Lee Jaemin, Wang Xiaofan, Di Jeso Bruno, Arvan Peter
Cell and Molecular Biology Program, University of Michigan Medical Center, Ann Arbor, MI 48109, USA.
J Biol Chem. 2009 May 8;284(19):12752-61. doi: 10.1074/jbc.M806898200. Epub 2009 Mar 9.
The carboxyl-terminal cholinesterase-like (ChEL) domain of thyroglobulin (Tg) has been identified as critically important in Tg export from the endoplasmic reticulum. In a number of human kindreds suffering from congenital hypothyroidism, and in the cog congenital goiter mouse and rdw rat dwarf models, thyroid hormone synthesis is inhibited because of mutations in the ChEL domain that block protein export from the endoplasmic reticulum. We hypothesize that Tg forms homodimers through noncovalent interactions involving two predicted alpha-helices in each ChEL domain that are homologous to the dimerization helices of acetylcholinesterase. This has been explored through selective epitope tagging of dimerization partners and by inserting an extra, unpaired Cys residue to create an opportunity for intermolecular disulfide pairing. We show that the ChEL domain is necessary and sufficient for Tg dimerization; specifically, the isolated ChEL domain can dimerize with full-length Tg or with itself. Insertion of an N-linked glycan into the putative upstream dimerization helix inhibits homodimerization of the isolated ChEL domain. However, interestingly, co-expression of upstream Tg domains, either in cis or in trans, overrides the dimerization defect of such a mutant. Thus, although the ChEL domain provides a nidus for Tg dimerization, interactions of upstream Tg regions with the ChEL domain actively stabilizes the Tg dimer complex for intracellular transport.
甲状腺球蛋白(Tg)的羧基末端胆碱酯酶样(ChEL)结构域已被确定在Tg从内质网输出过程中至关重要。在一些患有先天性甲状腺功能减退症的人类家系以及cog先天性甲状腺肿小鼠和rdw大鼠侏儒模型中,由于ChEL结构域中的突变阻碍了蛋白质从内质网输出,甲状腺激素合成受到抑制。我们推测,Tg通过非共价相互作用形成同型二聚体,这种相互作用涉及每个ChEL结构域中两个预测的α螺旋,它们与乙酰胆碱酯酶的二聚化螺旋同源。这一点已通过对二聚化伙伴进行选择性表位标记以及插入一个额外的未配对半胱氨酸残基以创造分子间二硫键配对的机会进行了探索。我们发现,ChEL结构域对于Tg二聚化是必要且充分的;具体而言,分离的ChEL结构域可以与全长Tg或其自身二聚化。在假定的上游二聚化螺旋中插入一个N - 连接聚糖会抑制分离的ChEL结构域的同型二聚化。然而,有趣的是,上游Tg结构域的共表达,无论是顺式还是反式,都能克服这种突变体的二聚化缺陷。因此,尽管ChEL结构域为Tg二聚化提供了一个核心,但上游Tg区域与ChEL结构域的相互作用积极地稳定了用于细胞内运输的Tg二聚体复合物。