Cohen L, Sekler I, Hershfinkel M
Department of Physiology and Cell Biology, Faculty of Health Science, Ben-Gurion University of the Negev, Beer-Sheva, Israel.
Cell Death Dis. 2014 Jun 26;5(6):e1307. doi: 10.1038/cddis.2014.262.
The intestinal epithelium is a renewable tissue that requires precise balance between proliferation and differentiation, an essential process for the formation of a tightly sealed barrier. Zinc deficiency impairs the integrity of the intestinal epithelial barrier and is associated with ulcerative and diarrheal pathologies, but the mechanisms underlying the role of Zn(2+) are not well understood. Here, we determined a role of the colonocytic Zn(2+) sensing receptor, ZnR/GPR39, in mediating Zn(2+)-dependent signaling and regulating the proliferation and differentiation of colonocytes. Silencing of ZnR/GPR39 expression attenuated Zn(2+)-dependent activation of ERK1/2 and AKT as well as downstream activation of mTOR/p70S6K, pathways that are linked with proliferation. Consistently, ZnR/GPR39 silencing inhibited HT29 and Caco-2 colonocyte proliferation, while not inducing caspase-3 cleavage. Remarkably, in differentiating HT29 colonocytes, silencing of ZnR/GPR39 expression inhibited alkaline phosphatase activity, a marker of differentiation. Furthermore, Caco-2 colonocytes showed elevated expression of ZnR/GPR39 during differentiation, whereas silencing of ZnR/GPR39 decreased monolayer transepithelial electrical resistance, suggesting compromised barrier formation. Indeed, silencing of ZnR/GPR39 or chelation of Zn(2+) by the cell impermeable chelator CaEDTA was followed by impaired expression of the junctional proteins, that is, occludin, zonula-1 (ZO-1) and E-cadherin. Importantly, colon tissues of GPR39 knockout mice also showed a decrease in expression levels of ZO-1 and occludin compared with wildtype mice. Altogether, our results indicate that ZnR/GPR39 has a dual role in promoting proliferation of colonocytes and in controlling their differentiation. The latter is followed by ZnR/GPR39-dependent expression of tight junctional proteins, thereby leading to formation of a sealed intestinal epithelial barrier. Thus, ZnR/GPR39 may be a therapeutic target for promoting epithelial function and tight junction barrier integrity during ulcerative colon diseases.
肠上皮是一种可再生组织,其增殖与分化需要精确平衡,这是形成紧密密封屏障的关键过程。锌缺乏会损害肠上皮屏障的完整性,并与溃疡性和腹泻性疾病相关,但锌离子(Zn²⁺)发挥作用的潜在机制尚不完全清楚。在此,我们确定了结肠细胞锌离子感应受体ZnR/GPR39在介导锌离子依赖性信号传导以及调节结肠细胞增殖和分化中的作用。沉默ZnR/GPR39表达会减弱锌离子依赖性的ERK1/2和AKT激活以及mTOR/p70S6K的下游激活,这些通路与增殖相关。一致地,沉默ZnR/GPR39会抑制HT29和Caco-2结肠细胞增殖,但不会诱导半胱天冬酶-3切割。值得注意的是,在分化的HT29结肠细胞中,沉默ZnR/GPR39表达会抑制碱性磷酸酶活性,这是分化的一个标志。此外,Caco-2结肠细胞在分化过程中ZnR/GPR39表达升高,而沉默ZnR/GPR39会降低单层跨上皮电阻,表明屏障形成受损。实际上,沉默ZnR/GPR39或用细胞不可渗透的螯合剂CaEDTA螯合锌离子后,会导致紧密连接蛋白,即闭合蛋白、紧密连接蛋白1(ZO-1)和E-钙黏蛋白的表达受损。重要的是,与野生型小鼠相比,GPR39基因敲除小鼠的结肠组织中ZO-1和闭合蛋白的表达水平也有所降低。总之,我们的结果表明,ZnR/GPR39在促进结肠细胞增殖和控制其分化方面具有双重作用。后者伴随着紧密连接蛋白的ZnR/GPR39依赖性表达,从而导致形成密封的肠上皮屏障。因此,ZnR/GPR39可能是促进溃疡性结肠疾病期间上皮功能和紧密连接屏障完整性的治疗靶点。