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

by E. S. Russell

The fullest development of Geoffroy's views on evolution is found in his memoir "Le degré d'influence du monde ambiant pour modifier les formes animales."[100] Here the relation of his evolution-theory to his morphology is pointed out. The principle of unity of plan and composition cannot be the final goal of zoology; there must follow on it a philosophical study of the differences between organic forms. The causes of these differences are to be found in the environment (pp. 66-7). Geoffroy seems here to be moving from a pure to a causal morphology. It is probable, he continues, that living species have descended by uninterrupted generation from the antediluvian species (p. 74), and that they have in the process become modified through external influences.

Now of all functions respiration is the most important, and upon respiration everything is regulated. "If it be admitted that the slow progression of the centuries has brought in its train successive changes in the proportion of the different elements of the atmosphere, it follows as a rigorously necessary consequence that the organisation has been proportionately influenced by them" (p. 76). The respiratory milieu changes, the species change with it, or are eliminated (p. 79). We may see, perhaps, in the stress which Geoffroy lays upon respiration and the respiratory milieu a result of his constant obsession with the comparison of fish with air-breathing Vertebrates.

In the first geological period, we read in another Memoir of the same year,[101] when ammonites and Gryphæa flourished, hot-blooded animals with lungs could not exist. "A lung constructed like that of mammals and birds would not have been adapted to the essence of the respiratory element such as in my conception of it the system of the environing air used to be"[102] (p. 58).

Geoffroy does not tell us exactly how the milieu is to act upon the organism; the whole theory is little more than a sketch and a pointing out of the way for future research—and in this prophetic enough. The action of external agents was apparently considered as physical, and no power of active adaptation was ascribed to the organism.

From a passage in the memoir "Sur la Vertèbre" we may perhaps infer that he believed increasing complexity of structure to be due to a realisation of potentialities, to the development of parts present in the lower animals only in potency—"the organisation ... only awaits favourable conditions to rise, by addition of parts, from the simplicity of the first formations to the complication of the creatures at the head of the scale" (p. 112). Evolution takes place as the environment allows, and in a sense in opposition to the environment.

He believed in saltatory evolution, for he considered that the lower oviparous Vertebrates could not be transformed into birds by slow modification, but only by a sudden transformation of their lungs, which would bring about the other characteristics of birds (p. 80). He considered, too, that transformations could arise by means of monstrous development (p. 86). In this connection the experiments which he made on the hen's egg[103] in order to produce artificial monstrosities are significant, though his purpose was rather to obtain proof of the inadequacy of the preformation hypothesis.[104]

It seems probable enough that if Geoffroy had developed his views on evolution he would finally have been led to interpret unity of plan in terms of genetic relationship. But as it was he remained at his morphological standpoint. He did not interpret rudimentary organs as useless heritages of the past; he preferred to think that Nature had prepared double means for the same function, one or other being predominant according as the animal lived in the water or on the land. "To the animal that lives exclusively in the air Nature has granted an organisation suited to this mode of respiration, without however suppressing the other corresponding means, that is to say, without depriving it of a second system which is applicable only to the mode of respiration by the intermediary of water, and vice versa."[105]

He seems, in one instance at least, to have hit upon the root-idea of the biogenetic law, but he was far from appreciating its significance. He recognised that an amphibian in its development passed through a stage when it was in all essentials similar to a fish, and he saw in this visible transformation a picture of the evolutionary transformation. "An amphibian," he writes,[106] "is at first a fish under the name of tadpole, and then a reptile [sic] under that of frog.... In this observed fact is realised what we have above represented as an hypothesis, the transformation of one organic stage into the stage immediately superior." But it is not clear that he considered the development of the amphibian to be a repetition of its ancestral history.

He went, however, a certain length towards recognising the main principle of a law which was a commonplace of German transcendental thought, and was developed later by his disciple E. Serres, the law that the higher animals repeat during their development the main features of the adult organisation of animals lower in the scale. Thus he compared fish as regards certain parts of their structure with the fœtus of mammals. He compared also Articulates with embryonic Vertebrates in respect of their vertebræ, for in the higher Vertebrates the body of the vertebra is tubular at an early stage of development, and in Articulates the body of the vertebra remains tubular permanently (supra, p. 61). As regards their vertebræ, "insects occupy a place in the series of the ages and developments of the vertebrate animals, that is to say, they realise one of the states of their embryo, as fishes do one of the states of their fœtal condition."[107]

This idea was destined to exercise a great influence upon the development of morphology. A further development of the thought is that certain abnormalities in the higher animals, resulting from arrest of development, represent states of organisation which are permanent in the lower animals.[108]