1200 



HORMONAL REGULATION 



time of sexual maturation, abundant evi- 

 dence of ovarian dysfunction. Tests given 

 after these female hermaphrodites had been 

 gonadectomized and injected with estrogen 

 and progesterone, or with an androgen, re- 

 vealed that striking modifications of the be- 

 havior pattern had been produced. 



Much less of the feminine measures of 

 behavior was displayed; there was a de- 

 crease in the percentage of tests positive for 

 estrus, in the duration of estrus, and in the 

 duration of maximal lordosis. An effect on 

 the tissues mediating the masculine compo- 

 nent of the pattern was revealed l)y the 

 greater amount of male-like mounting. The 

 hermaphroditic females had become more 

 responsive to the androgen than ovariecto- 

 mized but otherwise normal females. The 

 effects on the females receiving the andro- 

 gen prenatally were permanent. During 

 pregnancy there were no detectable effects 

 on the "mothers" into which the hormone 

 was being injected (Diamond, 1960), and 

 there were no lasting effects. Any effects 

 on normal young females treated with tes- 

 tosterone propionate postnatally from day 

 1 to day 80 were transitory (Phoenix, Goy, 

 Gerall and Young, 1959). 



The ovarian dysgenesis, clearly apparent 

 only after sexual maturity, was manifested 

 by disparities in the process of folliculo- 

 genesis. They varied from retardation of 

 follicular development, but with eventual 

 ovulation and formation of normal-appear- 

 ing corpora lutea, to anovulatory follicular 

 development. The dysgenesis is believed 

 to reflect damage, possibly in the nature of 

 a masculinization, to the hypothalamico- 

 pituitary gonadal axis and to provide evi- 

 dence for a central as opposed to an exclu- 

 sively peripheral action of the androgen 

 (Tedford and Young, 1960) . 



Some precocity of behavior may have 

 been shown by the male siblings; the pos- 

 sibility is being tested by Dr. A. A. Gerall. 

 However, after sexual maturity the behav- 

 ior of the male siblings was not significantly 

 different from that of the untreated controls 

 (Phoenix, Goy, Gerall and Young, 1959). 



The portion of the embryonic and fetal 

 periods during which the androgen must be 

 administered in order to masculinize the 

 female is being determined. At the time of 



the present writing it is clear that 40 mg. 

 testosterone propionate given between days 

 15 and 20 of gestation were without any de- 

 tectable effect on the behavior displayed 

 after the attainment of adulthood. How- 

 ever, the period 30 to 65 days of gestation 

 is critical in the sense that injections of 

 androgen need not be started iDefore this 

 time in order to suppress lordosis and in- 

 tensify masculinity (Goy, Bridson and 

 Young, 1961). The results from experiments 

 in progress may reveal that the critical 

 period is even shorter. 



The injection of estrogens into most preg- 

 nant female mammals that have been 

 studied soon terminates the pregnancy by 

 resorption or abortion, but some work has 

 been done with the chicken. Domm and 

 Davis (1948) observed the behavior of in- 

 tersexual brown Leghorn fowls resulting 

 from a single injection of each of several 

 estrogens on or before the 4th day of incu- 

 bation. vSome individuals were similar to 

 normal males in their general appearance, 

 but others were nearly indistinguishable 

 from females. The sexual reactions of the 

 intersexual males are described as varying 

 from essentially masculine in the case of 

 those transformed the least to neutral in 

 the case of those whose general appearance 

 was altered the most. Confusion and the 

 ability to perform only certain preliminary 

 phases of masculine behavior are noted, but 

 there was no squatting when they were 

 placed with normal males. 



The action of the exogenous hormone in 

 this experiment can be thought of as de- 

 pending on the interpretation of what hap- 

 pened. If the changes are regarded as hav- 

 ing been in the direction of feminization, 

 it could be considered that the estrogen left 

 its imprint on the tissues mediating mating 

 behavior and that a modification of organi- 

 zation occurred. But Domm and Davis 

 seem to have regarded the change as a 

 "displacement reaction," displayed when 

 the normal male pattern was impossible, 

 rather than as feminization. The abnormal 

 behavior was attributed to the low level of 

 hormone concentration. 



Use of the Sweltzer technique (Pincus 

 and Hopkins, 1958), which involves dipping 

 the eggs into solutions of steroids (or other 



