Contrariwise, there is a no less wide foundation for the generalisation that animals, as Cuvier puts it, depend directly or indirectly upon plants for the materials of their bodies; that is, either they are herbivorous, or they eat other animals which are herbivorous.
But for what constituents of their bodies are animals thus dependent upon plants? Certainly not for their horny matter; nor for chondrin, the proximate chemical element of cartilage; nor for gelatine; nor for syntonin, the constituent of muscle; nor for their nervous or biliary substances; nor for their amyloid matters; nor, necessarily, for their fats.
It can be experimentally demonstrated that animals can make these for themselves. But that which they cannot make, but must, in all known cases, obtain directly or indirectly from plants, is the peculiar nitrogenous matter, protein. Thus the plant is the ideal prolétaire of the living world, the worker who produces; the animal, the ideal aristocrat, who mostly occupies himself in consuming, after the manner of that noble representative of the line of Zähdarm, whose epitaph is written in Sartor Resartus.
Here is our last hope of finding a sharp line of demarcation between plants and animals; for, as I have already hinted, there is a border territory between the two kingdoms, a sort of no-man’s-land, the inhabitants of which certainly cannot be discriminated and brought to their proper allegiance in any other way.
Some months ago, Professor Tyndall asked me to examine a drop of infusion of hay, placed under an excellent and powerful microscope, and to tell him what I thought some organisms visible in it were. I looked and observed, in the first place, multitudes of Bacteria moving about with their ordinary intermittent spasmodic wriggles. As to the vegetable nature of these there is now no doubt. Not only does the close resemblance of the Bacteria to unquestionable plants, such as the Oscillatoriæ, and lower forms of Fungi, justify this conclusion, but the manufacturing test settles the question at once. It is only needful to add a minute drop of fluid containing Bacteria, to water in which tartrate, phosphate, and sulphate of ammonia are dissolved; and, in a very short space of time, the clear fluid becomes milky by reason of their prodigious multiplication, which, of course, implies the manufacture of living Bacterium-stuff out of these merely saline matters.
But other active organisms, very much larger than the Bacteria, attaining in fact the comparatively gigantic dimensions of 1/3000 of an inch or more, incessantly crossed the field of view. Each of these had a body shaped like a pear, the small end being slightly incurved and produced into a long curved filament, or cilium, of extreme tenuity. Behind this, from the concave side of the incurvation, proceeded another long cilium, so delicate as to be discernible only by the use of the highest powers and careful management of the light. In the centre of the pear-shaped body a clear round space could occasionally be discerned, but not always; and careful watching showed that this clear vacuity appeared gradually, and then shut up and disappeared suddenly, at regular intervals. Such a structure is of common occurrence among the lowest plants and animals, and is known as a contractile vacuole.
The little creature thus described sometimes propelled itself with great activity, with a curious rolling motion, by the lashing of the front cilium, while the second cilium trailed behind; sometimes it anchored itself by the hinder cilium and was spun round by the working of the other, its motions resembling those of an anchor buoy in a heavy sea. Sometimes, when two were in full career towards one another, each would appear dexterously to get out of the other’s way; sometimes a crowd would assemble and jostle one another, with as much semblance of individual effort as a spectator on the Grands Mulets might observe with a telescope among the specks representing men in the valley of Chamounix.
The spectacle, though always surprising, was not new to me. So my reply to the question put to me was, that these organisms were what biologists call Monads, and though they might be animals, it was also possible that they might, like the Bacteria, be plants. My friend received my verdict with an expression which showed a sad want of respect for authority. He would as soon believe that a sheep was a plant. Naturally piqued by this want of faith, I have thought a good deal over the matter; and as I still rest in the lame conclusion I originally expressed, and must even now confess that I cannot certainly say whether this creature is an animal or a plant, I think it may be well to state the grounds of my hesitation at length. But, in the first place, in order that I may conveniently distinguish this “Monad” from the multitude of other things which go by the same designation, I must give it a name of its own. I think (though, for reasons which need not be stated at present, I am not quite sure) that it is identical with the species Monas lens, as defined by the eminent French microscopist Dujardin, though his magnifying power was probably insufficient to enable him to see that it is curiously like a much larger form of monad which he has named Heteromita. I shall, therefore, call it not Monas, but Heteromita lens.
I have been unable to devote to my Heteromita the prolonged study needful to work out its whole history, which would involve weeks, or it may be months, of unremitting attention. But I the less regret this circumstance, as some remarkable observations recently published by Messrs. Dallinger and Drysdale34 on certain Monads, relate, in part, to a form so similar to my Heteromita lens, that the history of the one may be used to illustrate that of the other. These most patient and painstaking observers, who employed the highest attainable powers of the microscope and, relieving one another, kept watch day and night over the same individual monads, have been enabled to trace out the whole history of their Heteromita; which they found in infusions of the heads of fishes of the Cod tribe.
Of the four monads described and figured by these investigators, one, as I have said, very closely resembles Heteromita lens in every particular, except that it has a separately distinguishable central particle or “nucleus,” which is not certainly to be made out in Heteromita lens; and that nothing is said by Messrs. Dallinger and Drysdale of the existence of a contractile vacuole in this monad, though they describe it in another.
Their Heteromita, however, multiplied rapidly by fission. Sometimes a transverse constriction appeared; the hinder half developed a new cilium, and the hinder cilium gradually split from its base to its free end, until it was divided into two; a process which, considering the fact that this fine filament cannot be much more than 1/100000 of an inch in diameter, is wonderful enough. The constriction of the body extended inwards until the two portions were united by a narrow isthmus; finally, they separated and each swam away by itself, a complete Heteromita, provided with its two cilia. Sometimes the constriction took a longitudinal direction, with the same ultimate result. In each case the process occupied not more than six or seven minutes. At this rate, a single Heteromita would give rise to a thousand like itself in the course of an hour, to about a million in two hours, and to a number greater than the generally assumed number of human beings now living in the world in three hours; or, if we give each Heteromita an hour’s enjoyment of individual existence, the same result will be obtained in about a day. The apparent suddenness of the appearance of multitudes of such organisms as these, in any nutritive fluid to which one obtains access, is thus easily explained.
During these processes of multiplication by fission, the Heteromita remains active; but sometimes another mode of fission occurs. The body becomes rounded and quiescent, or nearly so; and, while in this resting state, divides into two portions, each of which is rapidly converted into an active Heteromita.
A still more remarkable phenomenon is that kind of multiplication which is preceded by the union of two monads, by a process which is termed conjugation. Two active Heteromitæ become applied to one another, and then slowly and gradually coalesce into one body. The two nuclei run into one; and the mass resulting from the conjugation of the two Heteromitæ, thus fused together, has a triangular form. The two pairs of cilia are to be seen, for some time, at two of the angles, which answer to the small ends of the conjoined monads; but they ultimately vanish, and the twin organism, in which all visible traces of organisation have disappeared, falls into a state of rest. Sudden wave-like movements of its substance next occur; and, in a short time, the apices of the triangular mass burst, and give exit to a dense yellowish, glairy fluid, filled with minute granules. This process, which, it will be observed, involves the actual confluence and mixture of the substance of two distinct organisms, is effected in the space of about two hours.
The authors whom I quote say that they “cannot express” the excessive minuteness of the granules in question, and they estimate their diameter at less than 1/200000 of an inch. Under the highest powers of the microscope at present applicable such specks are hardly discernible. Nevertheless, particles of this size are massive when compared to physical molecules; whence there is no reason to doubt that each, small as it is, may have a molecular structure sufficiently complex to give rise to the phenomena of life. And, as a matter of fact, by patient watching of the place at which these infinitesimal living particles were discharged, our observers assured themselves of their growth and development into new monads. These, in about four hours from their being set free, had attained a sixth of the length of the parent, with the characteristic cilia, though at first they were quite motionless; and, in four hours more, they had attained the dimensions and exhibited all the activity of the adult. These inconceivably minute particles are therefore the germs of the Heteromita; and from the dimensions of these germs it is easily shown that the body formed by conjugation may, at a low estimate, have given exit to thirty thousand of them; a result of a matrimonial process whereby the contracting parties, without a metaphor, “become one flesh,” enough to make a Malthusian despair of the future of the Universe.
I am not aware that the investigators from whom I have borrowed this history have endeavoured to ascertain whether their monads take solid nutriment or not; so that though they help us very much to fill up the blanks in the history of my Heteromita, their observations throw no light on the problem we are trying to solve—Is it an animal or is it a plant?
Undoubtedly it is possible to bring forward very strong arguments in favour of regarding Heteromita as a plant.
For example, there is a Fungus, an obscure and almost microscopic mould, termed Peronospora infestans. Like many other Fungi, the Peronosporæ are parasitic upon other plants; and this particular Peronospora happens to have attained much notoriety and political importance, in a way not without a parallel in the career of notorious politicians, namely, by reason of the frightful mischief it has done to mankind. For it is this Fungus which is the cause of the potato disease; and, therefore, Peronospora infestans (doubtless of exclusively Saxon origin, though not accurately known to be so) brought about the Irish famine. The plants afflicted with the malady are found to be infested by a mould, consisting of fine tubular filaments, termed hyphæ, which burrow through the substance of the potato plant, and appropriate to themselves the substance of their host; while, at the same time, directly or indirectly, they set up chemical changes by which even its woody framework becomes blackened, sodden, and withered.
In structure, however, the Peronospora is as much a mould as the common Penicillium; and just as the Penicillium multiplies by the breaking up of its hyphæ into separate rounded bodies, the spores; so, in the Peronospora, certain of the hyphæ grow out into the air through the interstices of the superficial cells of the potato plant, and develop spores. Each of these hyphæ usually gives off several branches. The ends of the branches dilate and become closed sacs, which eventually drop off as spores. The spores falling on some part of the same potato plant, or carried by the wind to another, may at once germinate, throwing out tubular prolongations which become hyphæ, and burrow into the substance of the plant attacked. But, more commonly, the contents of the spore divide into six or eight separate portions. The coat of the spore gives way, and each portion then emerges as an independent organism, which has the shape of a bean, rather narrower at one end than the other, convex on one side, and depressed or concave on the opposite. From the depression, two long and delicate cilia proceed, one shorter than the other, and directed forwards. Close to the origin of these cilia, in the substance of the body, is a regularly pulsating, contractile vacuole. The shorter cilium vibrates actively, and effects the locomotion of the organism, while the other trails behind; the whole body rolling on its axis with its pointed end forwards.
The eminent botanist, De Bary, who was not thinking of our problem, tells us, in describing the movements of these “Zoospores,” that, as they swim about, “Foreign bodies are carefully avoided, and the whole movement has a deceptive likeness to the voluntary changes of place which are observed in microscopic animals.”
After swarming about in this way in the moisture on the surface of a leaf or stem (which, film though it may be, is an ocean to such a fish) for half an hour, more or less, the movement of the zoospore becomes slower, and is limited to a slow turning upon its axis, without change of place. It then becomes quite quiet, the cilia disappear, it assumes a spherical form, and surrounds itself with a distinct, though delicate, membranous coat. A protuberance then grows out from one side of the sphere, and rapidly increasing in length, assumes the character of a hypha. The latter penetrates into the substance of the potato plant, either by entering a stomate, or by boring through the wall of an epidermic cell, and ramifies, as a mycelium, in the substance of the plant, destroying the tissues with which it comes in contact. As these processes of multiplication take place very rapidly, millions of spores are soon set free from a single infested plant; and, from their minuteness, they are readily transported by the gentlest breeze. Since, again, the zoospores set free from each spore, in virtue of their powers of locomotion, swiftly disperse themselves over the surface, it is no wonder that the infection, once started, soon spreads from field to field, and extends its ravages over a whole country.
However, it does not enter into my present plan to treat of the potato disease, instructively as its history bears upon that of other epidemics; and I have selected the case of the Peronospora simply because it affords an example of an organism, which, in one stage of its existence, is truly a “Monad,” indistinguishable by any important character from our Heteromita, and extraordinarily like it in some respects. And yet this “Monad” can be traced, step by step, through the series of metamorphoses which I have described, until it assumes the features of an organism, which is as much a plant as is an oak or an elm.
Moreover, it would be possible to pursue the analogy farther. Under certain circumstances, a process of conjugation takes place in the Peronospora. Two separate portions of its protoplasm become fused together, surround themselves with a thick coat, and give rise to a sort of vegetable egg called an oospore. After a period of rest, the contents of the oospore break up into a number of zoospores like those already described, each of which, after a period of activity, germinates in the ordinary way. This process obviously corresponds with the conjugation and subsequent setting free of germs in the Heteromita.
But it may be said that the Peronospora is, after all, a questionable sort of plant; that it seems to be wanting in the manufacturing power, selected as the main distinctive character of vegetable life; or, at any rate, that there is no proof that it does not get its protein matter ready made from the potato plant.
Let us, therefore, take a case which is not open to these objections.
There are some small plants known to botanists as members of the genus Coleochæte, which, without being truly parasitic, grow upon certain water-weeds, as lichens grow upon trees. The little plant has the form of an elegant green star, the branching arms of which are divided into cells. Its greenness is due to its chlorophyll, and it undoubtedly has the manufacturing power in full degree, decomposing carbonic acid and setting oxygen free, under the influence of sunlight. But the protoplasmic contents of some of the cells of which the plant is made up occasionally divide, by a method similar to that which effects the division of the contents of the Peronospora spore; and the severed portions are then set free as active monad-like zoospores. Each is oval and is provided at one extremity with two long active cilia. Propelled by these, it swims about for a longer or shorter time, but at length comes to a state of rest and gradually grows into a Coleochæte. Moreover, as in the Peronospora, conjugation may take place and result in an oospore; the contents of which divide and are set free as monadiform germs.
If the whole history of the zoospores of Peronospora, and of Coleochæte were unknown, they would undoubtedly be classed among “Monads” with the same right as Heteromita; why then may not Heteromita be a plant, even though the cycle of forms through which it passes shows no terms quite so complex as those which occur in Peronospora and Coleochæte? And, in fact, there are some green organisms, in every respect characteristically plants, such as Chlamydomonas, and the common Volvox, or so-called “Globe animalcule,” which run through a cycle of forms of just the same simple character as those of Heteromita.
The name of Chlamydomonas is applied to certain microscopic green bodies, each of which consists of a protoplasmic central substance invested by a structureless sac. The latter contains cellulose, as in ordinary plants; and the chlorophyll which gives the green colour enables the Chlamydomonas to decompose carbonic acid and fix carbon as they do. Two long cilia protrude through the cell-wall, and effect the rapid locomotion of this “monad,” which, in all respects except its mobility, is characteristically a plant. Under ordinary circumstances, the Chlamydomonas multiplies by simple fission, each splitting into two or into four parts, which separate and become independent organisms. Sometimes, however, the Chlamydomonas divides into eight parts, each of which is provided with four instead of two cilia. These “zoospores” conjugate in pairs, and give rise to quiescent bodies, which multiply by division, and eventually pass into the active state.
Thus, so far as outward form and the general character of the cycle of modifications, through which the organism passes in the course of its life, are concerned, the resemblance between Chlamydomonas and Heteromita is of the closest description. And on the face of the matter there is no ground for refusing to admit that Heteromita may be related to Chlamydomonas, as the colourless fungus is to the green alga. Volvox may be compared to a hollow sphere, the wall of which is made up of coherent Chlamydomonads; and which progresses with a rotating motion effected by the paddling of the multitudinous pairs of cilia which project from its surface. Each Volvoxmonad, moreover, possesses a red pigment spot, like the simplest form of eye known among animals. The methods of fissive multiplication and of conjugation observed in the monads of this locomotive globe are essentially similar to those observed in Chlamydomonas; and, though a hard battle has been fought over it, Volvox is now finally surrendered to the Botanists.
Thus there is really no reason why Heteromita may not be a plant; and this conclusion would be very satisfactory, if it were not equally easy to show that there is really no reason why it should not be an animal. For there are numerous organisms presenting the closest resemblance to Heteromita, and, like it, grouped under the general name of “Monads,” which, nevertheless, can be observed to take in solid nutriment, and which, therefore, have a virtual, if not an actual, mouth and digestive cavity, and thus come under Cuvier’s definition of an animal. Numerous forms of such animals have been described by Ehrenberg, Dujardin, H. James Clark, and other writers on the Infusoria. Indeed, in another infusion of hay in which my Heteromita lens occurred, there were innumerable infusorial animalcules belonging to the well-known species Colpoda cucullus.35
Full-sized specimens of this animalcule attain a length of between 1/300 or 1/400 of an inch, so that it may have ten times the length and a thousand times the mass of a Heteromita. In shape, it is not altogether unlike Heteromita. The small end, however, is not produced into one long cilium, but the general surface of the body is covered with small actively vibrating ciliary organs, which are only longest at the small end. At the point which answers to that from which the two cilia arise in Heteromita, there is a conical depression, the mouth; and, in young specimens, a tapering filament, which reminds one of the posterior cilium of Heteromita, projects from this region.
The body consists of a soft granular protoplasmic substance, the middle of which is occupied by a large oval mass called the “nucleus;” while, at its hinder end, is a “contractile vacuole,” conspicuous by its regular rhythmic appearances and disappearances. Obviously, although the Colpoda is not a monad, it differs from one only in subordinate details. Moreover, under certain conditions, it becomes quiescent, incloses itself in a delicate case or cyst, and then divides into two, four, or more portions, which are eventually set free and swim about as active Colpodæ.
But this creature is an unmistakable animal, and full-sized Colpodæ may be fed as easily as one feeds chickens. It is only needful to diffuse very finely ground carmine through the water in which they live, and, in a very short time, the bodies of the Colpodæ are stuffed with the deeply-coloured granules of the pigment.
And if this were not sufficient evidence of the animality of Colpoda, there comes the fact that it is even more similar to another well-known animalcule, Paramæcium, than it is to a monad. But Paramæcium is so huge a creature compared with those hitherto discussed—it reaches 1/120 of an inch or more in length—that there is no difficulty in making out its organisation in detail; and in proving that it is not only an animal, but that it is an animal which possesses a somewhat complicated organisation. For example, the surface layer of its body is different in structure from the deeper parts. There are two contractile vacuoles, from each of which radiates a system of vessel-like canals; and not only is there a conical depression continuous with a tube, which serve as mouth and gullet, but the food ingested takes a definite course, and refuse is rejected from a definite region. Nothing is easier than to feed these animals, and to watch the particles of indigo or carmine accumulate at the lower end of the gullet. From this they gradually project, surrounded by a ball of water, which at length passes with a jerk, oddly simulating a gulp, into the pulpy central substance of the body, there to circulate up one side and down the other, until its contents are digested and assimilated. Nevertheless, this complex animal multiplies by division, as the monad does, and, like the monad, undergoes conjugation. It stands in the same relation to Heteromita on the animal side, as Coleochæte does on the plant side. Start from either, and such an insensible series of gradations leads to the monad that it is impossible to say at any stage of the progress—here the line between the animal and the plant must be drawn.
There is reason to think that certain organisms which pass through a monad stage of existence, such as the Myxomycetes, are, at one time of their lives, dependent upon external sources for their protein matter, or are animals; and, at another period, manufacture it, or are plants. And seeing that the whole progress of modern investigation is in favour of the doctrine of continuity, it is a fair and probable speculation—though only a speculation—that, as there are some plants which can manufacture protein out of such apparently intractable mineral matters as carbonic acid, water, nitrate of ammonia, metallic and earthy salts; while others need to be supplied with their carbon and nitrogen in the somewhat less raw form of tartrate of ammonia and allied compounds; so there may be yet others, as is possibly the case with the true parasitic plants, which can only manage to put together materials still better prepared—still more nearly approximated to protein—until we arrive at such organisms as the Psorospermiæ and the Panhistophyton, which are as much animal as vegetable in structure, but are animal in their dependence on other organisms for their food.
The singular circumstance observed by Meyer, that the Torula of yeast, though an indubitable plant, still flourishes most vigorously when supplied with the complex nitrogenous substance, pepsin; the probability that the Peronospora is nourished directly by the protoplasm of the potato-plant; and the wonderful facts which have recently been brought to light respecting insectivorous plants, all favour this view; and tend to the conclusion that the difference between animal and plant is one of degree rather than of kind; and that the problem whether, in a given case, an organism is an animal or a plant, may be essentially insoluble.
In all the commentaries upon the “Historia Animalium” which I have met with, Aristotle’s express and repeated statement, that the heart of man and the largest animals contains only three cavities, is noted as a remarkable error. Even Cuvier, who had a great advantage over most of the commentators in his familiarity with the subject of Aristotle’s description, and whose habitual caution and moderation seem to desert him when the opportunity of panegyrising the philosopher presents itself, is betrayed into something like a sneer on this topic.
“Du reste il n’attribue à cet organe que trois cavités, erreur qui prouve au moins qu’il en avait regardé la structure.”36
To which remark, what follows will, I think, justify the reply, that it “prouve au moins” that Cuvier had not given ordinary attention, to say nothing of the careful study which they deserve, to sundry passages in the first and the third books of the “Historia” which I proceed to lay before the reader.
For convenience of reference these passages are marked A, B, C, etc.37
Book i. 17.—(A) “The heart has three cavities, it lies above the lung on the division of the windpipe, and has a fatty and thick membrane where it is united with the great vein and the aorta. It lies upon the aorta, with its point down the chest, in all animals that have a chest. In all, alike in those that have a chest and in those that have none, the foremost part of it is the apex. This is often overlooked through the turning upside down of the dissection. The rounded end of the heart is uppermost, the pointed end of it is largely fleshy and thick, and in its cavities there are tendons. In other animals which have a chest the heart lies in the middle of the chest; in men, more to the left side, between the nipples, a little inclined to the left nipple in the upper part of the chest. The heart is not large, and its general form is not elongated but rounded, except that the apex is produced into a point.
(B) “It has, as already stated, three cavities, the largest of them is on the right, the smallest on the left, the middle-sized one in the middle; they have all, also the two small ones, passages (τετρημένας) towards the lung, very evidently as respects one of the cavities. In the region of the union [with the great vein and the aorta] the largest cavity is connected with the largest vein (near which is the mesentery); the middle cavity with the aorta.
(C) “Canals (πόροι) from the heart pass to the lung and divide in the same fashion as the windpipe does, closely accompanying those from the windpipe through the whole lung. The canals from the heart are uppermost.
(D) “No canal is common [to the branches of the windpipe and those of the vein] (οὐδεὶς δ’ ἐστὶ κοινὸς πόρος) but through those parts of them which are in contact (τὴν σύναψιν) the air passes in and they [the πόροι] carry it to the heart.
(E) “One of the canals leads to the right cavity, the other to the left.
(F) “Of all the viscera, the heart alone contains blood [in itself]. The lung contains blood, not in itself but in the veins, the heart in itself; for in each of the cavities there is blood; the thinnest is in the middle cavity.”
Book iii. 3.—(G) “Two veins lie in the thorax alongside the spine, on its inner face; the larger more forwards, the smaller behind; the larger more to the right, the smaller, which some call aorta (on account of the tendinous part of it seen in dead bodies), to the left. These take their origin from the heart; they pass entire, preserving the nature of veins, through the other viscera that they reach; while the heart is rather a part of them, and more especially of the anterior and larger one, which is continued into veins above and below, while between these is the heart.
(H) “All hearts contain cavities, but, in those of very small animals, the largest [cavity] is hardly visible, those of middling size have another, and the biggest all three.
(I) “The point of the heart is directed forwards, as was mentioned at first; the largest cavity to the right and upper side of it, the smallest to the left, and the middle-sized one between these; both of these are much smaller than the largest.
(K) “They are all connected by passages (συντέτρηνται) with the lung, but, on account of the smallness of the canals, this is obscure except in one.
(L) “The great vein proceeds from the largest cavity which lies upwards and to the right; next through the hollow middle part (διὰ τοῦ κοίλου τοῦ μέσου) it becomes vein again, this cavity being a part of the vein in which the blood stagnates.
(M) “The aorta [proceeds from] the middle [cavity], but not in the same way, for it is connected [with the middle cavity] by a much more narrow tube (σύριγγα).
(N) “The [great] vein extends through the heart, towards the aorta from the heart.
(O) “The great vein is membranous like skin, the aorta narrower than it and very tendinous, and as it extends towards the head and the lower parts it becomes narrow and altogether tendinous.
(P) “In the first place, a part of the great vein extends upwards from the heart towards the lung and the attachment of the aorta, the vein being large and undivided. It divides into two parts, the one to the lung, the other to the spine and the lowest vertebra of the neck.
(Q) “The vein which extends to the lung first divides into two parts for the two halves of it and then extends alongside each tube (σύριγγα) and each passage (τρῆμα), the larger beside the larger and the smaller beside the smaller, so that no part [of the lung] can be found from which a passage (τρῆμα) and a vein are absent. The terminations are invisible on account of their minuteness, but the whole lung appears full of blood. The canals from the vein lie above the tubes given off from the windpipe.”
The key to the whole of the foregoing description of the heart lies in the passages (G) and (L). They prove that Aristotle, like Galen, five hundred years afterwards, and like the great majority of the old Greek anatomists, did not reckon what we call the right auricle as a constituent of the heart at all, but as a hollow part, or dilatation, of the “great vein.” Aristotle is careful to state that his observations were conducted on suffocated animals; and if any one will lay open the thorax of a dog or a rabbit, which has been killed with chloroform, in such a manner as to avoid wounding any important vessel, he will at once see why Aristotle adopted this view.
For, as the subjoined figure (p. 185) shows, the vena cava inferior (b), the right auricle (R.a.), and the vena cava superior and innominate vein (V.I.) distended with blood seem to form one continuous column, to which the heart is attached as a sort of appendage. This column is, as Aristotle says, vein above (a) and vein below (b), the upper and the lower divisions being connected διὰ τοῦ κοίλου τοῦ μέσου—or by means of the intervening cavity or chamber (R.a.)—which is that which we call the right auricle.
A dog having been killed by chloroform, enough of the right wall of the thorax was removed, without any notable bleeding, to expose the thoracic viscera. A carefully measured outline sketch of the parts in situ was then made, and on dissection, twenty-four hours afterwards, the necessary anatomical details were added. The woodcut is a faithfully reduced copy of the drawing thus constructed; and it represents the relations of the heart and great vessels as Aristotle saw them in a suffocated animal.
All but the inner lobe of the right lung has been removed; as well as the right half of the pericardium and the right walls of the right auricle and ventricle. It must be remembered that the thin transparent pericardial membrane appears nothing like so distinct in nature.
a.b., Aristotle’s “great vein”; V.I., right vena innominata and vena cava superior; b, the inferior vena cava; R.a., the “hollow middle” part of the great vein or the right auricle; R.v′, the prolongation of the cavity of the right ventricle R.v towards the pulmonary artery; tr, one of the tricuspid valves; Pc, the pericardium; I.sv, superior intercostal vein; Az, vena azygos; P.A., right pulmonary artery; Br, right bronchus; L, inner lobe of the right lung; Œ, œsophagus; Ao, descending aorta; H, liver, in section, with hepatic vein, vena portæ, and gall-bladder, gb, separated by the diaphragm, also seen in section, from the thoracic cavity.
But when, from the four cavities of the heart recognised by us moderns, one is excluded, there remain three—which is just what Aristotle says. The solution of the difficulty is, in fact, as absurdly simple as that presented by the egg of Columbus; and any error there may be, is not to be put down to Aristotle, but to that inability to comprehend that the same fact may be accurately described in different ways, which is the special characteristic of the commentatorial mind. That the three cavities mentioned by Aristotle are just those which remain if the right auricle is omitted, is plain enough from what is said in (B), (C), (E), (I), and (L). For, in a suffocated animal, the “right cavity” which is directly connected with the great vein, and is obviously the right ventricle, being distended with blood, will look much larger than the middle cavity, which, since it gives rise to the aorta, can only be the left ventricle. And this, again, will appear larger than the thin and collapsed left auricle, which must be Aristotle’s left cavity, inasmuch as this cavity is said to be connected by πόροι with the lung. The reason why Aristotle considered the left auricle to be a part of the heart, while he merged the right auricle in the great vein, is, obviously, the small relative size of the venous trunks and their sharper demarcation from the auricle. Galen, however, perhaps more consistently, regarded the left auricle also as a mere part of the “arteria venosa.” The canal which leads from the right cavity of the heart to the lung (or, as Aristotle puts it (E), from the lung to the heart) is, without doubt, the pulmonary artery. But it may be said that, in this case, Aristotle contradicts himself, inasmuch as in (P) and (Q) a vessel, which is obviously the pulmonary artery, is described as a branch of the great vein. However, this difficulty also disappears, if we reflect that, in Aristotle’s way of looking at the matter, the line of demarcation between the great vein and the heart coincides with the right auriculo-ventricular aperture; and that, inasmuch as the conical prolongation of the right ventricle which leads to the pulmonary artery (R.v′ in the Figure), lies close in front of the auricle, its base may very easily (as the figure shows) be regarded as part of the general opening of the great vein into the right ventricle. In fact, it is clear that Aristotle, having failed to notice the valves of the heart, did not distinguish the part of the right ventricle from which the pulmonary artery arises (R.v′) from the proper trunk of the artery on the one hand, and from the right auricle (R.a) on the other. Thus the root, as we may call it, of the pulmonary artery and the right auricle, taken together, are spoken of as the “part of the great vein which extends upwards” (P); and, as the vena azygos (Az) was one branch of this, so the “vein to the lung” was regarded as another branch of it. But the latter branch, being given off close to the connection of the great vein with the ventricle, was also counted as one of the two πόροι by which the “heart” (that is to say the right ventricle, the left ventricle, and the left auricle of our nomenclature) communicates with the lung.
The only other difficulty that I observe is connected with (K). If Aristotle intended by this to affirm that the middle cavity (the left ventricle), like the other two, is directly connected with the lung by a πόρος, he would be in error. But he has excluded this interpretation of his words by (E), in which the number and relations of the canals, the existence of which he admits, are distinctly defined. I can only imagine then, that, so far as this passage applies to the left ventricle, it merely refers to the indirect communication of that cavity with the vessels of the lungs, through the left auricle.
On this evidence I submit that there is no escape from the conclusion that, instead of having committed a gross blunder, Aristotle has given a description of the heart which, so far as it goes, is remarkably accurate. He is in error only in regard to the differences which he imagines to exist between large and small hearts (H).
Cuvier (who has been followed by other commentators) ascribes another error to Aristotle:—
“Aristote suppose que la trachée-artère se prolonge jusqu’au cœur, et semble croire, en conséquence, que l’air y pénétre (l. c. p. 152).”
Upon what foundation Cuvier rested the first of these two assertions, I am at a loss to divine. As a matter of fact, it will appear from the following excerpts that Aristotle gives an account of the structure of the lungs which is almost as good as that of the heart, and that it contains nothing about any prolongation of the windpipe to the heart.
“Within the neck lie what is called the œsophagus (so named on account of its length and its narrowness) and the windpipe (ἀρτηρία). The position of the windpipe in all animals that have one, is in front of the œsophagus. All animals which possess a lung have a windpipe. The windpipe is of a cartilaginous nature and is exsanguine, but is surrounded by many little veins....
“It goes downwards towards the middle of the lung, and then divides for each of the halves of the lung. In all animals that possess one, the lung is divided into two parts; but, in those which bring forth their young alive, the separation is not equally well marked, least of all in man.
“In oviparous animals, such as birds, and in quadrupeds which are oviparous, the one half of the lung is widely separated from the other; so that it appears as if they had two lungs. And from being single, the windpipe becomes (divided into) two, which extend to each half of the lung. It is fastened to the great vein, and to what is called the aorta. When the windpipe is blown up, the air passes into the hollow parts of the lung. In these, are cartilaginous tubes (διαφύσεις) which unite at an angle; from the tubes passages (τρήματα) traverse the whole of the lung; they are continually given off, the smaller from the larger.” (Book i. 16.)
That Aristotle should speak of the lung as a single organ divided into two halves, and should say that the division is least marked in man, is puzzling at first; but the statement becomes intelligible, if we reflect upon the close union of the bronchi, the pulmonary vessels and the mediastinal walls of the pleuræ, in mammals;38 and it is quite true that the lungs are much more obviously distinct from one another in birds.
Aubert and Wimmer translate the last paragraph of the passage just cited as follows:—
“Diese haben aber knorpelige Scheidewände, welche unter spitzen Winkeln zusammentreten, und aus ihnen führen Oeffnungen durch die ganze Lunge, indem sie sich in immer kleineren verzweigen.”
But I cannot think that by διαφύσεις and τρήματα, in this passage, Aristotle meant either “partitions” or openings in the ordinary sense of the latter word. For, in Book iii. Cap. 3, in describing the distribution of the “vein which goes to the lung” (the pulmonary artery), he says that it
“extends alongside each tube (σύριγγα) and each passage (τρῆμα), the larger beside the larger, and the smaller beside the smaller; so that no part (of the lung) can be found from which a passage (τρῆμα) and a vein are absent.”
Moreover, in Book i. 17, he says—
“Canals (πόροι) from the heart pass to the lung and divide in the same fashion as the windpipe does, closely accompanying those from the windpipe through the whole lung.”
And again in Book i. 17—