LECTURE ON THE PHYSIOLOGY OF THE HEART, And its Connections with the Brain, DELIVERED AT A MEETING AT THE SORBONNE, THE 27tii MARCH, 1865, BY CLAIJDE BERNARD, Member of the Institute of France, of the Imperial Academy of Med- icine at Paris, of the Royal Society of London; Professor of the College of France and the Sorbonne. TRANSLATED BY J. S. MOREL, M. D., SAVANNAH, GA. SAVANNAH: PURSE & SON. 1867. TABLE OF FIGURES. FIGURE 1.—Longitudinal section of the human heart, showing its four cavities, ° 12 FIGURE 2.—Diagram intended to illustrate the mechanism of the circulation, 14 FIGURE 3.—Longitudinal section of the human heart, to show the mechanism of the circulation in its four cavities, . . 17 FIGURE 4.—Cardiographic or Sphygmographic Apparatus of Mr. Marey, for directly inscribing the pulsations of the heart, . 20 FIGURE 5.—Tracing of the pulsations of the heart of a young man in the normal state, . 22 FIGURE G.—Tracing of the pulsations of the heart of a rabbit, . 22 FIGLTRE 7.—Tracing of the pulsations of the heart of a frog, . 23 FIGURE 8.—Tracing of the pulsations of the heart of an eel, . 23 FIGURE 9.—Tracing of the pulsations of the heart of a tortoise, 24 FIGURE 10.—Tracing of the pulsations of the heart of a crab, . 24 FIGURE 11.—Area vasculosa, or germinative field, representing the primitive circulation of the chick in the egg, . . . . 2G FIGURE 12,—Successive periods in the development of the heart of the chicken, 28 FIGURE 13.—The heart with the nervous branches which it re- ceives from the brain, 31 FIGURE 14.—Tracing representing the beating of the heart of the horse, to show the paralysing influence which irritation of the pneumogastric nerve exercises over the contractions of the heart, 33 FIGURE 15.—Tracing representing the contractions of the muscles of the foot of a frog, to show that the irritation of the nerves provoke these contractions, 33 FIGURE 16.—Anatomical connections of the heart to the brain, by the carotid arteries and the pneumogastric nerves, to explain the reactions of these two organs on each other, . . .44 ERRATA. Page 15, 20th line—for realised, read realized. Page 28, 10th line—for bulba, read bulb. Page 32, 23d line—for entrance towards, read commencement of. Page 33, 3d line in the description of Fig. 14—semi-colon (;) after pneumogastric. Page 34, lGtli line—semi-colon (;) after arrested. Page 45, 6th line—comma (,) after souvenir. In the description of the 2d Plate, higher should read lighter. On the Physiology of the Heart ami its Con- nections with the Brain. Gentlemen : I intend to entertain you in a general manner with the physiology of the heart, hut will dwell more particu- larly upon those points which sefcm necessary to clear up the physiology of the heart of man. To the physiologist the heart is the central organ of the circulation of the blood, and in this respect it is an organ essential to life. But by a singular privilege, en- joyed by no other organic apparatus, the word heart has passed, like the ideas one has of its functions, with very different acceptations, from the language of the physi- ologist into the language of the poet, the romancer and the man of the world. The heart may not only be a vital motor apparatus, pumping the sanguineous liquid into every part of our body, which it animates, it may also be the seat and symbol of the noblest and ten- derest sentiments of our soul. The study of the human heart may be pursued not only by the physiologist, it may also serve as the basis of the conceptions of the phi- losopher and the inspirations of the poet and the artist. The question now before us concerns the anatomical heart—that is to saj, the heart studied from a physio- logical, scientific, and purely experimental point of view. 10 ON THE MY3IOLOGY OF THE IIEAlit But will this rapid examination which we are about to make of the functions of the heart overthrow the ideas generally entertained respecting it ? Ought physiology to dissipate the illusions, and show us that the senti- mental role at all times attributed to the heart, is but a fiction purely arbitrary ? In a word, shall we be obliged to acknowledge a complete and peremptory con- tradiction between science and art, between sentiment and reason ? As to myself, I do not believe in the pos- sibility of that contradiction. Truth cannot differ from itself, and the truth of the man of science will not be contradicted by the truth of the artist. I believe, on the contrary, that the science which Hows from a pure source will become luminous to all, and that science and art should give their aid in explaining and interpreting for each other. I think, indeed, that in its elevated regions, human knowledge forms an atmosphere com- mon to all cultivated intelligences, in which the man of the world, the artist and the savant, will, as a matter of course, meet and fully comprehend each other. In that which follows, I will not attempt to deny, in the name of science, all that may be said, in the name of art, upon the heart considered as an organ intended to express our affections and sentiments. On the con- trary, I would like to be able to confirm art by science, in trying to explain, by means of physiology, that which, until now, has been but a simple intuition of the mind. I know that I have undertaken a very difficult task, it may be one of temerity, because of the slight advance that is yet made in the science of the pheno- mena of life. However, the beauty of the subject, and the light that physiology already seems to shed upon it, determine and encourage me to the investigation. Be- AND ITS CONNECTIONS WITH THE BRAIN. 11 sides, in speaking here upon the physiology of the heart, it will not be necessary to go into the details of a com- plete experimental and analytical investigation ; at this time, this would be impossible—it is only a simple test that I wish to make, and it will suffice me in express- ing my physiological ideas to support them by the clearest and best ascertained facts in science. The heart is truly a living motor machine, a forcing pump, intended to distribute to all of the organs of our body the nutritious fluid, the excitor of their functions. This mechanical role characterises the heart in an absolute manner, and wherever the heart exists, however simple it may be, or whatever complication it may present in the animal series, it constantly and ne- cessarily performs this function of organic irrigator. The anatomist regards the human heart as a viscus, that is, as one of the organs which forms a part of the apparatus of nutrition, situated in the splanchnic cavities. Everyone knows that the heart is placed in the tho- rax between the lungs—that it has the form of a cone, whose base is fixed by large blood vessels; whose point is free, inclining downwards to the left, in such a way as to place it between the fifth and sixth ribs below the left nipple. As to the nature of its tissue, the heart belongs to the muscular system; it is divided into cavities, which arc reservoirs for the blood—this is the reason why anatomists still' call it a hollow muscle. In the human heart are four compartments or cavities; two form the superior part, or the base of the heart, and are called the auricles—they receive the blood from all parts of the body, by means of large tubes, called veins; the other cavities form the lower portion, or apex of the heart; they are the ventricles, and are intended to force 12 ON T11E PHYSIOLOGY OF THE HEART the blood into all parts of the body, by means of large vessels, called arteries (Fig. 1). Fig. i.—Longitudinal section of the heart of man, showing its four cavities. od.—Right auricle, vd.—Right ventricle, or/.—Left auricle. tvg.—Left ven- tricle. The arrows placed at the entrance of the different vessels ter* ininating at the cavities of the heart, indicate the direction of the current of the blood in these vessels. Each auricle communicates with the ventricle which is below it on the same side, but a longitudinal septum separates laterally the auricles and ventricles in such a way, that the human heart, which is really double, is divided into two simple hearts, each formed of one auri- cle and one ventricle, situated, one on the right and the other on the left of the middle septum. Each ventricu- lar cavity is furnished with two valves. The one placed AND ITS CONNECTIONS WITH THE BE AIN. 13 where the blood passes from the auricle to the ventricle, is called the ouriculo ventricular valve ; the other, situated where the blood leaves the ventricle by the artery, is called the sigmoid valve. The human heart, as Avell as that of mammals and birds, is anatomically, double, and composed of two sin- gle hearts, called the right heart and the left heart. Each of these hearts play a very different part in the economy (Fig. 2). The left heart, called the heart of red blood, is destined to receive into its auricle, by the pulmonary veins, the pure and lively red blood which comes from the lungs; this blood passes into its ventricle, which distributes it by the arteries to every part of the body, where it becomes impure and black. The right heart, called the heart of black blood, receives into its auricle, by the vena cava, the impure blood which comes from every part of the body, and passes it afterwards into its ventricle, to be distributed to the lungs, where it becomes pure and of a lively red color. In a word, the left heart presides over the distribution of the vital liquid to all of our organs, and to all of our tissues; and the right heart presides over the revivification of the blood in the lungs, to restore it to the left heart. These premises being established, we shall consider the heart as an organ distributing life to all parts of our body, sending to them the nutritious liquid which is indispensible to their life, and the manifestation of their functions. As to the nutritious liquid itself, it is repre- sented by the blood, which is identical in all vertebrated animals, whatever may be their diversity in the animal species, and the variety of their food. In the external phenomenon of the mode of feeding, the zoologist distinguishes the ferocious carnivora, which 14 ON THE PHYSIOLOGY OF TIIE HEART Fig. 2—Intended to explain the mechanism of the circulation o.—Left auricle, vg.—Left ventricle, o.—right auricle, vd.—Right ven- tricle. The circulating system of red blood consists of arteries, over which the left heart presides—oog. is represented by the higher portion of the figure. The circulating system of dark blood consists of the veins over wldch the right heart presides—ood. is represented by the greyish tint. The arrows indicate the direction of the current of the different vessels. Above fcnd below are seen the capillary network of the supe- rior and inferior portions of the body, where the arterial blood is trans- formed into venous blood, and in the middle, the capillary network of the lungs, where the venous blood is revivified, to become arterial. And its connections with the brain. 15 feed upon palpitating flesh, the peaceful ruminant, which lives upon the herbs of the field, the frugivora and gramenivora, which are nourished more particularly by fruits and grain ; but when we descend into the inti- mate phenomenon of nutrition, general physiology teaches us that that which, properly speaking, is nour- ished in animals, is not the specific and individual type, which varies to infinity, but the elementary organs and tissues only, which are everywhere destroyed and live in the same manner. According to Goethe, nature is a great artist. Animals are constituted of the same or- ganic materials; it is the arrangement and relative dis- position of these materials which determine the variety of forms and the specific animal properties of these truly organised monuments. In like manner in the monuments of men, the materials are alike in all of their physical properties, and however different their arrange- ment, the various ideas may be realized, and give rise either to a palace or a cottage. In one word, the spe- cific type exists, but only in the state of an idea realised. Physiologically, it is not the animal type which lives and dies, but the organic materials or tissues which compose it; in the same way with the edifice which de- cays, it is not the ideal type of the monument which deteriorates, but only the stones of which it is built. We cannot, then, deduce from general physiology any essential difference in the organic nutrition, from the great variety of the food of animals. In man and all other animals, the elementary organs and living tissues are sanguinary; that is to say, they feed upon the blood in which they are immersed. They live there, as aquatic animals live in water, and in the same way that it be- comes necessary to renew the water which changes and 16 ON TIIF. niYSIOLOOV OF THE HEART loses its nutritive elements, so it becomes necessary to renew, by means of tlie circulation, the blood which loses its oxygen and is charged with carbonic acid gas. This, then, is precisely the duty devolving upon the heart. The system of the left heart carries the blood to the organs which animate them ; and the system of the right heart carries away the blood which has given them mo- mentary life. When we wish to understand the func- tions of an organ physiologically, it is always necessary to trace the vital properties of the substance of which it is 'Composed to their origin—it is consequently in the properties of the tissue of the heart that we will be able to find the explanation of its functions. No difficulty will be experienced in making this investigation, for as we have already said, the heart is a muscle possessing all of the physiological properties of that tissue. It will suf- fice, then, to call to your notice that this fleshy or mus- cular tissue consists of fibres which possess the property of contracting. When the muscular fibres are disposed so as to form a long muscle, whose extremities are inserted into two bones, forming a joint, the necessary effect of the con- traction or shortening of the fibre is to cause the two bones to move on each other, thus approximating; but when the muscular fibres are disposed so as to constitute the walls of a muscular pouch, as is the case in the heart, the necessary effect of the contraction of the mus- cular tissue is to shut up or to cause the cavity to dis- appear more or less completely, by expelling its contents. This enables us to understand how the blood is expelled in a direction determined by the disposition of the cardiac valves at each contraction of the cavities of the heart (Fig. 3). AND ITS CONNECTIONS WITH THE BRAIN. Fig. 3.—Longitudinal section of the heart of man, to show the mech anism of the circulation in the different cavities. od.—Right auricle, vd.—Right ventricle, ot7.—Left auricle, vg.—Left ven- tricle. The arrows placed at the opening of the different vessels ter- minating at the cavities of the heart, indicate the current of the blood in these vessels. When the auricle contracts, the blood is sent into the ventricle, because the auriculo ventricular valve is re- laxed ; when the ventricle contracts, the blood is sent into the arteries, because the sigmoid, or arterial valves, arc relaxed, to-allow the blood to pass at the same time that the auriculo ventricular valve contracts, to prevent the blood from flowing back into the auricle. The con- traction of the cavities of the heart which empties them is followed by a dilatation, during which they are again filled with blood, then by a new contraction which again empties them, and thus the circle is continued. It 18 ON THE PHYSIOLOGY OF T1IE HEART follows, that the motion of the heart consists of a succes- sion of alternate movements of contraction and dilata- tion of these cavities. The contractile movement is called the systole; the diastole is the act of dilatation. The four cavities of the heart contract and dilate succes- sively, two and two, first, the two auricles, then the two ventricles. A very short interval of repose separates the contraction of the auricles from the contraction of the ventricles ; a little longer interval then succeeds the contraction of the ventricles. It would be entirely for- eign to our subject to describe, in detail, the mechanism of the circulation in the different cavities of the heart. In our explanations further on, wo shall have to take notice only of the function of the left ventricle, which, as we have -already said, is the ventricle that supports, feeds and animates all the organs of the body. It will suffice us, then, to say, .that at the moment of the con- traction of this ventricle, the heart projects itself for- ward and strikes, like the beating of a clock, between the fifth and sixth ribs, below the left breast. This is called the beating of the heart. At the same moment of the contraction of the left ventricle, the blood is sent into the aorta, and by this channel, into all of the arte- ries of the body, with a pressure capable of supporting a oolumn of mercury of about five inches in height. It is that which produces the disturbance observed in all of the arteries, and which is called the .pulse. All of the mechanical movements of the heart have been the object of laborious investigation, and modern science has studied the phenomena of the circulation by the aid of graphic processes, which give to these re- searches great exactness. The only point to which we will call your attention, is that the heart is a true living 19 AND ITS CONNECTIONS WITH THE BRAIN. machine, which acts like a forcing pump, in which the piston is replaced by the muscular contraction. The heart itself can trace upon paper each of its contractions with their slightest modifications, and it may then he said, without metaphor, that we read the human heart. Dr. Marey, to whom the Academy of Sciences has ordered a premium, this year, for his important works on the circulation of the blood, will give you an idea of these graphic means, by exhibiting before you experi- ments with the apparatus which he has invented, or brought to its present perfection (Fig. 4). The cardiographic apparatus for the examination of the human heart, is composed of two parts—a stethc- scopic drum (A), intended to detect the beatings of the heart, and a registry drum (B), intended to write them off. The stethescopic drum (A) intended to receive the impulses, should be applied overlie region of the heart; it is formed by a funnel, the broad portion of which is exactly closed by two membranes of caoutchouc (/, /,) betwcecn which, water is introduced, so as to make them swell out in the form of a double convex. The registering drum (B) is formed by another funnel, the broadest part of which is closed by a single membrane of caoutchouc, on which rests a small lever (L), as light as a feather; these two funnels are connected by a tube of caoutchouc (C), which transmits the vibrations of the receiving'drum (A) to the curegistcring drum (B). Each pulsation of the heart reverberates against the water drum of the stethescopic funnel; the air behind these distended membranes vibrates—these vibrations arc transmitted by means of the air which fills the caout- chouc tube, to the membrane of the registering drum, which, vibrating in its turn, causes the little lever to 20 OST THE PHYSIOLOGY OF THE HEART Fig. 4.—Cardiographic or Sphygmograpliic Apparatus of Mr. Marer, for directly inscribing the pulsations of the heart. A. Stethescopic drum, which is placed on the chest, to detect the beatings of the heart. ir i. Two Membranes of caoutchouc, protruding in opposite directions and having between them a lenticular cavity filled with water. B. Registering drum, covered by a caoutchouc membrane slightly protru- ding h, on which is a small prism, which supports the lever L. C. Caout- chouc tube, intended to transmit the vibrations of the receiving drum A, to the registering drum B. c. Valve, for regulating the quantity of air contained in the caoutchouc tube C. L. A very light lever, jointed at one of its extremities so as to move freely around in a vertical plane ; it rests upon a small prism, fixed to the caoutchouc membrane A, of the register- ing membrane B, and is raised or lowered with it; its free extremity rests against the blackened paper ab: nb. Blackened tablet, moving regularly on a railway from b to «, and which receives the inscriptions of the mo- tions of the lever L, representing the beatings of the, heart. move; its point writes, upon tlie moving paper, black- ened with lamp-black, the pulsations of the heart. In this way is traced a figure written by'the heart itself, and which permits us to read its most intimate func- tions. The cardiograph is an instrument so much the more delicate and correct as it can be more closely applied over the heart, and as it is the least separated by the walls of the chest. Without explanation, it AND ITS CONNECTIONS WITH THE BRAIN. 21 will readily appear, why it is more easy to read the heart of children than that of adults, and why, also, it is naturally more difficult to read the heart of women than that of men. It is not my intention to initiate you into the interpre- tation of all of the cardiac writings in the normal and pathological state : I wish simply to exhibit to you a few tracings representing the beatings of the heart either in man or different animals, so as to give you an exact idea of what may he obtained by these graphic means. Here is traced the beatings or pulsations of the heart of a young man in the natural or normal state (Fig. 5). Here is traced the pulsations of the heart of a rabbit (Fig. 0). If we pass on now to cold-blooded animals, whose hearts are simple, we will see the tracings of the pul- sations also simplified in a corresp<®ding manner.* * To obtain a tracing of cold-blooded animals, the cardiographic instru- ment. is simplified: it consists simply of an enregistering lerer, which rests directly over the heart of the animal. ON THE PHYSIOLOGY OF THE HEART A. Systole of the auricle. B. Beginning of the systole of the ventricle. C. End of the systole of the ven- tricle. D. Particular motion, due, perhaps, to the sudden arrest of the blood in the ventricle during its relaxation. Fig. 5—Tracings of the pulsations of the heart of a young man in the normal state. Fig. G.—Tracings of the pulsation of the heart of a rabbit. AND ITS CONNECTIONS WITH THE BRAIN. The tracing of the pulsations of the heart of a frog (Fif. 7). Fit;. 7.—Tracings of the pulsations of the heart of a frog. Fn:. S.—Tracings of the pulsation of the heart of an eel. A. Systole of the auricle. B. Systole of the ventricle. A. Svstolo of the auricle. B. Systole of the ventricle. The tracing of the heart of an eel (Fig. 8). ON THE PHYSIOLOGY OF THE HEART The tracing of the heart of a tortoise (Fig. 9). Fig. 9.—Tracings of the pulsations of the heart of a loitoise. Fig. 10.—Tracings of the pulsations of the heait ( f a crab. A. Systole of the aurie’e. It. C. S.sto'c of the ventriel*. Lastly, is the tracing of the beatings of the heart of an invertebrate animal, a crab (Fig. 10). AND ITS CONNECTIONS WITH THE BRAIN. 25 question which we now wish to examine more particularly in this lecture, is how the heart, that simple motor of the circulation of the blood, can, by re-acting under the influence of the nervous system, co-operate with that exceedingly delicate mechanism of the senti- ments which are produced in us. The heart strikes us at once, as a strange organ, on account of its exceptional activity. In the development of the animal body, each vital organ enters upon the performance of its function, only after it has finished its evolution and acquired its definite texture. There are organs even, particularly those which are destined for the propagation of the species, which take no part in the organic scene until a long time after birth, they then dis- appear, to return again to a state of torpor during the last period of the life of the individual. The heart, on the contrary, manifests its activity from the beginning of life, long before it possesses its perfect form and its characteristic structure. This fact is not only remarka- ble in showing the precocity of the functions of the heart, but it is also of a nature to cause the physiolo- gist to reflect deeply upon the real connection which may exist between anatomical forms, and the vital pro- perties of the tissues. Nothing is so beautiful as to watch the birth of the heart. In the chick, at the twenty-sixth or thirtieth hour of incubation, there appears upon the germinal plane (or field) a very minute point, the punctum saliens, upon which at last may be seen movements, at first unfrequent and hardly perceptible. These motions grad- ually become more decided, and more frequent; the heart is better defined ; arteries and veins pierce it; the sanguineous liquid manifests itself more distinctly, ON THE PHYSIOLOGY OF TH,E HEART and a provisory vascular system (area vasculosa) is dis- played, radiating around the heart, now physiologically 'constituted the organ of embryonic circulation (Fig. 11). Fio. 11.—Area vasculosa, or germinal field, representing the primitive circulation of the chick in the egg. a. Primitive heart, central apparatus of the circulation. At this period, the fundamental lineaments of the body of the animal have already appeared: the heart, now in full activity, presents an isolated, moving blood point, anterior to the organization, and destined to transport on the highway of life, the materials necessary for the formation of the animal body. "While this organ is engaged in the construction and development of the whole body, its own volume is in- creased and developed. At its beginning, it was a small vesicle, obscurely contractile, like the circulatory vesicle A.ND ITS CONNECTIONS WITH THE BRAIN. 27 of an infusorial; but this vesicle soon lengthens itself, and beats with rapidity; the inferior portion receives the blood, and represents an auricle; the superior part constitutes a true ventricle, which sends the blood into the aortic bulb, dividing into the bronchial arcs—it is then the heart of a fish. After a while this heart under- goes a motion combined of a twist and see-saw action, which brings up the auricular part, and lowers the ven- tricular. Before the completion of this see-saw motion, the organ presents a heart with three cavities—it is now the heart of a reptile—and from the time this movement is completed, it possesses the four cavities of the heart of a bird or mammal. The various phases in the de- velopment of the heart, show us, then, that this organ does not reach the most perfect state of organization— as in birds, mammals and man—until it has passed through the forms which remain definite in the inferior classes (Fig. 12). It is the observation of these, and many other facts of the same kind, which has given birth to the idea, philosophically true, that every animal, in its embryonic condition, reflects the organizations which are inferior to it.* The heart differs, also, from all the muscles of the body, in this : that it acts from its first appearance in the embryo, and before it is fully developed. Its organiza- tion once completed, it still continues to form an excep- tion in the muscular system. Indeed, the whole muscu- * It is true that the heart of a chicken, during its development, has three cavities, and possesses an interventricular septum, ■which does not exist in the adult reptile. We must then compare the heart of a chick, during development, with the heart of a reptile undergoing development; for it may be possible that this interventricular septum existed in the reptile at a certain period of development, and that it disappears after a while. 28 ON THE PHYSIOLOGY OF THE HEART lar apparatus presents us with alternations of activity and repose ; the heart, on the contrary, is never at rest; of all the organs of the body, it acts the longest; it Fig. 12.'—Successive periods of the development of the heart of a chick. A. Primitive heart, formed of but one cavity, v. The arterial and the venous heart, not yet being distinct. B. Second period of the development of the heart, corresponding to the heart of a fish. v. The arterial heart, making tire auricular portion placed behind, b. b. The bronchial arteries emanating from the aortic bulba. C. Same figure as the preceding, seen in profile, v. Arterial heart, o. Ve- nous hep,rt. b. Bronchial arcs. D. Third period of the development of the heart, corresponding to the heart with the three cavities of reptiles, v. v. The two artei’ial hearts, right and left, corresponding to the two ventricles, right and left, of the fully developed heart, o. o. The two venous hearts, right and left, corres- ponding to the two auricles, the right and the left, of the fully developed heart, a. Aortic bulb. exists before the organism, it survives it, and in the suc- cessive and natural death of the organs, it is the last to continue to manifest its functions. In a word, to use AND ITS CONNECTIONS WITH THE BKAIN. the expression of the great Haller, the heart is the first to live (primum vivens) and the last to die (ultimum mo- riens). In that extinction of the life of the organism, the heart continues to act when the organs are silent around it. It is the last awake, as if it awaited the end of the struggle between life and death; for so long as it beats, life may be re-established ; 'when the heart iias ceased to beat, it is irrecoverably lost, and as its first pulsation was the certain sign of life, so its last pul- sation is the certain sign of death. The preceding notions are necessary, as they will assist us the better to comprehend the action of the nervous system over the heart. We have already per- ceived that this muscular organ possesses the property of contracting without the intervention of the nervous influence. It begins its functions long before the ner- vous system has given any sign of life; there is even more than this: the nerves may be well developed and anatomically constituted, yet without acting on any of the muscular organs which are already devel- oped. Indeed, I have proved, by direct experiments, that the nervous extremities are not physiologically fused into the muscular system, except in the last stages of embryonic life. After birth, when the nervous sys- tem has assumed the control over all of the muscular organs of the body; the heart, nevertheless, passes from its control, to perform its functions of motor power over the central circulation. The muscles of the members of the body are paralysed by cutting the nerves which ani- mate them : the movements of the heart are never para- lysed by dividing the nerves which are spent upon its tissue, on .the contrary, its movements become more rapid. Those poisons which destroy the properties of 30 ON THE PHYSIOLOGY OF THE HEART the motor nerves, abolish the movements of all of the muscular organs of the body, whilst they exert no influ* ence over the pulsations of the heart. We have here a frog poisoned with curare ; this poison paralyses, par ex- cellence, the motor nervous system : you sec that the heart continues to beat, and causes the blood to circulate through the body of this animal, which is absolutely deprived of all motor nervous influence. From all of this, shall we conclude that the heart pos- sesses no nerves ? This opinion, entertained by the older physiologists, is at this day contradicted by anato- my. That science shows us that the tissue of the heart is abundantly supplied with nerves (Fig. 13). It is not, then, to the absence of nerves that we are to attribute all of the anomilies that the heart presents ; it depends upon a very particular mechanism of the nerves, which remains still to be examined. The well known reaction of the motor nerves on the muscles in general, resolves itself into this fundamental principle: so long as the nerve is not excited, the muscle remains in a state of relaxation and repose, but when the nerve becomes natu- rally or artificially excited, the muscle is thrown into action—contraction. The observation of the influence of our will over the movements of our limbs, will be suflicient to prove what I have advanced, but really nothing is more easy of demonstration, by direct experi- ments, made on living animals, or on those but recently dead (Fig. 13). If a frog be prepared by vivisection, so as to insulate a nerve which is expended upon the muscles pf a limb, we see, that so long as this nerve is not touched, the muscles of the limb remain relaxed and in repose, but as soon as the nerve is excited by pinching, or better still, 31 AND ITS CONNECTIONS WITH THE BRAIN. by passing a current of electricity through it, the mus- cles become energetically and rapidly contracted; here is a general fact, which may be experimentally proved Fio. 13—The heart, with the nervous branches which it receives from the brain. C. The heart, a. The carotid artery going to the brain, n. Pneumogaatric nerve, the branches of which are distributed to the heart. iii man, and all vertebrated animals, either during life or immediately after death, so long as the muscular and nervous systems preserve their respective vital properties. 32 ON THE PHYSIOLOGY OF THE IIEAliT If now, we act upon the nerves of the heart by analo- gous processes, we will see that this paradoxical muscu- lar organ presents, in this point of view, an exception ; and, to he more explicit, I will even say that it presents phenomena absolutely in opposition to those which we have observed in the muscles of the extremities. To be within the truth, it will suffice to reverse the terms'of the proposition, and say, that so long as the nerves of the heart arc not excited, it beats and remains in a func- tional state ; but if the nerves are excited naturally or artificially, it becomes relaxed, and is reduced to a state of repose. If a frog, or any other animal, either living or recently dead, is prepared, by vivisection, so as to observe the heart, and to insulate the pneumogastric nerves which are expended in .its tissue, we prove that so long as these nerves are not acted upon, the heart continues to beat as in ordinary, and as soon as they are excited by a powerful electric current, the diastole of the heart is arrested, that is to say, relaxation takes place. Fig. 14 is a tracing which shows the beginning of re- pose of the heart of a frog,* when its pneumogastric nerve is excited by electricity. Fig. 15 is another tracing showing the entrance to- wards the contraction of the muscle on the limb of the frog, when the nerve which is expended upon it is ex- cited successively and alternately. * The paralysing influence of the pneumogastric upon the beatings of the heart, is not perfectly expressed in the frog. We here exhibit a tracing ob- tained under the same conditions, by Mr. Marey, with the heart of a horse, and which very nearly declares an almost absolute arrest of the heart under the influence of the irritation of the pneumogastric nerve. But it is under- stood, that at the conference in the amphitheatre of the Sorbonue, it was impossible to make the experiment upon a horse. AN!) ttS CONNECTIONS \VlTIl THE BRAIN. 33 Fig. 14.—Tracing representing the beatings of the heart of a horse, to show the paralysing influ- ence of the irritation of the pneumogastric nerve on the contractions of the heart. Excitation of the nerve: repose of the heart. Fig. 15.—Tracings representing the contractions of the muscles of the foot of a frog, to show that the irritation of the nerves provokes these contractions. Excitation of tlio nerve: contraction of the mnsolos. Repose of the pneumogastric contraction of the heart. Repose of the nerve : repose of the muscles The result is equally general ; it is produced m all vertebrated animals, from the frog to man. It will be 34 OX THE PHYSIOLOGY OF THE HEART necessary, tlien, to have the fact of this singular and paradoxical influence of the nerves on the heart always before our minds, because it is this result, which will serve us as a starting point, in explaining hereafter, how the central organ of the circulation can react on our perceptions. But before considering this point, it is necessary to examine more closely the various forms which the stop- page of the heart presents under the influence of the gal- vanic excitement of the nerves. The excitation of the pneumogastric nerves, or the nerves of the heart, by a very active electric current, soon arrests the heatings of that organ. However, there are always some varieties in the phenomena which depend upon the sensitiveness of the animal. If very sensitive mammifers be acted upon, the heart is instantly' arrested, whilst in cold- blooded animals, and especially in the winter, the heart does not immediately feel the nervous influence; many pulsations may still take place before it is arrested. After the cessation of violent galvanic excitation of the nerves, the pulsations no longer appear; the arrest of the heart is then definite, and death immediately takes place. Galvanic excitation of the pneumogastric nerves arrests the heart so much the more energetically, as the application is more sudden, and if it has been seldom repeated. When the excitation is repeated many times, or when it has been continued for a long time, the sen- sibility of the heart and nerves are blunted to such a degree that electricity loses its power to arresj; its pulsa- tions ; the same takes place when the nerves are gradu- ally irritated; very violent currents may thus be succes- sively employed without causing arrest of the heart’s AND ITS CONNECTIONS WITH THE BRAIN. 35 action. When feeble excitations are applied to the nerves of the heart, the results, in the main, are always the same; only, the difference of intensity gives them another appearance. Indeed, feeble and instantaneous galvanic excitation of the pneumogastrics, in a healthy and very sensitive animal, produces a sudden arrest of the heart; but this arrest is of so short duration, that it would often be imperceptible to an unsuspecting ob- server. Besides, after these light or moderate actions, the cardiac pulsations soon reappear, with more energy and rapidity. It is thus seen that the energetic ex- citation of the nerves of the heart, brings on a pro- longed arrest of that organ, with a return of its beat- ings more or less difficult and slow, whilst the mode- rate actions only provoke a very fleeting arrest of the heart’s action, immediately followed by an acceleration in the pulsations, with an increase of the energy of the ventricular contractions. All of the results which we have mentioned, whether relating to the excitation of the nerves which are dis- tributed to the muscles of the limbs, or to the excitation of the nerves of the heart, have been furnished by ex- periments of vivisection, in which the excitant was applied to the motor nerves themselves. But in the natural state, things do not go on in this way; there are physiological excitants which irritate the motor nerves so as to determine their reaction on the muscles. These physiological excitants are two in number, the will and sensation, or sensibility. The will cannot exercise its influence on all of the motor nerves of the body; the nerves of the heart, for example, are beyond its influ- ence. Sensation, on the contrary, exercises an influence which is general; and all of the motor nerves, be they ON THE PHYSIOLOGY OK THE HEART voluntary or involuntary, submit to its reflex action. All sensitive actions, which react on the motor nerves, and give rise to involuntary movements, are called reflex; because it is supposed that the sensitive impression, coming from the periphery, is reflected at the nervous center upon the motor nerve. It will be useless to extend our remarks any farther’ on the mechanism of reflex nervous actions, which, at this day, forms one of the most important liases of the physiology of the nervous system. It will sufflee us to know, that all involuntary movements are the result of the simple action of sensation, or of the sensitive nerve upon the motor nerve, which then reacts on the muscle. All of the involuntary movements of tho heart that we shall have to observe, have no other origin than the re- action of sensation on the motor pneumogastric nerves of that organ ; and when we say, for example, that a painful impression arrests the movements of the heart, we simply signify that a sensitive nerve, primitively excited, has transmitted its impressions to the heart by exciting the pneumogastric, which, in its turn, has caused its motor influence to be felt by tho heart, ab- solutely, as when wo act in our experiments with the galvanic current. When the physiologist provokes a motor nerve to react on the muscles by means of a gal- vanic current, or by pinching, he substitutes an artificial for a natural excitant, which is the will, or the sensi- bility; but the results of the nervous motor action are always the same. It will soon be seen, that all of the forms of arrest of the heart’s action which we have observed, by acting directly with the galvanic agent on the pneumogastric, reproduce themselves, in conse- quence of various sensitive influences. As we now AND ITS CONNECTIONS WITH THE BRAIN. know that sensitive influences can act on the heart, by exciting its motor nerves, we can then understand that intermedium of language, and when we say: that the sensibility, or the sentiments, react on the heart, you will know what that signifies, physiologically. Our direct experiments on the excitation of the pneu- raogastric nerves, have shown us, that the more sensi- tive the animal is, so much the more prompt is the heart in receiving nervous impressions, and to be arrested in its actions ; it is the same for the reactions of the sensitive nerves on the heart. In the frog the heart is not arrested by pinching the skin; more energetic means are necessary to produce this effect; hut in animals higher in the scale—in certain races of dogs, for exam- ple—the slightest excitations of the sensitive nerves show their effect upon the heart. If a hoemamometer be placed on the artery of one of these animals, so as to have the expression of the pulsations under our eyes, by the oscillation of the mercurial column, we will ob- serve, at the moment when a sensitive nerve is rapidly excited, that there is a stoppage in the diastole of the heart; this brings about a suspension of the oscillation, with a slight lowering of the mercurial column. Soon afterwards, the pulsations reappear, considerably accele- rated, and more energetic; for the mercury is raised, sometimes many centimetres, to return to its primitive point; when the heart becomes calm, and resumes its natural rythm. The heart is sometimes so sensitive in certain animals, that very slight excitations of the sensi- tive nerves may bring on reactions, even when the animal manifests no sign of pain. These are some of the ex- periments performed by my piaster, Magendie, and my-, 38 ON THE PHYSIOLOGY OF THE HEART self, many years ago, and which have been often repeated and verified by various processes. In proportion as the animal is elevated in the scale of organization, the heart becomes a more and more deli- cate reagent, to convey the sensitive impressions which take place in the body ; and it is natural to think that man, in this connection, should hold the first rank. In him the heart is not only the central organ of the cir- culation of the blood, but it becomes also a center where all of the sensitive nervous actions expend themselves. The nervous influences which react on the heart, come either from the periphery by the cerebro-spinal system, or from the internal organs by the great sympathetic, or from the cerebral center itself; for in a physiological point of view, the brain should be considered as the most delicate of all the neivous surfaces: whence it results, that the sensitive actions which are derived from this source, are those which exercise the most energetic influence over the heart. How is it possible to conceive of the physiological mechanism, by the aid of which, the heart is connected with the manifestation of our sentiments? AVe know that this organ can receive the contre coup of all the sen- sitive vibrations which take place in us ; and that some- times, if the impression has been very strong, there results from it a violent arrest, with momentary suspension and feebleness of the circulation: if the impression has been slight or moderate, a slight arrest, with reaction and increase in the number and the energy of the cardiac pulsations. But how, then, can this state translate our sentiments ? This is the question which needs an ex- planation. Let us remember that the heart never ceases to be a AND ITS CONNECTIONS WITH THE BfcAIN. forcing pump; that is to say, a motive power, which distributes the blood to all of the organs of the body. If it stops, there is necessarily a suspension or diminu- tion in the supply of the vital liquid to the organs, and as a consequence, suspension or decrease in their func- tions. If, on the other hand, the slight arrest of the heart’s action is followed by an increased intensity in its action, a large quantity of blood is distributed to the organs, and as a consequence, there is an exaggeration of their functions. However, all of the organs of the body, and all of the organic tissues, are not equally sen- sitive to these variations of the arterial circulation, which may quickly diminish or increase the quantity of the nutritious liquid which they receive. The nervous organs, and above all, the brain, which constitutes the apparatus whose texture is the most delicate, and the superior of all in the physiological order, receives the first contre coup of these circulatory disturbances. It is a general law for all animals, from the frog to man; that suspension of the circulation of the blood, in the first place, brings about loss of the cerebral and nervous func- tions, in the same way, as an exaggeration of the circu- lation exalts the central and nervous manifestations. Nevertheless, these reactions of a modified circulation on the nervous organs demand very different periocfi, according to the species; the time it takes to affect cold- blooded animals, is very long, especially in the winter; a frog remains several hours before he experiences the consequences of the arrest of the circulation ; the heart may be taken from it, and for four or five hours after, it continues to jump and swim, without appearing to have had either its will or its movements disturbed in the slightest degree. In warm-blooded animals it is alto- 40 on The physiology of the Heart getlier different; cessation of the heart’s action causes the cerebral phenomena to disappear very rapidly, and this takes place much more easily, as the animal is high in the scale of organization ; that is to say, as it pos- sesses a more delicate nervous organization. Reason and experiment show us that in this respect man ought to hold the first rank; his brain is so delicate that he will expe- rience, in a few seconds, almost instantaneously, the reverberation of the nervous influences exercised on the central organ of the circulation; influences which, you will soon see, translate themselves sometimes by an emotion, and sometimes by syncope. Physiological phenomena everywhere pursue one identical law in all animals; but the more or less delicate nature of the living organism gives them an entirely different expres- sion ; thus the law of the reaction of the heart on the brain, is the same in the frog and in the man ; the frog, however, can never experience an emotion or a syncope, because the time necessary for its heart to feel the ner- vous influence, and its brain to experience the circula- tory impulse, is so long, that the physiological relation between the two disappear. In man, the influence of the heart on the brain is translated by two principal con- ditions, between which many intermediate ones may be Apposed: these are syncope and emotion. Syncope is due to a momentary cessation of the cere- bral functions, in consequence of an interruption in the supply of arterial blood to the brain. Syncope may be produced by tying, or directly compressing all of the arteries which go to the brain; but we will occupy our- selves at this time only with syncope which occurs in consequence of a sensitive influence carried to the heart, and energetic enough to arrest its beatings. The arrest AND ITS CONNECTIONS WITH THE TRAIN. of the heart’s action which produces loss of conscious- ness, by depriving the brain of blood, also brings about pallor of the countenance, and a crowd of other acces- sory effects, which have nothing to do with the question now under consideration. All energetic and sudden sensitive impressions, whatever else may be their na- ture, may bring on syncope. Physical impressions on the sensitive nerves, moral impressions, or painful and voluptuous sensations, conduce to the same result, and bring on the arrest of the heart’s action. The duration of syncope is naturally connected with the duration of the stoppage of the heart. The more intense the arrest has been, the longer, in general, will the syncope last; and the more difficult will it be to re-establish the car- diac pulsations, which at first, return irregularly, and but slowly resume their normal rythm. Sometimes the arrest of the heart is final, and the syncope is mor- tal—this may take place in very feeble, and at the same time very sensitive, individuals. It has been proved, experimentally, that at times, it is only necessary to pro- duce an acute pain, by pinching a sensitive nerve, to bring about a definite arrest of the heart, and mortal syn- cope in doves, wasted by starvation. Emotion arises from the same physiological mech- anism as syncope, but it has an entirely different manifestation. Syncope, which takes the blood from the brain, gives a negative expression, proving that a violent nervous impression has gone to reflect itself on the heart, to rebound on the brain. Emotion, on the contrary, sends to the brain a more active circulation, and gives a positive expression; in this sense the cerebral organ receives a functional super- excitement, in harmony with the nature of the nervous ON THE PHYSIOLOGY OF THE HEART influence which has brought it on. In emotion, there is always an initial impression, which in some way sur- prises, and very slightly arrests, the heart’s action, and consequently excites a feeble cerebral shock, producing a fleeting pallor of the countenance ; the heart, like an animal pricked by a spur, soon reacts, quickens its pace, and sends the blood in full tide through the aorta, and all the other arteries. The brain, the most sensitive of all of the organs, immediately, and before all of the others, feels the effect of this modification of the circulation. The brain is doubtless the starting point of the nervous sensitive impression; but b} the reflex action on the motor nerves of the heart, the sensitive influence has provoked the conditions in the brain which are con- nected to the manifestation of sentiment. The heart is no more the seat of our sentiments than the hand is the seat of our will; but the heart is an instrument which concurs in the expression of our sentiments, as the hand concurs in the dictate of the will. In fine, the heart of man is the most sensitive of all the organs of vegetative life; it is the first to re- ceive the cerebral nervous influence. The brain is the most sensitive of the organs of animal life: it receives the first fruits of the circulation of the blood. The result is, that these two culminating organs of the living machine are connected in ceaseless action and reaction. The heart and brain are, then, in solidarity of reciprocal actions of the most intimate character; actions which increase and decrease in proportion as the organism be- comes more delicate and more developed. These con- nections may be constant or fleeting, varying with the sex and age of the animal. It is thus, that at the period of puberty, when the organs, until then imperfect and AND ITS CONNECTIONS WITH THE BRAIN. inert, are awakened and developed, giving birth to sen- timents in the brain, until now unknown, and bearing to the heart new impressions. The Sentiments we expe- rience are always accompanied by reflex actions of the heart; it is from this organ that the condition necessary for the manifestation of sentiments proceeds, although the brain may be their exclusive seat. In the more elevated organisms, life is but a continual exchange between the sanguineous and nervous systems. The expression of our sentiments is made by an exchange between the heart and the brain, the two most perfect wheels of the living machine. This exchange is carried on by well known anatomical mechanism; by the pneumogastric nerves, which transmit the nervous influ- ence to the heart; and by the carotid and vertebral arteries, which convey the blood to the brain (Fig. 16). The whole of this wonderful mechanism depends, then, upon a thread, and if the nerves which unite the heart to the brain should be destroyed, this reciprocity of action would be interrupted, and the manifestation of our sentiments very seriously disturbed. All of these explanations, I may be told, bear the impress of materialism. To that I will reply, that it is not the question under consideration. If it did not take pae too far from our inquiries, I could easily show you, that in physiology, materialism leads to nothing, and proves nothing; but I will ask ; is a concert of musical instru- ments less ravishing, because the philosopher calculates mathematically all of its vibrations ? Is a physiological phenomenon less wonderful because the physiologist has analysed all of its material conditions ? It is very ne- cessary, then, that these analyses should be made, that these calculations should be pursued, for without them ON THE PHYSIOLOGY OF THE HEART there would be no science. Then physiological science teaches us that outlie one side, the brain really recci ves tlie impression of all of our sentiments, and on the Fig. lti.—The anatomical connections of the heart with the brain, by the carotid arteries and the pneumogastric nerves, to explain the reactions of these two organs upon each other. C. The heart, a. 1 he carotid artery going to the brain, n. Pneun nerve, th e branches of which arc distributed to the heart. other side, that the heart reacts by sending to the brain the necessary conditions for the manifestation of these AND ITS CONNECTIONS WITH THE DRAIN. 45 sentiments, lienee it results that the poet, and the man of romance, appeal to our hearts to affect us, that the man of the world, expressing his sentiments at the same time by calling- upon his heart, uses metaphors which correspond to physiological realities. Sometimes a word, a souvenir the sight of a passing occurrence, awaken in us a deep and intense suffering; this word, this memory, may not be painful in themselves, but are so onty in the phenomenon they excite. When any one says that his heart is broken with grief, real phenomena are produced in the heart—its action has been arrested— if the painful impression has been too sudden, the blood does not reach the brain—syncope and nervous crises are the consequence. There is, then, good reason, when any one has terrible news to impart, which could cause great mental distress, why it should be conveyed with the utmost caution. By our experiments on the nerves of the heart, we know that excitants gradually applied to the heart, blunt or decrease the sensibility of that organ, without pro- ducing arrest of its pulsations. When any one says that he has a full heart, after having suffered agony for a long time, or having expe- rienced painful emotions, this also responds to particular physiological conditions of that organ ; prolonged pain- ful impressions, become incapable of arresting the heart’s action ; they fatigue and wear it out, retarding its pulsations, prolonging its diastole, and producing in the precordial region a sense of fulness or contraction. Agreeable impressions also correspond to particular states of the heart. When a female is surprised by a pleasant emotion, the words which have given it birth have flashed through the mind like lightning, without ON THE PHYSIOLOGY OF THE HEART stopping there ; the heart has been momentarily attacked in advance of all reasoning or reflection. Sentiment begins to manifest itself after a slight arrest of the heart, which may be imperceptible to all save the physi- ologist; the heart, pierced by a nervous impression, reacts in palpitations, which cause it to bound and beat more violently against the breast, at the same time that it sends the blood to the brain, whence results the blush of the face, and the peculiar expression of the features, expressive of pleasure. To say that love causes the heart to palpitate, is not only a poetic form of speech, it is also a physiological reality. When any one says that lie loves with all of his heart, this physiologically signifies that somebody’s presence, or his memory, awakens a nervous impression, which, transmitted to the heart by the pneumogastric nerves, causes that organ to react in a way most conduciye to arouse in the brain a sentiment or emotion of affection. I suppose, be it well understood, that the avowal is sincere ; without this, the heart will not be affected, and the sentiment would only be lip-service. When it is said that great thoughts spring from the heart, it is equivalent to saying that great thoughts come from sentiment, for our sentiments, which have their physiological starting point in the nervous centers, act on the heart as peripheric sensations. The brain of man should have the heart at its service to express its senti- ments, Two hearts united, are hearts which beat in unison under the influence of the same nervous impres- sions; hence result the harmonic expression of similar sentiments. Philosophers say that one can rnaster his heart and silence his passions. These are expressions which physi- AND ITS CONNECTIONS WITH THE BRAIN. 47 ology can interpret: It is known that, by the power of will, man may control many of the reflex actions due to sensations produced by physical causes. Reason, with- out doubt, exercises the same empire over the moral sentiments. Man may, then, become able, through the power of reason#to prevent certain reflex actions from being effected upon the heart; but the greater the tri- umph of pure reason, the weaker will sentiment become. The nervous power capable of arresting reflex ac- tions in general, is less in woman than it is in man: it is this which has given her the supremacy* in the do- main of physical and moral sensibility ; it is this which lias celebrated her as having a heart more tender than that of man. But I must bring these considerations, which would involve too much time, to a close, by a general con- clusion. Science does not contradict the observations and data of art, and I cannot subscribe to the opinion of those who believe that the positive character of science must prove fatal to inspiration. According to my views, I am obliged to come to another and contrtiry conclusion. The artist may find in science the most solid founda- tions, and the savant may draw from art the most assured intuition. Science, although advanced, is still very imperfect, and critical periods may arise, which harrass and disturb, rather than aid the artist. This may be the case with physiology, at the present time, as respects the poet and the philosopher. This is but a transitory state, and I am convinced, that when physiology shall be sufficiently advanced, the poet, the philosopher and the physiologist will understand each other.