Miething A
University of Bonn, Institute of Anatomy, Germany.
Adv Anat Embryol Cell Biol. 1998;140:1-92. doi: 10.1007/978-3-642-72005-5.
The present study deals with initial spermatogenesis in the pubertal golden hamster (Mesocricetus auratus), in the course of which both the complex process of germ cell proliferation and differentiation and the cyclic pattern of organization of spermatogenesis are established. Germ cells pass through the complete sequence of spermatogenic differentiation from the A-spermatogonium up to the mature spermatid between days 12 and 38. During this period, germ cell development is already based on the pattern of organization and synchronization characteristic of adult spermatogenesis: 1. Judged by the germ cell types present, each of the successively appearing germ cell associations conforms to one of the adult stages of the seminiferous epithelium (stage conformity of the developmental steps of the pubertal seminiferous epithelium). 2. The overall intratesticular variance of the developmental levels reached in different tubular segments each covers almost exactly one cycle length of spermatogenesis. 3. There is a regular, continuous succession of developmental steps along the longitudinal course of the tubules which, in every single tubule, comprises the interval of one spermatogenic cycle ("wave" of the pubertal seminiferous epithelium). 4. Taken together, the latter two results indicate that the tubules of one given testis are equal to one another with respect to their developmental level, i.e., they develop synchronously. Germ cell propagation follows the clonal mode of proliferation. In quantitative terms, stage-related S-phase indices largely parallel the values of adult spermatogenesis. Both the frequently found germ cell apoptosis and degenerating intrameiotic spermatocytes correspond to degenerative phenomena in the adult seminiferous epithelium with respect to degenerative mode and stage-related occurrence. Germ cell development is closely related to the maturation of both Sertoli cells and Leydig cells. At two developmental phases, localized disturbances of germ cell differentiation occur which are specific to the pubertal period and which correlate with a maturational delay of Sertoli cells and Leydig cells, respectively: 1. Completion of Sertoli cell barrier formation may temporarily follow the initial onset of meiosis in the neighboring germ cells. As a result, mitotically dividing germ cells fall under the influence of a (hitherto hypothetic) "meiotic milieu" and characteristically degenerate (arrested germ cell mitoses). This relationship emphasizes the functional significance of the separation of the two seminiferous compartments and, in addition, heightens the understanding of the necessity of normal termination of Sertoli cell proliferation before germ cells initially enter meiosis. 2. The Leydig cell enzyme 11 beta-HSD, which is known to promote testosterone output, is established several days after the leading spermatids initially reach step 6 of spermiogenesis (which is immediately prior to the particularly testosterone-dependent start of elongation). The resulting selective developmental arrest of round spermatids of steps 6-6/7 between days 26 and 30 leads to a localized, partial desynchronization of the seminiferous epithelium, which can be traced for several days after the above-mentioned maturational delay of Leydig cells has been overcome and which is accompanied by an extensive degenerative disposition of the spermatids involved. The present investigation documents an organizational and synchronizational level of initial spermatogenesis that is similar to the situation in the adult animal. The analysis of temporarily occurring perturbations in the process of establishing the pubertal seminiferous epithelium emphasizes and specifies the close structural and functional relationships between the maturational state of somatic cells and germ cell differentiation.
本研究探讨了青春期金黄仓鼠(Mesocricetus auratus)的初始精子发生过程,在此过程中,生殖细胞增殖与分化的复杂过程以及精子发生的周期性组织模式得以确立。在第12天至38天期间,生殖细胞经历了从A型精原细胞到成熟精子的完整精子发生分化序列。在此期间,生殖细胞的发育已基于成年精子发生的组织模式和同步特征:1. 根据存在的生殖细胞类型判断,相继出现的每个生殖细胞组合都符合生精上皮的一个成年阶段(青春期生精上皮发育步骤的阶段一致性)。2. 不同管状段所达到的发育水平的总体睾丸内差异几乎恰好覆盖一个精子发生周期的长度。3. 沿着小管的纵向过程存在发育步骤的规则、连续序列,在每个小管中,该序列包括一个精子发生周期的间隔(青春期生精上皮的“波”)。4. 综合起来,后两个结果表明,一个给定睾丸的小管在发育水平上彼此相等,即它们同步发育。生殖细胞增殖遵循克隆增殖模式。从数量上看,与阶段相关的S期指数在很大程度上与成年精子发生的值平行。经常发现的生殖细胞凋亡和减数分裂期内退化的精母细胞在退化模式和与阶段相关的发生方面都与成年生精上皮中的退化现象相对应。生殖细胞的发育与支持细胞和间质细胞的成熟密切相关。在两个发育阶段,会出现特定于青春期的生殖细胞分化局部紊乱,分别与支持细胞和间质细胞的成熟延迟相关:1. 支持细胞屏障形成的完成可能暂时滞后于相邻生殖细胞减数分裂的初始开始。结果,进行有丝分裂的生殖细胞受到(迄今假设的)“减数分裂环境”的影响并发生特征性退化(生殖细胞有丝分裂停滞)。这种关系强调了两个生精区室分离的功能意义,此外,还加深了对生殖细胞最初进入减数分裂之前支持细胞增殖正常终止的必要性的理解。2. 已知促进睾酮分泌的间质细胞酶11β - HSD在领先的精子细胞最初达到精子形成第6步(紧接在特别依赖睾酮的伸长开始之前)几天后才确立。在第26天至30天期间,由此导致的第6 - 6/7步圆形精子细胞的选择性发育停滞导致生精上皮的局部、部分不同步,在上述间质细胞成熟延迟被克服后的几天内都可以追踪到这种不同步,并且伴随着所涉及精子细胞的广泛退化倾向。本研究记录了初始精子发生的组织和同步水平与成年动物的情况相似。对青春期生精上皮建立过程中暂时出现的扰动的分析强调并明确了体细胞成熟状态与生殖细胞分化之间密切的结构和功能关系。