Regulation of germ cell development by intercellular signaling in the mammalian ovarian follicle.

Regulation of germ cell development by intercellular signaling in the mammalian ovarian follicle.
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DOI:
10.1002/wdev.294
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发表时间:
2018-01
期刊:
Wiley interdisciplinary reviews. Developmental biology
影响因子:
--
通讯作者:
Clarke HJ
Clarke HJ
中科院分区:
其他
文献类型:
--
作者:
Clarke HJ

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在排卵之前,哺乳动物的卵母细胞在卵巢卵泡内经历一个分化过程,这使其具备产生胚胎的能力。分化包括两个阶段——生长阶段,在此期间卵母细胞体积增大100多倍,因为它积累了维持早期胚胎发生的大分子和细胞器;减数分裂成熟阶段,在此期间卵母细胞进行第一次减数分裂并为第二次分裂做准备。当相邻的颗粒细胞产生特定的生长因子时,卵母细胞似乎就会进入生长阶段。随着卵母细胞的生长,它会形成一层厚厚的细胞外被,称为透明带。然而,相邻颗粒细胞的细胞质突起,称为穿通突起(TZPs),使它们能够与卵母细胞保持接触依赖性的通讯。通过位于穿通突起末端与卵母细胞膜相接处的间隙连接,它们为卵母细胞提供维持其代谢活动的产物以及调节其分化的信号。相反,卵母细胞分泌可扩散的生长因子,调节颗粒细胞的增殖和分化。颗粒细胞的可通过间隙连接的产物可防止减数分裂成熟的过早启动,并且间隙连接还能使卵母细胞根据颗粒细胞接收到的激素信号开始成熟。卵母细胞或体细胞区室的发育也可能受到卵泡液中和穿通突起末端新发现的细胞外囊泡的调节,这些囊泡可介导大分子的细胞间转移。因此,卵母细胞的分化依赖于与卵泡体细胞的持续信号相互作用。 出生后卵母细胞发育的所有阶段都依赖于与卵巢卵泡相邻的体颗粒细胞的通讯。卵母细胞发出的信号也调节颗粒细胞的分化,并确保它们为生殖细胞提供一个健康的环境。
Prior to ovulation, the mammalian oocyte undergoes a process of differentiation within the ovarian follicle that confers on it the ability to give rise to an embryo. Differentiation comprises two phases – growth, during which the oocyte increases >100-fold in volume as it accumulates macromolecules and organelles that will sustain early embryogenesis; and meiotic maturation, during which the oocyte executes the first meiotic division and prepares for the second division. Entry of an oocyte into the growth phase appears to be triggered when the adjacent granulosa cells produce specific growth factors. As the oocyte grows, it elaborates a thick extracellular coat termed the zona pellucida. Nonetheless, cytoplasmic extensions of the adjacent granulosa cells, termed transzonal projections (TZPs), enable them to maintain contact-dependent communication with the oocyte. Via gap junctions located where the TZP tips meet the oocyte membrane, they provide the oocyte with products that sustain its metabolic activity and signals that regulate its differentiation. Conversely, the oocyte secretes diffusible growth factors that regulate proliferation and differentiation of the granulosa cells. Gap junction-permeable products of the granulosa cells prevent precocious initiation of meiotic maturation, and the gap junctions also enable oocyte maturation to begin in response to hormonal signals received by the granulosa cells. Development of the oocyte or the somatic compartment may also be regulated by extracellular vesicles newly identified in follicular fluid and at TZP tips, which could mediate intercellular transfer of macromolecules. Oocyte differentiation thus depends on continuous signaling interactions with the somatic cells of the follicle. All stages of post-natal oocyte development depend on communication with the neighbouring somatic granulosa cells of the ovarian follicle. Signals sent by the oocyte also regulate differentiation of the granulosa cells and ensure that they provide a healthy environment for the germ cell.
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