374 



PHYSIOLOGY OF GONADS 



there are many potentialities for forming 

 accessory glands with varied anatomic 

 structure, histologic characteristics, and 

 functional activities. 



Female prostate. In fetuses of many fe- 

 male mammals, small cords of cells which 

 represent the homologues of the male pros- 

 tate bud off from the epithelial lining of the 

 urethra. These primordia normally retro- 

 gress or remain vestigial and only rarely 

 continue to develop after birth. In the hu- 

 man female, these rudimentary structures 

 are known as para-urethral glands of Skene. 

 They have also been referred to as peri- 

 urethral glands. However, it seems advis- 

 able, as Witschi, Mahoney and Riley (1938) 

 suggested, to restrict the usage para-ure- 

 thral and peri-urethral to the aggregations 

 of mucus-secreting glands that have short 

 ducts opening into the urethra. These clearly 

 differ from the true female prostate glands. 



In contrast to the rudimentary prostate 

 glands which are retained postnatally by 

 some female mammals, relatively large, well 

 developed female prostates have been re- 

 ported postnatally in some insectivores, 

 chiropterans, rodents, and lagomorphs. The 

 male accessory glands of many species in 

 these orders are exceptionally well devel- 

 oped and the prostates are usually lobed. 

 Female prostates are tubulo-alveolar glands, 

 as are their male homologues, and they too 

 form lobes, but the glands are never as 

 large as those of the male. Their secretory 

 activity is apparently dependent mainly on 

 ovarian androgens, but the function, if any, 

 of the secretion is obscure. Extensive re- 

 search has shown that the administration of 

 androgens to rodents, either to pregnant fe- 

 males or fetuses, to fetal lagomorphs, and 

 to pouch-young oppossums, results in the 

 formation and retention of prostates in fe- 

 males which normally do not have such 

 glands (see chapter by Burns). 



Deanesly (1934) described vaginal glands 

 in the female hedgehog and suggested that 

 one pair is homologous with the external 

 prostates of the male. The female glands ex- 

 tend dorsolaterally on either side of the ure- 

 thra and a single duct from each lobe opens 

 into the vagina. They seem to be active 

 during the breeding season and to retrogress 

 in the anestrum. In another insectivore, 



Hemicentetes, there is a pair of large 

 "paravaginal glands" which are function- 

 ally active in the mature female and have 

 large acini filled with secretion. They re- 

 semble the male prostate in histologic struc- 

 ture and anatomic position, but have no 

 ducts (Lehmann, 1938). In adult European 

 moles, most females have bilobed ventral 

 prostate glands which undergo cyclic 

 changes in the epithelium. The prostate of 

 the male is also bilobed and ventral in posi- 

 tion and the homology in the two sexes is 

 clear (Godet, 1949). 



Mathews (1941) studied the anatomy and 

 histophysiology of the male and female gen- 

 ital tracts of nine species of African bats. 

 Female prostates are well developed in four 

 species, less conspicuous in a fifth, and ab- 

 sent from the remaining four. There is a 

 marked tendency for greater development 

 of the glands in pregnant and lactating fe- 

 males. In three species, the female prostates 

 surround the urethra (as do their male ho- 

 mologues), but in Nycteris luteola the fe- 

 male prostate appears ventrally, whereas 

 the male prostate in this species is limited 

 to the dorsal aspect of the urethra. Mathews 

 considered that the female prostates repre- 

 sent greatly enlarged female urethral glands 

 which are homologous with the male ])yos- 

 tate. 



The occurrence of female prostates and 

 their relation to hormones have been most 

 extensively studied in rodents. The first de- 

 scription of a well developed female pros- 

 tate gland seems to be that of Rauther 

 (1909), who found such a gland ventral to 

 the neck of the bladder in the African field 

 rat, Arvicanthis cinereus. In Rattus nor- 

 vegicus, Marx (1931, 1932) reported the 

 sporadic occurrence of female prostates. 

 Korenchevsky and Dennison (1936) and 

 Korenchevsky (1937) found prostates in 9 

 of 56 females and stated that the glands 

 wei'c atrophic, but when androgens were ad- 

 ministered, these glands resembled the 

 male ventral prostate. On this basis, the 

 homology of the glands of the female with 

 the \-entral prostate of the male was sug- 

 gested. Further studies (Witschi, Mahoney 

 and Riley, 1938; Mahoney, 1940, 1942; Ma- 

 lioney and Witschi, 1947) showed that the 

 female prostate of the rat is homologous 



