The corresponding changes in the thumb-pads are as follows. Immediately after the breeding season the horny epidermis of the pad with its deeply pigmented papillae is cast off, and the thumb remains comparatively smooth from April or May until August or September. When the large papillae are shed, smaller papillae remain beneath, and are gradually obliterated by the epidermis growing up between them. The epidermis is therefore growing while the spermatogenesis is taking place. In August and September the epidermic papillae begin to be obvious, and from this time till February a continuous increase in the papillae and their pigmentation occur. Geoffrey Smith argues that the development of this somatic character occurs while the testes are inactive and unchanged. Considering that the testes throughout the winter months are crammed with spermatozoa, which must require some nourishment, and which may be giving off a hormone all the time, the argument has very little weight. Smith and Schuster found that ovariotomy, with or without subsequent implantation of testes or injection of testis extract, had no effect in causing the thumb of the female to assume any male characters.
Castration during the breeding season causes the external pigmented layer with its papillae to be cast off very soon—that is to say, it has the same effect as the normal discharge of the spermatozoa. Smith and Schuster found that castration at other seasons caused the pad to remain in the condition in which it was at the time, that there was no reduction or absorption as Nussbaum and Meisenheimer found, and that allo-transplantation of testes—that is, the introduction of testes from other frogs either into the dorsal lymph-sacs or into the abdominal cavity—or the injection of testis extract, had no effect in causing growth or development of the thumb-pad.
There seems to be one defect in the papers of both Nussbaum and Smith and Schuster—namely, that neither of them mentions or apparently appreciates the fact that the thumb-pads, apart from the dermal glands, consist of horny epidermis developed from the living epidermis beneath. The horny layer is not shown clearly in the figures of Smith and Schuster. It seems impossible that the horny layer or its papillae could atrophy in consequence of castration, or be absorbed. The horny part of the frog's thumb-pad is comparable with the horny sheath of the horns in the mammalian Prong-buck (Antilocapra) which are shed after the breeding season and annually redeveloped. Meisenheimer claims that he produced development of papillae on the thumb-pad, not only by implantation of pieces of testis, but also by implantation of pieces of ovary. This seems so very improbable that it suggests a doubt whether the same investigator was not mistaken with regard to the results of his experiments in transplanting gonads in Moths.
Smith and Schuster conclude that the normal development of the thumb-pad depends on the presence of normal testes, but that there is no sufficient evidence that the effect is due to a hormone derived from the testis. It is equally probable, according to Smith, that the testicular cells take up some substance or substances from the blood, thus altering the composition of the latter and perhaps stimulating the production of these substances in some other organ of the body. These substances may be provisionally called sexual formative substances. Smith's theory therefore is that the action of the testes in metabolism is rather to take something from the blood than to add something to it, and that it is this subtractive effect which influences the development of somatic sexual organs.
Geoffrey Smith in fact, in the paper above considered, attempts to apply to the frog the views he put forward [Footnote: Fauna und Flora des Golfes van Neapel, 29 Monographie Rhizocephala.] in relation to the effect of the parasite Sacculina on the sexual organs of crabs. The species in which he made the most complete investigation of the influence of the parasite was Inachus scorpio (or dorsettensis). Figures showing the changes in the abdomen produced by the presence of Sacculina are given in Doncaster's Determination of Sex, Pl. xv. Sacculina is one of the Cirripedia, and therefore allied to the Barnacles. It penetrates into the crab in its larval stage, and passes entirely into the crab's body, where it develops a system of branching root-like processes. When mature the body of the Sacculina containing its generative organs forms a projection at the base of the abdomen of the crab on its ventral surface, and after this is formed the crab does not moult. Crabs so affected do not show the usual somatic sexual characters, and at one time it was supposed that only females were attacked. It is now known that both sexes of the host may be infected by the parasite, but the presence of the latter causes suppression of the somatic sex-differences. The entry of the parasite is effected when the crab is young and small, before the somatic sex-characters are fully developed. The gonads are not actually penetrated, at least in some cases, by the fibrous processes of the parasite, but nevertheless they are atrophied and almost disappear. In Inachus the abdomen of the normal male is very narrow and has no appendages except two pairs of copulatory styles. The abdomen of the female is very broad, and has four pairs of biramous appendages covered with hairs, the normal function of which is to carry the eggs. The effect of the parasite in the male is that the abdomen is broader, the copulatory styles reduced, and biramous hairy appendages are developed similar to those of the female, but smaller. In the female the abdomen remains broad, but the appendages are much smaller than in the normal female, about equal in size to those of the 'sacculinised' male. Smith interpreted the alteration in the male as a development of female secondary characters, but it is obvious from the condition in Macrura or tailed Decapods, like the lobster or crayfish, that the abdomen or tail of the male originally carried appendages similar to those of the female, and that the male character is a loss of these appendages. The absence of the male character therefore necessarily involves a development of these appendages, and there is not much more reason for saying that the male under the influence of the parasite develops female characters, than for saying that the male character is absent. There is no evidence in the facts concerning parasitic castration for Geoffrey Smith's conclusion that the female characters are latent in the male, but the male characters not latent in the female: both return to a condition in which they resemble each other, and the primitive form from which they were differentiated.
By his studies of parasitic castration Geoffrey Smith was led to formulate a theory for the explanation of somatic sex-characters different from that of hormones. He found that in the normal female crab the blood contained fatty substances which were absorbed by the ovaries for the production of the yolk of the ova. When Sacculina is present these substances are absorbed by the parasite; the ovary is deprived of them, and therefore atrophies. In the male the parasite requires similar substances, and its demand on the blood of the host stimulates the secretion of such substances, so that the whole metabolism is altered and assimilated to that of the female. It is this physiological change which causes the development of female secondary characters. He describes this change as the production of a hermaphrodite sexual formative substance, on the ground that in at least one case eggs were found in the testis of a male Inachus which had been the host of a Sacculina, but had recovered. It must however be noted that the Sacculina itself is hermaphrodite, with ovaries much larger than the testes. It is possible that while the parasite prevents the development of testis or ovary in the host, it gives up to the body of the host a hormone from its own ovaries which tends to develop the female secondary characters: for the parasite is itself a Crustacean, and therefore the hormone from its ovaries would not be of too different a nature to act upon the tissues of the host.
The observation of Geoffrey Smith that eggs may occur in the testis of a crab after recovery from the parasite appears of more importance than his peculiar theoretical suggestions, for it tends to show that sex is not always unalterably fixed at fertilisation. In this case the influence of a parasite predominantly female would seem to be the real cause of the development of eggs in the testis of the host. Geoffrey Smith does not discuss the origin of the somatic sexual characters in evolution, or attempt to show how his theories of sexual formative substance, and of the influence of the gonads by subtraction rather than addition, would bear upon the problem.
CHAPTER VI
Origin Of Non-Sexual Characters: The Phenomena Of Mutation
According to the theory here advocated, modifications produced by external stimuli in the soma will also be inherited in some slight degree in each generation when they have no relation to sex or reproduction. In this case the habits and the stimuli which they involve will be common to both sexes, and the hormones given off by the hypertrophied tissues will act upon the corresponding determinants in the gametocytes. The modifications thus produced will therefore be related to habits, and the theory will include all adaptations of structure to function, but other characters may also be included which are the result of stimuli and yet have no function or utility.