Pulse Aids Epochal Discoveries

Tuesday, 3 June 2014

Images of veins from On the Motion of the Heart and Blood in Animals

William Harvey (1578–1657). On the Motion of the Heart and Blood in Animals.
Vol. 38, pp. 75-86 of The Harvard Classics

Galileo, by holding his pulse while watching a swinging cathedral lamp, evolved a theory that made clocks possible. Harvey, by feeling his pulse, educed that arteries carry blood.
(Dr. William Harvey died June 3, 1657.)

I. The Author’s Motives for Writing

WHEN I first gave my mind to vivisections, as a means of discovering the motions and uses of the heart, and sought to discover these from actual inspection, and not from the writings of others, I found the task so truly arduous, so full of difficulties, that I was almost tempted to think, with Fracastorius, that the motion of the heart was only to be comprehended by God. For I could neither rightly perceive at first when the systole and when the diastole took place, nor when and where dilatation and contraction occurred, by reason of the rapidity of the motion, which in many animals is accomplished in the twinkling of an eye, coming and going like a flash of lightning; so that the systole presented itself to me now from this point, now from that; the diastole the same; and then everything was reversed, the motions occurring, as it seemed, variously and confusedly together. My mind was therefore greatly unsettled nor did I know what I should myself conclude, nor what believe from others. I was not surprised that Andreas Laurentius should have written that the motion of the heart was as perplexing as the flux and reflux of Euripus had appeared to Aristotle.

  At length, by using greater and daily diligence and investigation, making frequent inspection of many and various animals, and collating numerous observations, I thought that I had attained to the truth, that I should extricate myself and escape from this labyrinth, and that I had discovered what I so much desired, both the motion and the use of the heart and arteries. From that time I have not hesitated to expose my views upon these subjects, not only in private to my friends, but also in public, in my anatomical lectures, after the manner of the Academy of old.

  These views as usual, pleased some more, others less; some chid and calumniated me, and laid it to me as a crime that I had dared to depart from the precepts and opinions of all anatomists; others desired further explanations of the novelties, which they said were both worthy of consideration, and might perchance be found of signal use. At length, yielding to the requests of my friends, that all might be made participators in my labors, and partly moved by the envy of others, who, receiving my views with uncandid minds and understanding them indifferently, have essayed to traduce me publicly, I have moved to commit these things to the press, in order that all may be enabled to form an opinion both of me and my labours. This step I take all the more willingly, seeing that Hieronymus Fabricius of Aquapendente, although he has accurately and learnedly delineated almost every one of the several parts of animals in a special work, has left the heart alone untouched. Finally, if any use or benefit to this department of the republic of letters should accrue from my labours, it will, perhaps, be allowed that I have not lived idly, and as the old man in the comedy says:

For never yet hath any one attained
To such perfection, but that time, and place,
And use, have brought addition to his knowledge;
Or made correction, or admonished him,
That he was ignorant of much which he
Had thought he knew; or led him to reject
What he had once esteemed of highest price.

  So will it, perchance, be found with reference to the heart at this time; or others, at least, starting hence, with the way pointed out to them, advancing under the guidance of a happier genius, may make occasion to proceed more fortunately, and to inquire more accurately.

II. On the Motions of the Heart, as Seen in the Dissection of Living Animals

IN THE FIRST place, then, when the chest of a living animal is laid open and the capsule that immediately surrounds the heart is slit up or removed, the organ is seen now to move, now to be at rest; there is a time when it moves, and a time when it is motionless.

  These things are more obvious in the colder animals, such as toads, frogs, serpents, small fishes, crabs, shrimps, snails, and shellfish. They also become more distinct in warm-blooded animals, such as the dog and hog, if they be attentively noted when the heart begins to flag, to move more slowly, and, as it were, to die: the movements then become slower and rarer, the pauses longer, by which it is made much more easy to perceive and unravel what the motions really are, and how they are performed. In the pause, as in death, the heart is soft, flaccid, exhausted, lying, as it were, at rest.

  In the motion, and interval in which this is accomplished, three principal circumstances are to be noted:

  1. That the heart is erected, and rises upwards to a point, so that at this time it strikes against the breast and the pulse is felt externally.

  2. That it is everywhere contracted, but more especially towards the sides so that it looks narrower, relatively longer, more drawn together. The heart of an eel taken out of the body of the animal and placed upon the table or the hand, shows these particulars; but the same things are manifest in the hearts of all small fishes and of those colder animals where the organ is more conical or elongated.

  3. The heart being grasped in the hand, is felt to become harder during its action. Now this hardness proceeds from tension, precisely as when the forearm is grasped, its tendons are perceived to become tense and resilient when the fingers are moved.

  4. It may further be observed in fishes, and the colder blooded animals, such as frogs, serpents, etc., that the heart, when it moves, becomes of a paler color, when quiescent of a deeper blood-red color.

  From these particulars it appears evident to me that the motion of the heart consists in a certain universal tension—both contraction in the line of its fibres, and constriction in every sense. It becomes erect, hard, and of diminished size during its action; the motion is plainly of the same nature as that of the muscles when they contract in the line of their sinews and fibres; for the muscles, when in action, acquire vigor and tenseness, and from soft become hard, prominent, and thickened: and in the same manner the heart.

  We are therefore authorized to conclude that the heart, at the moment of its action, is at once constricted on all sides, rendered thicker in its parietes and smaller in its ventricles, and so made apt to project or expel its charge of blood. This, indeed, is made sufficiently manifest by the preceding fourth observation in which we have seen that the heart, by squeezing out the blood that it contains, becomes paler, and then when it sinks into repose and the ventricle is filled anew with blood, that the deeper crimson colour returns. But no one need remain in doubt of the fact, for if the ventricle be pierced the blood will be seen to be forcibly projected outwards upon each motion or pulsation when the heart is tense.

  These things, therefore, happen together or at the same instant: the tension of the heart, the pulse of its apex, which is felt externally by its striking against the chest, the thickening of its parietes, and the forcible expulsion of the blood it contains by the constriction of its ventricles.

  Hence the very opposite of the opinions commonly received appears to be true; inasmuch as it is generally believed that when the heart strikes the breast and the pulse is felt without, the heart is dilated in its ventricles and is filled with blood; but the contrary of this is the fact, and the heart, when it contracts (and the impulse of the apex is conveyed through the chest wall), is emptied. Whence the motion which is generally regarded as the diastole of the heart, is in truth its systole. And in like manner the intrinsic motion of the heart is not the diastole but the systole; neither is it in the diastole that the heart grows firm and tense, but in the systole, for then only, when tense, is it moved and made vigorous.

  Neither is it by any means to be allowed that the heart only moves in the lines of its straight fibres, although the great Vesalius giving this notion countenance, quotes a bundle of osiers bound in a pyramidal heap in illustration; meaning, that as the apex is approached to the base, so are the sides made to bulge out in the fashion of arches, the cavities to dilate, the ventricles to acquire the form of a cupping-glass and so to suck in the blood. But the true effect of every one of its fibres is to constringe the heart at the same time they render it tense; and this rather with the effect of thickening and amplifying the walls and substance of the organ than enlarging its ventricles. And, again, as the fibres run from the apex to the base, and draw the apex towards the base, they do not tend to make the walls of the heart bulge out in circles, but rather the contrary; inasmuch as every fibre that is circularly disposed, tends to become straight when it contracts; and is distended laterally and thickened, as in the case of muscular fibres in general, when they contract, that is, when they are shortened longitudinally, as we see them in the bellies of the muscles of the body at large. To all this let it be added, that not only are the ventricles contracted in virtue of the direction and condensation of their walls, but farther, that those fibres, or bands, styled nerves by Aristotle, which are so conspicuous in the ventricles of the larger animals, and contain all the straight fibres (the parietes of the heart containing only circular ones), when they contract simultaneously by an admirable adjustment all the internal surfaces are drawn together as if with cords, and so is the charge of blood expelled with force.

  Neither is it true, as vulgarly believed, that the heart by any dilatation or motion of its own, has the power of drawing the blood into the ventricles; for when it acts and becomes tense, the blood is expelled; when it relaxes and sinks together it receives the blood in the manner and wise which will by-and-by be explained.

III. Of the Motions of the Arteries, as seen in the Dissection of Living Animals

IN CONNEXION with the motions of the heart these things are further to be observed having reference to the motions and pulses of the arteries.

  1. At the moment the heart contracts, and when the breast is struck, when in short the organ is in its state of systole, the arteries are dilated, yield a pulse, and are in the state of diastole. In like manner, when the right ventricle contracts and propels its charge of blood, the pulmonary artery is distended at the same time with the other arteries of the body.

  2. When the left ventricle ceases to act, to contract, to pulsate, the pulse in the arteries also ceases; further, when this ventricle contracts languidly, the pulse in the arteries is scarcely perceptible. In like manner, the pulse in the right ventricle failing, the pulse in the pulmonary artery ceases also.

  3. Further, when an artery is divided or punctured, the blood is seen to be forcibly propelled from the wound the moment the left ventricle contracts; and, again, when the pulmonary artery is wounded, the blood will be seen spouting forth with violence at the instant when the right ventricle contracts.

  So also in fishes, if the vessel which leads from the heart to the gills be divided, at the moment when the heart becomes tense and contracted, at the same moment does the blood flow with force from the divided vessel.

  In the same way, when we see the blood in arteriotomy projected now to a greater, now to a less distance, and that the greater jet corresponds to the diastole of the artery and to the time when the heart contracts and strikes the ribs, and is in its state of systole, we understand that the blood is expelled by the same movement.

  From these facts it is manifest, in opposition to commonly received opinions, that the diastole of the arteries corresponds with the time of the heart’s systole; and that the arteries are filled and distended by the blood forced into them by the contraction of the ventricles; the arteries, therefore, are distended, because they are filled like sacs or bladders, and are not filled because they expand like bellows. It is in virtue of one and the same cause, therefore, that all the arteries of the body pulsate, viz., the contraction of the left ventricle; in the same way as the pulmonary artery pulsates by the contraction of the right ventricle.

  Finally, that the pulses of the arteries are due to the impulses of the blood from the left ventricle, may be illustrated by blowing into a glove, when the whole of the fingers will be found to become distended at one and the same time, and in their tension to bear some resemblance to the pulse. For in the ratio of the tension is the pulse of the heart, fuller, stronger, and more frequent as that acts more vigorously, still preserving the rhythm and volume, and order of the heart’s contractions. Nor is it to be expected that because of the motion of the blood, the time at which the contraction of the heart takes place, and that at which the pulse in an artery (especially a distant one) is felt, shall be otherwise than simultaneous: it is here the same as in blowing up a glove or bladder; for in a plenum (as in a drum, a long piece of timber, etc.) the stroke and the motion occur at both extremities at the same time. Aristotle, 1  too, has said, “the blood of all animals palpitates within their veins (meaning the arteries), and by the pulse is sent everywhere simultaneously.” And further, 2 thus do all the veins pulsate together and by successive strokes, because they all depend upon the heart; and, as it is always in motion, so are they likewise always moving together, but by successive movements.” It is well to observe with Galen, in this place, that the old philosophers called the arteries veins.

  I happened upon one occasion to have a particular case under my care, which plainly satisfied me of the truth: A certain person was affected with a large pulsating tumour on the right side of the neck, called an aneurism, just at that part where the artery descends into the axilla, produced by an erosion of the artery itself, and daily increasing in size; this tumour was visibly distended as it received the charge of blood brought to it by the artery, with each stroke of the heart; the connexion of parts was obvious when the body of the patient came to be opened after his death. The pulse in the corresponding arm was small, in consequence of the greater portion of the blood being diverted into the tumour and so intercepted.

  Whence it appears that whenever the motion of the blood through the arteries is impeded, whether it be by compression or infarction, or interception, there do the remote divisions of the arteries beat less forcibly, seeing that the pulse of the arteries is nothing more than the impulse or shock of the blood in these vessels.

Note 1. De Anim., iii, cap. 9.
Note 2. De Respir., cap. 20.

IV. Of the Motion of the Heart and Its Auricles, as Seen in the Bodies of Living Animals

BESIDES the motions already spoken of, we have still to consider those that appertain to the auricles.

  Caspar Bauhin and John Riolan,  1 most learned men and skilful anatomists, inform us that from their observations, that if we carefully watch the movements of the heart in the vivisection of an animal, we shall perceive four motions distinct in time and in place, two of which are proper to the auricles, two to the ventricles. With all deference to such authority I say that there are four motions distinct in point of place, but not of time; for the two auricles move together, and so also do the two ventricles, in such wise that though the places be four, the times are only two. And this occurs in the following manner:

  There are, as it were, two motions going on together: on of the auricles, another of the ventricles; these by no means taking place simultaneously, but the motion of the auricles preceding, that of the heart following; the motion appearing to begin from the auricles and to extend to the ventricles. When all things are becoming languid, and the heart is dying, as also in fishes and the colder blooded animals there is a short pause between these two motions, so that the heart aroused, as it were, appears to respond to the motion, now more quickly, now more tardily; and at length, when near to death, it ceases to respond by its proper motion, but seems, as it were, to nod the head, and is so slightly moved that it appears rather to give signs of motion to the pulsating auricles than actually to move. The heart, therefore, ceases to pulsate sooner than the auricles, so that the auricles have been said to outlive it, the left ventricle ceasing to pulsate first of all; then its auricle, next the right ventricle; and, finally, all the other parts being at rest and dead, as Galen long since observed, the right auricle still continues to beat; life, therefore, appears to linger longest in the right auricle. Whilst the heart is gradually dying, it is sometimes seen to reply, after two or three contractions of the auricles, roused as it were to action, and making a single pulsation, slowly, unwillingly, and with an effort.

  But this especially is to be noted, that after the heart has ceased to beat, the auricles however still contracting, a finger placed upon the ventricles perceives the several pulsations of the auricles, precisely in the same way and for the same reason, as we have said, that the pulses of the ventricles are felt in the arteries, to wit, the distension produced by the jet of blood. And if at this time, the auricles alone pulsating, the point of the heart be cut off with a pair of scissors, you will perceive the blood flowing out upon each contraction of the auricles. Whence it is manifest that the blood enters the ventricles, not by any attraction or dilatation of the heart, but by being thrown into them by the pulses of the auricles.

  And here I would observe, that whenever I speak of pulsations as occurring in the auricles or ventricles, I mean contractions: first the auricles contract, and then and subsequently the heart itself contracts. When the auricles contract they are seen to become whiter, especially where they contain but little blood; but they are filled as magazines or reservoirs of the blood, which is tending spontaneously and, by its motion in the veins, under pressure towards the centre; the whiteness indicated is most conspicuous towards the extremities or edges of the auricles at the time of their contractions.

  In fishes and frogs, and other animals which have hearts with but a single ventricle, and for an auricle have a kind of bladder much distended with blood, at the base of the organ, you may very plainly perceive this bladder contracting first, and the contraction of the heart or ventricle following afterwards.

  But I think it right to describe what I have observed of an opposite character: the heart of an eel, of several fishes, and even of some (of the higher) animals taken out of the body, pulsates without auricles; nay, if it be cut in pieces the several parts may still be seen contracting and relaxing; so that in these creatures the body of the heart may be seen pulsating and palpitating, after the cessation of all motion in the auricle. But is not this perchance peculiar to animals more tenacious of life, whose radical moisture is more glutinous, or fat and sluggish, and less readily soluble? The same faculty indeed appears in the flesh of eels, which even when skinned and embowelled, and cut into pieces, are still seen to move.

  Experimenting with a pigeon upon one occasion, after the heart had wholly ceased to pulsate, and the auricles too had become motionless, I kept my finger wetted with saliva and warm for a short time upon the heart, and observed that under the influence of this fomentation it recovered new strength and life, so that both ventricles and auricles pulsated, contracting and relaxing alternately, recalled as it were from death to life.

  Besides this, however, I have occasionally observed, after the heart and even its right auricle had ceased pulsating,—when it was in articulo mortis in short,—that an obscure motion, an undulation or palpitation, remained in the blood itself, which was contained in the right auricle, this being apparent so long as it was imbued with heat and spirit. And, indeed, a circumstance of the same kind is extremely manifest in the course of the generation of animals, as may be seen in the course of the first seven days of the incubation of the chick: A drop of blood makes its appearance which palpitates, as Aristotle had already observed; from this, when the growth is further advanced and the chick is fashioned, the auricles of the heart are formed, which pulsating henceforth give constant signs of life. When at length, and after the lapse of a few days, the outline of the body begins to be distinguished, then is the ventricular part of the heart also produced, but it continues for a time white and apparently bloodless, like the rest of the animal; neither does it pulsate or give signs of motion. I have seen a similar condition of the heart in the human fœtus about the beginning of the third month, the heart then being whitish and bloodless, although its auricles contained a considerable quantity of purple blood. In the same way in the egg, when the chick was formed and had increased in size, the heart too increased and acquired ventricles, which then began to receive and to transmit blood.

  And this leads me to remark that he who inquires very particularly into this matter will not conclude that the heart, as a whole, is the primum vivens, ultimum moriens,—the first part to live, the last to die,—but rather its auricles, or the part which corresponds to the auricles in serpents, fishes, etc., which both lives before the heart and dies after it.

  Nay, has not the blood itself or spirit an obscure palpitation inherent in it, which it has even appeared to me to retain after death? and it seems very questionable whether or not we are to say that life begins with the palpitation or beating of the heart. The seminal fluid of all animals—the prolific spirit, as Aristotle observed, leaves their body with a bound and like a living thing; and nature in death, as Aristotle  2 further remarks, retracing her steps, reverts to where she had set out, and returns at the end of her course to the goal whence she had started. As animal generation proceeds from that which is not animal, entity from non-entity, so, by a retrograde course, entity, by corruption, is resolved into non-entity, whence that in animals, which was last created, fails first and that which was first, fails last.

  I have also observed that almost all animals have truly a heart, not the larger creatures only, and those that have red blood, but the smaller, and pale-blooded ones also, such as slugs, snails, scallops, shrimps, crabs, crayfish, and many others; nay, even in wasps, hornets, and flies, I have, with the aid of a magnifying glass, and at the upper part of what is called the tail, both seen the heart pulsating myself, and shown it to many others.

  But in the pale-blooded tribes the heart pulsates sluggishly and deliberately, contracting slowly as in animals that are moribund, a fact that may readily be seen in the snail, whose heart will be found at the bottom of that orifice in the right side of the body which is seen to be opened and shut in the course of respiration, and whence saliva is discharged, the incision being made in the upper aspect of the body, near the part which corresponds to the liver.

  This, however, is to be observed: that in winter and the colder season, exsanguine animals, such as the snail, show no pulsation; they seem rather to live after the manner of vegetables, or of those other productions which are therefore designated plant-animals.

  It is also to be noted that all animals which have a heart have also auricles, or something analogous to auricles; and, further, that whenever the heart has a double ventricle, there are always two auricles present, but not otherwise. If you turn to the production of the chick in ovo, however, you will find at first no more a vesicle or auricle, or pulsating drop of blood; it is only by and by, when the development has made some progress, that the heart is fashioned; even so in certain animals not destined to attain to the highest perfection in their organization, such as bees, wasps, snails, shrimps, crayfish, etc., we only find a certain pulsating vesicle, like a sort of red or white palpitating point, as the beginning or principle of their life.

  We have a small shrimp in these countries, which is taken in the Thames and in the sea, the whole of whose body is transparent; this creature, placed in a little water, has frequently afforded myself and particular friends an opportunity of observing the motions of the heart with the greatest distinctness, the external parts of the body presenting no obstacle to our view, but the heart being perceived as though it had been seen through a window.

  I have also observed the first rudiments of the chick in the course of the fourth or fifth day of the incubation, in the guise of a little cloud, the shell having been removed and the egg immersed in clear tepid water. In the midst of the cloudlet in question there was a bloody point so small that it disappeared during the contraction and escaped the sight, but in the relaxation it reappeared again, red and like the point of a pin; so that betwixt the visible and invisible, betwixt being and not being, as it were, it gave by its pulses a kind of representation of the commencement of life.

Note 1. Bauhin, lib. ii, cap. 21. Riolan, lib. viii, cap. 1.
Note 2. De Motu Animal., cap. 8.


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