PART IV. SUMMARY AND CONCLUSIONS.

1. The two principal results obtained were: (a) that castration of hen-feathered Sebright males causes them to develop the full plumage characteristic of the cock-bird; (b) that complete hen-feathering is due to two dominant Mendelian genes.

2. A striking change takes place when the Sebright male is castrated (plate 1, figs. 3, 4; plate 3, fig. 1). The new feathers on the upper surface of the head, neck, back, wings, rump, and tail-coverts assume a different color and distribution of their pigment; they take on a new shape, and in those regions where in the cock the barbules are absent from a part of the margin of the feather, the same absence occurs in the castrated birds. Such feathers are present on the neck, back, wing-bow, and rump. The transition is shown in the figures in plate 6, where for comparison one of the old and one of the new feathers lie side by side. The tail-coverts in the hen-feathered bird are short, and like those in the hen do not cover the true tail. After castration they become excessively long—longer, in fact, than in many cocks—and cover the true tail feathers. The tail feathers themselves, moreover, become increased in length, as do the posterior row of feathers of the wing-coverts. On the breast and sides the change is less marked. The castrated Sebright loses his erect carriage, but how far this is due to the changes in his plumage and how far is real (as a result of a new balance due possibly to the lengthening tail and its coverts) I can not decide.

3. While castration causes the hen-feathered male to make additions in color, length, and size of many feathers, it causes at the same time the other retrogressive changes characteristic of the capon (a castrated cock-feathered bird); the comb and wattles shrink and become pale, the birds almost cease crowing, and become timid. They do not make much effort to mate with the hens, but when they do they show the usual copulatory reactions.

4. If feathers are removed at the time of castration, the new feathers show the full effect of the removal of the testes, although they must have begun to develop immediately afterward. It is suggested that by means of this delicate test the time relations of the internal secretion can be profitably studied.

5. Feathers that may have started their development at the time of the operation show the old influence at the tip of the feathers (plate 10) and the new one in the rest of the feather. The change is abrupt, although the transition is perfect.

6. Incomplete castration of the hen-feathered male leads to smaller changes in the same direction than those following complete castration.

Where such small pieces of the testis were left that complete cock-feathering followed, the bird slowly changed back to hen-feathering as the testes began to regenerate. When the regenerated pieces were removed the bird became cock-feathered again.

7. One Sebright male whose testes appear to have been completely removed did not change the character of the plumage. No testes were found on autopsy. It is suggested that some other endocrine organs have taken over the function of the testes, but as yet none such can be indicated.

8. In one case an old hen-feathered (F₁) male began to change over to cock-feathering. It was found that his testes had dwindled (probably through disease) to very small size (10 by 5 mm.).

9. The F₁ male of the cross between the Sebright and game is also hen-feathered (plate 2, fig. 1). After castration he becomes cock-feathered (plate 2, fig. 4) and shows thereby the genetic type of the heterozygous cock-feathered class in which his hen belongs. The change in this male is even more striking than that in the Sebright. The change in the individual feathers is shown in plate 7, figs. 1 and 1a.

10. Three types of F₂ hen-feathered castrated males are shown in plate 2, figure 3, and plate 3, figure 3 and figure 4. The first was a dark bird that changed to a lighter red above. The third a gray bird that became bright red; the second was a light yellow that became deep yellow, etc. The class of hens to which such males belong, as cock-feathered birds, can thus be found out by castration. In this way the F₂, and back-cross, hen-feathered cocks can be classified with the corresponding F₂ cock-feathered males.

11. In the F₂ generation, made up of birds from the direct and reciprocal crosses taken together, there were 29 hen-feathered and 26 cock-feathered males. In the back-cross (F₁ hen by game male) the classes were 2 and 7. The results seem in better accord with the assumption that two factors are present in the Sebright that stand for hen-feathering; that either alone will give hen-feathered birds (intermediate type?), but that both together give the extreme type of hen-feathering seen in the Sebright.

12. The difference in color in the two races (Sebright and Black Breasted Game bantams) is very great. The former have almost uniformly laced feathers, while the latter has the varied plumage of the jungle-fowl. The game is strongly dimorphic in color and color-pattern; the Sebright has the same type of coloration and pattern both in the male and female, but this is deceptive, as castration shows, because the castrated male is as strikingly different from the normal Sebright female as is the cock of other birds from the hen. The resemblance of male and female in this race is due to the suppression of the true male plumage by something produced in the testes. Therefore the heredity of dimorphism resolves itself here into the problem of the heredity of hen-feathering. That the female Sebright has the same genetic factors as the male is shown by the fact that she transmits hen-feathering in the same way as does the male, and also by the fact, as Darwin pointed out, that an old female Sebright whose ovaries had degenerated developed not the hen-feathered plumage of her own cock, but cock-feathered plumage like that of most male poultry.

13. The color of the F₁ birds is shown in plate 2, figs. 1 and 2. In general, the feathers are stippled, black and light yellow being the two most conspicuous ingredients. Since hen-feathering dominates, the dimorphism is absent, or at least is so slight as to not attract attention—little more, in fact, than in the Sebright race. The carriage of the male is like that of the Sebright male. The F₁ male and female are alike in the direct cross and the reciprocal, or at least no conspicuous difference is found between the two classes of hens, indicating that no important sex-linked factors are involved in the cross.

14. The F₂ birds show a great variety of color and pattern, but those obtained can be approximately grouped into 16 classes. The classes are, however, admittedly not uniform, indicating minor factors not here reckoned with. The classification of the hens is easiest; the F₂ hen-feathered males can then in many cases be referred to the proper classes; the F₂ cock-feathered males can not be accurately classified with their corresponding hens, except in the case of those that resemble the two P₁ males, the F₁ male, and those that castration experiments of the hen-feathered males have shown to belong to certain hen types.

15. Despite the admitted difficulties of classification, it is suggested that three factor-pairs of differences will cover the main color classes seen in the F₂ and in the back-cross. One or two of these seem to be incompletely dominant, since the F₁ birds are not like either parent in any single character, nor are they like the wild type in so far as this is represented by the game.

16. A histological examination of the testis of the male Sebright by Boring and Morgan has shown that it contains cells like those present in the ovary of all breeds of poultry. These cells are called luteal cells by Pearl and Boring, from their resemblance to the cells of that name found in the corpora lutea of mammals. In the mammals similar cells are supposed to produce internal secretions that act as hormones. Their function in the female bird is unknown, but the fact that after the removal of the ovary the female develops the secondary sexual plumage of the male suggests that some secretion from these cells performs this function. Their occurrence in the male Sebright and their complete absence, or paucity, in the males of other races supports strongly the view that these cells are concerned with the suppression of the secondary sexual plumage.

17. While in mammals the interstitial cells have been supposed to produce an internal secretion that causes the development of some of the secondary sexual characters of the male, and the fuller elaboration of others, in birds no such connection exists, if we except the case of the Sebright. Castration of ordinary males does not affect deleteriously the secondary sexual plumage (although it does the comb, behavior, etc.), in fact may even enhance their effects. But, while in the mammal a secretion is necessary for the full development of the secondary sexual characters, in the Sebright a secretion inhibits certain of them. What element in the ordinary bird and in the Sebright causes the full development of the comb, wattles, sexual behavior, etc., is not known. Possibly it is the sexual elements themselves, but possibly it is a secondary influence of the luteal cells producing a contrary effect on these parts from its effects on the feathers; but possibly more than one kind of secretory cell is present in the testis of the cock.

18. The causes of the development of the secondary sexual characters are seen to be of such diverse physiological kinds that one may well hesitate to apply the same explanation as to their evolution. In fact, it is pointed out that several of the theories that have been suggested run counter to the conditions that bring about the development of the secondary sexual characters.

19. An attempt is made to give a critical review of Darwin’s theory of sexual selection in the light of the modern genetic and operative results on the secondary sexual characters of the vertebrates. It is pointed out that far from extending the general theory in its applications, the modern work has shown in the first place that the underlying conditions that call forth the development of the secondary sexual differences are so diverse in the different groups of animals that it is a priori very unlikely that this evolution can have been directed by the same external agent, such as the choice of the female, for such an assumption carries with it in several cases other implications concerning the causes of the suppression of these same characters in the female herself, etc. In the second place, it is pointed out that the problem of the excessive development of certain characters in the male whose genes are present in both sexes no longer oppresses us as it did Darwin, for it has been shown both by the genetic and by the operative work that a single factorial difference may be at the root of exceedingly great differences in the individual. Such results, while they admittedly do not in most cases tell us that the differences involved have arisen at a single progressive step, show us nevertheless that such differences may depend on very simple initial differences, and if so, the entire problem becomes enormously simplified. To Darwin the excessive development of color and ornamentation appeared due to a long, slow process of evolution laboriously brought about by the female through selection of those males a little more ornamented than their fellows. To-day we have found out that in many cases the genetic composition of a male with such ornamentation and of a female without it may be almost identical, except that the genes in one chromosome are duplex in one sex and simplex in the other. Owing to this initial difference, the female in birds produces an internal secretion that suppresses in her the ornamentation shown by the male, and in the mammal an internal secretion produced by the testes causes the full development in the male of the secondary sexual characters. If, as seems probable, these secretions are some particular kind of substance, the condition that led to their appearance historically need not have been very complex; and if not, the problem appears simplified. It still remains to give some reasonable explanation as to why such substances should continue to be produced if their products—the secondary sexual characters—possess no “beauty” for the female. Here more work is necessary, but the modern genetic point of view may possibly give an important clue. We are coming to realize more fully that the hereditary genes generally have more than a single effect on the characters of the animal. The secondary sexual characters may, then, be only by-products of genes whose important function lies in some other direction. If, for example, the secretion produced by the cells of the male have an important influence on his output of energy, or strength, or activity, their secondary influence over certain parts of the body would not call for any further explanation on the modern view of natural selection. If the secretions of the ovary of the female bird have some direct relation to her physiological processes that are important in the development of the oviduct, for instance, it would be a matter of no importance from an evolutionary point of view if that same secretion suppresses in her the development of the high color shown by the male.