Jablonka-Shariff Albina, Boime Irving
Department of Developmental Biology, Washington University School of Medicine, 660 South Euclid Avenue, Campus Box 8103, St. Louis, Missouri 63110, USA.
Mol Endocrinol. 2009 Mar;23(3):316-23. doi: 10.1210/me.2008-0351. Epub 2009 Jan 8.
Although the LHbeta- and chorionic gonadotropin-beta- (CGbeta) subunits share a high degree of sequence identity (>85%) in the first 114 amino acids, there is considerable sequence divergence at their carboxy ends. The CGbeta-subunit terminates with a unique carboxyl-terminal extension (115-145; carboxyl-terminal peptide), which contains four O-linked oligosaccharides, whereas the LHbeta-subunit bears a hydrophobic heptapeptide (115-121) at its carboxy terminus. LH is released through the regulated pathway in the pituitary, whereas CG is secreted constitutively from the placenta. We previously demonstrated in rat somatotroph-derived GH(3) cells that the LH is associated primarily with a regulated routing, and although the majority of CG was released constitutively from the cells, there was a fraction that was segregated through the regulated pathway. Moreover, we showed that the LHbeta heptapeptide is a determinant for the regulated secretion of LH. Given that the primary evolutionary change between LHbeta and CGbeta occurred at the carboxy terminus, these data suggested that the presence of the CGbeta carboxyl-terminal peptide region is responsible for the constitutive secretion of CG. A CG114 mutant (CGDeltaT) was constructed and expressed in GH(3) cells. Steady-state labeling and pulse-chase experiments demonstrated that the CGDeltaT entered the regulated pathway resulting in over 4-fold increase in the intracellular pool. The secretagogue, forskolin, stimulated CGDeltaT release over 3-fold, which was accompanied by a parallel intracellular decrease, and only marginal stimulation of CG was seen. Immunofluorescence demonstrated a unique membrane pattern of staining for CGDeltaT compared with dispersed cytoplasmic puncta for CG. Stimulation with forskolin caused a significant reduction in the relative fluorescence of CGDeltaT cells compared with a minor reduction for CG. These data show that the CGDeltaT analog resembles LH in its intracellular trafficking, further supporting the hypothesis that determinants at the carboxyl-terminal end of the CGbeta-subunit evolved from the LHbeta-subunit primarily to overcome the slow release and intracellular storage of LH resulting in rapid secretion of CG from the placenta.
尽管促黄体生成素β亚基(LHβ)和绒毛膜促性腺激素β亚基(CGβ)在前114个氨基酸中具有高度的序列同一性(>85%),但其羧基末端存在相当大的序列差异。CGβ亚基以独特的羧基末端延伸序列(115 - 145;羧基末端肽)结尾,该序列包含四个O - 连接寡糖,而LHβ亚基在其羧基末端带有一个疏水七肽(115 - 121)。LH通过垂体中的调节途径释放,而CG则由胎盘持续分泌。我们之前在大鼠生长激素分泌细胞来源的GH(3)细胞中证明,LH主要与调节性转运相关,虽然大部分CG从细胞中持续释放,但有一部分是通过调节途径分离出来的。此外,我们表明LHβ七肽是LH调节性分泌的决定因素。鉴于LHβ和CGβ之间的主要进化变化发生在羧基末端,这些数据表明CGβ羧基末端肽区域的存在是CG持续分泌的原因。构建了CG114突变体(CGDeltaT)并在GH(3)细胞中表达。稳态标记和脉冲追踪实验表明,CGDeltaT进入调节途径导致细胞内池增加超过4倍。促分泌剂福斯可林刺激CGDeltaT释放超过3倍,同时细胞内相应减少,而对CG的刺激则很微弱。免疫荧光显示,与CG的分散细胞质斑点相比,CGDeltaT具有独特的膜染色模式。用福斯可林刺激后,CGDeltaT细胞的相对荧光显著降低,而CG细胞的相对荧光仅有轻微降低。这些数据表明,CGDeltaT类似物在细胞内运输方面类似于LH,进一步支持了以下假设:CGβ亚基羧基末端的决定因素主要从LHβ亚基进化而来,以克服LH的缓慢释放和细胞内储存,从而使CG从胎盘快速分泌。