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 treatise περὶ τροφῆς, On nourishment, which may perhaps be dated about 400 b. 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 work περὶ καρδίης, 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 390 b. c., is the treatise περὶ σαρκῶν, 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 eat in utero. The action of 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, περὶ γονῆς, On generation, that may perhaps be dated about 380 b. c.[8] 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 foretaste. 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 exfoliated membrana mucosa uteri which our author mistakes for an embryo, but his remarks at least exhibit the most eager curiosity.[9]

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 the membranes 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.’[10]

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 (= faculty ἡ δὗύναμις) diffused in the seed is compressed by living principle (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, 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 (δύναμις ὶσχυρὴ καὶ πιαρἀ) from which a seed can be precipitated. But when, with time, the plant becomes firmer and better rooted, it develops veins as passages both upwards 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.’[11]

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 with that 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 (πνεῦμα ἴσχει), 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.’[12]