William Harvey (1578–1657). On the Motion of the Heart and Blood in Animals.
The Harvard Classics. 1909–14.
IV. Of the Motion of the Heart and Its Auricles, as Seen in the Bodies of Living Animals
Caspar Bauhin and John Riolan, 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 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.