Liu Jing, Park Eun-Sil, Curry Thomas E, Jo Misung
Department of Obstetrics and Gynecology, Chandler Medical Center, University of Kentucky, Lexington, Kentucky 40536-0298, USA.
Mol Endocrinol. 2010 Jun;24(6):1203-17. doi: 10.1210/me.2009-0325. Epub 2010 Mar 25.
Periovulatory follicular matrix plays an important role in cumulus-oocyte complex (COC) expansion, ovulation, and luteal formation. Hyaluronan and proteoglycan link protein 1 (HAPLN1), a component of follicular matrix, was shown to enhance COC expansion in vitro. However, the regulatory mechanisms of periovulatory expression of Hapln1 and its role in periovulatory granulosa cells have not been elucidated. We first determined the periovulatory expression pattern of Hapln1 using pregnant mare serum gonadotropin/human chorionic gonadotropin (PMSG/hCG)-primed immature rat ovaries. Hapln1 expression was transiently induced both in intact ovaries and granulosa cells at 8 h and 12 h after hCG injection. This in vivo expression of Hapln1 was recapitulated by culturing preovulatory granulosa cells with hCG. The stimulatory effect of hCG was blocked by inhibition of protein kinase A, phosphatidylinositol-dependent kinase, p38 MAPK, epidermal growth factor signaling, and prostaglandin synthesis, revealing key mediators involved in LH-induced Hapln1 expression. In addition, knockdown of Runx1 and Runx2 expression by small interfering RNA or inhibition of RUNX activities by dominant-negative RUNX decreased hCG or agonist-induced Hapln1 expression. Chromatin immunoprecipitation assays verified the in vivo binding of RUNX1 and RUNX2 to the Hapln1 promoter in periovulatory granulosa cells. Luciferase reporter assays revealed that mutation of the RUNX binding sites completely obliterated the agonist-induced activity of the Hapln1 promoter. These data conclusively identified RUNX proteins as the crucial transcription regulators for LH-induced Hapln1 expression. Functionally, treatment with HAPLN1 increased the viability of cultured granulosa cells and decreased the number of the cells undergoing apoptosis, whereas knockdown of Hapln1 expression decreased granulosa cells viability. This novel finding indicates that HAPLN1 may promote periovulatory granulosa cell survival, which would facilitate their differentiation into luteal cells.
排卵周围卵泡基质在卵丘-卵母细胞复合体(COC)扩张、排卵和黄体形成中起重要作用。透明质酸和蛋白聚糖连接蛋白1(HAPLN1)是卵泡基质的一个组成部分,已被证明在体外可增强COC扩张。然而,Hapln1排卵周围表达的调控机制及其在排卵周围颗粒细胞中的作用尚未阐明。我们首先使用孕马血清促性腺激素/人绒毛膜促性腺激素(PMSG/hCG)预处理的未成熟大鼠卵巢确定了Hapln1的排卵周围表达模式。Hapln1表达在hCG注射后8小时和12小时在完整卵巢和颗粒细胞中均被短暂诱导。通过用hCG培养排卵前颗粒细胞可重现Hapln1的这种体内表达。hCG的刺激作用被蛋白激酶A、磷脂酰肌醇依赖性激酶、p38丝裂原活化蛋白激酶、表皮生长因子信号传导和前列腺素合成的抑制所阻断,揭示了参与LH诱导的Hapln1表达的关键介质作用。此外,小干扰RNA敲低Runx1和Runx2表达或显性负性RUNX抑制RUNX活性可降低hCG或激动剂诱导Hapln1表达。染色质免疫沉淀试验证实了RUNX1和RUNX2在排卵周围颗粒细胞中与Hapln1启动子的体内结合。荧光素酶报告试验表明,RUNX结合位点的突变完全消除了激动剂诱导的Hapln1启动子活性。这些数据最终确定RUNX蛋白是LH诱导的Hapln1表达的关键转录调节因子。在功能上,用HAPLN1处理可增加培养的颗粒细胞活力并减少发生凋亡的细胞数量,而敲低Hapln1表达则降低颗粒细胞活力。这一新发现表明HAPLN1可能促进排卵周围颗粒细胞存活,这将有助于它们分化为黄体细胞。