Kang L, Marin M, Kelley D
Department of Biological Sciences, Columbia University, New York, New York 10027, USA.
Gen Comp Endocrinol. 1995 Dec;100(3):293-307. doi: 10.1006/gcen.1995.1160.
In the frog, Xenopus laevis, castration or anti-androgen treatment during late tadpole or early juvenile stages blocks masculinization of vocal neuroeffectors in males while exogenous androgen or a testicular transplant masculinizes vocal neuroeffectors in females. To elucidate the relation between androgen secretion and the process of masculinization, we measured androgens by radioimmunoassay in adults, juveniles, and tadpoles. In adult females, values for both testosterone (T) and dihydrotestosterone (DHT) were essentially identical during the winter and the summer while in males, summer values for both androgens were 7 to 8x winter values. For DHT, circulating values in males significantly exceeded those in females in both winter and summer while for T, values differed significantly only in summer. Sex differences in circulating androgen arise at late juvenile stages (PM4 and PM5); T values are higher in males than in females and higher, for both sexes, in winter than in summer. During early juvenile stages (PM1-PM3), there were no seasonal or sex differences in circulating androgens. While androgens were detected in liver during late tadpole stages (stage 56-66), no sex differences were apparent. Androgens can be detected in tadpole tissues, as early as tadpole stage 47. delta 5-3 beta-Hydroxysteroid dehydrogenase (HSD) histochemistry indicates that tadpole gonads and interrenals (and, to a much lesser extent, kidneys) are the only tissues capable of steroid biosynthesis. Interrenal HSD activity was present throughout tadpole development; in gonads, activity began before sexual differentiation. Thin-layer chromatographic separation of steroid metabolites following in vitro incubation with radioactive precursors indicated that the interrenals were more active than the gonads in tadpoles but not in juveniles. Thus, interrenals and gonads of developing X. laevis can synthesize delta 4 steroids (which include androgens) and, from early tadpole stages, are capable of metabolizing pregnenolone and testosterone, early and late compounds in the steroid synthetic pathway, respectively. The absence of dramatic sex differences in T or DHT levels during late tadpole and early juvenile stages suggests that while necessary, androgen secretion is not sufficient for masculinization of vocal neuroeffectors.
在非洲爪蟾(Xenopus laevis)中,在蝌蚪晚期或幼蛙早期进行去势或抗雄激素处理会阻止雄性的发声神经效应器雄性化,而外源性雄激素或睾丸移植则会使雌性的发声神经效应器雄性化。为了阐明雄激素分泌与雄性化过程之间的关系,我们通过放射免疫分析法测量了成年蛙、幼蛙和蝌蚪体内的雄激素。成年雌性蛙在冬季和夏季的睾酮(T)和双氢睾酮(DHT)值基本相同,而成年雄性蛙的这两种雄激素在夏季的值是冬季的7至8倍。对于DHT,雄性蛙体内的循环值在冬季和夏季均显著超过雌性蛙;对于T,仅在夏季两性的值存在显著差异。循环雄激素的性别差异出现在幼蛙晚期(PM4和PM5);雄性蛙的T值高于雌性蛙,并且两性在冬季的T值均高于夏季。在幼蛙早期(PM1 - PM3),循环雄激素不存在季节性或性别差异。虽然在蝌蚪晚期(56 - 66期)肝脏中检测到了雄激素,但未发现明显的性别差异。早在蝌蚪47期就能在蝌蚪组织中检测到雄激素。δ5 - 3β - 羟基类固醇脱氢酶(HSD)组织化学表明,蝌蚪的性腺和肾上腺(以及程度较轻的肾脏)是唯一能够进行类固醇生物合成的组织。肾上腺HSD活性在整个蝌蚪发育过程中都存在;在性腺中,活性在性别分化之前就已开始。用放射性前体进行体外孵育后,通过薄层色谱法分离类固醇代谢产物表明,蝌蚪的肾上腺比性腺更活跃,但幼蛙中并非如此。因此,发育中的非洲爪蟾的肾上腺和性腺能够合成δ4类固醇(包括雄激素),并且从蝌蚪早期开始,就能够分别代谢孕烯醇酮和睾酮,它们分别是类固醇合成途径中的早期和晚期化合物。蝌蚪晚期和幼蛙早期T或DHT水平没有显著的性别差异,这表明虽然雄激素分泌是必要的,但对于发声神经效应器的雄性化来说并不充分。