Neveu M J, Hully J R, Babcock K L, Hertzberg E L, Nicholson B J, Paul D L, Pitot H C
McArdle Laboratory for Cancer Research, University of Wisconsin, Madison.
J Cell Sci. 1994 Jan;107 ( Pt 1):83-95. doi: 10.1242/jcs.107.1.83.
Although several abnormalities in gap junction (GJ) structure and/or function have been described in neoplasms, the molecular mechanisms responsible for many of the alterations remain unknown. The identification of a family of GJ proteins, termed connexins, prompted this study of connexin32 (Cx32), connexin26 (Cx26) and connexin43 (Cx43) expression during rat hepatocarcinogenesis. Using antibody, cDNA and cRNA probes, we investigated connexin mRNA and protein expression in preneoplastic and neoplastic rat livers. In normal liver, Cx32 is expressed in hepatocytes throughout the hepatic acinus, Cx26 is restricted to periportal hepatocytes, and Cx43 is expressed by mesothelial cells forming Glisson's capsule. Most preneoplastic altered hepatic foci generated by diethylnitrosamine (DEN) initiation and either phenobarbital (PB) or 2,3,7,8-dichlorodibenzo-p-dioxin (TCDD) promotion exhibited decreased Cx32 or increased Cx26 staining. Foci from either protocol failed to display Cx43 immunoreactivity. In the majority of PB-promoted foci, Cx32 immunoreactivity decreased independently of changes in mRNA abundance. Continuous thymidine labeling, following cessation of PB promotion, showed that downregulation of Cx32 staining is reversible in foci that are promoter-dependent for growth, but irreversible in lesions that are promoter-independent for growth. Hepatic neoplasms from rats initiated with DEN and promoted with PB or TCDD also displayed modified connexin expression. While all 24 neoplasms studied were deficient in normal punctate Cx32 and Cx26 staining, altered cellular localization of these proteins was apparent in some tumors. Immunoblotting of crude tissue extracts revealed that neoplasms with disordered Cx32 staining showed immunoreactive bands with altered electrophoretic mobility. These observations show that hepatomas may downregulate Cx32 expression through changes in the primary structure of Cx32 or by post-translational modifications. Northern blotting of total tumor mRNAs failed to demonstrate consistent changes in the abundance of Cx32, Cx26 or Cx43 transcripts. Some tumors expressed steady-state transcripts without observable immunoreactivity, indicating that some hepatomas downregulate connexin immunoreactivity independently of mRNA abundance. Increased levels of Cx43 mRNA and protein were found in several neoplasms, but immunostaining was always localized to nonparenchymal cells. Areas of bile duct proliferation and cholangiomas displayed Cx43 staining, whereas, cholangiocarcinomas were deficient in immunoreactivity. These findings show that alterations in the expression of connexins, by either downregulation or differential induction, represent common modifications during hepatocarcinogenesis. Although our results imply that connexins represent useful markers for the boundary between tumor promotion and progression, preneoplastic and neoplastic rat hepatocytes fail to use a common mechanism to modify connexin expression.
尽管在肿瘤中已描述了间隙连接(GJ)结构和/或功能的几种异常情况,但导致许多改变的分子机制仍不清楚。一类被称为连接蛋白的GJ蛋白的鉴定促使了本研究对大鼠肝癌发生过程中连接蛋白32(Cx32)、连接蛋白26(Cx26)和连接蛋白43(Cx43)表达的研究。使用抗体、cDNA和cRNA探针,我们研究了癌前和肿瘤大鼠肝脏中连接蛋白mRNA和蛋白的表达。在正常肝脏中,Cx32在整个肝腺泡的肝细胞中表达,Cx26局限于门静脉周围的肝细胞,Cx43由形成Glisson囊的间皮细胞表达。大多数由二乙基亚硝胺(DEN)启动并经苯巴比妥(PB)或2,3,7,8-二氯二苯并对二恶英(TCDD)促进产生的癌前改变肝灶显示Cx32染色减少或Cx26染色增加。两种方案产生的病灶均未显示Cx43免疫反应性。在大多数PB促进的病灶中,Cx32免疫反应性降低,与mRNA丰度的变化无关。PB促进停止后进行连续胸苷标记显示,在依赖启动子生长的病灶中,Cx32染色的下调是可逆的,但在不依赖启动子生长的病变中是不可逆的。用DEN启动并用PB或TCDD促进的大鼠肝脏肿瘤也显示连接蛋白表达改变。虽然所研究的所有24个肿瘤均缺乏正常的点状Cx32和Cx26染色,但这些蛋白的细胞定位改变在一些肿瘤中很明显。粗组织提取物的免疫印迹显示,Cx32染色紊乱的肿瘤显示免疫反应条带,其电泳迁移率改变。这些观察结果表明,肝癌可能通过Cx32一级结构的改变或翻译后修饰来下调Cx32表达。肿瘤总mRNA的Northern印迹未能证明Cx32、Cx26或Cx43转录本丰度的一致变化。一些肿瘤表达稳态转录本,但无明显免疫反应性,表明一些肝癌下调连接蛋白免疫反应性与mRNA丰度无关。在一些肿瘤中发现Cx43 mRNA和蛋白水平升高,但免疫染色始终定位于非实质细胞。胆管增生和胆管瘤区域显示Cx43染色,而胆管癌缺乏免疫反应性。这些发现表明,通过下调或差异诱导,连接蛋白表达的改变是肝癌发生过程中的常见修饰。虽然我们的结果表明连接蛋白是肿瘤促进和进展之间边界的有用标志物,但癌前和肿瘤大鼠肝细胞未能使用共同机制来修饰连接蛋白表达。