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

by E. S. Russell

In the memoir on the sternum Geoffroy's first care is to arrive at a definition of what a sternum is. He defines it partly by its functions, partly by its connections, as the system of bones which covers and protects the thorax, and gives attachment to certain groups of muscles.

The most highly developed sternum (according to this definition) is the plastron of the tortoise, whose structure it dominates (p. 103). It is important, therefore, to determine of how many bones the plastron is composed, since the full number of elementary parts of which an organ is composed is best seen when the organ is at the maximum of its development. There are nine bones in the plastron of the tortoise. "The conclusion to be drawn from this is that every sternum, provided that it is not inhibited in its development by some obstacle, is composed of nine elementary parts" (p. 105). These nine bones are in Geoffroy's nomenclature, the episternals, the hyosternals, the hyposternals, the xiphisternals, which are all paired bones, and the entosternal, which is unpaired. The arrangement of them is in the tortoise:—

The articulations in the tortoise are indicated by the connecting lines. Geoffroy tries to show that the sternum in other animals is composed of these nine bones, or at least of a certain number of them, always in the same invariable relative positions. Thus in birds the sternum consists of five pieces, of a huge keeled entosternal, and of two "annexes" on either side, which are the hyo-and hyposternals. These are separate only in young birds. Occasionally, especially in young birds, rudiments of episternals and xiphisternals also occur. The minuteness of the episternals and the xiphisternals

Fig 1.—Hyoid Arch of the Conger.(Original)

may be attributed to the gigantic size of the entosternal, in accordance with the Loi de balancement. In the other air-breathing Vertebrates the nine sternal elements can according to Geoffroy be discovered without great difficulty. But when we come to the determination of the sternum in fishes, difficulties abound, which Geoffroy solves in the following way. He points out that between the clavicles (cleithra) and the hyoid bone (basihyal) in fishes there is a long median bone (urohyal) which is attached in front by two strong tendons to the horns of the hyoid and is free behind (see [Fig. 1]). Gouan (1720) had seen in this bone the homologue of the sternum. Geoffroy adopts this view, but considers that this bone alone cannot represent the whole sternum. He finds the representatives of other bones of the sternum in the large bones (epihyal and ceratohyal, or the two pieces of the ceratohyal) which are comprised in the hyoid arch. But he is immediately met by the difficulty that this complex of bones is situated in front of the pectoral girdle, whereas the sternum in higher Vertebrates lies behind the pectoral girdle. He reflects, however, that the gills of fish, situated in front of the clavicles, are merely the lungs under another name. The gills have become shifted forward by a metastasis similar to that which brought the whole thoracic organs far forward in fish. This being so, their supporting elements, the sternum and the ribs, must have moved with them, and are hence to be found in front of the pectoral girdle.

Geoffroy's next step is to point out that the only possible homologues of sternal ribs are the branchiostegal rays, which arise from the large bones of the hyoid arch. If these are sternal ribs, the bones to which they are attached must be the hyo- and hyposternals or "annexes," the bones from which in birds the ribs take their origin.

The unpaired sternal bone (urohyal) cannot be homologous with the entosternal, for it has no connections with the annexes. He decides that it must represent the episternals, for in some young birds there is a two-headed episternal to which two strong tendons are attached, just in the same way as the unpaired piece in fish is bound to the bones of the hyoid by two tendons. "Thus it is not the sternum as a whole that has shifted in front of the clavicles and covered with its side pieces the gills placed there; it is a piece exclusively piscine, in the sense that it is only in the class of fishes that it reaches the maximum of its development" (p. 83).

To sum up, the sternum in all four vertebrate classes is composed of the same elements, arranged always in the same way. "One is ... led to the conception of an ideal type of sternum for all Vertebrates, which then, considered from a lower standpoint, resolves itself into several secondary forms according as the whole or the majority of the constituent materials are employed, or even as these elements come to change their respective dimensions or proportions" (p. 134). As to the elementary constituents, "they give proof of individuality, and sometimes even, in certain abnormalities, of independence, and rise to the level of primary organisatory materials" (p. 132). What holds good for the sternum holds good for other organs—and accordingly the unity of plan and composition can be demonstrated for all the organs of Vertebrates.