ms Spares ATeaetcti tet ey oe se 1H iphoto es itis take sires DAT ed aa Subsets sabe . fad & 9 harp a epely fas He . yy f he ahas ra ve an ii 4 r i ee “sree: a $i; if prt ;. : > ise Obs ui + var 4 4 re Ake Ly ‘ chi nts i eeth “4 pee uieten? 3 HEA tis Wi “USAT S78 debt Mee he : a} feet rei ha tee rh I nate aber ae cee je Babb ig hte : ot. ‘ vad anes o} iM 1 teu Aastt TI4E OT 404) pty Be ‘ “ ltr ¥ * Hy , ae bel ‘ bas ett Tai eal hk a: Legastt en by 1 4 4 x ot ‘ ; Seah Yi ‘ meats ace : + intel As . p, SS me pela ms es Pirishys inst eaese ; Mdedets Raval hte sti Bt vue ‘ Ny ae yal aight! 4 Mov #8 a4) Hab ot) A Bday ad ia ; ti rant ree $09 elgg dant byes ae ape easels 7 ? i oe Made parte tity ( +s AR iy AAD eriey 4 eA, HAF enh } a hed odode is far Wiese tin enon brett ge § {yeni Hie AS mb pare f ans 30 Ghee aaa bade ' by sieesses 3 ut ip vie ‘ 4 > ging i: i in bauer aide San tte ti dhey ye zi Reus opener ¥ Can aot Prat re sida (Al maak Spit ead vivrates pi Gholbestsranny Httelig uss iyrnipare ti HEAR ryt he i Ry wel, nde aie ettporst roy Hh Neagle r : Bea) rt Wes uy {tare " arent res ay on rata aea b iz Wey PR ¥ fe Peder Rhee s (uyy LPL bh ehinery arn i) bbs yeartalndy dye , : ‘ine Hove rt / Seen Ryan uta rate Haiidies Ossie rede redte sah : A ad ote 4 4G My Pra yaa othe ECU A™ iva te die HOES Ridiet ey iy Pishin wg srtet abe Hi Gepdieing 54> ‘ reo ane By pets pleted may iy, H ‘ f bere Ss, ey bs ’ hie, est era a tide att af Hit $ M a pital: 18 ate Ruler ee t nie ie lhe ; “ ree area ay oes ff his Aah f ; 3 Pah eaten i: ; ids Hine his WG Weta ; Tisarecaasen via | 1 HO Hs aie bord ' 1g + 4 pi oy Taga ' incl tthe grt el < : SM piad Wied Woredbey - Capteratt porary, Si rtag abe Ms syhen ce Weed . smiyte 44 Y EY ha) olhe ¥ is anit et 7 m9 ” - 4) Bie mike Ms ete ni bt Hy tA fie : , f weasaatess eye? fa 1g mn aay sets Wad ed b 4 ; 0 tara ae 725 yaa Ry hs ‘ ugh env) sk ei waster tates aveceie i pea toa eah aT See Hi oh ‘4: osc ag bathe ni ea “4 Mi tee! Ay batt Ms my iheayi ir 8 dain " iets ratty i os ¥ Sanit a) tf Ey t ( Liteea bh! ne 4 Peon Peay i rrreren) a : Ys 3 wyey Lj i} iy i 4 r ye be} hea) Pat Ne iy eh Fi pepe vu malian sa , pitts * iy et me ws gran enaet lined taf analy 17 we Me jue malas ay * he AA vad res ft r ye Logs Tet h z nae Poe ne traage is + Sengelo am ay via dde ‘ mihi O Minty tet el og sae Cares * ves eyes 4 4 ara “ " ohlied 4 onde ih A: Bia Hitt 4 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.