Plattner Helmut, Verkhratsky Alexei
Faculty of Biology, University of Konstanz, 78457 Konstanz, Germany.
Faculty of Biological Sciences, University of Manchester, Manchester M13 9PT, UK; Achucarro Centre for Neuroscience, IKERBASQUE, Basque Foundation for Science, 48011 Bilbao, Spain; University of Nizhny Novgorod, Nizhny Novgorod 603022, Russia.
Cell Calcium. 2015 Mar;57(3):123-32. doi: 10.1016/j.ceca.2014.12.004. Epub 2014 Dec 15.
Molecular cascades of calcium homeostasis and signalling (Ca(2+) pumps, channels, cation exchangers, and Ca(2+)-binding proteins) emerged in prokaryotes and further developed at the unicellular stage of eukaryote evolution. With progressive evolution, mechanisms of signalling became diversified reflecting multiplication and specialisation of Ca(2+)-regulated cellular activities. Recent genomic analysis of organisms from different systematic positions, combined with proteomic and functional probing invigorated expansion in our understanding of the evolution of Ca(2+) signalling. Particularly impressive is the consistent role of Ca(2+)-ATPases/pumps, calmodulin and calcineurin from very early stages of eukaryotic evolution, although with interspecies differences. Deviations in Ca(2+) handling and signalling are observed between vertebrates and flowering plants as well as between protists at the basis of the two systematic categories, Unikonta (for example choanoflagellates) and Bikonta (for example ciliates). Only the B-subunit of calcineurin, for instance, is maintained to regulate highly diversified protein kinases for stress defence in flowering plants, whereas the complete dimeric protein, in vertebrates up to humans, regulates gene transcription, immune-defence and plasticity of the brain. Calmodulin is similarly maintained throughout evolution, but in plants a calmoldulin-like domain is integrated into protein kinase molecules. The eukaryotic cell has inherited and invented many mechanisms to exploit the advantages of signalling by Ca(2+), and there is considerable overall similarity in basic processes of Ca(2+) regulation and signalling during evolution, although some details may vary.
钙稳态与信号传导的分子级联反应（钙泵、通道、阳离子交换体以及钙结合蛋白）在原核生物中出现，并在真核生物进化的单细胞阶段进一步发展。随着进化的推进，信号传导机制变得多样化，这反映了钙调节细胞活动的增加和特化。最近对来自不同系统位置的生物体进行的基因组分析，结合蛋白质组学和功能探测，增强了我们对钙信号传导进化的理解。特别令人印象深刻的是，从真核生物进化的早期阶段起，钙ATP酶/泵、钙调蛋白和钙神经磷酸酶就发挥着一致的作用，尽管存在种间差异。在脊椎动物和开花植物之间，以及在两个系统类别（单鞭毛生物，如领鞭毛虫；双鞭毛生物，如纤毛虫）基础上的原生生物之间，观察到钙处理和信号传导的偏差。例如，只有钙神经磷酸酶的B亚基在开花植物中得以保留，用于调节高度多样化的蛋白激酶以进行应激防御，而在脊椎动物直至人类中，完整的二聚体蛋白则调节基因转录、免疫防御和大脑可塑性。钙调蛋白在整个进化过程中同样得以保留，但在植物中，一个类钙调蛋白结构域被整合到蛋白激酶分子中。真核细胞继承并发明了许多机制来利用钙信号传导的优势，尽管一些细节可能有所不同，但在进化过程中钙调节和信号传导的基本过程总体上有相当大的相似性。
