植物叶片衰老过程中基因的表达与调控

从衰老相关基因的分离、克隆、表达、调控及叶片衰老延缓几个方面,介绍叶片衰老过程中基因表达调控的研究进展。     

叶片衰老的特性及分子调控（综述）

就叶片衰老研究在生理,生化及分子水平上的最新进展,以及有希望应用于农业,操纵叶片衰老的转基因手段作一简要综述。     

水稻叶片衰老相关基因的研究进展

水稻叶片的衰老是制约杂交稻产量提高的主要因素之一,有数据表明水稻籽粒灌浆所需营养物质的60%～80%来自叶片的光合作用,实践证明叶片每推迟1天衰老,产量可提高产1%左右.因此,对叶片衰老的形态、生理生化及其相关分子机理等进行研究具有重要的现实意义.近年来水稻叶片衰老的相关研究表明,叶片的衰老是一个受众多因素影响的复杂过程,在这个过程中叶片发生了巨大的形态与生理生化变化,而这些变化均离不开基因的调控作用.大量实验结果表明:在衰老过程中,叶片细胞有选择地启动或增强某些基因(叶片衰老相关基因)的表达,而关闭或减弱另一些基因(衰老下调基因)的表达,由此来调控叶片衰老的进程.目前研究者已在研究衰老突变体等相关的材料中发现了许多与水稻叶片衰老有关的基因.本文重点概述了近年来水稻叶片衰老相关基因的研究状况,并对未来研究方向等问题做了思考与探讨,以期能为开展进一步的研究工作提供参考.     

光质对设施葡萄叶片衰老与内源激素含量的影响

以设施延迟栽培条件下叶片衰老速度不同的意大利和无核白鸡心2个葡萄品种为试材,分别进行补充红光和蓝光处理,研究不同光质对叶片衰老过程中叶绿素含量、净光合速率和内源激素含量的影响.结果表明: 与未补光对照相比,红光能够显著提高叶片的叶绿素含量和净光合速率,降低了内源赤霉素(GA₃)含量,但明显减缓了脱落酸(ABA)含量的增加和玉米素核苷(ZR)总含量的减少,从而显著提高了(GA₃+ZR)/ABA值,延缓叶片衰老.叶片衰老前期,蓝光处理叶片叶绿素含量、净光合速率和(GA₃+ZR)/ABA值均低于对照,加速了植株的衰老进程;但在叶片衰老后期,蓝光处理叶绿素含量、净光合速率和(GA₃+ZR)/ABA值逐渐高于对照,在一定程度上延缓了叶片衰老.植物内源激素生长素(IAA)则表现出叶片衰老前期促进叶片生长发育、叶片衰老后期加速衰老的双重作用.意大利叶片衰老速度较无核白鸡心慢.在本试验条件下,红光处理效果最好,有效延缓了叶片衰老进程,延长了叶片的生理功能期.     

水稻叶片衰老过程中氨肽酶活性的变化

水稻叶片中存在着氨肽酶,其最适反应pH和最适反应温度分别为8.2℃和40℃,酶促反应的产物量在最初30min内与时间呈直线相关。 水稻叶片衰老过程中叶绿素和蛋白质含量下降,而氨肽酶比活上升;用植物激素延缓或促进叶片衰老蛋白质降解的同时也抑制或促进了氨肽酶比活的上升,说明氨肽酶在水稻叶片衰老蛋白质降解过程中起一定的作用。根据水稻叶片衰老过程中大分子化合物和叶片外部形态的变化,可将叶片衰老过程划分为缓衰期、急衰期和竭衰期。     

植物激素在叶片衰老中的作用机制研究进展

叶片衰老是叶片逐渐丧失光合作用并进入细胞死亡的一个复杂的叶片发育过程,与农作物的产量和品质密切相关。植物生长发育信号和外界环境因素通过多种激素信号途径调节叶片衰老的起始和进程。本文简要综述了八种常见激素在叶片衰老中的研究进展,并总结了这些激素在叶片衰老调控中的作用方式和机制。     

植物叶片在衰老期间膜结构变化研究进展

引起生物体的器官或某一部分最终走向死亡的变化即为衰老(senescence),所有高等植物在其生活史上的任何阶段,或在任何结构水平上,衰老均可发生。衰老作为植物生长发育中的重要事件,受到研究者们的极大关注。1908年minot首先对植物的衰老进行了研究,对植物器官衰老研究最多的是叶片和子叶,一般地,子叶的衰老与叶片的衰老没有本质的差异。     

苗期玉米叶片碳氮平衡与干旱诱导的叶片衰老之关系

为了探究干旱诱导的碳氮平衡破坏与干旱诱导的叶片衰老之间的关系,该实验以8个在干旱胁迫下叶片衰老进程有明显差异的玉米品种为实验材料,采用PEG模拟干旱处理,通过测定光合速率、叶绿素含量和叶绿素荧光参数等叶片衰老指标以及非结构性碳水化合物(可溶性糖、淀粉)和全氮含量等变化,分析玉米中干旱诱导的叶片衰老与叶片中碳氮平衡(碳氮比)之间的关系。结果显示:(1)干旱胁迫下,8个玉米品种叶片净光合速率受到严重抑制,Fv/Fm大幅下降,叶绿素含量显著降低,说明干旱诱导了玉米叶片的衰老;(2)干旱诱导玉米叶片衰老的同时,8个玉米品种的叶片中可溶性糖含量显著升高,淀粉含量小幅上升,全氮含量大幅降低,碳氮比显著升高,碳氮平衡遭到了破坏;(3)8个玉米品种叶片的叶绿素含量与非结构性碳水化合物含量以及碳氮比呈极显著负相关关系,与全氮含量呈极显著正相关关系。因此,碳氮代谢与干旱诱导的叶片衰老紧密联系,碳氮平衡可能参与了干旱诱导的叶片衰老调控。     

植物Whirly蛋白调控叶片衰老的研究进展

植物衰老是植物细胞生长发育的最后一个阶段,其启动的早晚对植物生物量和品质的形成有很大影响。叶片衰老是植物衰老的主要形式,受到内外环境因素的诱导,并被多种转录因子介导的信号传导途径所调控。对叶片衰老调控机制的研究一直是植物衰老研究中的重点。Whirly蛋白作为一类广泛存于植物中的特异转录因子小家族,能与单链DNA分子结合,双定位于细胞器(线粒体或叶绿体)与细胞核中,在植物细胞核和细胞器中发挥多种功能,参与对植物叶片衰老的调控。本文概述了植物Whirly蛋白的结构和定位,重点阐述了Whirly蛋白的功能与细胞衰老关系及其对叶片衰老调节机理的研究进展等,并对未来的研究方向进行了展望。     

植物叶片衰老的分子机制

文章就叶片衰老过程中基因表达调控机制的研究进展作了介绍     

Hormonal regulation of leaf senescence in Lilium

In addition to floral senescence and longevity, the control of leaf senescence is a major factor determining the quality of several cut flowers, including Lilium, in the commercial market. To better understand the physiological process underlying leaf senescence in this species, we evaluated: (i) endogenous variation in the levels of phytohormones during leaf senescence, (ii) the effects of leaf darkening in senescence and associated changes in phytohormones, and (iii) the effects of spray applications of abscisic acid (ABA) and pyrabactin on leaf senescence. Results showed that while gibberellin 4 (GA(4)) and salicylic acid (SA) contents decreased, that of ABA increased during the progression of leaf senescence. However, dark-induced senescence increased ABA levels, but did not affect GA(4) and SA levels, which appeared to correlate more with changes in air temperature and/or photoperiod than with the induction of leaf senescence. Furthermore, spray applications of pyrabactin delayed the progression of leaf senescence in cut flowers. Thus, we conclude that (i) ABA plays a major role in the regulation of leaf senescence in Lilium, (ii) darkness promotes leaf senescence and increases ABA levels, and (iii) exogenous applications of pyrabactin inhibit leaf senescence in Lilium, therefore suggesting that it acts as an antagonist of ABA in senescing leaves of cut lily flowers.     

Substantial variation in leaf senescence times among 1360 temperate woody plant species: implications for phenology and ecosystem processes

Background and Aims Autumn leaf senescence marks the end of the growing season in temperate ecosystems. Its timing influences a number of ecosystem processes, including carbon, water and nutrient cycling. Climate change is altering leaf senescence phenology and, as those changes continue, it will affect individual woody plants, species and ecosystems. In contrast to spring leaf out times, however, leaf senescence times remain relatively understudied. Variation in the phenology of leaf senescence among species and locations is still poorly understood.Methods Leaf senescence phenology of 1360 deciduous plant species at six temperate botanical gardens in Asia, North America and Europe was recorded in 2012 and 2013. This large data set was used to explore ecological and phylogenetic factors associated with variation in leaf senescence.Key Results Leaf senescence dates among species varied by 3 months on average across the six locations. Plant species tended to undergo leaf senescence in the same order in the autumns of both years at each location, but the order of senescence was only weakly correlated across sites. Leaf senescence times were not related to spring leaf out times, were not evolutionarily conserved and were only minimally influenced by growth habit, wood anatomy and percentage colour change or leaf drop. These weak patterns of leaf senescence timing contrast with much stronger leaf out patterns from a previous study.Conclusions The results suggest that, in contrast to the broader temperature effects that determine leaf out times, leaf senescence times are probably determined by a larger or different suite of local environmental effects, including temperature, soil moisture, frost and wind. Determining the importance of these factors for a wide range of species represents the next challenge for understanding how climate change is affecting the end of the growing season and associated ecosystem processes.     

Photosynthesis and leaf-nitrogen dynamics during leaf senescence of tropical maize cultivars in hydroponics in relation to N efficiency in the field

The selection process of nitrogen (N)-efficient cultivars during plant breeding could be simplified by a specification of secondary plant traits that are decisive for N efficiency. It was shown that leaf senescence under N deprivation of sixteen tropical maize cultivars in a short-term nutrient solution experiment was related to leaf senescence and grain yield under N deficiency (N efficiency) in field experiments. In this study we investigated if a quantification of leaf- and plant-N flows by ¹⁵N labelling can improve the evaluation of genotypic differences in leaf senescence in short-term experiments. Cultivars differed in leaf-N content prior to senescence; however, this appeared to have no significant impact on the development of leaf senescence. N import into senescing leaves was not related to total plant N uptake, but seems to have been regulated by leaf-inherent factors. Leaf N remaining in the leaf seems to have comprised inefficiently remobilized leaf N, at least during early senescence stages. Photosynthetic rate and chlorophyll contents at early senescence stages depended on additional factors to leaf-N content. Nevertheless, all parameters used to characterize leaf senescence were related to leaf senescence at anthesis in field experiments. However, only photosynthetic rate during late leaf senescence reflected cultivar differences in leaf senescence during reproductive growth and N efficiency in field experiments.     

Differences in Sugar Accumulation and Mobilization between Sequential and Non-Sequential Senescence Wheat Cultivars under Natural and Drought Conditions

Wheat leaf non-sequential senescence at the late grain-filling stage involves the early senescence of younger flag leaves compared to that observed in older second leaves. On the other hand, sequential senescence involves leaf senescence that follows an age-related pattern, in which flag leaves are the latest to undergo senescence. The characteristics of sugar metabolism in two sequential senescence cultivars and two non-sequential senescence cultivars under both natural and drought conditions were studied to elucidate the underlying mechanism of drought tolerance in two different senescence modes. The results showed that compared to sequential senescence wheat cultivars, under natural and drought conditions, non-sequential senescence wheat cultivars showed a higher leaf net photosynthetic rate, higher soluble sugar levels in leaves, leaf sheaths, and internodes, higher leaf sucrose synthase (SS) and sucrose phosphate synthase (SPS) activity, and higher grain SS activity, thereby suggesting that non-sequential senescence wheat cultivars had stronger source activity. Spike weight, grain weight per spike, and 100-grain weight of non-sequential senescence cultivars at maturity were significantly higher than those of sequential senescence cultivars under both natural and drought conditions. These findings indicate that the higher rate of accumulation and the higher mobilization of soluble sugar in the leaves, leaf sheaths and internodes of non-sequential senescence cultivars improve grain weight and drought tolerance. At the late grain-filling stage, drought conditions adversely affected leaf chlorophyll content, net photosynthetic rate, soluble sugar and sucrose content, SS and SPS activity, gain SS activity, and weight. This study showed that higher rates of soluble sugar accumulation in the source was one of the reasons of triggering leaf non-sequential senescence, and higher rates of soluble sugar mobilization during leaf non-sequential senescence promoted high and stable wheat yield and drought tolerance.     

Arabidopsis onset of leaf death mutants identify a regulatory pathway controlling leaf senescence

The onset of leaf senescence is controlled by leaf age and ethylene can promote leaf senescence within a specific age window. We exploited the interaction between leaf age and ethylene and isolated mutants with altered leaf senescence that are named as onset of leaf death (old) mutants. Early leaf senescence mutants representing three genetic loci were selected and their senescence syndromes were characterised using phenotypical, physiological and molecular markers. old1 is represented by three recessive alleles and displayed earlier senescence both in air and upon ethylene exposure. The etiolated old1 seedlings exhibited a hypersensitive triple response. old2 is a dominant trait and the mutant plants were indistinguishable from the wild-type when grown in air but showed an earlier senescence syndrome upon ethylene treatment. old3 is a semi-dominant trait and its earlier onset of senescence is independent of ethylene treatment. Analyses of the chlorophyll degradation, ion leakage and SAG expression showed that leaf senescence was advanced in ethylene-treated old2 plants and in both air-grown and ethylene-treated old1 and old3 plants. Epistatic analysis indicated that OLD1 might act downstream of OLD2 and upstream of OLD3 and mediate the interaction between leaf age and ethylene. A genetic model was proposed that links the three OLD genes and ethylene into a regulatory pathway controlling the onset of leaf senescence.