# The Works of William Harvey/An Anatomical Disquisition on the Motion of the Heart and Blood in Animals/Chapter 17

CHAPTER XVII.
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 no great bulk, of soft texture or of a certain uniform sameness or simplicity of structure; among the number I may instance grubs and earthworms, 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.
In some of the bloodless and colder animals, further, 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 save 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 made a question without any kind of impropriety.
In all the larger and warmer, because [red-] blooded animals, 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, that no [red-] blooded 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; it is 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 ever hotter and more spirituous, and possess bodies of greater size and consistency, they 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, another 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 principal 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; for 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; but 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, but flowing 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 in fact 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 begin to be apparent: in the altered circumstances the right has only to throw the blood through the lungs, whilst the left has to impel it through the whole body.
There are further within the heart numerous braces, so to speak, 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 some animals have them strongly marked, others have them less so; and, in all that have them, they are more numerous and stronger in the left than in the right ventricle; and whilst some have them in the left, there are yet none present in the right ventricle. In the human subject, again, these fleshy columns and braces are more numerous in the left than in the right ventricle, and they are more abundant in the ventricles than in the auricles; occasionally, indeed, in the auricles there appear to be none present whatsoever. In large, more muscular and hardier bodies, as of countrymen, they are numerous; in more slender frames and in females they are fewer.
In those animals in which the ventricles of the heart are smooth within, and entirely without fibres or muscular bands, or anything like foveae, as in almost all the smaller birds, the partridge and the common fowl, serpents, frogs, tortoises, and also fishes, for the major part, there are no chordæ tendineæ, 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 here is still the same ​as elsewhere: as the lungs are spongy, and loose, and soft, no great amount of force is required to force the blood through them; hence the right ventricle is either without the bundles in question, or they are fewer and weaker, 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 not otherwise—that are endowed with particularly strong frames, and that have large and fleshy limbs at a great distance from the heart, have this central organ of greater thickness, strength, and muscularity. And this is both obvious and necessary. Those, on the contrary, that are of softer and more slender make have the heart more flaccid, softer, and internally either sparely 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 venæ cavæ and pulmonary veins, lest the blood when most forcibly impelled should flow back; and it is for this reason that they are not found in all animals; neither do they appear to have been constructed with equal care in all the 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, and 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, ​indeed, 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, too, it is essential that there can be no heart without a ventricle, since this must be the source and storehouse 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 \vhich 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, however, 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 larger 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 nothing to do save to contain? No; the auricles are prime movers of the blood, especially the right auricle, which is "the first to live, the last to die;" as already said; whence they are subservient to sending the blood into the ventricle, which, contracting incontinently, more readily and forcibly expels 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, since neither the heart nor anything else can dilate or distend itself so as to ​draw aught into its cavity during the diastole, unless, like a sponge, it has been first compressed, and as it is returning to its primary condition; but in animals all local motion proceeds from, and has its original in the contraction of some part: it is consequently by the contraction of the auricles that the blood is thrown into the ventricles, as I have already shown, and from thence, by the contraction of the ventricles, it is propelled and distributed. Which truth concerning local motions, and how the immediate moving organ in every motion of an animal primarily endowed with a motive spirit (as Aristotle has it,) is contractile; and in what way the word νενρον is derived from νεύω, nuto, contraho; and how Aristotle was acquainted with the muscles, and did not unadvisedly refer all motion in animals to the nerves, or to the contractile element, and therefore called those little bands in the heart nerves—all this, 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.
But that we may go on with the subject 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 auricle, 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 within of bands and variously interlacing fibres, that it seems to equal 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 [the ventricular portion] 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 fœtus 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 fœtus is yet in the guise 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; by and by, when the outline of the fœtus is distinctly indicated, and it begins to have greater bodily consistence, the vesicle in question having become more fleshy and stronger, and changed its position, passes into the auricles, over or upon which the body of the heart begins to sprout, though as yet it apparently performs no duty; but when the fœtus is farther advanced, when the bones can be distinguished from the soft parts, and movements take place, then it has also a heart internately 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 constitutions of all, as I may say (ovum, worm, fœtus), it acquires perfection in each. These points will be found elsewhere confirmed by numerous observations on the formation of the fœtus.
Finally, it was not without good grounds that Hippocrates, in his book, 'De Corde,' intitles it a muscle; as its action is the same, so is its function, viz., to contract and move something else, in this case, the charge of blood.
Farther, as in muscles at large, so can we infer the action and use of the heart from the arrangement of its fibres and its general structure. 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 columnæ 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 columnæ, 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 sovereignty of the heart; nor are we to inquire whether it receives sense and motion from the brain? whether blood from the liver? whether it be the origin of the veins and of the blood? and more of the same description. They who affirm these propositions against Aristotle, 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, motion, before either the brain or the liver were in being, 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, is of a date anterior to the body: first formed, nature willed that it should afterwards fashion, nourish, preserve, complete the entire animal, as its work and dwelling place: the heart, like the prince in a kingdom, in whose hands lie the chief and highest authority, rules over all; it is the original and foundation from which all power is derived, on which all power depends in the animal body.
And many things having reference to the arteries farther illustrate and confirm this truth. Why does not the arteria venosa pulsate, seeing that it is numbered among the arteries? Or wherefore is there a pulse in the vena arteriosa? 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: for the more remote vessels are from the heart, with so much the less force are they impinged upon 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; for they have either no perceptible pulse, or they rarely exhibit one, and never save where the heart beats more violently than wont, or at a part where the minute vessel is more dilated or open than elsewhere. Hence it 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. Hence, too, by this single symptom I have ascertained for certain that young persons, whose pulses are naturally rapid, were labouring under fever; 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 at the wrist, and even at the temple; this is the case in persons afflicted with Iipothymiæ and asphyxia, and hysterical symptoms, as also in persons of very weak constitution and in the moribund.
And 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 per saltum, however, because the smaller arteries do not pulsate, especially if a tourniquet has been applied.
And then the reason is the same wherefore the pulmonary artery has not only the structure of an artery, but wherefore it does not differ so widely in the thickness of its tunics from the veins as the aorta: the aorta sustains a more powerful shock from the left ventricle than the pulmonary artery does from the right; and the tunics 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; and in like manner, in the same degree in which the lungs are softer and laxer in structure than the flesh and other constituents of the body at large, do the tunics of the branches of the pulmonary artery differ from the tunics of the vessels derived from the aorta. And the same proportion in these several particulars is universally preserved. The more muscular and powerful men are, the firmer their flesh, the stronger, thicker, denser, and more fibrous their heart, in the same proportion are the auricles and arteries in all respects thicker, closer, and stronger. And again, and on the other hand, in those animals the ventricles of whose heart are smooth within, without villi or valves, and the walls of which are thinner, as in fishes, serpents, birds, and very many genera of animals, in all of them the arteries differ little or nothing in the thickness of their coats from the veins.
Farther, the reason why the lungs have such ample vessels, both arteries and veins, (for the capacity of the pulmonary veins exceeds that of both the 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 arteria venosa and left ventricle so full of blood, of the same black colour and clotted character, too, as that with which the right ventricle and pulmonary artery are filled, inasmuch as the blood is incessantly passing from one side of the heart to the other through the lungs. Wherefore, in fine, the pulmonary artery or vena arteriosa has the constitution of an artery, and the pulmonary veins or arterise venosse have the structure of veins; because, in sooth, in function and constitution, and everything else, the first is an artery, the others are veins, in opposition to what is commonly believed; and why the pulmonary artery has so large an orifice, 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 stand in opposition to 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.
* ↑ [i.e. Not having red blood.—Ed..]
* ↑ De Part. Animal, lib. iii.
* ↑ In the book, de Spiritu, and elsewhere.