Chapter II: Part 2

Some approach to evolutionary doctrine is also foreshadowed by Aristotle in his theories of the development of the individual. This is obscured, however, by his peculiar view of the nature of procreation. On this topic his general conclusion is that the material substance of the embryo is contributed by the female, but that this is mere passive formable material, almost as though it were the soil in which the embryo grows. The male by giving the principle of life, the soul, contributes the essential generative agency. But this _soul_ is not material and it is, therefore, not theoretically necessary for anything material to pass from male to female. The material which does in fact so pass with the seed of the male is an accident, not an essential, for the essential contribution of the male is not matter but _form_ and _principle_. The female provides the _material_, the male the _soul_, the _form_, the _principle_, that which makes life. Aristotle was thus prepared to accept instances of fertilization without material contact.

‘The female does not contribute semen to generation but does contribute something ... for there must needs be that which generates and that from which it generates.... If, then, the male stands for the effective and active, and the female, considered as female, for the passive, it follows that what the female would contribute to the semen of the male would not be semen but material for the semen to work upon....

‘How is it that the male contributes to generation, and how is it that the semen from the male is the cause of the offspring? Does [the semen] exist in the body of the embryo as a part of it from the first, mingling with the material which comes from the female? Or does the semen contribute nothing to the material body of the embryo but only to the power and movement in it?... The latter alternative appears to be the right one both _a priori_ and in view of the facts.’[30]

[30] _De generatione animalium_, i. 21; 729ᵅ 21.

This discussion leads to the question of the natural process of generation itself. It is a topic that we have seen discussed by an earlier writer who had set forth a sort of doctrine of pangenesis (see p. 14). His view Aristotle declines to share. ‘We must’, he says, ‘say the opposite of what the ancients said. For whereas they said that semen is that which _comes from all_ the body, we shall say that it is that whose nature is to _go to all_ of it, and what they thought a waste-product seems rather to be a secretion.’ According to Aristotle semen is derived from the same nutritive material in the blood vessels that is distributed to the rest of the body. The semen, however, is strained or secreted off from this nutritive material—as being its most essential and representative portion—before the distribution actually takes place.[31] But why, it may be asked, if the semen does not come from the various parts of the body, is it yet able to reproduce those various parts? The answer, on the Aristotelian view, seems to be that the semen contains special and peculiar fractions of the nutritive fluid which have been so modified and adapted that, if not secreted off as semen, they would be distributed to the different parts of the body to nourish each of these various parts. These substances have been elaborated by the _soul_ or vital principle in a manner that is specifically suited for each organ, hand, liver, face, heart, &c., and from each of these specific substances a specific essence is separated off into the semen corresponding to hand, liver, face, heart, &c., of the offspring.

The next question that arises is the mechanism by which the offspring come to resemble their parents. The mechanism in the case of inheritance from the father is comprehensible when we consider the origin and nature of the semen, but the inheritance from the mother requires further explanation. The view of Aristotle is based upon the nature of the catamenia and their disappearance during gestation. ‘The catamenia’, in his view, ‘are a secretion as the semen is.’[32] The female contributes the material by which the embryo grows and she does this through the catamenia which are suspended during gestation for this very purpose. The matter is thus summed up by Aristotle.

[31] _De generations animalium_, i. 18; 725ᵅ 22.

[32] _De generatione animalium_, i. 19; 727ᵅ 31.

‘The male does not emit semen at all in some animals, and where he does, this is no part of the resulting embryo; just so no material part comes from the carpenter to the material, i.e. to the wood in which he works, nor does any part of the carpenter’s art exist within what he makes, but the shape and the form are imparted from him to the material by means of the motion he sets up. It is his hands that move his tools, his tools that move the material; it is his knowledge of his art, and his _soul_, in which is the form, that move his hands or any other part of him with a motion of some definite kind, a motion varying with the varying nature of the object made. In like manner, in the male of those animals which emit semen, Nature uses the semen as a tool and as possessing motion in actuality, just as tools are used in the products of any art, for in them lies in a certain sense the motion of the art.’[33]

‘For the same reason the development of the embryo takes place in the female; neither the male himself nor the female emits semen into the female, but the female receives within herself the share contributed by both, because in the female is the material from which is made the resulting product. Not only must the mass of material from which the embryo is in the first instance formed exist there, but further material must constantly be added so that the embryo may increase in size. Therefore the birth must take place in the female. For the carpenter must keep in close connexion with his timber and the potter with his clay, and generally all workmanship and the ultimate movement imparted to matter must be connected with the material concerned, as, for instance, architecture is _in_ the buildings it makes.’[34]

[33] _De generatione animalium_, i. 22; 730ᵇ 10.

[34] _De generatione animalium_, i. 22; 730ᵅ 34.

The problem of the nature of generation is one in which Aristotle never ceased to take an interest, and among the methods by which he sought to solve it was embryological investigation. In his ideas on the methods of reproduction we must seek also the main bases of such classification of animals as he exhibits. His most important embryological researches were made upon the chick. He asserts that the first signs of development are noticeable on the third day, the heart being visible as a palpitating blood-spot whence, as it develops, two meandering blood vessels extend to the surrounding tunics.

‘Generation from the egg’, he says, ‘proceeds in an identical manner with all birds.... With the common hen after three days and nights there is the first indication of the embryo.... The heart appears like a speck of blood in the white of the egg. This point beats and moves as though endowed with life, and from it two vessels with blood in them trend in a convoluted course ... and a membrane carrying bloody fibres now envelops the yolk, leading off from the vessels.’[35]

Aristotle lays considerable stress on the early appearance of the heart in the embryo. Corresponding to the general gradational view that he had formed of Nature, he held that the most primitive and fundamentally important organs make their appearance before the others. Among the organs all give place to the heart, which he considered ‘the first to live and the last to die’.[36]

A little later he observed that the body had become distinguishable, and was at first very small and white.

[35] _Historia animalium_, vi. 3; 561ᵅ 4.

[36] _Cor primum movens ultimum moriens._ This famous sentence is the sense though not the phrasing of _De generatione animalium_, ii. 1 and 4.

‘The head is clearly distinguished and in it the eyes, swollen out to a great extent.... At the outset the under portion of the body appears insignificant in comparison with the upper portion....

‘When an egg is ten days old the chick and all its parts are distinctly visible. The head still is larger than the rest of the body and the eyes larger than the head. At this time also the larger internal organs are visible, as also the stomach and the arrangement of the viscera; and the vessels that seem to proceed from the heart are now close to the navel. From the navel there stretch a pair of vessels, one [vitelline vein] towards the membrane that envelops the yolk, and the other [allantoic vein] towards that membrane which envelops collectively the membrane wherein the chick lies, the membrane of the yolk and the intervening liquid.... About the twentieth day, if you open the egg and touch the chick, it moves inside and chirps; and it is already coming to be covered with down when, after the twentieth day, the chick begins to break the shell.’[37]

[37] _Historia animalium_, vi. 3; 561ᵅ 18.

Aristotle recognized a distinction in the mode of development of mammals from that of all other viviparous creatures. Having divided the apparently viviparous animals into two groups, one of which is truly and internally and the other only externally viviparous, he pointed out that in the mammalia, the group regarded by him as internally viviparous, the foetus is connected until birth with the wall of the mother’s womb by the navel-string. These animals, in his view, produce their young without the intervention of an ovum, the embryo being ‘living from the first’. Such non-mammals, on the other hand, as are viviparous are so in the external sense only, that is, the young which he considered to arise in this group from ova may indeed develop within the mother’s womb and be born alive, but they go through their development without organic connexion with the mother’s body, so that her womb acts but as a nursery or incubator for her eggs. It was indeed a sort of accident among the ovipara whether in any particular species the ovum went through its development inside or outside the mother’s body. ‘Some of the ovipara’, he says, ‘produce the egg in a perfect, others in an imperfect state, but it is perfected outside the body as has been stated of fish.’[38]

Yet though Aristotle regarded fish as an oviparous group, he knew also of kinds of fish that were externally viviparous. It is most interesting to observe, moreover, that he was acquainted with one particular instance among fish in which matters were less simple and in which the development bore an analogy to that of the mammalia, his true internal vivipara. ‘Some animals’, he says, ‘are viviparous, others oviparous, others vermiparous. Some are viviparous, such as man, the horse, the seal and all other animals that are hair-coated, and, of marine animals, the Cetaceans, as the dolphin, _and the so-called Selachia_.’[39]

Aristotle tells us elsewhere that a species of these Selachia which he calls _galeos_—a name still used for the dog-fish by Greek fishermen—‘has its eggs in betwixt the [two horns of the] womb; these eggs shift into each of the two horns of the womb and descend, and the young develop with the navel-string attached to the womb, so that, as the egg-substance gets used up, the embryo is sustained to all appearances just as in quadrupeds. The navel-string is ... attached as it were by a sucker, and also to the centre of the embryo in the place where the liver is situated.... Each embryo, as in the case of quadrupeds, is provided with a chorion and separate membranes.’[40]

[38] _De generatione animalium_, iii. 9; 758ᵅ 37.

[39] _Historia animalium_, i. 5; 489ᵅ 35.

[40] _Historia animalium_, vi. 10; 565ᵇ 2.

The remarkable anatomical relationship of the embryo of _Galeus_ (_Mustelus_) _laevis_ to its mother’s womb was little noticed by naturalists until the whole matter was taken up by Johannes Müller about 1840.[41] That great observer demonstrated the complete accuracy of Aristotle’s description and the justice of his comparison to and contrast with the mammalian mode of development.[42] The work of Johannes Müller at once had the effect of drawing the attention of naturalists to the importance and value of the Aristotelian biological observations.

[41] The history of this discovery is given by Charles Singer, _Studies in the History and Method of Science_, vol. ii, Oxford, 1921, pp. 32 ff.

[42] Johannes Müller, _Ueber den glatten Hai des Aristoteles_, Berlin, 1842.

Aristotle attempts to explain the viviparous character of the Selachians. His explanation has perhaps little meaning for the modern biologist, just as many of our scientific explanations will seem meaningless to our successors. But such explanations are often worth consideration not only as stages in the historical development of scientific thought, but also as illustrating the fact that while the ultimate object of science is a _description_ of nature, the immediate motive of the best scientific work is usually an _explanation_ of nature. Yet it is usually the descriptive, not the explanatory element that bears the test of time.

‘Birds and scaly reptiles’, says Aristotle, ‘because of their heat produce a perfect egg, but because of their dryness it is only an egg. The cartilaginous fishes have less heat than these but more moisture, so that they are intermediate, for they are both oviparous and viviparous within themselves, the former because they are cold, the latter because of their moisture; for moisture is vivifying, whereas dryness is farthest removed from what has life. Since they have neither feathers nor scales such as either reptiles or other fishes have, all of which are signs rather of a dry and earthy nature, the egg they produce is soft; for the earthy matter does not come to the surface in their eggs any more than in themselves. That is why they lay eggs in themselves, for if the egg were laid externally it would be destroyed, having no protection.’[43]

[43] _De generatione animalium_, ii. 1; 733ᵅ 6.

This explanation is based on Aristotle’s fundamental doctrine of the opposite _qualities_, heat, cold, wetness, and dryness, that are found combined in pairs in the four _elements_, earth, air, fire, and water. The theory was of the utmost importance for the whole subsequent development of science and was not displaced until quite modern times. It was not an original conception of Aristotle, for something resembling it had been set forth long before his time in figurative language by Empedocles (_c._ 500-_c._ 430 B. C.), as Aristotle himself tells us.[44] The same view had been foreshadowed by Pythagoras (_c._ 580-_c._ 490 B. C.) at an even earlier date and was perhaps of much greater antiquity. But Aristotle developed the doctrine and was the main channel for its conveyance to later ages, so that his name will always be associated with it. Matter in general and living matter in particular was held by him to be composed of these four essential so-called _elements_ (στοιχεῑ), each of which is in turn compounded from two of the primary _qualities_ (δυνάμεις) which Aristotle brought into relation with the elements. Thus earth was cold and dry, water cold and wet, air hot and wet, and fire hot and dry (Fig. 7b).

[44] _Metaphysics_, i. 4. _De generatione et corruptione_, ii. 1.

The theory of the elements and qualities is applicable to all matter and not specially to living things. The distinction between the living and not-living is to be sought not so much in its material constitution, but in the presence or absence of ‘soul’, and his teaching on that topic is to be found in his great work περὶ ψυχῆς, _On Soul_. He does not think of matter as organic or inorganic—that is a distinction of the seventeenth century physiologists—nor does he think of things as divided into animal, vegetable, and mineral—that is a distinction of the mediaeval alchemists,—but he thinks of things as either with soul or without soul (ἔμψυχα or ἄαψυχα).

His belief as to the relationship of this soul to material things is a difficult and complicated subject which would take us far beyond the topics included in biological writings to-day, but he tells us that ‘there is a class of existent things which we call substance, including under that term, firstly, matter, which in itself is not this nor that; secondly, shape or form, in virtue of which the term this or that is at once applied; thirdly, the whole made up of matter and form. Matter is identical with potentiality, form with actuality,’ the soul being, in living things, that which gives the form or actuality. ‘Of natural bodies’, he continues, ‘some possess life and some do not: where by life we mean the power of self-nourishment and of independent growth and decay’.[45] It should here be noted that in the Aristotelian sense the ovum is not at first a living thing, for in its earliest stage and before fertilization it does not possess soul even in its most elementary form.

[45] _De anima_, ii. 1, ii.

‘The term life is used in various senses, and, if life is present in but a single one of these senses, we speak of a thing as living. Thus there is intellect, sensation, motion from place to place and rest, the motion concerned with nutrition, and, further, [there are the processes of] decay and growth,’ all various meanings or at least exhibitions of some form of life. Hence even ‘plants are supposed to have life, for they have within themselves a faculty and principle whereby they grow and decay.... They grow and continue to live so long as they are capable of absorbing nutriment. This form of life can be separated from the others ... and plants have no other faculty of soul at all,’ but only this lowest vegetative soul. ‘It is then in virtue of this principle that all living things live, whether animals or plants. But it is sensation which primarily constitutes the animal. For, provided they have sensation, even those creatures that are devoid of movement and do not change their place are called animals.... As the nutritive faculty may exist without touch or any form of sensation, so also touch may exist apart from other senses.’[46] Apart from these two lower forms of soul, the _vegetative_ or nutritive and reproductive and the _animal_ or sensitive, stands the _rational_ or intellectual soul peculiar to man, a form of soul with which we shall here hardly concern ourselves.[47]

[46] _De anima_, ii. 2, ii; 413ᵅ 22.

[47] The question of Aristotle’s meaning in connexion with this topic, of primary importance for all thought, has a vast literature. An authoritative work is R. D. Hicks, _Aristotle, De anima_, Cambridge, 1907.

The possession of one or more of the three types of soul, vegetative, sensitive, and rational, provides in itself a basis for an elementary form of arrangement of living things in an ascending scale. We have already seen that Aristotle certainly describes something resembling a ‘Scala Naturae’ and that such a scheme can easily be drawn up from passages in his works. It may, however, be doubted whether his phraseology is capable of extension so as to include a true _classification_ of animals in any modern sense. It is true that he repeatedly divides animals into classes, _Sanguineous_ and _Non-sanguineous_, _Oviparous_ and _Viviparous_, _Terrestrial_ and _Aquatic_, &c., but his divisions are for the most part simply dichotomic. He certainly defines a few groups of animals as the Lophura (_Equidae_), the Cete (_Cetacea_), and the Selache (_Elasmobranchiae_ together with the _Lophiidae_) in a way that fairly corresponds to similar groups in later systems. In most cases, however, his definitions are not exact enough for modern needs, for the same animal may fall into more than one of his classes and widely different animals into the same class. Thus he invents a category _Carcharodonta_ for animals with sharp interlocking teeth and includes in it carnivores, reptiles, and fish; again, the horse kind must be included both among his _Anepallacta_ or animals having flat crowned teeth as well as among the _Amphodonta_ or animals with front teeth in both jaws. Such words as these are really terms of _description_, not of classification in the modern biological sense of that word.

There are, however, scattered through the biological works, certain terms which are applied to animal groups and organs and are defined in such a way as to suggest that they might ultimately have been developed for classificatory purposes. Thus his lowest group is the _species_. ‘The individuals comprised within a single _species_ (εîδος) ... are the real existences; but inasmuch as these individuals possess one common specific form, it will suffice to state the universal attributes of the species, that is, the attributes common to all its individuals, once and for all.’[48] This is surely not very far removed from the modern biological conception of a species.

[48] _De partibus animalium_, i. 4; 644ᵅ 22.

‘But as regards the larger groups—such as birds—which comprehend many species, there may be a question. For on the one hand it may be urged that as the ultimate species represent the real existences, it will be well, if practicable, to examine these ultimate species separately, just as we examine the species Man separately; to examine, that is, not the whole class Birds collectively, but the Ostrich, the Crane, and the other indivisible groups or species belonging to the class.

‘On the other hand, this course would involve repeated mention of the same attribute, as the same attribute is common to many species, and so far would be somewhat irrational and tedious. Perhaps, then, it will be best to treat generically the universal attributes of the groups that have a common nature and contain closely allied subordinate forms, whether they are groups recognized by a true instinct of mankind, such as Birds and Fishes, or groups not popularly known by a common appellation, but withal composed of closely allied subordinate groups; and only to deal individually with the attributes of a single species, when such species—man, for instance, and any other such, if such there be—stands apart from others, and does not constitute with them a larger natural group.

‘It is generally similarity in the shape of particular organs, or of the whole body, that has determined the formation of the larger groups. It is in virtue of such a similarity that Birds, Fishes, Cephalopoda, and Testacea have been made to form each a separate _genus_ (γένος). For within the limits of each such _genus_, the parts do not differ in that they have no nearer resemblance than that of analogy—such as exists between the bone of man and the spine of fish—but they differ merely in respect of such corporeal conditions as largeness smallness, softness hardness, smoothness roughness, and other similar oppositions, or, in one word, in respect of degree.’[49]

[49] _De partibus animalium_, i. 4; 644ᵅ 27.

The Aristotelian _genus_ thus differs widely from the term as used in modern biology. In another passage he comes nearer to defining it and the analogy of parts which extends from genus to genus.

‘Groups that differ only in the degree, and in the more or less of an identical element that they possess are aggregated together under a single _genus_; groups whose attributes are not identical but _analogous_ are separated. For instance, bird differs from bird by gradation, or by excess and defect; some birds have long feathers, others short ones, but all are feathered. Bird and Fish are more remote and only agree in having analogous organs; for what in the bird is feather, in the fish is scale. Such _analogies_ can scarcely, however, serve universally as indications for the formation of groups, for almost all animals present analogies in their corresponding parts.’[50]

Aristotle nowhere gives to his term _genus_ a rigid application that can be applied throughout the animal kingdom. He uses the word in fact much as we should use the conveniently flexible term _group_, now for a larger and less definite, now for a smaller and more definite collection of species. This varying use of a technical word makes it impossible to draw up a classification based on his _genera_ or indeed with any consistent use of the terms which he actually employs.

The difficulty or impossibility of drawing up a satisfactory classificatory system from the Aristotelian writings has not, however, deterred numerous naturalists and scholars from making the attempt, and the subject has in itself a considerable history and literature[51] extending from the days of Edward Wotton (1492-1555) downward.[52] The more recent efforts at drawing up an Aristotelian classificatory system have been based on the methods of reproduction to which he certainly attached very great importance.[53] Provided that it be remembered that Aristotle does not himself detail any such system there can be no harm in constructing one from his works. At worst it will serve as a _memoria technica_ for the extent and character of his knowledge of natural history, and at best it may represent a scheme to which he was tending.

[50] _De partibus animalium_, i. 4; 644ᵅ 16.

[51] The classificatory system of Aristotle and its history are discussed in great detail by J. B. Meyer, _Aristoteles’ Thierkunde: ein Beitrag zur Geschichte der Zoologie, Physiologie und alten Philosophie_, Berlin, 1855.

[52] The work by which Wotton is known is his _De differentiis animalium_, Paris, 1552.

[53] There is a valuable chapter on the subject of the Aristotelian classificatory system as based on the method of reproduction in W. Ogle, _Aristotle on the Parts of Animals_, London, 1882.

_ENAIMA_ (_Sanguineous and either viviparous or oviparous_)
= _vertebrates_.

{ 1. ἅνθρωπος. Man.
{ 2. κήτη. Cetaceans.
{ 3. ζῷα τετράποδα ζωοτόκα ὲν αὑτοῖς.
{ Viviparous quadrupeds.
{ (a) μὴ ἀμφώδοντα. Non-amphodonts
Viviparous in the { = Ruminants with incisor in
internal sense. { lower jaw only and with cloven
{ hoofs.
{ (b) μώνυχα. Solid-hoofed animals.
{ i. λόφουρα. Equidae.
{ ii. μώνυχα ἔτερα. Other
solid-hoofed animals.

{ 4. ὄρνιθες. Birds.
{ (a) γαμψώνυχα. Birds of prey with
{ talons.
{ (b) στεγανόποδες. Swimmers with
{ With { webbed feet.
{ perfect { (c) περιστεροειδῆ. Pigeons, doves, &c.
{ ovum. { (d) ἄποδες. Swifts, martins, &c.
{ { (e) ὄρνιθες ἕτεροι. Other birds.
Oviparous { { 5. ζὌῷα τετράποδα ῷοτόκα. Oviparous
though { { quadrupeds = Amphibians and most
sometimes { { reptiles.
_externally_ { { 6. ὀφιώδη. Serpents.
viviparous. {
{ { 7. ἰχθύες. Fishes.
{ With { (a) σελάχη. Selachians. Cartilaginous
{ imperfect { fishes and, doubtfully, the
{ { fishing-frog.
{ ovum. { (b) ιχθύες ἕτεροι. Other fishes.

_ANAIMA_ (_Non-sanguineous and either viviparous, vermiparous or
budding_) = _Invertebrates_.

With perfect ovum. { 8. μαλάκια. Cephalopods.
{ 9. μαλακόστρακα. Crustaceans.

With ‘scolex’. 10. ἔντομα. Insects, spiders,
scorpions, &c.
With generative }11. ὀστρακόδερμα. Molluscs (except
slime, buds or } Cephalopods), Echinoderms, &c.
spontaneous generation. }

With spontaneous }12. ζωόφυτα. Sponges, Coelenterates,
generation only. } &c.

Some of the elements in this classification are fundamentally unsatisfactory in that they are based on negative characters. Such is the group of _Anaima_ which is parallelled by our own equally convenient and negative though morphologically meaningless equivalent _Invertebrata_. Others, such as the subdivisions of the viviparous quadrupeds, can only be forcibly extracted out of Aristotle’s text. But there are yet others, such as the separation of the cartilaginous from the bony fishes, that exhibit true genius and betray a knowledge that can only have been reached by careful investigation. Remarkably brilliant too is his treatment of Molluscs. There can be no doubt that he dissected the bodies and carefully watched the habits of octopuses and squids, _Malacia_ as he calls them. He separates them too far from the other Molluscs, grouped by him as _Ostracoderma_, but his actual descriptions of the structure and sexual process of the cephalopods are exceedingly remarkable, and after being long disregarded or misunderstood were verified and repeated in the course of the nineteenth century.[54]

[54] The rediscovery and verification of this and other Aristotelian observations is detailed by C. Singer, ‘Greek Biology and the Rise of Modern Biology,’ _Studies in the History and Method of Science_, vol. ii, Oxford, 1921.

Passing from his general ideas on the nature and division of living creatures we may turn to some of the most noteworthy of his actual observations. In the realm of comparative anatomy proper we may instance that of the stomach of ruminants. He must have dissected these animals, for he gives a clear and correct account of the four chambers. ‘Animals’, he says, ‘present diversities in the structure of their stomachs. Of the viviparous quadrupeds, such of the horned animals as are not equally furnished with teeth in both jaws are furnished with four such chambers. These animals are those that are said to chew the cud. In these animals the oesophagus extends from the mouth downwards along the lung, from the midriff to the _big stomach_ [_rumen_, or paunch], and this stomach is rough inside and semi-partitional. And connected with it near to the entry of the oesophagus is what is called the _kekryphalos_ [_reticulum_, or honeycomb bag]; for outside it is like the stomach, but inside it resembles a netted cap; and the kekryphalos is a good deal smaller than the _big stomach_.’ The term _kekryphalos_ was applied to the net that women wore over their hair to keep it in order. ‘Connected with this kekryphalos,’ he continues, ‘is the _echinos_ [_psalterium_, or _manyplies_], rough inside and laminated, and of about the same size as the kekryphalos. Next after this comes what is called the _enystron_ [_abomasum_], larger and longer than the echinos, furnished inside with numerous folds or ridges, large and smooth. After all this comes the gut....’[55] ‘All animals that have horns, the sheep for instance, the ox, the goat, the deer and the like, have these several stomachs.... The several cavities receive the food one from the other in succession: the first taking the unreduced substances, the second the same when somewhat reduced, the third when reduction is complete, and the fourth when the whole has become a smooth pulp....’[56] ‘Such is the stomach of those quadrupeds that are horned and have an unsymmetrical dentition (μὴ ἀμφώδοντα); and these animals differ one from another in the shape and size of the parts, and in the fact of the oesophagus reaching the stomach central-wise in some cases and sideways in others. Animals that are furnished equally with teeth in both jaws (ἀμφώδοντα) have one stomach; as man, the pig, the dog, the bear, the lion, the wolf.’[57]

[55] _Historia animalium_, ii. 17; 507ᵅ 33.

[56] _De partibus animalium_, ii. 17; 507ᵇ 12.

[57] _Historia animalium_, ii. 17; 507ᵇ 12.

A very famous example in the Aristotelian works anticipating modern biological knowledge is afforded by his reference to the mode of reproduction of the cephalopods. ‘The Malacia such as the octopus, the sepia, and the calamary, have sexual intercourse all in the same way; that is to say, they unite at the mouth by an interlacing of their tentacles. When, then, the octopus rests its so-called head against the ground and spreads abroad its tentacles, the other sex fits into the outspreading of these tentacles, and the two sexes then bring their suckers into mutual connexion. Some assert that the male has a kind of penis in one of his tentacles, the one in which are the largest suckers; and they further assert that the organ is tendinous in character growing attached right up to the middle of the tentacle, and that the latter enables it to enter the nostril or funnel of the female.’[58]

[58] _Historia animalium_, v. 6; 541ᵇ 1. The hectocotylization of the cephalopod arm which is here recorded as an element in the reproductive process of these animals is denied in the _De generatione animalium_, i. 15; 720ᵇ 32, where we read that ‘the insertion of the arm of the male into the funnel of the female ... is only for the sake of attachment, and it is not an organ useful for generation, for it is outside the passage in the male and indeed outside the body of the male altogether.‘ Yet even here Aristotle knows of the physical relationship of the arm. See note on this point in the translation of the passage by A. Platt, Oxford, 1910.

The reproductive processes of the Cephalopods were unknown to modern naturalists until the middle of the nineteenth century. Before that time several observers had noted the occasional presence of a peculiar parasite in the mantle cavity of female cephalopods and had described its supposed structure without tracing any relationship to the process of generation. In 1851 it was first shown that this supposed parasite was the arm of the male animal specially modified for reproductive purposes and broken off on insertion into the mantle cavity of the female[59]. The actual process of reproduction does not seem to have been observed until 1894[60].

[59] J. B. Verany, _Mollusques méditerranéens_, Genoa, 1851.

[60] E. Racovitza. _Archives de zoologie experimentale_, Paris, 1894.

Aristotle is perhaps at his best and happiest when describing the habits of living animals that he has himself observed. Among his most pleasing accounts are those of the fishing-frog and torpedo. In these creatures he did not fail to notice the displacement of the fins associated with the depressed form of the body.

‘In marine creatures,’ he says, ‘one may observe many ingenious devices adapted to the circumstances of their lives. For the account commonly given of the frog-fish or angler is quite true; as is also that of the torpedo....

‘In the Torpedo and the Fishing-frog the breadth of the anterior part of the body is not so great as to render locomotion by fins impossible, but in consequence of it the upper pair [_pectorals_] are placed further back and the under pair [_ventrals_] are placed close to the head, while to compensate for this advancement they are reduced in size so as to be smaller than the upper ones.

‘In the Torpedo the two upper fins [pectorals] are placed in the tail, and the fish uses the broad expansion of its body to supply their place, each lateral half of its circumference serving the office of a fin.... The torpedo narcotizes the creatures that it wants to catch, overpowering them by the force of shock that is resident in its body, and feeds upon them; it also hides in the sand and mud, and catches all the creatures that swim in its way and come under its narcotizing influence. This phenomenon has been actually observed in operation.... The torpedo-fish is known to cause a numbness even in human beings.

‘The frog-fish has a set of filaments that project in front of its eyes; they are long and thin, like hairs, and are round at the tips; they lie on either side, and are used as baits.... The little creatures on which this fish feeds swim up to the filaments, taking them for bits of seaweed such as they feed upon. Accordingly, when the frog-fish stirs himself up a place where there is plenty of sand and mud and conceals himself therein, it raises the filaments, and when the little fish strike against them the frog-fish draws them in underneath into its mouth.... That the creatures get their living by this means is obvious from the fact that, whereas they are peculiarly inactive, they are often caught with mullets, the swiftest of fishes, in their interior. Furthermore, the frog-fish is usually thin when he is caught after losing the tips of his filaments.’[61]

[61] The paragraphs concerning the fishing-frog and torpedo are made up of sentences rearranged from the _De partibus animalium_, iv. 13; 696ᵅ 26, and the _Historia animalium_, ix. 37; 620ᵇ 15.

The modification of the musculature of the torpedo-fish for electric purposes and the fishing habits of the fishing-frog or _Lophius_ are now well known, but it was many centuries before naturalists had confirmed the observations of the father of biology.

When we turn from Aristotle’s observations in the department of natural history to his discussion of the actual mechanism of the living body, the subject now contained under the heading _Experimental Physiology_, we are in the presence of much less satisfactory material. Aristotle here exhibits his weakness in physics and not being endowed with any experimental knowledge of that subject his physiological development is very greatly handicapped. He seems often to accept fancies of his own in place of generalizations from collated observations. This tendency of his was conveyed to his successors and delayed physiological advance for many centuries. It forms a striking contrast to the method of certain of the Hippocratic works such as the _Epidemics_ and the _Aphorisms_ which exhibit an investigator intent on recording actual observations and on deducing general laws therefrom. Had the Hippocratic method been extended by Aristotle beyond the field of natural history, where he freely follows it, to that of physiology, the succeeding generations might have established medicine far more firmly as a science.

An important factor in Aristotle’s physical and physiological teaching is the doctrine that matter is continuous and not made up of indivisible parts. He thus rejected the atomic views of his predecessors Leucippus and Democritus which have been preserved for us by the poem of Lucretius. The different kinds of matter existing merely in their state of simple mixture formed various uniform or homogeneous substances, _homoeomeria_, of which the _tissues_ of living bodies provided one type. We now consider tissues as having structure made up of living cells or their products, but to Aristotle their structure was an essential fact following on their particular elemental constitution. The structure of muscle or flesh was perhaps comparable to that of a crystalline substance, for, as we have seen, Aristotle made no fundamental distinction between organic and inorganic _substances_, which are in his view alike subject to the processes of generation and corruption. The difference between them lies not in their structure but in their potential relation to the various degrees of soul, the vegetative, the animal, and the rational.

‘There are’, says Aristotle, ‘three degrees of composition, and of these the first in order is composition out of what some call the _elements_, earth, air, water, and fire....

‘The second degree of composition is that by which the _homogeneous_ parts of animals (ὁμοιομερῆ), such as bone, flesh, and the like, are constituted out of [these] primary substances.

‘The third and last stage is the composition which forms the _heterogeneous_ parts (ἀνομοιομερῆ) such as face, hand, and the rest.’[62]

[62] _De partibus animalium_, ii. 1; 646ᵅ 12.

The distinctions are not altogether clear but may perhaps be explained along such lines as the following. The division into homogeneous and heterogeneous corresponds in a general way to the later division into Tissues and Organs, the former, however, including much that we should not call tissue. The homogeneous parts were again of two kinds: (_a_) simple tissues or stuffs without any notion of size or shape, that is, mere substance capable of endowment with life or soul, e.g. cartilaginous or osseous tissues; and (_b_) simple structure, that is actual structure made of such a single tissue but with definite form and size, matter to which form had been added and which either was actually or had been endowed with soul, e.g. _a_ cartilage or _a_ bone.

As a physiologist Aristotle is, in fact, in much the same position as he is as a physicist. He never dissected the human body, he had only the roughest idea of the course of the vessels, and his description of the vascular system is so difficult and confused that a considerable literature has been written on its interpretation. He regarded the heart as the central organ of the body and the seat of sensation and he probably believed that the arteries contained air as well as blood. He made no adequate distinction between veins and arteries. He tells us that two great vessels arise from the heart and that the heart is, as it were, a part of these vessels. The two vessels are apparently the aorta and the vena cava, and a very elementary and not very accurate description is given of the branches of these vessels. He believed that the heart had three chambers or cavities and that it took in air direct from the lung.

The brain was for him mainly an organ by which were secreted certain cold humours which prevented any overheating of the body by the furnace of the heart under the action of the bellows of the lung. He formally rejected the older views of Diogenes of Apollonia, of Alcmaeon of Croton, and of the Hippocratic writings, that placed the seat of sensation in the brain.[63] He failed to trace any adequate relation of sense organs and nerves to brain. He considered that the spinal marrow served to hold the vertebrae together.

[63] _De partibus animalium_, ii. 10.

In general we may say that his physiology is on a much lower plane than his natural history, since in dealing with physiological questions he always seems to have in mind the body as a whole and seldom pauses for any detailed investigation of a particular part. The physiological views of Aristotle were far from being fully accepted even by the generation which followed him. There was already growing up a school of physiologists whose work culminated five centuries later in that of Galen, where we find quite other views of the bodily functions. It is these views which we may take as more typical of the bases of Greek physiology (see p. 66).

In much of the Aristotelian material that we have discussed we have seen the development of a class of interests very foreign to those of the modern biologist, in whose work the general discussion of the ultimate nature and origin of life seldom plays a large part. The business of the modern biologist is mainly with vital phenomena as he encounters them and he is not concerned with the deeper philosophical problems. The man of science considers a part of the Universe where the philosopher makes it his business to regard the whole. With Aristotle this modern scientific process of taking a part of the sensible Universe, such as a particular group of animals or the particular action of a particular organ, and considering it in and by and for itself without reference to other things, had not yet fully emerged. Philosophy and science are still inextricably linked and there is no clear demarcation between them.

This is at least his theoretical view. But besides being a philosopher by choice he was a supreme naturalist by his natural endowments and he cannot suppress his love for nature and his capacity for observation. We see Aristotle the naturalist at his greatest as a direct observer or when reasoning directly about the observations that he has made. When he disregards his own observations and begins to erect theories on the observations or the views of others, he becomes weaker and less comprehensible.

§ 3. _After Aristotle_

All Aristotle’s surviving biological works refer primarily to the animal creation. His work on plants is lost or rather has survived as the merest corrupted fragment. We are fortunate, however, in the possession of a couple of complete works by his pupil and successor Theophrastus (372-287), which may not only be taken to represent the Aristotelian attitude towards the plant world, but also give us an inkling of the general state of biological science in the generation which succeeded the master.

These treatises of Theophrastus are in many respects the most complete and orderly of all ancient biological works that have reached our time. They give an idea of the kind of interest that the working scientist of that day could develop when inspired rather by the genius of a great teacher than by the power of his own thoughts. Theophrastus is a pedestrian where Aristotle is a creature of wings, he is in a relation to the master of the same order that the morphologists of the second half of the nineteenth century were to Darwin. For a couple of generations after the appearance of the _Origin of Species_ in 1859 the industry and ability of naturalists all over the world were occupied in working out in detail the structure and mode of life of living things on the basis of the Evolutionary philosophy. Nearly all the work on morphology and much of that on physiology since his time might be treated as a commentary on the works of Darwin. These volumes of Theophrastus give the same impression. They represent the remains—alas, almost the only biological remains—of a school working under the impulse of a great idea and spurred by the memory of a great teacher. As such they afford a parallel to much scientific work of our own day, produced by men without genius save that provided by a vision and a hope and an ideal. Of such men it is impossible to write as of Aristotle. Their lives are summed up by their actual achievement, and since Theophrastus is an orderly writer whose works have descended to us in good state, he is a very suitable instance of the actual standard of achievement of ancient biology. ‘Without vision the people perish’ and the very breath of life of science is drawn, and can only be drawn, from that very small band of prophets who from time to time, during the ages, have provided the great generalizations and the great ideals. In this light let us examine the work of Theophrastus.

In the absence of any adequate system of classification, almost all botany until the seventeenth century consisted mainly of descriptions of species. To describe accurately a leaf or a root in the language in ordinary use would often take pages. Modern botanists have invented an elaborate terminology which, however hideous to eye and ear, has the crowning merit of helping to abbreviate scientific literature. Botanical writers previous to the seventeenth century were substantially without this special mode of expression. It is partly to this lack that we owe the persistent attempts throughout the centuries to represent plants pictorially in herbals, manuscript and printed, and thus the possibility of an adequate history of plant illustration.

Theophrastus seems to have felt acutely the need of botanical terms, and there are cases in which he seeks to give a special technical meaning to words in more or less current use. Among such words are _carpos_ = fruit, _pericarpion_ = seed vessel = pericarp, and _metra_, the word used by him for the central core of any stem whether formed of wood, pith, or other substance. It is from the usage of Theophrastus that the exact definition of fruit and pericarp has come down to us.[64] We may easily discern also the purpose for which he introduces into botany the term _metra_, a word meaning primarily the _womb_, and the vacancy in the Greek language which it was made to fill. ‘_Metra_,’ he says, ‘is that which is in the middle of the wood, being third in order from the bark and [thus] like to the marrow in bones. Some call it the _heart_ (καρδίαν), others the _inside_ (ἐντεριώνην), yet others call only the innermost part of the metra itself the heart, while others again call this _marrow_.’[65] He is thus inventing a word to cover all the different kinds of core and importing it from another study. This is the method of modern scientific nomenclature which hardly existed for botanists even as late as the sixteenth century of our era. The real foundations of our modern nomenclature were laid in the later sixteenth and in the seventeenth century by Cesalpino and Joachim Jung.

[64] It is possible that Theophrastus derived the word pericarp from Aristotle. Cp. _De anima_, ii. 1, 412ᵇ 2. In the passage τὸ φύλλον περικαρπίου σκέπασμα, τὸ δὲ περικάρπιον καρποῦ, in the _De anima_ the word does not, however, seem to have the full technical force that Theophrastus gives to it.

[65] _Historia plantarum_, i. 2, vi.

Theophrastus understood the value of developmental study, a conception derived from his master. ‘A plant’, he says, ‘has power of germination in all its parts, for it has life in them all, wherefore we should regard them not for what they are but for what they are becoming.’[66] The various modes of plant reproduction are correctly distinguished in a way that passes beyond the only surviving earlier treatise that deals in detail with the subject, the Hippocratic work _On generation_. ‘The manner of generation of trees and plants are these: spontaneous, from a seed, from a root, from a piece torn off, from a branch or twig, from the trunk itself, or from pieces of the wood cut up small.’[67] The marvel of generation must have awakened admiration from a very early date. We have already seen it occupying a more ancient author, and it had also been one of the chief pre-occupations of Aristotle. It is thus not remarkable that the process should impress Theophrastus, who has left on record his views on the formation of the plant from the seed.

[66] _Ibid._ i. 1, iv.

[67] _Historia plantarum_, ii. 1, i.

‘Some germinate, root and leaves, from the same
point, some separately from either end of the seed.
Thus wheat, barley, spelt, and all such cereals
[germinate] from either end, corresponding to the
position [of the seed] in the ear, the root from the
stout lower part, the shoot from the upper; but the
two, root and stem, form a single continuous whole.
The bean and other leguminous plants are not so, but
in them root and stem are from the same point, namely,
their place of attachment to the pod, where, it is
plain, they have their origin. In some cases there is
a process, as in beans, chick peas, and especially
lupines, from which the root grows downward, the leaf
and stem upward.... In certain trees the bud first
germinates within the seed, and, as it increases in
size, the seeds split—all such seeds are, as it were,
in two halves; again, all those of leguminous plants
have plainly two lobes and are double—and then the
root is immediately thrust out. But in cereals, the
seeds being in one piece, this does not happen, but the
root grows a little before [the shoot].

‘Barley and wheat come up monophyllous, but peas,
beans, and chick peas polyphyllous. All leguminous
plants have a single woody root, from which grow
slender side roots ... but wheat, barley, and the other
cereals have numerous slender roots by which they are
matted together.... There is a contrast between these
two kinds; the leguminous plants have a single root and
have many side-growths above from the [single] stem
... while the cereals have many roots and send up many
shoots, but these have no side-shoots.’[68]

There can be no doubt that here is a piece of minute observation on the behaviour of germinating seeds. The distinction between dicotyledons and monocotyledons is accurately set forth, though the stress is laid not so much on the cotyledonous character of the seed as on the relation of root and shoot. In the dicotyledons root and shoot are represented as springing from the same point, and in monocotyledons from opposite poles in the seed.

No further effective work was done on the germinating seed until the invention of the microscope, and the appearance of the work of Highmore (1613-85),[69] and the much more searching investigations of Malpighi (1628-94)[70] and Grew (1641-1712)[71] after the middle of the seventeenth century. The observations of Theophrastus are, however, so accurate, so lucid, and so complete that they might well be used as legends for the plates of these writers two thousand years after him.

[68] _Historia plantarum_, viii. 1, i.

[69] Nathaniel Highmore, _A History of Generation_, London, 1651.

[70] Marcello Malpighi, _Anatome plantarum_, London, 1675.

[71] Nehemiah Grew, _Anatomy of Vegetables begun_, London, 1672.