1198 



HORMONAL REGULATION 



rudimentary. In Beach's experiments each 

 female received 24 mg. over a period of 

 2 weeks. This amount of androgen was 

 followed by a shift from a peak at the 

 level of "sexual clasp" (the measure of 

 lowest degree) to a peak at "palpation and 

 thrust," the next higher measure. The 

 complete pattern, i.e., palpation with pelvic 

 thrusts and a backward lunge (copulation), 

 composed 8 per cent of the total responses 

 compared with 1 per cent after ovariectomy, 

 and 60 to 65 per cent in normal males. It 

 is evident from a later publication (Beach 

 and Holz-Tucker, 1949) that 50 to 75 ^g- 

 testosterone propionate daily (0.7 to 1.0 

 mg. over a 2-week period) is sufficient to 

 maintain this level of behavior in the male. 

 We assume that the display of the copula- 

 tory response in 8 per cent of the tests 

 when females were injected required about 

 three times the amount of hormone that 

 was necessary for induction of the cor- 

 responding response in 60 to 65 per cent of 

 the tests when males were injected. 



Some female guinea pigs are responsive to 

 testosterone propionate. Frequent injections 

 or pellet implantation is followed by a 

 random display of mounting. In addition, 

 that displayed by spayed animals contain- 

 ing pellets and injected with estrogen and 

 progesterone exceeds by far that exhibited 

 in response to the same treatment before 

 pellet implantation or after the pellet has 

 been absorbed (Goy and Young, 1958). 

 On the other hand, the androgen given these 

 animals was not effective in strain 2 females 

 which do not show mounting behavior spon- 

 taneously. 



When explanation is sought for what 

 might seem to be anomalous effects it is 

 clear that more than one problem exists. 

 The estrogenic stimulation of masculine 

 behavior in the male (relationship 2) and 

 the androgenic stimulation of feminine re- 

 actions in the female (relationship 6) may 

 well be comparable with the many actions 

 of heterosexual hormones on the genital 

 tracts and accessories described by Bur- 

 rows (1949) in his review. A mimicking 

 action by heterosexual hormones is com- 

 mon and in studies involving the tissues 

 mediating behavior, as in studies of tissues 

 of the genital tract (Hisaw, 1943), the ef- 



fective quantity of the heterosexual hor- 

 mones tends to be large. This fact alone 

 attests to a considerable degree of speci- 

 ficity. 



The stimulation of feminine behavior in 

 the male (relationships 3 and 4) and of 

 masculine behavior in the female (relation- 

 ships 7 and 8) is another problem. It was 

 long ago suggested that each sex contains 

 the mechanism capable of mediating the 

 behavior of the opposite sex (Goodale, 

 1918; Sand, 1919; Pezard, 1922; Finlay, 

 1925; Zawadowsky, 1926; Beach, 1938, 

 1941, 1945a), and that under suitable stim- 

 ulation the behavior of the opposite sex is 

 disi^layed. We reaffirm the general truth of 

 this hypothesis, but will point out that 

 often in many lower vertebrates and mam- 

 mals the hormone of the opposite sex is not 

 a "suitable" stimulus to induce the full be- 

 havior of that sex. The point is illustrated 

 by what is seen in the relationships 3, 4, 

 7, and 8. The tissues mediating masculine 

 behavior in females are very responsive to 

 estrogens (relationship 7). They will re- 

 spond to androgens (relationship 8), but 

 not infrequently the behavior is obtained 

 only with difficulty and at a low level. 

 Rarely, as in the case described by Beach 

 (1945aj, the tissues mediating feminine be- 

 havior in males are very responsive to an- 

 drogens (relationship 3) ; more often, how- 

 ever, they are refractory. They are also 

 refractory to estrogens (relationship 4) and 

 this fact is reflected by the difficulty in 

 stimulating feminine responses in males 

 when estrogens are administered. We feel 

 that the responses members of the different 

 sexes give to hormonal stimulation are pre- 

 determined; in this sense a specific rather 

 than a nonspecific relationship exists. This 

 view was well expressed by Boss (1943) 

 when he wrote: ". . . genetical factors are 

 strongly involved, not only in determining 

 the specific behavior pattern of the taxo- 

 nomic unit, but also of the sex. It is a widely 

 if not a generally valid rule that the same 

 amounts of a hormone do not produce iden- 

 tical reactions in the two sexes of a given 

 species." The view is also consistent with 

 the conclusions reached by Burns (1942, 

 1949) during his studies of the role of fetal 

 gonadal hormones in sexual differentiation. 



