Around the middle of the 4th century bce, ancient Greek science reached a climax with Aristotle, who was interested in all branches of knowledge, including biology. Using his observations and theories, Aristotle was the first to attempt a system of animal classification, in which he contrasted animals containing blood with those that were bloodless. The animals with blood included those now grouped as mammals (except the whales, which he placed in a separate group), birds, amphibians, reptiles, and fishes. The bloodless animals were divided into the cephalopods, the higher crustaceans, the insects, and the testaceans, the last group being a collection of all the lower animals. His careful examination of animals led to the understanding that mammals have lungs, breathe air, are warm-blooded, and suckle their young. Aristotle was the first to show an understanding of an overall systematic taxonomy and to recognize units of different degrees within the system.

The most-important part of Aristotle’s work was that devoted to reproduction and the related subjects of heredity and descent. He identified four means of reproduction, including the abiogenetic origin of life from nonliving mud, a belief held by Greeks of that time. Other modes of reproduction recognized by him included budding (asexual reproduction), sexual reproduction without copulation, and sexual reproduction with copulation. Aristotle described sperm and ova and believed that the menstrual blood of viviparous organisms (those that give birth to living young) was the actual generative substance.

Although Aristotle recognized that species are not stable and unalterable and although he attempted to classify the animals he observed, he was far from developing any pre-Darwinian ideas concerning evolution. In fact, he rejected any suggestion of natural selection and sought teleological explanations (i.e., all phenomena in nature are shaped by a purpose) for any given observation. Nevertheless, many important scientific principles, some of which are often thought of as 20th-century concepts, can be ascribed to Aristotle. The following are a few such: (1) Using birds as an example, he formulated the principle that all organisms are structurally and functionally adapted to their habits and habitats. (2) Nature is parsimonious; it does not expend unnecessary energy. (3) In classifying animals, Aristotle rejected the idea of dividing them solely by their external structures (e.g., animals with wings and those without wings). He recognized instead a basic unity of plan among diverse organisms, a principle that is still conceptually and scientifically sound. Further, Aristotle also believed that the entire living world could be described as a unified organization rather than as a collection of diverse groups. (4) By his observations, Aristotle realized the importance of structural homology, basically similar organs in different animals, and functional analogy, different structures that serve somewhat the same function—e.g., the hand, the claw, and the hoof are analogous structures. Those principles constitute the basis for the biological field of study known as comparative anatomy. (5) Aristotle’s observations also led to the formulation of the principle that general structures appear before specialized ones and that tissues differentiate before organs.

Botanical investigations

Of all the works of Aristotle that have survived, none deals with what was later differentiated as botany, although it is believed that he wrote at least two treatises on plants. Fortunately, however, the work of Theophrastus, one of Aristotle’s students, has been preserved to represent plant science of the Greek period. Like Aristotle, Theophrastus was a keen observer, although his works do not express the depth of original thought exemplified by his teacher. In his great work, De historia et causis plantarum (The Calendar of Flora, 1761), in which the morphology, natural history, and therapeutic use of plants are described, Theophrastus distinguished between the external parts, which he called organs, and the internal parts, which he called tissues. This was an important achievement because Greek scientists of that period had no established scientific terminology for specific structures. For that reason, both Aristotle and Theophrastus were obliged to write very long descriptions of structures that can be described rapidly and simply today. Because of that difficulty, Theophrastus sought to develop a scientific nomenclature by giving special meaning to words that were then in more or less current use; for example, karpos for fruit and perikarpion for seed vessel.

Although he did not propose an overall classification system for plants, more than 500 of which are mentioned in his writings, Theophrastus did unite many species into what are now considered genera. In addition to writing the earliest detailed description of how to pollinate the date palm by hand and the first unambiguous account of sexual reproduction in flowering plants, he also recorded observations on seed germination and development.

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Post-Grecian biological studies

With Aristotle and Theophrastus, the great Greek period of scientific investigation came to an end. The most famous of the new centres of learning were the library and museum in Alexandria. From 300 bce until around the time of Christ, all significant biological advances were made by physicians at Alexandria. One of the most outstanding of those individuals was Herophilus, who dissected human bodies and compared their structures with those of other large mammals. He recognized the brain, which he described in detail, as the centre of the nervous system and the seat of intelligence. On the basis of his knowledge, he wrote a general anatomical treatise, a special one on the eyes, and a handbook for midwives.

Erasistratus, a younger contemporary and reputed rival of Herophilus who also worked at the museum in Alexandria, studied the valves of the heart and the circulation of blood. Although he was wrong in supposing that blood flows from the veins into the arteries, he was correct in assuming that small interconnecting vessels exist. He thus suspected (but did not see) the presence of capillaries; he thought, however, that the blood changed into air, or pneuma, when it reached the arteries, to be pumped throughout the body.

Perhaps the last of the ancient biological scientists of note was Galen of Pergamum, a Greek physician who practiced in Rome during the middle of the 2nd century ce. His early years were spent as a surgeon at the gladiatorial arena, which gave him the opportunity to observe details of human anatomy. At that time in Rome, however, it was considered improper to dissect human bodies, and, as a result, a detailed study of human anatomy was not possible. Thus, though Galen’s research on animals was thorough, his knowledge of human anatomy was faulty. Because his work was extensive and clearly written, Galen’s writings, nevertheless, dominated medicine for centuries.