Iida Kei, Go Mitiko
Faculty of Bio-Science, Nagahama Institute of Bio-Science and Technology, Siga, Japan.
Mol Biol Evol. 2006 May;23(5):1085-94. doi: 10.1093/molbev/msj118. Epub 2006 Mar 6.
The serine/arginine-rich (SR) protein family plays an important role in constitutive and alternative splicing (AS). These proteins regulate AS in a tissue-specific and stress-responsive manner. Pre-mRNAs encoding SR proteins are often alternatively spliced, and these AS events may be important for the regulation of AS events of other pre-mRNAs. In this study, we analyzed AS events of SR proteins in Arabidopsis thaliana and Oryza sativa (rice). We found three sets of AS events conserved between Arabidopsis and rice. These conserved AS events were found in the plant-novel-SR protein, SC35-like (SCL), and two-Zn-knuckles-type 9G8 subfamilies. Each member of these subfamilies has at least one RNA recognition motif (RRM) and at least one intron in the RRM-encoded region. We found that the conserved AS events occurred in these introns and, in each case, the conserved AS events resulted in mature mRNAs encoding proteins with incomplete RRMs. To search for the evolutionary origin of these AS events, we analyzed SR proteins in Physcomitrella patens (moss) in addition to those in Arabidopsis and rice. We found moss homologues of the plant-novel-SR protein, SCL, and the two-Zn-knuckles-type 9G8 subfamilies in silico, and these homologues have long introns at the same location of the conserved AS sites in Arabidopsis and rice. Such long introns are quite specific for alternatively spliced introns concerning the Arabidopsis SR protein genes. The long introns found in the moss SR protein genes strongly suggested that conserved AS events in moss SR protein genes might be similar to those in Arabidopsis and rice. We traced the evolutionary origin of the conserved AS events to 400 MYA, when plants first invaded land. These events are likely important in the regulation of whole AS events and likely contribute to the complicated transcriptome described by AS. The complicated transcriptome created by regulated AS events might have provided plants tolerance against droughts or temperature shifts and given them the ability to live on land.
富含丝氨酸/精氨酸(SR)的蛋白质家族在组成型剪接和可变剪接(AS)中发挥着重要作用。这些蛋白质以组织特异性和应激反应性的方式调节可变剪接。编码SR蛋白的前体mRNA常常发生可变剪接,而这些可变剪接事件可能对其他前体mRNA的可变剪接事件的调节很重要。在本研究中,我们分析了拟南芥和水稻中SR蛋白的可变剪接事件。我们发现了三组在拟南芥和水稻之间保守的可变剪接事件。这些保守的可变剪接事件存在于植物新型SR蛋白、类SC35(SCL)和双锌指型9G8亚家族中。这些亚家族的每个成员在RRM编码区域至少有一个RNA识别基序(RRM)和至少一个内含子。我们发现这些内含子中发生了保守的可变剪接事件,并且在每种情况下,保守的可变剪接事件都导致成熟mRNA编码具有不完整RRM的蛋白质。为了寻找这些可变剪接事件的进化起源,除了拟南芥和水稻中的SR蛋白外,我们还分析了小立碗藓(苔藓)中的SR蛋白。我们在计算机上发现了植物新型SR蛋白、SCL和双锌指型9G8亚家族的苔藓同源物,并且这些同源物在拟南芥和水稻中保守可变剪接位点的相同位置具有长内含子。就拟南芥SR蛋白基因而言,这种长内含子对于可变剪接内含子来说是相当特异的。在苔藓SR蛋白基因中发现的长内含子强烈表明,苔藓SR蛋白基因中的保守可变剪接事件可能与拟南芥和水稻中的相似。我们将保守可变剪接事件的进化起源追溯到4亿年前,即植物首次登陆陆地的时候。这些事件可能在整个可变剪接事件的调节中很重要,并且可能对可变剪接所描述的复杂转录组有贡献。由受调控的可变剪接事件产生的复杂转录组可能为植物提供了对干旱或温度变化的耐受性,并赋予它们在陆地上生存的能力。