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

by E. S. Russell

"Can the organisation of vertebrated animals be referred to one uniform type?" This is the question with which the Philosophie anatomique opens, the question to which the whole book is an answer. But is it not generally acknowledged by naturalists that Vertebrates are built upon one uniform plan, that, for instance, the fore limb may be modified for running, climbing, swimming, or flying, yet the arrangement of the bones remain the same? How else could there be a "natural method" of classification?[87]

But the homologies so drawn repose upon a vague and confused feeling for likenesses; they are not based upon an explicit principle. What general principle can be applied? "Now it is evident that the sole general principle one can apply is given by the position, the relations, and the dependencies of the parts, that is to say, by what I name and include under the term of connections." For instance, the part known as the hand in man and generally as the fore foot in other Vertebrates, is the fourth part in order in the anterior member, and its homologue can always be recognised by this fact of its connections (p. xxvi.). The principle of connections serves as a guide in tracing an organ through all its functional transformations, for "an organ can be deteriorated, atrophied, annihilated, but not transposed" (p. xxx.).

It is this principle which enables one to follow out in detail the further fundamental conception that in every Vertebrate there are found the same "organic materials," or units of construction. This conception, which Geoffroy calls the Théorie des analogues (p. xxxii.), is clearly one part of the old idea of the unity of type; it teaches the unity of composition of organic beings, while the Principe des connexions adds the unity of plan.

Both conceptions are logically implicit in the vague notion of unity of type; Geoffroy disengaged them, and pushed each to its logical extreme.

Most of the ordinary homologies of structure in air-breathing Vertebrates have already been seized, he continues, for they are more or less obvious, and many intermediate states exist (p. xxxiv.). But ordinary methods of comparison fail when the attempt is made to homologise the structure of fishes with that of air-breathing Vertebrates, for the homologies are anything but obvious and no intermediate organs are found.

Most air-breathing Vertebrates have a larynx, a trachea, and bronchi, which are absent in fish; and fish have many parts which seem to be absent in higher Vertebrates. But apply the "Theory of Analogues"; it teaches that there can be no organ peculiar to fish and not found in other Vertebrates; apply the "Principle of Connections," it will show which organs are homologous in the two types (p. xxxv.).

Comparative anatomists, with few exceptions, had hitherto taken man as the type, and referred all structure to his; Geoffroy's principles led him to give preference to no one animal in particular, but to seize upon each part in the species in which it reaches the maximum of its development (p. xxxvi.). He is thus led to refer all structures to a generalised abstract type. In this abstract type each organ exists at the maximum of its development, each organ shows all its potentialities realised. In a way, therefore, this type, this abstraction, gives the scheme of the possible transformations of each organ.

It is true Geoffroy does not refer to this "Archetype" in so many words, but it must always have been vaguely present in his mind. He has this idea in his head when he says in one of his later works, "There is, philosophically speaking, only a single animal."[88] The "single animal" is simply the generalised type.

Having laid down his two principles Geoffroy goes on to apply them to the difficult case of the comparison of the skeleton of fish with the skeleton of the higher Vertebrates. "My present task is to demonstrate that there is no part of the bony framework of fishes that cannot find its analogue in the other vertebrated animals."[89] It seems at first sight that many bones are peculiar to fish, formed expressly for performing the functions which fish do not share with higher animals. These are the bones connected with respiration—the operculum, the branchiostegal rays, the branchial arches, and others. That the peculiar bones should be connected with the respiratory functions is only natural, for the contrast between fish and higher Vertebrates is essentially a contrast between water-breathing and air-breathing animals. Considering first the general form of the skeleton in fish, we are met at once with a difficulty; there is no obvious homologue in fishes of the neck, the trunk, and the abdomen of higher animals. What apparently corresponds to the trunk is in fishes crowded close up under the head. But, after all, it is not of the essence of the vertebrate type to have the trunk and the abdomen attached at definite and invariable distances along the vertebral column—that is a notion surviving from the anatomy which made man its type. The "trunk" differs in position according to the class, in quadrupeds, birds, and fishes (p. 9). Now, says Geoffroy, allow me this one hypothesis, that the trunk with its organs can, as it were, move bodily along the vertebral column, so as to be found in one class near the front end of the vertebral column, in another about the middle, and in a third near the end, then I can show you in detail that the constituent parts of this trunk are found in all classes to be invariably in the same positions relatively to one another (p. 10). It is important to note this hypothesis of a "metastasis" which Geoffroy makes, for it is the key to the understanding of many of the far-fetched homologies which he tries to establish. It is, of course, clear that this hypothesis is in formal contradiction with his principal hypothesis of the invariability of connections, and that he, so to speak, gets a hold on his fish to apply his principle of connections only by admitting at the very outset an exception to his primary principle. A further application of the hypothesis of metastasis will be noticed below in connection with the determination of the sternum of fishes. We note here an interpretation of the first metastasis in terms of functional adaptation. "The constant and violent action of the tail, if it does not go so far as actually to displace and move forward the internal organs, at least fits in well with an arrangement in which the organs are so disposed" (p. 99).

The first memoir deals with the homologies of the opercular bones. Geoffroy considers that the external opening of the ear corresponds to the external opening of the gill-chamber, which lies between the operculum and the pectoral girdle. The ear communicates with the buccal cavity by the Eustachian tube, so does the branchial chamber by means of the gill-slits. The auditory chamber of higher Vertebrates is, therefore, the homologue of the branchial chamber in fish; the opercular bones in fish and the ossicles of the ear in other Vertebrates stand in close relation to this chamber; therefore the opercular bones are the homologues of the ossicles of the ear, the interoperculum corresponding to the malleus, the suboperculum to the lenticular, the minute lower part of the suboperculum to the incus, the operculum to the stapes, and the pre-operculum to the tympanic ring. In making these particular determinations Geoffroy professes to be led by his principle of connections. The pre-operculum has, he says, the same connections with neighbouring bones as the tympanic bone in other Vertebrates, and the other pieces of the gill-cover are homologised with particular ear-ossicles according to the order in which they stand to one another. The second memoir in the book deals with the sternum, and affords a very good example of Geoffroy's method of dealing with the facts of structure. We shall omit here any detailed reference to the other three memoirs, which deal with the hyoid, with the branchial arches and the structures which correspond in air-breathing Vertebrates, and with the bones of the shoulder-girdle.