On the Motion of the Heart and Blood in Animals, by William Harvey

Chapter XVII

The Motion and Circulation of the Blood are Confirmed from the Particulars Apparent in the Structure of the Heart, and From Those Things which Dissection Unfolds

I do not find the heart as a distinct and separate part in all animals; some, indeed, such as the zoophytes, have no heart; this is because these animals are coldest, of one great bulk, of soft texture, or of a certain uniform sameness or simplicity of structure; among the number I may instance grubs and earth-worms, and those that are engendered of putrefaction and do not preserve their species. These have no heart, as not requiring any impeller of nourishment into the extreme parts; for they have bodies which are connate and homogeneous and without limbs; so that by the contraction and relaxation of the whole body they assume and expel, move and remove, the aliment. Oysters, mussels, sponges, and the whole genus of zoophytes or plant-animals have no heart, for the whole body is used as a heart, or the whole animal is a heart. In a great number of animals — almost the whole tribe of insects — we cannot see distinctly by reason of the smallness of the body; still in bees, flies, hornets, and the like we can perceive something pulsating with the help of a magnifying-glass; in pediculi, also, the same thing may be seen, and as the body is transparent, the passage of the food through the intestines, like a black spot or stain, may be perceived by the aid of the same magnifying-glass.

But in some of the pale-blooded and colder animals, as in snails, whelks, shrimps, and shell-fish, there is a part which pulsates, — a kind of vesicle or auricle without a heart — slowly, indeed, and not to be perceived except in the warmer season of the year. In these creatures this part is so contrived that it shall pulsate, as there is here a necessity for some impulse to distribute the nutritive fluid, by reason of the variety of organic parts, or of the density of the substance; but the pulsations occur unfrequently, and sometimes in consequence of the cold not at all, an arrangement the best adapted to them as being of a doubtful nature, so that sometimes they appear to live, sometimes to die; sometimes they show the vitality of an animal, sometimes of a vegetable. This seems also to be the case with the insects which conceal themselves in winter, and lie, as it were, defunct, or merely manifesting a kind of vegetative existence. But whether the same thing happens in the case of certain animals that have red blood, such as frogs, tortoises, serpents, swallows, may be very properly doubted.

In all the larger and warmer animals which have red blood, there was need of an impeller of the nutritive fluid, and that, perchance, possessing a considerable amount of power. In fishes, serpents, lizards, tortoises, frogs, and others of the same kind there is a heart present, furnished with both an auricle and a ventricle, whence it is perfectly true, as Aristotle has observed, 13 that no sanguineous animal is without a heart, by the impelling power of which the nutritive fluid is forced, both with greater vigour and rapidity, to a greater distance; and not merely agitated by an auricle, as it Is in lower forms. And then in regard to animals that are yet larger, warmer, and more perfect, as they abound in blood, which is always hotter and more spirituous, and which possess bodies of greater size and consistency, these require a larger, stronger, and more fleshy heart, in order that the nutritive fluid may be propelled with yet greater force and celerity. And further, inasmuch as the more perfect animals require a still more perfect nutrition, and a larger supply of native heat, in order that the aliment may be thoroughly concocted and acquire the last degree of perfection, they required both lungs and a second ventricle, which should force the nutritive fluid through them.

Every animal that has lungs has, therefore, two ventricles to its heart — one right, the other left; and wherever there is a right, there also is there a left ventricle; but the contrary of this does not hold good: where there is a left there is not always a right ventricle. The left ventricle I call that which is distinct in office, not in place from the other, that one, namely, which distributes the blood to the body at large, not to the lungs only. Hence the left ventricle seems to form the principle part of the heart; situated in the middle, more strongly marked, and constructed with greater care, the heart seems formed for the sake of the left ventricle, and the right but to minister to it. The right neither reaches to the apex of the heart nor is it nearly of such strength, being three times thinner in its walls, and in some sort jointed on to the left (as Aristotle says), though, indeed, it is of greater capacity, inasmuch as it has not only to supply material to the left ventricle, but likewise to furnish aliment to the lungs.

It is to be observed, however, that all this is otherwise in the embryo, where there is not such a difference between the two ventricles. There, as in a double nut, they are nearly equal in all respects, the apex of the right reaching to the apex of the left, so that the heart presents itself as a sort of double-pointed cone. And this is so, because in the foetus, as already said, whilst the blood is not passing through the lungs from the right to the left cavities of the heart, it flows by the foramen ovale and ductus arteriosus directly from the vena cava into the aorta, whence it is distributed to the whole body. Both ventricles have, therefore, the same office to perform, whence their equality of constitution. It is only when the lungs come to be used and it is requisite that the passages indicated should be blocked up that the difference in point of strength and other things between the two ventricles begins to be apparent. In the altered circumstances the right has only to drive the blood through the lungs, whilst the left has to propel it through the whole body.

There are, moreover, within the heart numerous braces, in the form of fleshy columns and fibrous bands, which Aristotle, in his third book on “Respiration,” and the “Parts of Animals,” entitles nerves. These are variously extended, and are either distinct or contained in grooves in the walls and partition, where they occasion numerous pits or depressions. They constitute a kind of small muscles, which are superadded and supplementary to the heart, assisting it to execute a more powerful and perfect contraction, and so proving subservient to the complete expulsion of the blood. They are, in some sort, like the elaborate and artful arrangement of ropes in a ship, bracing the heart on every side as it contracts, and so enabling it more effectually and forcibly to expel the charge of blood from its ventricles. This much is plain, at all events, that in some animals they are less strongly marked than in others; and, in all that have them, they are more numerous and stronger in the left than in the right ventricle; and while some have them present in the left, yet they are absent in the right ventricle. In man they are more numerous in the left than in the right ventricle, more abundant in the ventricles than in the auricles; and occasionally there appear to be none present in the auricles. They are numerous in the large, more muscular and hardier bodies of countrymen, but fewer in more slender frames and in females.

In those animals in which the ventricles of the heart are smooth within and entirely without fibres of muscular bands, or anything like hollow pits, as in almost all the smaller birds, the partridge and the common fowl, serpents, frogs, tortoises, and most fishes, there are no chordae tendineae, nor bundles of fibres, neither are there any tricuspid valves in the ventricles.

Some animals have the right ventricle smooth internally, but the left provided with fibrous bands, such as the goose, swan, and larger birds; and the reason is the same here as elsewhere. As the lungs are spongy and loose and soft, no great amount of force is required to force the blood through them; therefore the right ventricle is either without the bundles in question, or they are fewer and weaker, and not so fleshy or like muscles. Those of the left ventricle, however, are both stronger and more numerous, more fleshy and muscular, because the left ventricle requires to be stronger, inasmuch as the blood which it propels has to be driven through the whole body. And this, too, is the reason why the left ventricle occupies the middle of the heart, and has parietes three times thicker and stronger than those of the right Hence all animals — and among men it is similar — that are endowed with particularly strong frames, and with large and fleshy limbs at a great distance from the heart, have this central organ of greater thickness, strength, and muscularity. This is manifest and necessary. Those, on the contrary, that are of softer and more slender make have the heart more flaccid, softer, and internally either less or not at all fibrous. Consider, farther, the use of the several valves, which are all so arranged that the blood, once received into the ventricles of the heart, shall never regurgitate; once forced into the pulmonary artery and aorta, shall not flow back upon the ventricles. When the valves are raised and brought together, they form a three-cornered line, such as is left by the bite of a leech; and the more they are forced, the more firmly do they oppose the passage of the blood. The tricuspid valves are placed, like gate-keepers, at the entrance into the ventricles from the venae cavae and pulmonary veins, lest the blood when most forcibly impelled should flow back. It is for this reason that they are not found in all animals, nor do they appear to have been constructed with equal care, in all animals in which they are found. In some they are more accurately fitted, in others more remissly or carelessly contrived, and always with a view to their being closed under a greater or a slighter force of the ventricle. In the left ventricle, therefore, in order that the occlusion may be the more perfect against the greater impulse, there are only two valves, like a mitre, and produced into an elongated cone, so that they come together and touch to their middle; a circumstance which perhaps led Aristotle into the error of supposing this ventricle to be double, the division taking place transversely. For the same reason, and that the blood may not regurgitate upon the pulmonary veins, and thus the force of the ventricle in propelling the blood through the system at large come to be neutralized, it is that these mitral valves excel those of the right ventricle in size and strength and exactness of closing. Hence it is essential that there can be no heart without a ventricle, since this must be the source and store-house of the blood. The same law does not hold good in reference to the brain. For almost no genus of birds has a ventricle in the brain, as is obvious in the goose and swan, the brains of which nearly equal that of a rabbit in size; now rabbits have ventricles in the brain, whilst the goose has none. In like manner, wherever the heart has a single ventricle, there is an auricle appended, flaccid, membranous, hollow, filled with blood; and where there are two ventricles, there are likewise two auricles. On the other hand, some animals have an auricle without any ventricle; or, at all events, they have a sac analogous to an auricle; or the vein itself, dilated at a particular part, performs pulsations, as is seen in hornets, bees, and other insects, which certain experiments of my own enable me to demonstrate, have not only a pulse, but a respiration in that part which is called the tail, whence it is that this part is elongated and contracted now more rarely, now more frequently, as the creature appears to be blown and to require a large quantity of air. But of these things, more in our “Treatise On Respiration.”

It is in like manner evident that the auricles pulsate, contract, as I have said before, and throw the blood into the ventricles; so that wherever there is a ventricle, an auricle is necessary, not merely that it may serve, according to the general belief, as a source and magazine for the blood: for what were the use of its pulsations had it only to contain?

The auricles are prime movers of the blood, especially the right auricle, which, as already said, is “the first to live, the last to die”; whence they are subservient to sending the blood into the ventricles, which, contracting continuously, more readily and forcibly expel the blood already in motion; just as the ball-player can strike the ball more forcibly and further if he takes it on the rebound than if he simply threw it. Moreover, and contrary to the general opinion, neither the heart nor anything else can dilate or distend itself so as to draw anything into its cavity during the diastole, unless, like a sponge, it has been first compressed and is returning to its primary condition. But in animals all local motion proceeds from, and has its origin in, the contraction of some part; consequently it is by the contraction of the auricles that the blood is thrown into the ventricles, as I have already shown, and from there, by the contraction of the ventricles, it is propelled and distributed. Concerning local motions, it is true that the immediate moving organ in every motion of an animal primarily endowed with a motive spirit (as Aristotle has it14) is contractile; in which way the word veopou is derived from veuw, nuto, contraho; and if I am permitted to proceed in my purpose of making a particular demonstration of the organs of motion in animals from observations in my possession, I trust I shall be able to make sufficiently plain how Aristotle was acquainted with the muscles, and advisedly referred all motion in animals to the nerves, or to the contractile element, and, therefore, called those little bands in the heart nerves.

But that we may proceed with the subject which we have in hand, viz., the use of the auricles in filling the ventricles, we should expect that the more dense and compact the heart, the thicker its parietes, the stronger and more muscular must be the auricle to force and fill it, and vice versa. Now this is actually so: in some the auricle presents itself as a sanguinolent vesicle, as a thin membrane containing blood, as in fishes, in which the sac that stands in lieu of the auricles is of such delicacy and ample capacity that it seems to be suspended or to float above the heart. In those fishes in which the sac is somewhat more fleshy, as in the carp, barbel, tench, and others, it bears a wonderful and strong resemblance to the lungs.

In some men of sturdier frame and stouter make the right auricle is so strong, and so curiously constructed on its inner surface of bands and variously interlacing fibres, that it seems to equal in strength the ventricle of the heart in other subjects; and I must say that I am astonished to find such diversity in this particular in different individuals. It is to be observed, however, that in the foetus the auricles are out of all proportion large, which is because they are present before the heart makes its appearance or suffices for its office even when it has appeared, and they, therefore, have, as it were, the duty of the whole heart committed to them, as has already been demonstrated. But what I have observed in the formation of the foetus, as before remarked (and Aristotle had already confirmed all in studying the incubated egg), throws the greatest light and likelihood upon the point. Whilst the foetus is yet in the form of a soft worm, or, as is commonly said, in the milk, there is a mere bloody point or pulsating vesicle, a portion apparently of the umbilical vein, dilated at its commencement or base. Afterwards, when the outline of the foetus is distinctly indicated and it begins to have greater bodily consistence, the vesicle in question becomes more fleshy and stronger, changes its position, and passes into the auricles, above which the body of the heart begins to sprout, though as yet it apparently performs no office. When the foetus is farther advanced, when the bones can be distinguished from the fleshy parts and movements take place, then it also has a heart which pulsates, and, as I have said, throws blood by either ventricle from the vena cava into the arteries.

Thus nature, ever perfect and divine, doing nothing in vain, has neither given a heart where it was not required, nor produced it before its office had become necessary; but by the same stages in the development of every animal, passing through the forms of all, as I may say (ovum, worm, foetus), it acquires perfection in each. These points will be found elsewhere confirmed by numerous observations on the formation of the foetus.

Finally, it is not without good grounds that Hippocrates in his book, “De Corde,” entitles it a muscle; its action is the same; so is its functions, viz., to contract and move something else — in this case the charge of the blood.

Farther, we can infer the action and use of the heart from the arrangement of its fibres and its general structures, as in muscles generally. All anatomists admit with Galen that the body of the heart is made up of various courses of fibres running straight, obliquely, and transversely, with reference to one another; but in a heart which has been boiled, the arrangement of the fibres is seen to be different. All the fibres in the parietes and septum are circular, as in the sphincters; those, again, which are in the columns extend lengthwise, and are oblique longitudinally; and so it comes to pass that when all the fibres contract simultaneously, the apex of the cone is pulled towards its base by the columns, the walls are drawn circularly together into a globe — the whole heart, in short, is contracted and the ventricles narrowed. It is, therefore, impossible not to perceive that, as the action of the organ is so plainly contraction, its function is to propel the blood into the arteries.

Nor are we the less to agree with Aristotle in regard to the importance of the heart, or to question if it receives sense and motion from the brain, blood from the liver, or whether it be the origin of the veins and of the blood, and such like. They who affirm these propositions overlook, or do not rightly understand, the principal argument, to the effect that the heart is the first part which exists, and that it contains within itself blood, life, sensation, and motion, before either the brain or the liver were created or had appeared distinctly, or, at all events, before they could perform any function. The heart, ready furnished with its proper organs of motion, like a kind of internal creature, existed before the body. The first to be formed, nature willed that it should afterwards fashion, nourish, preserve, complete the entire animal, as its work and dwelling-place: and as the prince in a kingdom, in whose hands lie the chief and highest authority, rules over all, the heart is the source and foundation from which all power is derived, on which all power depends in the animal body.

Many things having reference to the arteries farther illustrate and confirm this truth. Why does not the pulmonary vein pulsate, seeing that it is numbered among the arteries? Or wherefore is there a pulse in the pulmonary artery? Because the pulse of the arteries is derived from the impulse of the blood. Why does an artery differ so much from a vein in the thickness and strength of its coats? Because it sustains the shock of the impelling heart and streaming blood. Hence, as perfect nature does nothing in vain, and suffices under all circumstances, we find that the nearer the arteries are to the heart, the more do they differ from the veins in structure; here they are both stronger and more ligamentous, whilst in extreme parts of the body, such as the feet and hands, the brain, the mesentery, and the testicles, the two orders of vessels are so much alike that it is impossible to distinguish between them with the eye. Now this is for the following very sufficient reasons: the more remote the vessels are from the heart, with so much the less force are they distended by the stroke of the heart, which is broken by the great distance at which it is given. Add to this that the impulse of the heart exerted upon the mass of blood, which must needs fill the trunks and branches of the arteries, is diverted, divided, as it were, and diminished at every subdivision, so that the ultimate capillary divisions of the arteries look like veins, and this not merely in constitution, but in function. They have either no perceptible pulse, or they rarely exhibit one, and never except where the heart beats more violently than usual, or at a part where the minute vessel is more dilated or open than elsewhere. It, therefore, happens that at times we are aware of a pulse in the teeth, in inflammatory tumours, and in the fingers; at another time we feel nothing of the sort. By this single symptom I have ascertained for certain that young persons whose pulses are naturally rapid were labouring under fever; and in like manner, on compressing the fingers in youthful and delicate subjects during a febrile paroxysm, I have readily perceived the pulse there. On the other hand, when the heart pulsates more languidly, it is often impossible to feel the pulse not merely in the fingers, but the wrist, and even at the temple, as in persons afflicted with lipothymiae asphyxia, or hysterical symptoms, and in the debilitated and moribund.

Here surgeons are to be advised that, when the blood escapes with force in the amputation of limbs, in the removal of tumours, and in wounds, it constantly comes from an artery; not always indeed per saltum, because the smaller arteries do not pulsate, especially if a tourniquet has been applied.

For the same reason the pulmonary artery not only has the structure of an artery, but it does not differ so widely from the veins in the thickness of its walls as does the aorta. The aorta sustains a more powerful shock from the left than the pulmonary artery does from the right ventricle, and the walls of this last vessel are thinner and softer than those of the aorta in the same proportion as the walls of the right ventricle of the heart are weaker and thinner than those of the left ventricle. In like manner the lungs are softer and laxer in structure than the flesh and other constituents of the body, and in a similar way the walls of the branches of the pulmonary artery differ from those of the vessels derived from the aorta. And the same proportion in these particulars is universally preserved. The more muscular and powerful men are, the firmer their flesh; the stronger, thicker, denser, and more fibrous their hearts, the thicker, closer, and stronger are the auricles and arteries. Again, in those animals the ventricles of whose hearts are smooth on their inner surface, without villi or valves, and the walls of which are thin, as in fishes, serpents, birds, and very many genera of animals, the arteries differ little or nothing in the thickness of their coats from the veins.

Moreover, the reason why the lungs have such ample vessels, both arteries and veins (for the capacity of the pulmonary veins exceeds that of both crural and jugular vessels), and why they contain so large a quantity of blood, as by experience and ocular inspection we know they do, admonished of the fact indeed by Aristotle, and not led into error by the appearances found in animals which have been bled to death, is, because the blood has its fountain, and storehouse, and the workshop of its last perfection, in the heart and lungs. Why, in the same way, we find in the course of our anatomical dissections the pulmonary vein and left ventricle so full of blood, of the same black colour and clotted character as that with which the right ventricle and pulmonary artery are filled, is because the blood is incessantly passing from one side of the heart to the other through the lungs. Wherefore, in fine, the pulmonary artery has the structure of an artery, and the pulmonary veins have the structure of veins. In function and constitution and everything else the first is an artery, the others are veins, contrary to what is commonly believed; and the reason why the pulmonary artery has so large an orifice is because it transports much more blood than is requisite for the nutrition of the lungs.

All these appearances, and many others, to be noted in the course of dissection, if rightly weighed, seem clearly to illustrate and fully to confirm the truth contended for throughout these pages, and at the same time to oppose the vulgar opinion; for it would be very difficult to explain in any other way to what purpose all is constructed and arranged as we have seen it to be.

13De Part. Animal., lib. iii.

14In the book de Spiritu, and elsewhere.

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