水稻淀粉胚乳程序性细胞死亡中的去核化

THE STARCHY ENDOSPERM DENUCLEATION BY A PROCESS OF PROGRAMMED CELL DEATH DURING RICE GRAIN DEVELOPMENT

对水稻品种中籼8836淀粉胚乳细胞的去核化发育阶段的细胞超微结构变化和同期籽粒灌浆速率及相关酶活性的动态进行了观察和分析。开花受精后约在第3天胚乳完成细胞化,花后第5天少数淀粉胚乳细胞启动去核发育过程。核消亡是淀粉胚乳细胞程序性细胞死亡(PCD)的第一步。同一籽粒淀粉胚乳细胞的去核进程是不同步的。花后第13天所有淀粉胚乳细胞都已完成去核过程。在去核过程中,胚乳核的形态变化特征既有动植物PCD的共性又有其特殊性。伴随核降解过程,一部分线粒体解体,表明去核化与线粒体解体有一定联系。在去核化发育阶段,与PCD有关的酶类,如超氧化物歧化酶(SOD)过氧化氢酶(CAT)活性非常高;与淀粉合成有关的酶类,如ADPG焦磷酸化酶、可溶性淀粉合成酶(SSS酶)、淀粉分支酶(或Q酶)也表现出很高的活性。去核化发育阶段籽粒灌浆速率最高,籽粒增重亦最快。淀粉胚乳细胞去核之后,细胞并未立即死亡,这些无核的细胞仍维持正常有序的代谢活动,继续进行淀粉和贮藏蛋白的合成与积累,但上述酶类的活性明显降低,灌浆速率也明显趋缓。淀粉胚乳细胞最终被贮藏物质充满时成为死细胞,完成其程序性死亡过程。Evan’s blue染色鉴定表明淀粉胚乳细胞死亡不同步,细胞死亡在淀粉胚乳组织中是随机发生的。

The cellular ultrastructural morphology, grain filling ratio and activity of related enzymes during denucleation development stage in starchy endosperm cell of an Indica rice variety Zhongxian 8836 were observed and analyzed. The endosperm cellularization was completed at about the 3rd day after flowering (DAF). At 5 DAF a few endosperm cells initiated denucleation development. At 13 DAF almost all the starchy endosperm cells completed their denucleation. The nuclear degradation is the first stage of programmed cell death (PCD) in starchy endosperm. It occurred unsynchronously among different starchy endosperm cells. The nuclear degradation of starchy endosperm cell during denucleation development stage showed not only morphological features commonly observed in animal and plant PCD, but also some unique characteristics. Mitochondria degeneration was observed along with nuclear degradation, indicating there were interrelations between the two processes. Enzymes related to PCD, such as super-oxide dismutases (SOD) and catalase (CAT), as well as enzymes related to starch synthesis, such as ADP-glu-cose pyriphosphorylase (AGP), soluble starch synthase (SSS) and starch branching enzyme or Q- enzyme, showed very high activity during the denucleation development stage. Maximum grain filling rate and grain weight increase were also achieved in the denucleation developing stage of most starchy endosperm cells. The denucleated cell remained alive in the developing endosperm, keeping its normal metabolisms, the synthesis and accumulation of starch and storage proteins. However, enzyme activities and grain filling rate were apparently dropped to a lower level after denucleation. The starchy endosperm cell finally completed its PCD process when it was completely filled with reserves. Evan's blue staining indicated that cell death occurred unsynchronously among the starchy endosperm cells with initiation points randomly distributed in the endosperm tissue.