‘ Genito-urinary ”
Urine, 205
Uterus, see ‘‘ Genito-urinary ”
V.
Variations ;—of cranium, 136, 159, 386;
thorax, 321; upper limbs, 113, 383;
lower limbs, 134, 358; teeth, 133,
140; muscles, 150, 377; Nerves, 297;
arteries, 155, 321; veins, 149; lung,
eculsr system, see ‘‘ Circulatory ”
Venous system, see ‘‘ Circulatory”
Veratria, 228
Vierordt, C., absorption spectrum of
hydrobilirubin, 209
Viscera, see “ Digestive”
“ Genito-urinary ”
Viverra civetta, visceral anat. of, 176
Volkers, accommodation movement of
choroid in eye, 400
Vogel, reaction of milk to litmus, 427
Voit, feeding with flesh, &c., 204, 207,
415
Vulpian, nerve-lesions on muscles, 186;
union of cut ends of nerves, 186,
400; chorda tympani, 187, 399;
splanchnic nerve, 187; effect of ex-
tirpation of supr. cervical ganglion
on movements of iris, 398
and also
INDEX.
Wi.
Wagener, G. R., striated muscle, 161
Walker, R., lower jaw of porpoise, 176
Waller, regeneration of nerves under
paraplegia, 186
Wartmann, L., amorphous aconitia, 223
Water, phys. of, 204
Watson, M., anat. of Indian elephant,
85
Wegner, phosphorus, 217
Weise, glycogen in liver, 209; contents
of liver-cells, 209
Weiske, H., comp. of bone, &e., poor,
425
Wernich, A., ergotin, 228, 229
West, 8. H., peculiar digastric muscles,
150
Westphalen, H., biliary fistula, 418
Whales, finger-muscles in, 114; dol-
’ phins and, of New Zealand Seas, 176
White, J. C., Rhus venenata and KR.
toxicodendron, 224
Wildt, E., comp. of bone, &e., 425
Williams, H. S., muscles of shoulder-
girdles, 161
Wilson, A., Student's Guide to Zoology,
384
Wilson, E., Anatomist’s Vade Mecum,
157
Wilson, H. S., rete mirabile of the
Narwhal, 176
Winiwarter, F. V., vessels in inflamma-
tion, 403
Winkler, F., human placenta, 162
Wittich, Von, sugar-producing ferment
of blood, 199; pepsin on fibrin, 204;
conversion of starch into sugar by
bile, 209
Wolithiigel, digestion of fibrin without
pepsin, 204
Wood, H. C., Atropia, 225; convul-
sants, 230
Woroschiloff, food, &c., 205
Ve
Yule, C. J. F., opening and closing of
Eustachian tube, 127, 400
Z.
Zaaijer, T., seaphocephalic cranium, 386
Zmitz, expulsion of nitric oxide from
blood, 197
Zuppinger, H., spinal ganglion-cells,
392
oo
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