Department of Applied Plant Science, Graduate School of Agricultural Sciences, Tohoku University, Aoba-ku, Sendai, Japan.
Plant Physiol. 2010 Nov;154(3):1196-209. doi: 10.1104/pp.110.158519. Epub 2010 Aug 31.
Autophagy is an intracellular process facilitating the vacuolar degradation of cytoplasmic components and is important for nutrient recycling during starvation. We previously demonstrated that chloroplasts can be partially mobilized to the vacuole by autophagy via spherical bodies named Rubisco-containing bodies (RCBs). Although chloroplasts contain approximately 80% of total leaf nitrogen and represent a major carbon and nitrogen source for new growth, the relationship between leaf nutrient status and RCB production remains unclear. We examined the effects of nutrient factors on the appearance of RCBs in leaves of transgenic Arabidopsis (Arabidopsis thaliana) expressing stroma-targeted fluorescent proteins. In excised leaves, the appearance of RCBs was suppressed by the presence of metabolic sugars, which were added externally or were produced during photosynthesis in the light. The light-mediated suppression was relieved by the inhibition of photosynthesis. During a diurnal cycle, RCB production was suppressed in leaves excised at the end of the day with high starch content. Starchless mutants phosphoglucomutase and ADP-Glc pyrophosphorylase1 produced a large number of RCBs, while starch-excess mutants starch-excess1 and maltose-excess1 produced fewer RCBs. In nitrogen-limited plants, as leaf carbohydrates were accumulated, RCB production was suppressed. We propose that there exists a close relationship between the degradation of chloroplast proteins via RCBs and leaf carbon but not nitrogen status in autophagy. We also found that the appearance of non-RCB-type autophagic bodies was not suppressed in the light and somewhat responded to nitrogen in excised leaves, unlike RCBs. These results imply that the degradation of chloroplast proteins via RCBs is specifically controlled in autophagy.
自噬是一种细胞内过程,有助于溶酶体降解细胞质成分,在饥饿时对于营养物质的回收很重要。我们之前的研究表明,叶绿体可以通过自噬被部分转移到液泡中,形成一种被称为 Rubisco 体(RCB)的球形结构。虽然叶绿体含有叶片中大约 80%的总氮,是新生长的主要碳氮来源,但叶片养分状况与 RCB 形成之间的关系尚不清楚。我们研究了营养因素对表达质体靶向荧光蛋白的转基因拟南芥(Arabidopsis thaliana)叶片中 RCB 形成的影响。在离体叶片中,代谢糖的存在会抑制 RCB 的形成,这些糖可以从外部添加,也可以在光照下光合作用中产生。光合作用的抑制会缓解这种光介导的抑制。在一个昼夜周期中,在一天结束时淀粉含量较高的叶片中,RCB 的形成受到抑制。缺乏磷酸葡萄糖变位酶和 ADP-Glc 焦磷酸化酶 1 的淀粉缺陷突变体产生了大量的 RCB,而淀粉过量的突变体 starch-excess1 和麦芽糖过量的突变体 maltose-excess1 则产生较少的 RCB。在氮限制的植物中,随着叶片碳水化合物的积累,RCB 的形成受到抑制。我们提出,在自噬中,通过 RCB 降解叶绿体蛋白与叶片碳而不是氮的状态之间存在密切关系。我们还发现,与 RCB 不同的是,非 RCB 型自噬体的形成在光照下不受抑制,并且在离体叶片中对氮有一定的反应。这些结果表明,通过 RCB 降解叶绿体蛋白的自噬受到特异性调控。