BIOLOGY

Tibi res antiquae laudis et artisingredior, sanctos ausus recludere fontes.

But now I, who have dared to draw my tiny draft from Aristotle’s great well, seem after all to be seeking an excuse, seeking it in example and precept. Precept, at least, I know to be of no avail. My father spent all the many days of his life in the study of Greek; you might suppose it was for Wisdom’s sake,—but my father was a modest man. The fact is, he did it for a simpler reason still, a very curious reason, to be whispered rather than told: he did itfor love.

Nigh forty years ago, I first stepped out on the east-windy streets of a certain lean and hungry town (lean, I mean, as regards scholarship) where it was to be my lot to spend thereafter many and many a year. And the very first thing I saw there was an inscription over a very humble doorway, ‘Hic mecum habitant Dante, Cervantes, Molière’. It was the home of a poor schoolmaster, who as a teacher of languages eked out the scanty profits of his school. I was not a little comforted by the announcement. So the poor scholar, looking on the ragged regiment of his few books, is helped, consoled, exalted by the reflection:Hic mecum habitant ... Homerus, Plato, Aristoteles. And were one in a moment of inadvertence to inquire of him why he occupied himself with Greek, he might perchance stammer (like Dominie Sampson) an almost inarticulate reply; but more probably he would be stricken speechless by the enormous outrage of the request, and the reason of his devotion would be hidden from the questioner for ever.

D’Arcy Wentworth Thompson.

What is science? It is a question that cannot be answered easily, nor perhaps answered at all. None of the definitions seem to cover the field exactly; they are either too wide or too narrow. But we can see science in its growth and we can say that being a process it can exist only as growth. Where does the science of biology begin? Again we cannot say, but we can watch its evolution and its progress. Among the Greeks the accurate observation of living forms, which is at least one of the essentials of biological science, goes back very far. The wordBiology, used in our sense, would, it is true, have been an impossibility among them, forbiosrefers to the life of man and could not be applied, except in a strained or metaphorical sense, to that of other living things.[6]But theideaswe associate with the word are clearly developed in Greek philosophy and the foundations of biology are of great antiquity.

The Greek people had many roots, racial, cultural, and spiritual, and from them all they inherited various powers and qualities and derived various ideas and traditions. The most suggestive source for our purpose is that of the Minoan race whom they dispossessed and whose lands they occupied. That highly gifted people exhibited in all stages of its development a marvellous power of graphically representing animal forms, of which the famous Cretan friezes, Vaphio cups (Fig. 5), andMycenean lions provide well-known examples. It is difficult not to believe that the Minoan element, entering into the mosaic of peoples that we call the Greeks, was in part at least responsible for the like graphic power developed in the Hellenic world, though little contact has yet been demonstrated between Minoan and archaic Greek Art.

For the earliest biological achievements of Greek peoples we have to rely largely on information gleaned from artistic remains. It is true that we have a few fragments of the works of both Ionian and Italo-Sicilian philosophers, and in them we read of theoretical speculation as to the nature of life and of the soul, and we can thus form some idea of the first attempts of such workers as Alcmaeon of Croton (c.500B. C.) to lay bare the structure of animals by dissection.[7]The pharmacopœia also of some of the earliest works of the Hippocratic collection betrays considerable knowledge of both native and foreign plants.[8]Moreover, scattered through the pages of Herodotus and other early writers is a good deal of casual information concerning animals and plants, though such material is second-hand and gives us little information concerning the habit of exact observation that is the necessary basis of science.

Something more is, however, revealed by early Greek Art. We are in possession of a series of vases of the seventh and sixth centuries before the Christian era showing a closeness of observation of animal forms that tells of a people awake to the study of nature. We have thus portrayed for us a number of animals—plants seldom or never appear—and among the best rendered are wild creatures: we see antelopes quietly feeding or startled at a sound, birds flying or picking worms from the ground, fallowdeer forcing their way through thickets, browsing peacefully, or galloping away, boars facing the hounds and dogs chasing hares, wild cattle forming their defensive circle, hawks seizing their prey. Many of these exhibit minutely accurate observation. The very direction of the hairs on the animals’ coats has sometimes been closely studied, and often the muscles are well rendered. In some cases even the dentition has been foundaccurately portrayed, as in a sixth-century representation on an Ionian vase of a lioness—an animal then very rare on the Eastern Mediterranean littoral, but still known in Babylonia, Syria, and Asia Minor. The details of the work show that the artist must have examined the animal in captivity (Figs. 1 and 2).

Fig. 1. Lioness and young from an Ionian vase of the sixth century B. C. found at Caere in Southern Etruria (Louvre, Salle E, No. 298), from Le Dessin des Animaux en Grèce d’après les vases peints, by J. Morin, Paris (Renouard), 1911. The animal is drawing itself up to attack its hunters. The scanty mane, the form of the paws, the udders, and the dentition are all heavily though accurately represented.Fig. 1.Lioness and young from an Ionian vase of the sixth century B. C. found at Caere in Southern Etruria (Louvre, Salle E, No. 298), from Le Dessin des Animaux en Grèce d’après les vases peints, by J. Morin, Paris (Renouard), 1911. The animal is drawing itself up to attack its hunters. The scanty mane, the form of the paws, the udders, and the dentition are all heavily though accurately represented.

Fig. 2. A, Jaw bones of lion; B, head of lioness from Caere vase (Fig. 1), after Morin. Note the careful way in which the artist has distinguished the molar from the cutting teeth.Fig. 2.A, Jaw bones of lion; B, head of lioness from Caere vase (Fig. 1), after Morin. Note the careful way in which the artist has distinguished the molar from the cutting teeth.

Fig. 3. Paintings of fish on plates.Fig. 3.Paintings of fish on plates. Italo-Greek work of the fourth century B. C. From Morin.Sargus vulgaris.Crenilabrus mediterraneus.Uranoscopus scaber?

Fig. 3.Paintings of fish on plates. Italo-Greek work of the fourth century B. C. From Morin.

Animal paintings of this order are found scattered over the Greek world with special centres or schools in such places as Cyprus, Boeotia, or Chalcis. The very name for a painter in Greek,zoographos, recalls the attention paid to living forms. By the fifth century, in painting them as in other departments of Art, the supremacy of Attica had asserted itself, and there are many beautiful Attic vase-paintings of animals to place by the side of the magnificent horses’ heads of the Parthenon (Fig. 6). In Attica, too, was early developed a characteristic and closely accurate type of representation of marine forms, and this attained a wider vogue in Southern Italy in the fourth century. From the latter period a number of dishes and vases have come down to us bearing a large variety of fish forms, portrayed with an exactness that is interesting in view of the attention to marine creatures in the surviving literature of Aristotelian origin (Fig. 3).

These artistic products are more than a mere reflex of thedaily life of the people. The habits and positions of animals are observed by the hunter, as are the forms and colours of fish by the fisherman; but the methods of huntsman and fisher do not account for the accurate portrayal of a lion’s dentition, the correct numbering of a fish’s scales or the close study of the lie of the feathers on the head, and the pads on the feet, of a bird of prey (Fig. 4). With observations such as these we are in the presence of something worthy of the nameBiology. Though but little literature on that topic earlier than the writings of Aristotle has come down to us, yet both the character of his writings and such paintings and pictures as these, suggest the existence of a strong interest and a wide literature, biological in the modern sense, antecedent to the fourth century.

Fig. 4. Head and talons of the Sea-eagle, Haliaëtus albicillaFig. 4.Head and talons of the Sea-eagle,Haliaëtus albicilla:From an Ionic vase of the sixth century B. C.Drawn from the object.From Morin.

Fig. 4.Head and talons of the Sea-eagle,Haliaëtus albicilla:

From Morin.

Greek science, however, exhibits throughout its history a peculiar characteristic differentiating it from the modern scientific standpoint. Most of the work of the Greek scientist was done in relation to man. Nature interested him mainly in relation to himself. The Greek scientific and philosophic world was an anthropocentric world, and this comes out inthe overwhelming mass of medical as distinct from biological writings that have come down to us. Such, too, is the sentiment expressed by the poets in their descriptions of the animal creation:

Many wonders there be, but naught more wondrous than man:…………………The light-witted birds of the air, the beasts of the weald and the woodHe traps with his woven snare, and the brood of the briny flood.Master of cunning he: the savage bull, and the hartWho roams the mountain free, are tamed by his infinite art.And the shaggy rough-maned steed is broken to bear the bit.Sophocles,Antigone, verses 342 ff.(Translation of F. Storr.)

It is thus not surprising that our first systematic treatment of animals is in a practical medical work, theπερι διαιτης,On diet, of the Hippocratic Collection. This very peculiar treatise dates from the later part of the fifth century. It is strongly under the influence of Heracleitus (c.540-475) and contains many points of view which reappear in later philosophy. All animals, according to it, are formed of fire and water, nothing is born and nothing dies, but there is a perpetual and eternal revolution of things, so that change itself is the only reality. Man’s nature is but a parallel to that of the universal nature, and the arts of man are but an imitation or reflex of the natural arts or, again, of the bodily functions. The soul, a mixture of water and fire, consumes itself in infancy and old age, and increases during adult life. Here, too, we meet with that singular doctrine, not without bearing on the course of later biological thought, that in the foetus all parts are formed simultaneously. On the proportion of fire and water in the body all depends, sex, temper, temperament, intellect. Such speculative ideas separate this book from the sober method of the more typical Hippocratic medical works with which indeed it has little in common.

After having discussed these theoretical matters the work turns to its own practical concerns, and in the course of setting out the natures of foods gives in effect a rough classification of animals. These are set forth in groups, and from among the larger groups only the reptiles and insects are missing. The list has been described, perhaps hardly with justification, as theCoan classificatory system. We have here, indeed, nosystemin the sense in which that word is now applied to the animal kingdom, but we have yet some sort of definite arrangement of animals according to their supposed natures. The passage opens with mammals, which are divided into domesticated and wild, the latter being mentioned in order according to size, next follow the land-birds, then the water-fowl, and then the fishes. These fish are divided into (1) the haunters of the shore, (2) the free-swimming forms, (3) the cartilaginous fishes or Selachii, which are not so named but are placed together, (4) the mud-loving forms, and (5) the fresh-water fish. Finally come invertebrates arranged in some sort of order according to their structure. The characteristic feature of the ‘classification’ is the separation of the fish from the remaining vertebrates and of the invertebrates from both. Of the fifty animals named no less than twenty are fish, about a fifth of the number studied by Aristotle, but we must remember that here only edible species are mentioned. The existence of the work shows at least that in the fifth century there was already a close and accurate study of animal forms, a study that may justly be called scientific. The predominance of fish and their classification in greater detail than the other groups is not an unexpected feature. The Mediterranean is especially rich in these forms, the Greeks were a maritime people, and Greek literature is full of imagery drawn from the fisher’s craft. From Minoan to Byzantine times the variety, beauty, and colour of fish made a deep impression on Greek minds as reflected in their art.

Much more important, however, for subsequent biologicaldevelopment than such observations on the nature and habits of animals, is the service that the Hippocratic physicians rendered to Anatomy and to Physiology, departments in which the structure of man and of the domesticated animals stands apart from that of the rest of the animal kingdom. It is with the nature and constitution of man that most of the surviving early biological writings are concerned, and in these departments are unmistakable tendencies towards systematic arrangement of the material. Thus we have division and description of the body in sevens from the periphery to the centre and from the vertex to the sole of the foot,[9]or a division into four regions or zones.[10]The teaching concerning the four elements and four humours too became of great importance and some of it was later adopted by Aristotle. We also meet numerous mechanical explanations of bodily structures, comparisons between anatomical conditions encountered in related animals, experiments on living creatures,[11]systematic incubation of hen’s eggs for the study of their development, parallels drawn between the development of plants and of human and animal embryos, theories of generation, among which is that which was afterwards called ‘pangenesis’—discussion of the survival of the stronger over the weaker—almost our survival of the fittest—and a theory of inheritance of acquired characters.[12]All these things show not only extensive knowledge but also an attempt to apply such knowledge to human needs. When we consider how even in later centuries biology was linked with medicine, and how powerful and fundamental was the influence of the Hippocratic writings, not only on their immediate successors in antiquity, but also on the Middle Ages and right into the nineteenth century, we shall recognize the significance of these developments.

Fig. 5. MINOAN GOLD CUP. SIXTEENTH CENTURY B. C.Fig. 5.MINOAN GOLD CUP. SIXTEENTH CENTURY B. C.

Fig. 6. HORSE’S HEAD. FROM PARTHENON. 440 B. C.Fig. 6.HORSE’S HEAD. FROM PARTHENON. 440 B. C.

Such was the character of biological thought within the fifth century, and a generation inspired by this movement produced some noteworthy works in the period which immediately followed. In the treatiseπερι τροφης,On nourishment, which may perhaps be dated about 400B. C., we learn of the pulse for the first time in Greek medical literature, and read of a physiological system which lasted until the time of Harvey, with the arteries arising from the heart and the veins from the liver. Of about the same date is a workπερι καρδιης,On the heart, which describes the ventricles as well as the great vessels and their valves, and compares the heart of animals with that of man.

A little later, perhaps 390B. C., is the treatiseπερι σαρκων,On muscles, which contains much more than its title suggests. It has the old system of sevens and, inspired perhaps by the philosophy of Heracleitus (c.540-475), describes the heart as sending air, fire, and movement to the different parts of the body through the vessels which are themselves constantly in movement. The infant in its mother’s womb is believed to draw in air and fire through its mouth and to eatin utero. The action of the air on the blood is compared to its action on fire. In contrast to some of the other Hippocratic treatises the central nervous system is in the background; much attention, however, is given to the special senses. The brain resounds during audition. The olfactory nerves are hollow, lead to the brain, and, convey volatile substances to it which cause it to secrete mucus. The eyes also have been examined, and their coats and humours roughly described; an allusion, the first in literature, is perhaps made to the crystalline lens, and the eyes of animals are compared with those of man. There is evidence not only of dissection but of experiment, and in efforts to compare the resistance of various tissues to such processes as boiling, we may see the small beginning of chemical physiology.

An abler work than any of these, but exhibiting less power of observation is a treatise,περι γονης,On generation, that may perhaps be dated about 380B. C.[13]It exhibits a writer of much philosophic power, very anxious for physiological explanations, but hampered by ignorance of physics. He has, in fact, the weaknesses and in a minor degree the strength of his successor Aristotle, of whose great work on generation he gives us a fore-taste. He sets forth in considerable detail a doctrine of pangenesis, not wholly unlike that of Darwin. In order to explain the phenomena of inheritance he supposes that vessels reach the seed, carrying with them samples from all parts of the body. He believes that channels pass from all the organs to the brain and then to the spinal marrow (or to the marrow direct), thence to the kidneys and on to the genital organs; he believes, too, that he knows the actual location of one such channel, for he observes, wrongly, that incision behind the ears, by interrupting the passage, leads to impotence. As an outcome of this theory he is prepared to accept inheritance of acquired characters. The embryo develops and breathes by material transmitted from the mother through the umbilical cord. We encounter here also a very detailed description of a specimen of exfoliatedmembrana mucosa uteriwhich our author mistakes for an embryo, but his remarks at least exhibit the most eager curiosity.[14]

The author of this work on generation is thus a ‘biologist’ in the modern sense, and among the passages exhibiting him in this light is his comparison of the human embryo with the chick. ‘The embryo is in a membrane in the centre of which is the navel through which it draws and gives its breath, and themembranes arise from the umbilical cord.... The structure of the child you will find from first to last as I have already described.... If you wish, try this experiment: take twenty or more eggs and let them be incubated by two or more hens. Then each day from the second to that of hatching remove an egg, break it, and examine it. You will find exactly as I say, for the nature of the bird can be likened to that of man. The membranes [you will see] proceed from the umbilical cord, and all that I have said on the subject of the infant you will find in a bird’s egg, and one who has made these observations will be surprised to find an umbilical cord in a bird’s egg.’[15]

The same interest that he exhibits for the development of man and animals he shows also for plants.

‘A seed laid in the ground fills itself with the juices there contained, for the soil contains in itself juices of every nature for the nourishment of plants. Thus filled with juice the seed is distended and swells, and thereby the power (= facultyἡ δυναμις) diffused in the seed is compressed by pneuma and juice, and bursting the seed becomes the first leaves. But a time comes when these leaves can no longer get nourished from the juices in the seed. Then the seed and the leaves erupt below, for urged by the leaves the seed sends down that part of its power which is yet concentrated within it and so the roots are produced as an extension of the leaves. When at last the plant is well rooted below and is drawing its nutriment from the earth, then the whole grain disappears, being absorbed, save for the husk, which is the most solid part; and even that, decomposing in the earth, ultimately becomes invisible. In time some of the leaves put forth branches. The plant being thus produced by humidity from the seed is still soft and moist. Growing actively both above and below, it cannot as yet bear fruit, for it has not the quality of force and reserve (δυναμις ισχυρη και πιαρα) from which a seed can be precipitated. But when, with time, the plant becomes firmer and better rooted, it develops veins as passages bothupwards and downwards, and it draws from the soil not only water but more abundantly also substances that are denser and fatter. Warmed, too, by the sun, these act as a ferment to the extremities and give rise to fruit after its kind. The fruit thus develops much from little, for every plant draws from the earth a power more abundant than that with which it started, and the fermentation takes place not at one place but at many.’[16]

Nor does our author hesitate to draw an analogy between the plant and the mammalian embryo. ‘In the same way the infant lives within its mother’s womb and in a state corresponding to the health of the mother ... and you will find a complete similitude between the products of the soil and the products of the womb.’

The early Greek literature is so scantily provided with illustrations drawn from botanical study, that it is worth considering the remarkable comparison of generation of plants from cuttings and from seeds in the same work.

‘As regards plants generated from cuttings ... that part of a branch where it was cut from a tree is placed in the earth and there rootlets are sent out. This is how it happens: The part of the plant within the soil draws up juices, swells, and develops a pneuma (πνευμα ισχει), but not so the part without. The pneuma and the juice concentrate the power of the plant below so that it becomes denser. Then the lower end erupts and gives forth tender roots. Then the plant, taking from below, draws juices from the roots and transmits them to the part above the soil which thus also swells and develops pneuma; thus the power from being diffused in the plant becomes concentrated and budding, gives forth leaves.... Cuttings, then, differ from seeds. With a seed the leaves are borne first, then the roots are sent down; with a cutting the roots form first and then the leaves.’[17]

But with these works of the early part of the fourth century the first stage of Greek biology reaches its finest development.Later Hippocratic treatises which deal with physiological topics are on a lower plane, and we must seek some external cause for the failure. Nor have we far to seek. This period saw the rise of a movement that had the most profound influence on every department of thought. We see the advent into the Greek world of a great intellectual movement as a result of which the department of philosophy that dealt with nature receded before Ethics. Of that intellectual revolution—perhaps the greatest the world has seen—Athens was the site and Socrates (470-399) the protagonist. With the movement itself and its characteristic fruit we are not concerned. But the great successor and pupil of its founder gives us in theTimaeusa picture of the depth to which natural science can be degraded in the effort to give a specific teleological meaning to all parts of the visible Universe. The book and the picture which it draws, dark and repulsive to the mind trained in modern scientific method, enthralled the imagination of a large part of mankind for wellnigh two thousand years. Organic nature appears in this work of Plato (427-347) as the degeneration of man whom the Creator has made most perfect. The school that held this view ultimately decayed as a result of its failure to advance positive knowledge. As the centuries went by its views became further and further divorced from phenomena, and the bizarre developments of later Neoplatonism stand to this day as a warning against any system which shall neglect the investigation of nature. But in its decay Platonism dragged science down and destroyed by neglect nearly all earlier biological material. Mathematics, not being a phenomenal study, suited better the Neoplatonic mood and continued to advance, carrying astronomy with it for a while—astronomy that affected the life of man and that soon became the handmaid of astrology; medicine, too, that determined the conditions of man’s life was also cherished, though often mistakenly, but pure science was doomed.

But though the ethical view of nature overwhelmed science in the end, the advent of the mighty figure of Aristotle (384-322) stayed the tide for a time. Yet the writer on Greek Biology remains at a disadvantage in contrast with the Historian of Greek Mathematics, of Greek Astronomy, or of Greek Medicine, in the scantiness of the materials for presenting an account of the development of his studies before Aristotle. The huge form of that magnificent naturalist completely overshadows Greek as it does much of later Biology.

Charles Singer.

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.

Fig. 7. ARISTOTLE From HERCULANEUM Probably work of fourth century B. C.Fig. 7. ARISTOTLEFrom HERCULANEUMProbably work of fourth century B. C.

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 theOrigin of Speciesin 1859 the industry and ability of naturalists all over the world were occupied in working out in detail the structureand 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 botanicalterms, 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 arecarpos= fruit,pericarpion= seed vessel = pericarp, andmetra, 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.[18]We may easily discern also the purpose for which he introduces into botany the termmetra, a word meaning primarily thewomb, 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 theheart(καρδιαν), others theinside(εντεριωνην), yet others call only the innermost part of the metra itself the heart, while others again call thismarrow.’[19]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.

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.’[20]The various modes of plant reproduction are correctly distinguished in a way that passes beyond the only surviving earlier treatise that deals indetail with the subject, the Hippocratic workOn 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.’[21]The marvel of germination 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 preoccupations 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.

‘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 havea 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.’[22]

‘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 havea 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.’[22]

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),[23]and the much more searching investigations of Malpighi (1628-94[24]and Grew (1641-1712)[25]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.

Much has been written as to the knowledge of the sex of plants among the ancients. It may be stated that of the sexual elements of the flower no ancient writer had any clear idea. Nevertheless, sex is often attributed to plants, and the simile of theLoves of Plantsenters into works of the poets. Plants are frequently described as male and female in ancient biological writings also, and Pliny goes so far as to say that some students considered that all herbs and trees were sexual.[26]Yet when such passages can be tested it will be found that these so-called males and females are usually different species. Ina few cases a sterile variety is described as the male and a fertile as the female. In a small residuum of cases diœcious plants or flowers are regarded as male and female, but with no real comprehension of the sexual nature of the flowers. There remain the palms, in which the knowledge of plant sex had advanced a trifle farther. ‘With dates’, says Theophrastus, ‘the males should be brought to the females; for the males make the fruit persist and ripen, and this some call by analogyto use the wild fig(ολυνθαζειν).[27]The process is thus: when the male is in the flower they at once cut off the spathe with the flower and shake the bloom, with its flower and dust, over the fruit of the female, and, if it is thus treated, it retains the fruit and does not shed it.’[28]The fertilizing character of the spathe of the male date palm was familiar in Babylon from a very early date. It is recorded by Herodotus[29]and is represented by a frequent symbol on the Assyrian monuments.

The comparison of the fertilization of the date palm to the use of the wild fig refers to the practice of Caprification. Theophrastus tells us that there are certain trees, the fig among them, which are apt to shed their fruit prematurely. To remedy this ‘the device adopted is caprification. Gall insects come out of the wild figs which are hanging there, eat the tops of the cultivated figs, and so make them swell’.[30]These gall-insects ‘are engendered from the seeds’.[31]Theophrastus distinguished between the process as applied to the fig and the date, observing that ‘in both [fig and date] themale aids the female—for they call the fruit-bearing [palm]female—but whilst in the one there is a union of the two sexes, in the other things are different’.[32]

Theophrastus was not very successful in distinguishing the nature of the primary elements of plants, though he was able to separate root, stem, leaf, stipule, and flower on morphological as well as to a limited extent on physiological grounds. For the root he adopts the familiar definition, the only one possible before the rise of chemistry, that it ‘is that by which the plant draws up nourishment’,[33]a description that applies to the account given by the pre-Aristotelian author of the workπερι γονης,On generation. But Theophrastus shows by many examples that he is capable of following out morphological homologies. Thus he knows that the ivy regularly puts forth roots from the shoots between the leaves, by means of which it gets hold of trees and walls,[34]that the mistletoe will not sprout except on the bark of living trees into which it strikes its roots, and that the very peculiar formation of the mangrove tree is to be explained by the fact that ‘this plant sends out roots from the shoots till it has hold on the ground and roots again: and so there comes to be a continuous circle of roots round the tree, not connected with the main stem, but at a distance from it’.[35]He does not succeed, however, in distinguishing the real nature of such structures as bulbs, rhizomes, and tubers, but regards them all as roots. Nor is he more successful in his discussion of the nature of stems. As to leaves, he is more definite and satisfactory, though wholly in the dark as to their function; he is quite clear that the pinnate leaf of the rowan tree, for instance, is a leaf and not a branch.

Notwithstanding his lack of insight as to the nature of sex in flowers, he attains to an approximately correct idea of therelation of flower and fruit. Some plants, he says, ‘have [the flower] around the fruit itself as vine and olive; [the flowers] of the latter, when they drop, look as though they had a hole through them, and this is taken for a sign that it has blossomed well; for if [the flower] is burnt up or sodden, the fruit falls with it, and so it does not become pierced. Most flowers have the fruit case in the middle, or it may be the flower is on the top of the pericarp as in pomegranate, apple, pear, plum, and myrtle ... for these have their seeds below the flower.... In some cases again the flower is on top of the seeds themselves as in ... all thistle-like plants’.[36]Thus Theophrastus has succeeded in distinguishing between the hypogynous, perigynous, and epigynous types of flower, and has almost come to regard its relation to the fruit as the essential floral element.

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