Dr. Wilson, T. H. Morgan, and Richard Hartwig have therefore suggested that the sex-difference as regards gametes is not a qualitative but a quantitative one. In certain cases there is no evident quantitative difference of chromatin as a whole, but there may in all cases be a difference in the quantity of special sex-chromatin contained in the X element. The theory put forward by Wilson then is that a single X element means per se the male condition, while the addition of a second element of the same kind produces the female condition. Such a theory might apply even to cases where no sex-chromosomes can be distinguished by the eye: the ova, in such cases (probably the majority), might also have a double dose of sex-chromatin, the males a single dose. This theory, however, is still open to the objection that the female gametes before fertilisation, and half the male gametes, have the half quantity of sex-chromatin which by hypothesis determines the male condition, so that here again we have the male condition as something which is distinct from the characteristics of the spermatozoon. But if this is the case, what is the male condition? The parthenogenetic ovum of the bee is male, and yet it is an ovum capable only of producing spermatozoa. If the single X chromosomes is the cause of the development of spermatozoa in the male bee, why does it not produce spermatozoa in the gametes of the female bee, since when reduction takes place all these gametes have a single X chromosome?
In biology, as in every other science, we must admit facts even when we cannot explain them. The facts of what we call gravitation are obvious, and any attempt to disregard them would result in disaster, yet no satisfactory explanation of gravitation has yet been discovered: many theories have been suggested, but no theory has yet been proved to be true. In the same way it may be necessary to admit that two X chromosomes result in the development of a female, and one X, or XY chromosomes result in the development of a male. But Mendelians have omitted to consider what is meant by male and female. The soma with its male and female somatic characters has nothing to do with the question, since somatic sex-differences may be altogether wanting, and moreover, the essential male character, the formation of spermatozoa, is by the Mendelian hypothesis due to descent of the male gametes from the original fertilised or unfertilised ovum. The Mendelian theory therefore is that when an ovum has two X sex-chromosomes it can only after a number of cell-divisions, at the following reduction division, give rise to ova, while an ovum containing one X sex-chromosome, or two different, XY, chromosomes, at the next reduction division gives rise to spermatozoa. The X sex-chromosome is not in itself either female or male, since, as we have seen, either ovum or spermatozoon may contain a single X chromosome. The ovum then with one X chromosome or one X and one Y changes its sex at the next reduction division and becomes male. In parthenogenetic ova this happens without conjugation with a spermatozoon at all: in other cases, since the zygote is compounded of spermatozoon and ovum, we can only say that in the XX zygote, the ovum developing only ova, the female is dominant, in the X or XY zygote developing only spermatozoa the male is dominant. Hermaphrodite animals, as has been pointed out by Correns and Wilson, cannot be brought under this scheme at all. In the earthworms, for instance, we have, in every individual developed from a zygote, ova and spermatozoa developing in different gonads in different parts of the body. The differentiation here, therefore, must occur in some cell-division preceding the reduction divisions. Every zygote must have the same composition, and yet give rise to two sexes in the same individual.
Further light on the sex problem, as in many other problems in biology, can only be obtained by more knowledge of the physical and chemical processes which take place in the chromosomes and in the relations of these structures to the rest of the cell. The recent advances in cytology, remarkable as they are, consist almost entirely of observations of microscopic structure. They may be said to reveal the statics of the cell rather than its dynamics. Cytology is in fact a branch of anatomy, and in the anatomy of the cell we have made some progress, but our knowledge of the physiology of the cell is still infinitesimal. The nucleus, and especially the chromosomes, are supposed in some unknown way to influence or govern the metabolism of the cytoplasm. From this point of view the hypothesis mentioned above that the sex-difference in the gametes is not qualitative but quantitative is probably nearer to the truth. Geddes and Thomson and others have maintained that the sex-difference is one of metabolism, the ovum being more anabolic, the sperm more katabolic. A double quantity of special chromatin may be the cause of the greater anabolism of the ovum. In that case the difficulty indicated in a previous part of this chapter, that the ovum after reduction resembles the sperm in having only one X chromosome, may be explained by the fact that the growth of the ovum and its accumulation of yolk substances has been already accomplished under the influence of the two chromosomes before reduction. Other difficulties previously discussed also appear to be diminished if we adopt this point of view. We need not regard maleness and femaleness as unit characters in heredity of the same kind as Mendelian characters of the soma. Instead of saying that the zygote composed of ovum and spermatozoon is incapable of giving rise in the male to ova, or in the female to sperms, we should hold that the gametocytes ultimately give rise to ova or to sperms according to the metabolic processes set up and maintained in them through their successive cell-divisions under the influence of the double or single X chromosome. There still remains the difficulty of explaining why the male gametocytes after reduction develop into similar sperms, with their heads and long flagella, although half of them possess one X chromosome each and the other half none. We can only suppose that the final development of the sperms is the result of the presence of the single X chromosome in the successive generations of male gametocytes before the reduction divisions.
The Mendelian theory of sex-heredity assumed that in the reduction divisions the two sex-characters, maleness and femaleness, were segregated in the same way as a pair of somatic allelomorphs, but the words maleness and femaleness expressed no real conceptions. The view above suggested merely attempts to bring our real knowledge of the difference between ovum and sperm into relation with our real knowledge of the sex-chromosomes and their behaviour in reduction and fertilisation.
CHAPTER III
Influence Of Hormones On Development Of Somatic Sex-Characters
We have next to consider what are commonly called secondary sexual characters. These are characters or organs more or less completely limited to one sex. When we distinguish in the higher animals the generative organs or gonads on the one hand from the body or soma on the other, we see that all differences between the sexes, except the gonads, are somatic, and we may call them somatic sexual characters. The question at once arises whether the soma itself is sexual, that is to say, whether on the assumption that the sex of the zygote is already determined before it begins to develop, the somatic cells as well as the gametocytes are individually and collectively either male or female. In previous discussions of the subject I have urged that the only meaning of sex was the difference between the megagamete or ovum, and the microgamete or sperm. But if the zygote, although compounded of ovum and sperm, is predestined to give rise in the gametes descended from it, either to sperms only or to ova only, it may be suggested that all the somatic cells descended from the zygote are likewise either male or female, although they do not give rise to gametes. It is evident, however, that the somatic cells, organs, and characters do not differ necessarily or universally in the two sexes. On the one hand, we have extraordinary and very conspicuous peculiarities in the male, entirely absent in the female, such as the antlers of stags, and the vivid plumage of the gold pheasant; on the other we have the sexes externally alike and only distinguished by their sexual organs, as in mouse, rabbit, hare, and many other Rodents, most Equidae, kingfisher, crows and rooks, many parrots, many Reptiles, Amphibia, Fishes, and invertebrate animals. In the majority of fishes, in which fertilisation is external and no care is taken of the eggs or young, there are no somatic sexual differences. Moreover, somatic sexual characters where they do occur have no common characteristics either in structure or position in the body. It may be said that any part of the soma may in different cases present a sex-limited development. In the stag the male peculiarity is an enormous development of bone on the head, in the peacock it is the enlargement of the feathers of the tail. In some birds there are spurs on the legs, in others spurs on the wings. It is no explanation, therefore, to say that these various organs and characters are the expression of sex in the somatic cells.
As I pointed out in my Sexual Dimorphism (1900), the common characteristic of somatic sexual characters is their adaptive relation to some function in the sexual habits of the species in which they occur. There is no universal characteristic of sex except the difference between the gametes and the reproductive organs (gonads) in which they are produced. All other differences, therefore, including genital ducts and copulatory or intromittent organs, are somatic. When we examine these somatic differences we find that they can be classified according to their relation to fertilisation and reproduction, including the care or protection of the offspring. The precise classification is of no great importance, but we may distinguish the following kinds to show the chief functions to which the characters or organs are adapted.
1. GENITAL DUCTS AND INTROMITTENT ORGANS.—According to the theory of the coelom which we owe to Goodrich, in all the coelomata the coelom is primarily the generative cavity, on the walls of which the gametocytes are situated, and the coelomic ducts are the original genital ducts. In Vertebrates we find two such ducts in both sexes in the embryo, originally formed apparently by the splitting of a single duct. In the male one of these ducts becomes connected with the testis while the other degenerates: the one which degenerates in the male forms the oviduct in the female, while the one which is functional in the male degenerates in the female.
Intromittent organs are formed in all sorts of different ways in different animals. In Elasmobranchs (sharks and skates) they are enlarged portions of the pelvic fins, and therefore paired. In Lizards they are pouches of the skin at the sides of the cloacal opening. In Mammals the single penis is developed from the ventral wall of the cloaca. In Crustacea certain appendages are used for this function. There are a great many animals, from jelly-fishes to fishes and frogs, in which fertilisation is external, and there are no intromittent organs at all.