1046 



SUBMAMMALIAN VERTEBRATES 



terone) when fed to Lebistes interfered with 

 normal body growth in both sexes (Scott, 

 1944). The liver of Oryzias is sexually di- 

 morphic (luring the breeding season; if a 

 testosterone propionate pellet is implanted 

 in the female at this time, her liver acciuires 

 a typical male appearance (Egami, 1955a). 



It has already been shown that skin color 

 may be under the control of androgens, and 

 it is therefore not surprising that adminis- 

 tered male sex hormone may conspicuously 

 affect coloration. Thus, the injection of tes- 

 tosterone propionate into hypophysecto- 

 mized Fundulus or into Fimdulus during the 

 nonbreeding period evoked the yellow body 

 color typical of the spawning season (Bur- 

 ger, 1942). Nuptial coloration was produced 

 in Rhodeus, Acheilogriathus, Oryzias, and an 

 unspecified genus of bitterling by adminis- 

 tration of male hormone in various forms; 

 the androgen was given during the nonbreed- 

 ing season, or to castrated males, or to fe- 

 males. ^*^ The addition of pregneninolone to 

 a(|uarium water induced the ai)pearance of 

 male coloration, and also caused the disajv 

 pearance of the black "gravid spot" from 

 the tail fin of the female, in Lebistes (Ever- 

 sole, 1941; Regnier, 1941). 



Gonopodia and other male secondary sex 

 characters have also been studied as indi- 

 cators of androgen action. Gonopodia have 

 been produced in female Platypoecilus, Le- 

 bistes, Gmnbusia, and Molliensia by injec- 

 tions of testosterone propionate or by the 

 addition to the aquarium water of pregneni- 

 nolone (in some cases, 1 mg. in 14,000,000 

 ml. water) or methyl testosterone ( 1 mg. in 

 as much as 25,000,000 ml.)i" Treatment of 

 castrated male Gmnbusia with pregnenino- 

 lone or with testis grafts permits the normal 

 development of the gonopodium (St. Amant, 

 1941). If the anal fin, the female homologue 

 of the gonopodium of adult female Platy- 

 poecilus maculatus, is amputated and if any 

 of several androgens (androsterone and tes- 

 tosterone propionate are most effective) is 

 then injected, a male-like but atypical go- 



'" Gliiser and Haempel, 1932 ; Owen, 1937 ; Glaser 

 and Ranftl, 1938; Havas, 1939; Niwa, 1955. 



'"Eversole, 1941; Regnier, 1941; Turner, 1941b, 

 1942a, b, c; Cummings, 1943b; Hamon, 1945; 

 Gallien, 1948; Hopper, 1949; Tavolga, 1949; 

 Egami, 1954a. 



noiiodium is regenerated (Grobstein, 1940, 

 1942a, b, 1947). 



When adult lampreys become sexually 

 mature, ducts develop for the escape of ma- 

 ture germ cells from the mesonephros to the 

 exterior, and the cloacal labia undergo vas- 

 cular distention. Administration of either 

 testosterone or of estrone, a female sex hor- 

 mone, to adult but sexually immature 1am- 

 i:)reys will evoke the same changes (Knowles, 

 1939). 



Further evidence for the control of sexual 

 accessory structures by androgens is sum- 

 marized by Pickford and Atz (1957). 



A)nphibians 



Androgen has been supplied experimen- 

 tally by injection and by the transplantation 

 of testes. Testicular homotransplants failed 

 to restore male sex accessories in castrate 

 Rana in an early experiment (Smith and 

 Schuster, 1912), perhaps because the grafts 

 did not survive. Amplexus occurred promptly 

 after injection of a suspension of dried bull 

 testes into the dorsal lymph sacs of adult 

 male frogs ( Brossard and Gley, 1929) . Ponse 

 (1922a, b, 1930) restored male sex charac- 

 ters by testis grafting in castrate Bujo vul- 

 garis, as did Moszkowska (1932) in Bombi- 

 nator. The injection of androgens caused 

 development of the nuptial pads in Disco- 

 glossus (Kehl, 1944a) but not in Xenopus 

 (Berk, 1939). Curiously, ovulation could be 

 induced in the latter genus both by injecting 

 the intact animal with any of several andro- 

 gens or other steroids (Shapiro, 1939) or by 

 adding testosterone and androstenedione to 

 a frog Ringer solution in which an excised 

 ovary was suspended (Shapiro and Zwaren- 

 stein, 1937). Implantation of testosterone 

 propionate tablets into castrate male and fe- 

 male toads (Bufo vulgaris) resulted in the 

 development of male accessories in both 

 sexes (Harms, 1950). Similar treatment be- 

 fore the breeding season of both male and 

 female cricket frogs {Acris gryllus) evoked 

 male skin coloration and hypertrophy of 

 Wolffian ducts (both sexes), oviducts, and 

 seminal vesicles (Greenberg, 1942). Injec- 

 tions of the same hormone into castrate male 

 Rana pipiens resulted in growth of the nup- 

 tial pads and vestigial oviducts (Wolf, 1939) . 



Bresca (1910) made an interesting ob- 



