Jiang Zhi, Song Jianbo, Qi Fei, Xiao An, An Xizhou, Liu Ning-ai, Zhu Zuoyang, Zhang Bo, Lin Shuo
Key Laboratory of Cell Proliferation and Differentiation, Center of Developmental Biology and Genetics, College of Life Sciences, Peking University, Ministry of Education, Beijing, People's Republic of China.
PLoS Biol. 2008 Nov 25;6(11):e293. doi: 10.1371/journal.pbio.0060293.
Both endocrine and exocrine pancreatic cells arise from pancreatic-duodenal homeobox 1 (pdx1)-positive progenitors. The molecular mechanisms controlling cell fate determination and subsequent proliferation, however, are poorly understood. Unlike endocrine cells, less is known about exocrine cell specification. We report here the identification and characterization of a novel exocrine cell determinant gene, exocrine differentiation and proliferation factor (exdpf), which is highly expressed in the exocrine cell progenitors and differentiated cells of the developing pancreas in zebrafish. Knockdown of exdpf by antisense morpholino caused loss or significant reduction of exocrine cells due to lineage-specific cell cycle arrest but not apoptosis, whereas the endocrine cell mass appeared normal. Real-time PCR results demonstrated that the cell cycle arrest is mediated by up-regulation of cell cycle inhibitor genes p21(Cip), p27(Kip), and cyclin G1 in the exdpf morphants. Conversely, overexpression of exdpf resulted in an overgrowth of the exocrine pancreas and a severe reduction of the endocrine cell mass, suggesting an inhibitory role for exdpf in endocrine cell progenitors. We show that exdpf is a direct target gene of pancreas-specific transcription factor 1a (Ptf1a), a transcription factor critical for exocrine formation. Three consensus Ptf1a binding sites have been identified in the exdpf promoter region. Luciferase assay demonstrated that Ptf1a promotes transcription of the exdpf promoter. Furthermore, exdpf expression in the exocrine pancreas was lost in ptf1a morphants, and overexpression of exdpf successfully rescued exocrine formation in ptf1a-deficient embryos. Genetic evidence places expdf downstream of retinoic acid (RA), an instructive signal for pancreas development. Knocking down exdpf by morpholino abolished ectopic carboxypeptidase A (cpa) expression induced by RA. On the other hand, exdpf mRNA injection rescued endogenous cpa expression in embryos treated with diethylaminobenzaldehyde, an inhibitor of RA signaling. Moreover, exogenous RA treatment induced anterior ectopic expression of exdpf and trypsin in a similar pattern. Our study provides a new understanding of the molecular mechanisms controlling exocrine cell specification and proliferation by a novel gene, exdpf. Highly conserved in mammals, the expression level of exdpf appears elevated in several human tumors, suggesting a possible role in tumor pathogenesis.
胰腺的内分泌细胞和外分泌细胞均起源于胰腺十二指肠同源框1(pdx1)阳性祖细胞。然而,控制细胞命运决定及后续增殖的分子机制仍知之甚少。与内分泌细胞不同,外分泌细胞分化的相关情况了解较少。我们在此报告了一种新型外分泌细胞决定基因——外分泌分化与增殖因子(exdpf)的鉴定与特性,该基因在斑马鱼发育中的胰腺外分泌细胞祖细胞和分化细胞中高度表达。通过反义吗啉代寡核苷酸敲低exdpf会导致外分泌细胞因谱系特异性细胞周期停滞而缺失或显著减少,但不会引发细胞凋亡,而内分泌细胞团看起来正常。实时定量PCR结果表明,细胞周期停滞是由exdpf morphants中细胞周期抑制基因p21(Cip)、p27(Kip)和细胞周期蛋白G1的上调介导的。相反,exdpf的过表达导致外分泌胰腺过度生长,内分泌细胞团严重减少,这表明exdpf对内分泌细胞祖细胞具有抑制作用。我们发现exdpf是胰腺特异性转录因子1a(Ptf1a)的直接靶基因,Ptf1a是外分泌形成的关键转录因子。在exdpf启动子区域已鉴定出三个一致的Ptf1a结合位点。荧光素酶测定表明Ptf1a促进exdpf启动子的转录。此外,在ptf1a morphants中,外分泌胰腺中exdpf的表达缺失,exdpf的过表达成功挽救了ptf1a缺陷胚胎中的外分泌形成。遗传学证据表明expdf位于视黄酸(RA)下游,RA是胰腺发育的诱导信号。用吗啉代寡核苷酸敲低exdpf可消除由RA诱导的异位羧肽酶A(cpa)表达。另一方面,exdpf mRNA注射挽救了用RA信号抑制剂二乙氨基苯甲醛处理的胚胎中的内源性cpa表达。此外,外源性RA处理以类似模式诱导exdpf和胰蛋白酶的前部异位表达。我们的研究为一种新型基因exdpf控制外分泌细胞分化和增殖的分子机制提供了新的认识。exdpf在哺乳动物中高度保守,其表达水平在几种人类肿瘤中似乎升高,提示其在肿瘤发病机制中可能发挥作用。