2. Endocrines and Populations 219 



the cortieoids, although with a marked increase in the secretion of C19 

 steroids (Dorfman and Shipley, 1956; Bradlow and Gallagher, 1957, Dorf- 

 man, 19G0) . It is also possible that these effects arc a result of aberrant bio- 

 chemical pathways in the synthesis of adrenal steroids which produce 

 greatly increased amounts of weakly androgenic steroids or possibly some 

 compounds with more strongly androgenic activity (Dorfman and Shipley, 

 1956; Bradlow and Gallagher, 1957; Gallagher, 1958, Dorfman, 1960). 

 Finally there may be abnormal or greatly increased androgenic metabolites 

 of adrenal steroids (Gallagher et at., 1958). It is well known that gonadec- 

 tomy is followed by a marked increase in the production of gonadotropins ; 

 therefore if the production of sex steroids by the adrenal cortex normally 

 was controlled by gonadotropins, one would expect an increased secretion 

 of these sex steroids to follow gonadectomy and to be reflected by con- 

 tinued maintenance of the sex accessories. This is not the case, as the sex 

 accessories involute nearly completely following gonadectomy (Burrows, 

 1949; Jones, 1957). However, it has been shown that increased secretion 

 of ACTH is followed after a time by a marked increase in the secretion of 

 Ci9 steroids (Bush, 1953; Dorfman and Shipley, 1956; Bradlow and Gal- 

 lagher, 1957; Jones, 1957; Wilson et al, 1958, Vande Wiele and Lieberman, 

 1960). Apparently slight stimulation of the sex accessories by adrenocorti- 

 cal androgens occurs following ACTH treatment of gonadectomized rats, 

 but these effects can be detected only histologically (Davidson and Moon, 

 1936; Davidson, 1937; Lostroh and Li, 1957, Desclin, 1959) . In marked con- 

 trast to the general tenor of these results, Delost (1951) reports that the 

 adrenal cortex secretes hormones with androgenic actions on the sex acces- 

 sories of male Microtus arvalis, as determined by morphologic criteria, but 

 this work has not been confirmed. Delost (1956a) maintains that cortisone 

 will stimulate some of the secondary sex organs in these mammals, but 

 confirmation of these res ilts also is needed before acceptance of his conclu- 

 sions is warranted. It is best for the present to assume that the adrenal 

 cortex of most species normally does not secrete appreciable amounts of 

 steroids with pronounced androgenic or estrogenic activity and that, it 

 usually represents abnormal function when appreciable quantities of hor- 

 mones with these activities are secreted by the cortex, (Bradlow and Gal- 

 lagher, 1957). The ability of the adrenal cortical and gonadal endocrine 

 cells to secrete steroids with overlapping activity is an intriguing problem, 

 and the reader is referred to Dorfman (1960) for a more detailed discussion 

 of the steroidal biosynthetic pathways involved and the elucidation of some 

 of these paradoxes. There is certainly variation between species in the types 

 of adrenal steroids normally secreted. Under abnormal conditions there is 

 no doubt that the adrenals produce increased amounts of steroids which may 

 have sex steroidal activity and that the ovaries, for example, can be induced 



