Section of Cell and Developmental Biology, Division of Biological Sciences, University of California, San Diego, La Jolla, CA 92093, USA.
Dev Genes Evol. 2012 Mar;222(1):29-44. doi: 10.1007/s00427-011-0387-z. Epub 2012 Feb 23.
Gap junctional proteins are important components of signaling pathways required for the development and ongoing functions of all animal tissues, particularly the nervous system, where they function in the intracellular and extracellular exchange of small signaling factors and ions. In animals whose genomes have been sufficiently sequenced, large families of these proteins, connexins, pannexins, and innexins, have been found, with 25 innexins in the nematode Caenorhabditis elegans Starich et al. (Cell Commun Adhes 8: 311-314, 2001) and at least 37 connexins in the zebrafish Danio rerio Cruciani and Mikalsen (Biol Chem 388:253-264, 2009). Having recently sequenced the medicinal leech Hirudo verbana genome, we now report the presence of 21 innexin genes in this species, nine more than we had previously reported from the analysis of an EST-derived transcriptomic database Dykes and Macagno (Dev Genes Evol 216: 185-97, 2006); Macagno et al. (BMC Genomics 25:407, 2010). Gene structure analyses show that, depending on the leech innexin gene, they can contain from 0 to 6 introns, with closely related paralogs showing the same number of introns. Phylogenetic trees comparing Hirudo to another distantly related leech species, Helobdella robusta, shows a high degree of orthology, whereas comparison to other annelids shows a relatively low level. Comparisons with other Lophotrochozoans, Ecdyzozoans and with vertebrate pannexins suggest a low number (one to two) of ancestral innexin/pannexins at the protostome/deuterostome split. Whole-mount in situ hybridization for individual genes in early embryos shows that ∼50% of the expressed innexins are detectable in multiple tissues. Expression analyses using quantitative PCR show that ∼70% of the Hirudo innexins are expressed in the nervous system, with most of these detected in early development. Finally, quantitative PCR analysis of several identified adult neurons detects the presence of different combinations of innexin genes, a property that may underlie the participation of these neurons in different adult coupling circuits.
缝隙连接蛋白是所有动物组织(尤其是神经系统)发育和持续功能所必需的信号通路的重要组成部分,它们在细胞内和细胞外小分子信号因子和离子的交换中发挥作用。在基因组测序足够充分的动物中,已经发现了这些蛋白的大家族,包括连接蛋白、pannexin 和 innexin,在秀丽隐杆线虫 Caenorhabditis elegans 中发现了 25 个 innexin,在斑马鱼 Danio rerio 中发现了至少 37 个连接蛋白 Cruciani 和 Mikalsen (Biol Chem 388:253-264, 2009)。最近测序了医用水蛭 Hirudo verbana 的基因组,我们现在报告在该物种中存在 21 个 innexin 基因,比我们之前从 EST 衍生的转录组数据库分析中报告的数量多 9 个 Dykes 和 Macagno (Dev Genes Evol 216: 185-97, 2006); Macagno 等人 (BMC Genomics 25:407, 2010)。基因结构分析表明,根据水蛭 innexin 基因的不同,它们可以包含 0 到 6 个内含子,密切相关的同源基因具有相同数量的内含子。将 Hirudo 与另一种亲缘关系较远的水蛭 Helobdella robusta 进行比较的系统发育树表明,它们具有高度的同源性,而与其他环节动物的比较表明,它们的同源性相对较低。与其他 Lophotrochozoans、Ecdyzozoans 和脊椎动物 pannexin 的比较表明,在原口动物/后口动物分裂时,仅有一个到两个祖先 innexin/pannexin。在早期胚胎中对单个基因进行全胚胎原位杂交显示,在多个组织中可以检测到约 50%的表达 innexin。使用定量 PCR 进行的表达分析表明,约 70%的 Hirudo innexin 在神经系统中表达,其中大多数在早期发育中检测到。最后,对几种鉴定的成年神经元进行定量 PCR 分析检测到不同组合的 innexin 基因的存在,这一特性可能是这些神经元参与不同成年耦合回路的基础。