Translations of foreign biological memoirs.

I. Memoirs

ON THE

PHYSIOLOGY OP NERVE, OP MUSCLE

AND OF THE

ELECTRICAL ORGAN

EDITED BY

J. BURDON-SANDERSON, M.D., F.R. SS. L. & E.

WAYNFLETE PEOFESSOE OF PHYSIOLOGY IN THE HNIYEESITY OF OXFOED

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AT THE CLARENDON PRESS 1887

[ All rights reserved ]

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'GRAG1

Preface

The following pages are intended to give the reader such preliminary hints as to the subjects of the Memoirs which are included in the present series of translations, as may aid him in using \he volume either for the purpose of extending his knowledge of the branch of physiology to which they relate, or of obtaining information on special questions. It will be seen by glancing at the Table of Contents, that the Memoirs refer to three principal subjects, and that with one exception, that of Prof, du Bois-Reymond’s research on Malapterurus, all have appeared within the last five years.

The purpose of the investigation recorde'd in the first Paper is to obtain a new proof of the well-known law established by Pfliiger in his classical research on Electrotonus, published in 1859, relating to the changes of excitability of a nerve p :oduced by the flow along it of a voltaic current. In 1881 the Author invented and described a new instrument, by which it is possible to subject a nerve to instantaneous mechanical excitation, of which the intensity can be measured and varied at will. When this is accompli shed by the falling on the nerve of a weighted lever, it is found that the nerve is so little damaged thereby, that the excitation may be repeated as often as necessary without impairment of the effect, and that its strength may be graduated by the observer with an exactitude scarcely inferior to that which is attainable in the use of induction currents. As a means of measuring the excitability of a nerve, mechanical excitation has this great advantage as compared with excitation by induction currents, that it is free from those ‘ after-effects ’ of which the nature is so fully studied in the series of researches contained in Part II

A

255219

IV

PREFACE.

of this volume; and secondly, that by the mechanical method it is possible to explore the condition of the nerve in the ‘ intra-polar ’ region, i. e. in that ’ part of its course which lies between the inflow and outflow of the current. It may be stated generally that the observations of Tigerstedt made by so different a method, while they extend those of Pfluger, afford additional evidence of their accuracy, the only difference between them being that the state of diminished excitability at the cathode after opening of the current is found not to be so marked when tested mechanically as when tested electrically.

The second paper contains the results of a new investigation of the £ Law of Contraction,’ with special reference to the conditions which determine the state of excitation that occurs at the anode at the moment that a current led through a motor nerve is broken. The author shows that these conditions correspond with those which produce ordinary negative polarisation in the nerve, that is, a state of the nerve in which, if the seat of outflow of the current be investigated immediately after the circuit is broken, it is found to be negative to all other parts ; and he thinks that this correspondence is so close, that the two concomitant phenomena, viz. the contraction of the muscle and the polarisation of its nerve, must have the same cause. He is thus led to regard the make and break contractions as phenomena which are alike produced by the closing of a current, but which differ from each other as regards the way in which this current itself originates. This research should be read along with that of Griitzner (No. 4) on the same subject, who by different experimental methods has arrived at the same explanation of the anodic contraction. He finds that by far the greater number of the known instances in which the phenomenon can be experimentally investigated admit of more complete and satisfactory explanation as resulting from negative polarisation than on Pfluger’ s theory. The special cases which he has investigated are — (1) that in which the anodic excitation produced by breaking a current which is led through a nerve in a direction opposed to the nerve-current, is found to be dependent on certain complicated but

PREFACE.

V well-ascertained relations between the strength of the former and that of the latter (see Introduction to Griitzner’s paper); and (2) the numerous cases in which excitation at the anode results, not from break of the current, but from sudden diminution of its strength. In these the varieties of effect which are observed are, in Griitzner’s opinion, such as point to polarisation, not subsidence of anelectrotonus. How far this way of looking at the subject is justified, the reader will be able to judge after studying the criticisms of Prof. Hermann and Prof, du Bois- Beymond.

In No. 5, Prof. Hering has investigated the excitation of motor nerves by voltaic currents from an entirely different point of view. More than thirty years ago du Bois-Beyinond showed by a remarkable experiment (see p. 138), in which he used no apparatus excepting ‘physiological clay,’ that is, clay kneaded with ‘normal’ salt solution instead of water, that if the cut and natural surfaces of a nerve-muscle preparation are bridged, the muscle contracts when the bridge is completed, and again when it is broken. Starting from this observation, the author proceeds to illustrate, by a variety of new modes of experiment, the exciting influence exercised on nerve by its own current. He shows, for example, that the mere falling of the end of a nerve on a moist conductor (p. 130) produces contraction when the contact is of such a nature as to bridge the two surfaces, that by this means, if the contact is repeatedly made and broken, a tetanus ‘without metals ’ may be produced, and that a real ‘ secondary twitch from nerve ’ is obtainable when one of two nerves is excited, of which the two ends are applied to each other with their cut surfaces in the same plane and looking in the same direction. When this is the case, the muscle to which the unexcited nerve belongs responds to excitations of a distant part of the other by currents of minimal intensity. In this respect the phenomenon may be contrasted with du Bois’ ‘paradoxical twitch,’ which, as is well known, only appears when the exciting currents are strong, and led in in pretty close proximity to the contact between the two nerves. He finally shows how the effects of opening and

VL

PREFACE.

closing of external currents are modified by the excitatory influence of the nerve-current, and how impossible it is to judge of or interpret these effects correctly without taking this ‘interference 5 into account. It is to be noted that Prof. Hering is led by his observations to a conclusion as to the nature of the opening contraction, which, although by no means identical with that arrived at by Tigerstedt and Griitzner, is in intelligible relation with it.

The second Part begins with three independent investigations of the same subject by three of the most distinguished living physiologists — the first by Prof, du Bois-Reymond, of Berlin; the second by Prof Hering, of Prague ; the other by Prof. Hermann, of Zurich, now of Konigsberg — all of which were published within a few months of each other. The purpose of du Bois’ paper is to give a more full account, based on new experimental investigations, of the phenomena which the author discovered many years before and designated by the term ‘positive polarisation.’ In muscle and nerve there are, according to him, two kinds of polarisation, of which one is identical with physical polarisation of porous bodies steeped with electrolytes, the other is purely physiological, that is, peculiar to living excitable structures. As it manifests itself in muscle and nerve, positive polarisation consists in this, that when either of these structures is traversed axially by a voltaic current of very short duration, the galvanoscopical after-effect observed on opening, instead of being, as in ordinary polarisation, opposed in direction to the led-in current, is in the same direction with it, varying according to the density of the external current and other well-defined conditions which need not here be referred to, but exhibiting throughout the most intimate association with the functional activity of the organ.

A few months later in the year in which du Bois presented his Memoir to the Berlin Academy, a communication on the same subject was submitted to the Academy of Vienna by Hering, in which he described the same phenomena under another designation, the main fact being that when a current

PREFACE.

Vll of considerable intensity and short duration is led through a muscle, and the condition of the muscle is immediately after investigated galvanoscopically, it is found that a sudden change as regards its electrical condition takes place, of such a kind that the current led off from the organ, at first feebly ‘negative/ that is, in the opposite direction to that of the external current, becomes strongly ‘ positive/ that is, in the same direction.

Similarly, Prof. Hermann, in a Paper published in Pfluger’s Archiv in January 1883, gave an account of the same phenomena, which, as regards muscle, closely accorded with that of Hering, adding new proof of the polar localisation of the electromotive changes observed ; ’ and he further extended his researches from muscle to nerve so as to include all the subjects investigated by du Bois, with the exception of that of the electrical organs of fish. Prof. Hering’s paper (‘Ueber Yeranderungen des electromotorischen Yerhalten der Muskeln in Folge electrischer Reizung’) we have not thought it necessary to translate, preferring to present to the reader the more advanced view of the subject contained in his second paper, which appeared at the same time with it, under the title, ‘Ueber du Bois-Reymond’s Untersuchungen der secundarelectromotorischen Erscheinungen am Muskel.5 The reader will find in the three papers satisfactory evidence that all these distinguished observers have had before them the same facts, though from difference of method, or from other causes, they may have been presented to them under different aspects. But in all that belongs to the doctrine of ‘ secondary electromotive phenomena ’ it will be equally clear to him, that while Hering and Hermann tend in the same direction, their views are utterly at variance with those of the great founder of Electro-physiology.

Du Bois holds that just as in the ordinary polarisation of porous bodies soaked with electrolytic liquids, the change affects all parts through which branches of current pass, so in polarisation of nerve and muscle, whether negative or positive, the change affects all the parts traversed in a degree which depends upon the current density, and that as regards positive polarisa-

Vlll

PREFACE.

tion, the action of the current is directly exerted on certain hypothetical carriers (Trager) of electromotive forces which are distributed throughout the polarisable structure, and which the current is capable of £ directing ’ (see concluding paragraph of No. 6).

Hering rejects this theory on the ground that the changes of which ‘ positive polarisation ’ is the expression, have their seat exclusively at the anode, and explains the experimental facts used by du Bois in proof of ‘internal polarisation 5 (that is of a polarisation affecting all traversed structures, and consisting in the coming into existence of electromotive forces all along the line of flow, and not merely at anodes or cathodes), on the supposition that he employs structures in which, in consequence of their heterogeneity, such sudden transitions from less to greater density occur in the course of the current-paths, that the effects of anodes are produced. On the assumption that positive polarisation has its seat at the anode exclusively, he has no difficulty in referring what is observed to the well-recognised facts — (i) that the effect of opening a current led through a muscle or nerve is to produce excitation at the anode ; and (2) that the characteristic expression of the excitatory state is negativity; and accordingly, while recognising the essential relation of the change in question to the functional activity of the organ in which it manifests itself, he prefers to speak of it as £ anodic after-effect’ rather than as £ positive polarisation.’

Hermann objects to the term ‘positive polarisation’ even more decidedly than Hering, on the ground that the phenomenon is not analogous to physical polarisation. He, like Hering, believes it to be excitatory, and holds that it is due to the setting up of a persistent change of this nature in the neighbourhood of the spot at which the current has entered. Among the various experiments which lead to this conclusion, those are of most importance which show (1) that in muscle it is confined to the anodic region ; (2) that it is annulled whenever the current is led into the muscle or nerve through a dead part ; and (3) that the anode becomes relatively negative not only

PREFACE.

IX to the £ intra-polar ’ region, but also to extra-polar parts in its neighbourhood. Hermann is accordingly led to regard all secondary electromotive phenomena as resulting from the interference of the excitatory change which occurs at the anode in consequence of the opening of the current, with ordinary galvanic polarisation, it being understood that the physical change always anticipates the vital one as regards the order of its occurrence ; so that whenever the conditions (strong current and short closure) are such as to enable these two antagonistic influences to manifest themselves together, the process consists of a more transitory physical effect, which is followed by a more lasting physiological one.

If the views of Hermann and Hering are accepted, the reader will see that the form given by Tigerstedt and Griitzner to their theory as to the essential nature of the opening contraction must be modified. Tigerstedt assumes that at the break of a current, the anode becomes, in consequence of negative polarisation, virtually a cathode, an assumption which takes no account of the facts of positive polarisation. On this point the reader will compare with advantage du Bois’ criticism of Tigerstedt in his sixteenth paragraph, with that of Hermann in the paragraph which precedes his conclusion.

If it is true that £ positive polarisation 5 of a nerve or of a muscle is in reality a purely physiological, or, in other words, an excitatory effect of the leading through it of a voltaic current, it becomes of interest to enquire what is the relation of the electrical after-effects so fully studied by du Bois-Reymond, Hering, and Hermann, to the more mechanical effects which form the subject of the first part of this series. In general it is, as every physiologist knows, extremely difficult to demonstrate in detail the correlation which undoubtedly exists between the electrotonus of Pfluger, and the galvanoscopic phenomena which were originally investigated by du Bois and treated of by him under the same title. The chief reason of the difficulty lies in the circumstance that until recently it has been possible to investigate the former only in muscle, the latter only in nerve. But

X

PKEFACE.

from the moment that it was shown by Hermann and Hering’s researches, that the well-known electromotive after-effects of the current in nerve (du Bois’ electrotonus) have their counterparts in muscle, that is, that the electrical state of a muscle which is traversed axially by a current is subject to polar variations which correspond to an- and cathelectrotonus, it at once became of interest to enquire how these variations are linked to concomitant changes in what must always be regarded as the essential function of muscle — that of contracting in response to stimulation.

In 1 88 1 it was discovered by Engelmann that the anodic and cathodic localisation of the closing and opening excitatory effects could be demonstrated in the heart of the frog, and Hering showed how, with the aid of the double myograph, as much could be done with a striated muscle. Electrotonus of muscle was thus in a certain sense demonstrated, but to complete the proof it was necessary to show that as cathelectrotonus is represented by contraction, so anelectrotonus is represented by relaxation. This was accomplished by Prof. Biedermann, Hering’s coadjutor at Prague, in 18831. It was necessary for the purpose to employ as subject of experiment a muscular structure capable of persistent tonus, for it was only in such a structure that relaxation could be observed. The required condition was found to be fulfilled by the heart of the snail (Helix pomatia), which, as Biedermann discovered, can generally be thrown into spasm of sufficient duration for the purpose, by the simple expedient of allowing the ventricle to contract against unwonted resistance. By the observation that the closure of a battery-current through the heart leads to relaxation which takes its start from the anode, just as in the relaxed heart, the closing contraction takes its start from the cathode, Biedermann was able to prove that in that structure, relaxation of muscle already contracted is as characteristic of anelectrotonus, as contraction of relaxed muscle is of cathelectrotonus.

1 W. Biedermann, Ueber das Herz von Helix pomatia. (Ein Beitrag zur vergleichenden Physiologie der Muskeln.) Wiener Sitzungsberichte, vol. lxxxix. 19-55.

PREFACE.

XI

In the paper of which a translation is introduced into our series of Memoirs, Biedermann extended the same mode of experiment to a striated muscle of the frog. In order to bring about a similar condition of permanent spasm to that which, in the heart of Helix, is produced so easily by mere distension, he availed himself of the action of veratrine, and found that in this, as in the other case, the cathodic contraction is replaced by anodic relaxation, and the anodic contraction by cathodic relaxation. Here, then, an opportunity seemed to be afforded for the first time, of determining experimentally whether electrical and functional electrotonus occur under the same conditions — whether, as the opening contraction corresponds with the anodic after-effect (of which, according to Hering, positive polarisation is but the expression), the cathodic break-relaxation is marked by a cathodic after-effect, which, although of opposite sign, i.e. positive, as regards the condition of the cathode itself, is of the same sign galvanoscopically. To ascertain whether it was so or not, Biedermann prepared the sartorius by subjecting one end of it to the action of veratrine, leaving the other intact, and then induced cathelectrotonus in the poisoned, that is, tonic end, leading a battery-current through the whole riband for a couple of seconds. The poisoned end was at once thrown into veratrine spasm and the cathode became negative ; but when (after compensating) the current was again closed, its after-effect was in the opposite direction, that is, in the direction of the exciting current. In other words, relaxation of the veratrine spasm was, as was expected, accompanied by diminution of the negativity which is its galvanoscopic expression. When the same observation was made with the modifying current in the opposite direction, that is, with the anode at the veratrine end, it was similarly found that a current led through during spasm was reversed, that is, that the anode became positive.

The general conclusion derived by Biedermann from these experiments is that the state of muscle varies, not merely between the state of excitation and the normal, or unexcited

Xll

PREFACE.

state (the former being indicated galvanoscopically by the presence in the tissue of a current from the excited to the unexcited part, and mechanically by the presence of more or less spasm, the latter by the opposite effects), but that muscle varies in the direction of greater or in that of less fitness for functional activity. By employing such terms as ‘up’ or ‘down’ to denote these two tendencies, it is possible to express his conception thus : — A voltaic current produces at the anode when it is closed an ‘ up ’ state, of which the galvanoscopic sign would be relatively positive, and the mechanical sign axial extension but aptness to contract ; at the cathode a ‘ down ’ state, associated conversely with ‘ negativity,’ and axial shortening. The assumption of a state intermediate between these two is suggested by the indications of what, for want of a better name, is called by Biedermann ‘ reaction,’ that is, of a tendency (of which the behaviour of all excitable tissues suggests the existence) to assume a condition opposite to that in which it actually is, by virtue of which the ‘ down ’ state is produced at the anode at make, the ‘ up ’ state by the cathode at break. In Biedermann’s experiments it was strikingly seen that the same external cause produces contraction and negativity when acting on muscle in the ‘ up ’ state, relaxation and positivity when acting on the same muscle in the ‘ down ’ state. This being so, there may be between contraction and complete relaxation an intermediate state in which the two opposite tendencies neutralise one another. If such a state exists, it does not follow that it is the normal state of the tissue, for by the word normal the physiologist understands simply the state which best fits the structure for its function, and cannot be guided in his use of it by the correlation of its elementary properties.

Part III contains Prof, du Bois-Reymond’s investigations in a field of research in which up to the present moment he has been almost the only worker — that of the electrical phenomena of fish which are possessed of special electrical apparatus. The Memoir on Malapterurus, which appeared a few years ago in the Author’s

PREFACE.

Xlll

Collected Papers, has been translated at his suggestion. It contains a full description of the methods by which the difficulties of the investigation were in the first instance overcome, and the reader will find it to be an indispensable introduction to the other two Memoirs on Torpedo, which contain his more recent observations. As each of the three memoirs is complete in itself, and is preceded by a summary of the subjects discussed in each paragraph, no introduction is needed. It may not, however, be out of place to draw attention to the very remarkable discoveries recorded in the concluding paper relating to the fact of ‘ irreciprocal conduction,’ which the author has proved to exist in a remarkable degree in the columns of the electrical organs of Torpedo.

The Editor regrets that the publication of the volume has been unduly delayed, in consequence of the unexpected difficulties experienced by the translators in their work. The subject is so new to English scientific literature, that much time is unavoidably lost in deciding between the two alternatives, of using Germanised phraseology and of coining English equivalents for the forms of expression used by German writers. In general, the latter alternative has been preferred, but not without a heavy sense of responsibility. Whether successfully or not, great pains have been taken to select such equivalents as appear most accurately to express the meaning of the original, and the translators have, as a rule, consented to employ the same equivalents for the same words.

PART I. Researches relating to the Law of Contraction

Author. 1. Tigerstedt

4. Grutzner .

5. Hering

Title.

Translator.

Page.

Changes of excitability produced in nerves by a constant current

Dr. James Niven .

Contribution to the theory of the Break Contraction . . .

yy yy

The Nature of the Electrical Break-contraction ....

Mr. J. S. Haldane .

• 93

Dr. Aug. Waller .

. 127

PART II. Researches relating to Secondary Electromotive Phenomena

6. Du Bois-Rey- On Secondary Electromotive

mond. Phenomena in Muscles,

Nerves, and Electrical Organs. Mrs. Lauder Brunton . 163

7. Hering . . On Du Bois-Reymond’s Re-

searches on Secondary Electromotive Phenomena of Muscle. Dr. Aug. Waller . . 229

8. „ On the Positive After-variation

of the Nerve-current consequent on Electrical Stimulation „ „ 255

9. Hermann. . On so-called Secondary Electro-

motive Phenomena in Muscle

and Nerve Mr. Francis Gotch . . 277

XVI

Contents

Title. Translator. Page.

On the Phenomena of Inhibition produced by the Electrical Excitation of Striated Muscles, and on Positive Cathodic

Polarisation Miss Edith Prance . 331

PART III. Researches on the Electrical Phenomena of Malapterurus and Torpedo

11. Du Bois-Rey- Observations and Experiments

mond. on Malapterurus Miss Edith Prance . 369

12. „ Living Torpedos at Berlin. First

Communication Editor . . . . . . 417

13. „ Living Torpedos at Berlin. Se-

cond Communication . . . Miss Edith Prance . 479

Author.

10. Biedermann .

CORRIGENDA.

P. 6, note 2, for fortschritte read Fortschritte.

P. 7, note 4 ,for nouvelles read nouvelle.

P. 23, § 2, for irritability read excitability.

P. 56, line 4 from top, for Marcy read Marey.

P. 56, Exp. II, beading, for .... read the break-contraction. P. 57, Exp. Ill, heading, for . . . .read the break-contraction. P. 57, Exp. IV, heading, for .... read the break-contraction. P. 94, line 13 from bottom, for break read make.

P. 123, line 18, for Thermopyle read Thermopile.

P. 166, line 15 ,for through read in.

P. 168, line 1 9, for polarisation-current read effects.

P. 187, line 13 from bottom, for would read should.

P. 203, line 19 from bottom, for Sect. 2 read p. 163.

P. 204, line 13 from bottom, for that read of.

P. 204, line 11 from bottom, for Sect. 9 read Sect. 10.

P. 249, line 4, for Sect. 9 read Sect. 8.

P. 280, line 2 from bottom, for Sect. 1 read Sect. 21.

P. 297, line 11, for or read in the.

P. 343, note, for No. V read No. VI.

P. 47.1, line 3, for positive read negative.

P. 490, line 6 from bottom, for No. V read No. VI.

Part I. RESEARCHES

RELATING TO THE

“LAW OF CONTRACTION.”

t£

B

CHANGES OF EXCITABILITY

PRODUCED IN

NERVES BY A CONSTANT CURRENT.

By Db. ROBERT TIGERSTEDT.

Communicated to the Swedish Royal Academy of Sciences, April 12, 1882.

I.

In his famous investigations into the physiology of electrotonus, Pfliiger, in the year 1859, submitted the changes of excitability in nerves elicited by a constant current of electricity to an experimental examination which will stand to all time as a model for similar investigations, as regards both the number and variety of the methods of research employed and the completeness of the experimental criticism. As test-stimulus he employed partly the constant current, partly the induction-shock, and in part also he used chemical stimuli. The most important actual relations established by this investigation are, very briefly, as follows. In the region of the positive pole of the current the excitability of the nerve is diminished, as well in the extrapolar as in the intrapolar direction ; and the diminution attains its definite amount gradually, being not so great for some moments after the current is closed as it afterwards becomes. When the current is opened there occurs immediately an increased excitability, which may last for a longer or shorter period. In the region of the negative pole of the current the excitability of the nerve is increased as well in the extrapolar as in the intrapolar direction. This increase occurs directly after contact is made. When contact is broken, diminished excitability is exhibited if the current is weak or moderate, which, however, after an exceedingly brief interval gives place to a condition of increased excitability lasting a longer or shorter time.

It is at the poles that the change in excitability is greatest, diminishing as we leave these both in the extrapolar and intrapolar directions. The greater the strength of the polarising current, the greater is the change of excitability and its extrapolar extension. Between the poles the diminution of excitability extends as the strength of the polarising current increases ; or, to put it otherwise, with increasing strength of current, the indifference-point moves gradually from the positive to the negative pole.

The greater the separation of the electrodes the greater is the tract of nerve over which the extrapolar change of excitability extends. The behaviour of the intrapolar tract is absolutely the same whatever the distance between the electrodes.

To test the local changes of excitability in the intrapolar tract Pfliiger used chemical stimuli ; the induction-coil he employed only to investigate the total excitability. He abstained from examining the changes of excitability with sharply localised electrical stimuli fearing an escape of electricity between the constant and test currents.

An important, though perhaps not unexpected, addition was made to these results two years after their publication by Obernier in a research carried on under Pfliiger’ s guidance. Obernier proved by experiments that, with the application of strong polarising currents (6-8 Groves), there occurs at the negative pole in the first moments after contact is broken a very considerable diminution of excitability1.

Before this, however, attacks had been made on the correctness of Pfliiger’ s results — attacks which have since been from time to time renewed by various enquirers. The most notable fact, however, is, that all those who have thus assailed Pfliiger, with few exceptions, have conducted their counter-researches with nothing like the same care as Pfliiger employed. What Meissner says of one of the first of them may be applied with justice to them all. £ Inasmuch as Pfliiger has grounded all his enunciations on investigations made with every kind of efficient stimulus, one has a right to demand from a criticism which is directed to prove these enunciations in part untenable, that it be pursued on a like basis and with a similar use of every species of stimulus2.

To this category of wrorks we must refer those of Schiff3, of

1 Obernier, Archiv fur Anatomie und Physiologie, 1861, pp. 268-278.