Form and Function: A Contribution to the History of Animal Morphology

by E. S. Russell

Now since the mode of development in each type is peculiar to that type, organs of the same name in different types must not necessarily be accounted homologous, even if they correspond exactly with one another in their general functional relations to the rest of the organs. Thus the central nervous system of Arthropods must not be homologised with the central nervous system of Vertebrates, for it develops in a different manner. So, too, the brain of Arthropods or of Mollusca is not strictly comparable with the brain of Vertebrates. Again, the air-tubes or tracheæ of insects are, like the trachea and bronchi of many Vertebrates, air-breathing organs. But the two organs are not homologous, for the air-tubes of Vertebrates are developed from the alimentary tube ("fundamental organ" of the alimentary system, developed from the vegetative layer), while the air-tubes of insects arise either by histological differentiation, or by invagination of the skin (p. 236). Organs can be homologous only within the limits of the big groups; there can be no question of homology between members of different types.

The development of plants, like the development of animals, is essentially a progress from the general to the special (p. 242). Botanists have not been troubled by any recapitulation theory, and in founding their big groups, Acotyledons, Monocotyledons, and Dicotyledons, upon embryological characters, they were guided by true principles, which ought indeed to be followed in zoology. If we knew the development of all kinds of animals sufficiently well, then the best way to classify them would be according to the characters they show in their early development, for it is in early development that they show the characters of the type in their most generalised form. As it is, we have in our ignorance to establish the big groups by the study of adult structure, but we find, on putting together all we know of comparative embryology, that a classification of animals according to the mode of their development gives, as is only natural, the same four groups as does the study of adult structure. The four types of development are thus:—

(1) The double-symmetrical, which is found in Vertebrates. It is called the double-symmetrical, because in Vertebrates development takes place from a central axis (notochord) in two directions, upwards and downwards, in such a way that two tubes are formed, one above and one below the axis. (2) The second type is the symmetrical, which is shown by Annulates. A primitive streak is formed on the ventral surface of the yolk; development proceeds symmetrically on both sides of the streak. (3) Radiate development is probably typical of the radiate structural type. (4) In the massive type, the development seems to be a spiral one.

Common to most modes is a separation of the germ into animal and plastic layers, a separation which seems to be conditioned largely by the presence of yolk. A classification based upon embryological characters ought to be applied even to the lesser groups and would here prove itself of service. Embryology, for instance, fully supports de Blainville's separation of Batrachia from true reptiles,[176] for reptiles develop an amnion and Batrachia do not.

We come now to the sixth and last Scholion. Development is a true evolution of the special from the general, so runs von Baer's most general law of all. This can be expressed in a slightly different way, and the words which he chooses in the sixth Scholion to express this final and most general result are these:—"The developmental history of the individual is the history of the growing individuality in every respect" (p. 263). The greatest modern treatise on embryology ends on a splendid note. One creative thought rules all the forms of life. And more—"It is this same thought that in cosmic space gathered the scattered masses into spheres and bound them together in the solar system, the same that from the weathered dust on the surface of the metallic planets brought forth the forms of life. And this thought is nought else but life itself, and the words and syllables in which life expresses itself are the varied forms of the living" (p. 264).

Von Baer reminds one greatly of Cuvier. There is the same sheer intellectual power, the same sanity of mind, the same synthetic grip. Von Baer, like Cuvier, never forgot that he was working with living things; he was saturated, like Cuvier, with the sense of their functional adaptedness. In his paper on the external and internal skeleton[177] he gives a masterly analysis of the functional modifications of the limbs in Vertebrates, and the whole paper indeed, with its sober attack on transcendentalism, is a vindication as much of the functional point of view as of the importance of embryology.

Both Cuvier and von Baer, by the very sanity of their views, found themselves in partial opposition to the theories current in their time. Cuvier was the critic of Geoffroy and the transcendentalists, of Lamarck and the believers in the Échelle des êtres, evolutionary or ideal. Von Baer also, though influenced greatly by Naturphilosophie, turned against the exaggerations of the transcendental school, and by his unanswerable criticism of the theory of parallelism took away the ground from those who too easily believed in an historical evolution.[178]

We have seen what were von Baer's criticisms of the theory of parallelism. If we turn to the later writings of Cuvier we find the essential criticism expressed in similar terms. Speaking of an attempt which had been made to show that fish were molluscs developed to a higher degree, he wrote in 1828,[179] "Let us draw the conclusion that even if these animals can be spoken of as ennobled molluscs, as molluscs raised to a higher power, or if they are embryos of reptiles, the beginnings of reptiles, this can be true of them only in an abstract and metaphysical sense, and that even this abstract statement would be very far from giving an accurate idea of their organisation." From the fact that the respiratory and circulatory organs of fish greatly resemble those of tadpoles the conclusion has been drawn that fish are in a sense embryos of Amphibia (p. 547). But this manner of viewing things is none the less vicious, "for this reason ... that it considers only one or two points and neglects all the others" (p. 548), and is directly contrary to common sense. There is never a recapitulation of total organisations, only at the most of single organs.

It will be remembered that Cuvier opposed and demolished the theory of the Échelle des êtres, not only by showing that there were in Nature four entirely different plans of animal structure, but also by demonstrating that even the animals of each single Embranchement could not readily be arranged in one series, that a serial arrangement was really valid only for their separate organs. Von Baer also held that there are four distinct types of structure; he, too, combated the idea of gradation within the limits of the type. In so far as species represent successive stages in the development, the Ausbildung, of the type, so far can the idea of a scale of beings be applied. But the members of a type follow not one line of evolution but several diverging lines, in direct adaptation to different environmental conditions, so that a serial arrangement of them is not as a rule possible. It may be possible to establish a serial arrangement of single organs from the simplest to the most complex. But each organ or organ-system will require a different serial arrangement, for the different systems vary on different lines and an animal may be highly developed in respect of one system and little developed in respect of all the others. Man, for instance, is the highest animal only in respect of his nervous system. The idea of the scale of beings has therefore only a very limited application even within the limits of the type. Applied to the whole animal kingdom it becomes merely absurd.

Another point of resemblance between Cuvier and von Baer was that Cuvier, though essentially a student of adult structure, did recognise the importance of embryology; following up some observations of Dutrochet he studied the fœtal membrane of mammals and tried to establish their homologies.[180] And in his criticism of the vertebral theory of the skull he advanced as an argument against the basisphenoid being a vertebral centrum the fact (established by Kerkring, 1670), that it develops from two centres.[181] Von Baer's relation to transcendental anatomy was in some ways a close one, though he was a trenchant critic of the extreme views of the school.[182] He took from Oken the idea that a simple fundamental plan rules the organisation of all Vertebrates; "That jaws and limbs are modifications of one fundamental form is readily apparent, and, after Oken, the fact ought to be accepted by the majority of those naturalists who do not refuse to admit the existence of a general type from which the diversity of structure is developed" (i., p. 192). He accepted the vertebral theory of the skull in its main lines, and used his embryological knowledge to support the idea that jaws correspond to limbs—the latter point as part of the transcendental idea that the hind end of the body repeats the organisation of the anterior part (i., p. 192). The particular form which his theory of the relation of jaws to limbs took is shown in the following passage:—"The maxillary bone has ... the significance of an extremity and at the same time that of a rib or lower arch of a vertebra, just as the pelvic bones unite in themselves the signification of ribs and proximal members of the hinder extremity" (Meckel's Archiv, p. 367, 1826).