Department of Molecular Biology, Princeton University, Princeton, NJ 08544, USA.
Curr Biol. 2011 Mar 22;21(6):439-48. doi: 10.1016/j.cub.2011.01.073. Epub 2011 Mar 3.
In many organisms, germ cells are segregated from the soma through the inheritance of the specialized germ plasm, which contains mRNAs and proteins that specify germ cell fate and promote germline development. Whereas germ plasm assembly has been well characterized, mechanisms mediating germ plasm inheritance are poorly understood. In the Drosophila embryo, germ plasm is anchored to the posterior cortex, and nuclei that migrate into this region give rise to the germ cell progenitors, or pole cells. How the germ plasm interacts with these nuclei for pole cell induction and is selectively incorporated into the forming pole cells is not known.
Live imaging of two conserved germ plasm components, nanos mRNA and Vasa protein, revealed that germ plasm segregation is a dynamic process involving active transport of germ plasm RNA-protein complexes coordinated with nuclear migration. We show that centrosomes accompanying posterior nuclei induce release of germ plasm from the cortex and recruit these components by dynein-dependent transport on centrosome-nucleated microtubules. As nuclei divide, continued transport on astral microtubules partitions germ plasm to daughter nuclei, leading to its segregation into pole cells. Disruption of these transport events prevents incorporation of germ plasm into pole cells and impairs germ cell development.
Our results indicate that active transport of germ plasm is essential for its inheritance and ensures the production of a discrete population of germ cell progenitors endowed with requisite factors for germline development. Transport on astral microtubules may provide a general mechanism for the segregation of cell fate determinants.
在许多生物中,生殖细胞通过专门的生殖质的遗传与体组织分离,生殖质包含指定生殖细胞命运并促进生殖细胞发育的 mRNA 和蛋白质。尽管生殖质的组装已得到很好的描述,但介导生殖质遗传的机制知之甚少。在果蝇胚胎中,生殖质锚定在后皮层,迁移到该区域的核产生生殖细胞前体,或极细胞。生殖质如何与这些核相互作用以诱导极细胞形成,以及如何被选择性地纳入正在形成的极细胞中尚不清楚。
对两种保守的生殖质成分,nanos mRNA 和 Vasa 蛋白的实时成像显示,生殖质的分离是一个动态过程,涉及到生殖质 RNA-蛋白复合物的主动运输,与核迁移协调。我们表明,伴随后核的中心体诱导生殖质从皮层释放,并通过中心体起源的微管上的动力蛋白依赖性运输来招募这些成分。随着核的分裂,在星体微管上的持续运输将生殖质分配到子核,导致其分离到极细胞中。这些运输事件的破坏阻止了生殖质被纳入极细胞,并损害了生殖细胞的发育。
我们的结果表明,生殖质的主动运输对于其遗传是必不可少的,确保了具有生殖系发育所需因素的离散的生殖细胞前体群体的产生。星体微管上的运输可能为命运决定因素的分离提供了一种普遍的机制。