Hahn T P, Ball G F
Department of Psychology, Johns Hopkins University, Baltimore, Maryland 21218, USA.
Gen Comp Endocrinol. 1995 Sep;99(3):349-63. doi: 10.1006/gcen.1995.1119.
Temperate zone birds terminate reproduction when they become photorefractory. In many species, refractoriness is "absolute" in that gonadal regression occurs before day length declines, and individuals are reproductively unresponsive even to continuous light. Based on studies of a few species, this form of refractoriness appears to be associated with a reduction (compared with breeding and/or photosensitive birds) in numbers of hypothalamic cells and fibers that are immunoreactive for gonadotropin releasing hormone (GnRH). Some species display "relative" refractoriness, in that day length must decline before gonadal regression occurs, and individuals never lose the capacity to respond to very long days. A few reports suggest that GnRH levels in the brain do not change in relatively refractory birds. House sparrows (Passer domesticus) regress their gonads sooner when day length declines during summer than they do if photoperiod is not permitted to decline after the summer solstice (i.e., they appear to be "relatively" refractory). However, if held on long days they eventually regress the gonads despite no decline in photoperiod (they appear to be "absolutely" refractory). In this experiment, we tested whether gonadal regression was associated with changes in hypothalamic GnRH in adult male house sparrows. We used immunocytochemistry (primary antibody sensitive to both forms of avian GnRH) to compare the distribution and number of GnRH-immunoreactive cells and fibers among reproductively active individuals on long days, "absolutely" refractory individuals on long days, and presumably "relatively" refractory individuals induced to regress by shortened days. Similar reductions in ir-GnRH cell number and cell size occurred in both groups of refractory birds compared with birds still in breeding condition. Gonadal regression also was correlated with reduced staining intensity of cell bodies and reduced fiber staining, but these reductions were somewhat more pronounced in the "absolutely" than the "relatively" refractory birds. We discuss these results in light of other studies of avian GnRH changes with reproductive stage. Our results are consistent with the idea that house sparrows become absolutely refractory, regardless of whether exposed to a decline in photoperiod. However, the results also suggest that relative refractoriness may induce gonadal regression through a cessation of GnRH secretion, whereas absolute refractoriness involves down-regulation of peptide production as well.
温带鸟类在进入光不应期时会终止繁殖。在许多物种中,不应期是“绝对的”,因为性腺退化发生在白昼长度下降之前,而且即使给予持续光照,个体在生殖方面也没有反应。基于对少数物种的研究,这种不应期形式似乎与下丘脑对促性腺激素释放激素(GnRH)免疫反应阳性的细胞和纤维数量减少(与繁殖期和/或光敏鸟类相比)有关。一些物种表现出“相对”不应期,即性腺退化必须在白昼长度下降之后才会发生,而且个体对很长的白昼长度始终保持反应能力。一些报告表明,相对不应期的鸟类大脑中的GnRH水平没有变化。家麻雀(Passer domesticus)在夏季白昼长度下降时比在夏至后不允许光周期下降时(即它们似乎是“相对”不应期)更快地使性腺退化。然而,如果将它们置于长日照条件下,尽管光周期没有下降,它们最终也会使性腺退化(它们似乎是“绝对”不应期)。在本实验中,我们测试了成年雄性家麻雀性腺退化是否与下丘脑GnRH的变化有关。我们使用免疫细胞化学方法(对两种形式的鸟类GnRH均敏感的一抗)比较了长日照下处于生殖活跃状态的个体、长日照下“绝对”不应期的个体以及因白昼缩短而诱导退化的推测“相对”不应期个体之间GnRH免疫反应阳性细胞和纤维的分布及数量。与仍处于繁殖状态的鸟类相比,两组不应期鸟类的免疫反应性GnRH细胞数量和细胞大小均出现类似程度的减少。性腺退化还与细胞体染色强度降低和纤维染色减少相关,但这些减少在“绝对”不应期鸟类中比在“相对”不应期鸟类中更为明显。我们结合其他关于鸟类GnRH随生殖阶段变化的研究来讨论这些结果。我们的结果与家麻雀无论是否暴露于光周期下降都会进入绝对不应期的观点一致。然而,结果还表明,相对不应期可能通过GnRH分泌停止诱导性腺退化,而绝对不应期还涉及肽产生的下调。