HORMONES AND MATING BEHAVIOR 



1199 



Before leaving the subject of the speci- 

 ficity of the relationship between hormone 

 and response, we would recall the results 

 from the numerous attempts to substitute 

 adrenal steroids and other compounds for 

 progesterone (Hertz, Meyer and Spielman, 

 1937; van Heuverswyn, Collins, Williams 

 and Gardner, 1939; Soderwall, 1940; Torst- 

 veit and Mellish, 1941; Isaacson, 1949; 

 Melampy, Emmcrson, Rakes, Hanka and 

 Eness, 1957 ) . Heat behavior was induced in 

 ostrogen-conditioned females by a number 

 of these substances, but their relative effec- 

 tiveness was much less than that of pro- 

 gesterone. In a study in which the female 

 guinea pig was used the relative effective- 

 ness of a number of adrenal steroids and 

 similar compounds was from !4 (desoxy- 

 corticosterone acetate) to 1/40 (11-keto- 

 progesterone, 11 -hydroxy progesterone, and 

 corticosterone). Hydrocortisone, cortisone, 

 17-hydroxy-desoxycorticosterone, 17-hy- 

 droxy progesterone, 21 -desoxy hydrocorti- 

 sone, adrenosterone, 4-androstene-3,17- 

 dione, and the amorphous fraction were 

 ineffective or had only slight activity 

 (Byrnes and Shipley, 1955 ) . 



B. THE MANNER OF HORMONE ACTION, I.E., 

 ORGANIZATION OR ACTIVATION 



An important problem pertaining to the 

 hormonal regulation of mating behavior has 

 to do with the manner of hormone action. 

 It has long been recognized that these sub- 

 stances might organize patterns of behavior 

 (Stcinach, 1912, 1913, 1916; Sand, 1919; 

 Lipschiitz, 1924; Nissen, 1929); in the 

 opinion of the writer, this would be in the 

 sense of producing changes in the resjionscs 

 to hormones different from those normally 

 associated with an individual, giving due 

 regard to age, sex, strain, and species. Or, 

 the hormones might simply activate an in- 

 dividual to respond in accordance with the 

 ciiaracter of a substrate already established 

 (Goodale, 1918; Ball, 1937a, 1939; Young, 

 Dem])sey, Myers and Hagcjuist, 1938; 

 Beach, 1941, 1945a). The po.s-<il)ility that 

 they might do both has received less em- 

 phasis; conceivably the action could be or- 

 ganizational before birth or hatching, or 

 before sexual maturation, and activational 

 in the adult. 



The behavior of the congenitally anom- 

 alous freemartin (Folley and Malpress, 

 1944; de Alba and Asdell, 1946) and sex- 

 intergrade pigs (Baker, 1925) provides 

 some support for the hypothesis that gon- 

 adal hormones organize patterns of behav- 

 ior. More direct evidence comes from a 

 number of experimental studies. 



Dantchakoff (1938a, b, 1947) stated that 

 female guinea pigs born to mothers given 

 intrauterine injections of testosterone pro- 

 pionate contained a normal female genital 

 tract with ovaries and a more or less well 

 developed epididymis, ductuli efferentes, 

 prostate, Cowper's gland, and penis. In their 

 behavior, when they were given male hor- 

 mone, they performed completely as males. 

 Alales receiving testosterone propionate 

 prenatally and after birth possessed only 

 the male genital tract and its accessories, 

 but the development of most parts was 

 precocious as was the display of masculine 

 behavior (Dantchakoff', 1938c). If such 

 transformations can be produced, the fact 

 would suggest strongly that male hormone 

 has an organizing action on the tissues 

 mediating mating behavior, at least when 

 the hormone is present in the early stages 

 of embryonic or fetal development. Unfor- 

 tunately, many details of Dantchakoff's 

 procedure are not clear from her articles, 

 and no controls were established in the form 

 of animals receiving androgen postnatally 

 only. Partly because of this circumstance, 

 new experiments were undertaken (Phoe- 

 nix, Goy, Gerall and Young, 1959; Tedford 

 and Young, 1960). From what follows it 

 will be clear that an androgen when given 

 prenatally does have an organizing action 

 on the tissues mediating mating behavior. 



Pregnant females were injected intra- 

 muscularly with 5 mg. of testosterone pro- 

 pionate on day 10 of the pregnancy and 

 with 1 mg. daily from day 11 to day 68 of 

 tiie gestation period. At birth the external 

 genitalia of the female offspring were indis- 

 tinguishable macroscopically from those of 

 the sibling males and examination of the 

 genital tracts by laparotomy was necessary 

 for identification of the sex. Internally there 

 were hypertrophied Wolffian ducts (detect- 

 able microscopically), failure of Miillerian 

 duct-urogenital sinus fusion, and, by the 



