水稻叶片早衰突变体的农艺与生理性状研究进展

叶片早衰严重影响水稻产量与品质。研究水稻叶片早衰突变体的农艺及生理性状,有助于了解水稻叶片衰老进程和机理。本文简述了叶片衰老特征和衰老机制,重点综述了水稻叶片早衰突变体的类型、农艺与生理性状研究进展及延缓水稻叶片衰老的栽培调控途径。可为水稻抗衰老品种选育及栽培调控提供理论依据。     

水稻叶片早衰突变体的农艺与生理性状研究进展

叶片早衰严重影响水稻产量与品质。研究水稻叶片早衰突变体的农艺及生理性状,有助于了解水稻叶片衰老进程和机理。本文简述了叶片衰老特征和衰老机制,重点综述了水稻叶片早衰突变体的类型、农艺与生理性状研究进展及延缓水稻叶片衰老的栽培调控途径。可为水稻抗衰老品种选育及栽培调控提供理论依据。     

4PU－30延缓杂交水稻叶片衰老的生理基础

以汕优63连体和离体叶片为材料,研究了4PU-30对水稻叶片衰老进程及叶片内蛋白质、核酸含量和相关酶活力的调控效应。4PU-30能显著地延缓水稻叶片衰老,保绿效果明显。叶片中蛋白质和核酸含量及相关酶活力的变化表明,4PU-30控制叶片衰老过程中蛋白质水解酶和核糖核酸酶比活上升,减缓蛋白质和核酸的降解,是其延缓杂交水稻叶片衰老的主要生理基础之一。     

4PU－30延缓杂交水稻叶片衰老的生理基础

 以汕优63连体和离体叶片为材料，研究了4PU-30对水稻叶片衰老进程及叶片内蛋白质、核酸含量和相关酶活力的调控效应。4PU-30能显著地延缓水稻叶片衰老，保绿效果明显。叶片中蛋白质和核酸含量及相关酶活力的变化表明，4PU-30控制叶片衰老过程中蛋白质水解酶和核糖核酸酶比活上升，减缓蛋白质和核酸的降解，是其延缓杂交水稻叶片衰老的主要生理基础之一。     

水稻剑叶衰老过程中内源IAA的轭合态变化及其氮素调控效应

目的 揭示水稻抽穗开花后剑叶衰老过程的内源自由态IAA和轭合态IAA含量的动态变化及其基因型间的差异,明确氮素营养供应对叶片衰老过程的延缓效应及其与两种化学态IAA含量之间的关系。方法 以剑叶早衰水稻突变体(psf)与其野生型(浙恢7954)为材料,比较了其在大田条件下剑叶衰老过程中的内源IAA含量、IAA类型和YUCCAs家族基因转录表达量差异,并利用水培种植试验设低氮(LN, 1.45 mmol/L)、正常氮(NN, 2.9 mmol/L)和高氮(HN, 5.8 mmol/L)3个氮浓度水平处理,对不同氮素水平下水稻叶片衰老相关指标与两类IAA含量间的关系进行了分析。结果 早衰突变体psf剑叶中的自由态IAA含量显著低于相同取样时期的野生对照,但前者剑叶中的轭合态IAA含量显著高于后者,且随叶片衰老进程呈较明显的上升趋势;在7个YUCCAs同工型基因中,YUCCA1、YUCCA2、YUCCA3、YUCCA4和YUCCA6)的转录表达量随叶片衰老而下降,但YUCCA7则随叶片衰老呈现出先上升后下降的变化趋势。相同灌浆时期相比,绝大多数YUCCAs在早衰突变体psf剑叶中的转录表达量要低于其野生型对照,说明IAA合成代谢较弱是psf剑叶叶绿素含量迅速下降和早衰症状加剧的重要原因之一。其中,YUCCA7转录表达量随叶片衰老而上升,可能对psf已衰老叶片中的IAA合成起重要调控作用;氮浓度对水稻叶片中的IAA含量及其自由态与共轭态的影响表现为,在叶片开始衰老的初期和前中期,低氮处理(LN)引起剑叶中的自由态IAA含量下降和轭合态IAA含量上升,而高氮处理(HN)可诱导叶片中的自由态IAA含量增加,并导致轭合态IAA含量明显降低,但对已严重衰老的叶片而言(psf在抽穗开花后28 d的剑叶),高氮处理下(HN)的自由态IAA和轭合态IAA含量均低于正常氮处理(NN)和低氮处理(LN)。结论 水稻叶片的衰老进程与其叶片器官中的IAA含量及其化学态之间存在较密切联系。其中,轭合态IAA含量在叶片衰老过程的大幅度提升,是早衰突变体psf 剑叶衰老进程加快的一个重要生理特征,而轭合态IAA含量的下降,可能是高氮处理对水稻叶片衰老进程起到延缓作用的一个重要生理调节因素。     

水稻叶片衰老与超氧化物歧化酶(SOD)

水稻早衰是限制产量和品质提高的最重要因素之一。叶片是水稻最重要的源器官,其功能及衰老对水稻产量与品质有重要影响。叶片衰老特性的研究对提高光合潜力、提高肥料等投入的利用率和经济系数有重要指导意义(Ｗａｄａｙ,1991;Ｔｈｏｍａｓ等,1993)。因此,人们很早以前便已开展水稻叶片衰老生理的研究,特别是在近十几年中发展尤为迅速。但是,有关叶片衰老的机制还远没有搞清楚,特别是对原初伤害机制了解更少。衰老是生物界的普遍现象,是生物发展进化过程中不可分割的一部分。在众多有关水稻叶片衰老的学说中,生物自由基伤害学说引起人们广泛…     

水稻叶片衰老的分子机理及研究进展

水稻作为我国最主要的粮食作物,已知在生育期允许的范围内,适当延缓水稻叶片早衰,对产量有很大的作用。中国是最早利用水稻杂种优势的国家,杂交稻尚存在结实率偏低、空秕率较高的问题。有研究表明,叶片和根系早衰是造成这一现象的主要原因。有些杂交稻通过推迟停水期来延缓衰老,并推迟收割期,有明显的增产效果,每延迟一天,平均增产1%左右。1　叶片衰老期间基因表达的变化基因表达参与叶片衰老的调控,在转录水平上,衰老期间的基因表达存在4种模式,即基因激活、表达增强、部分抑制和完全抑制。这4种模式在衰老期间体现在ｍＲＮＡ水平上就是上…     

不同基因型水稻抽穗后衰老进程的比较研究

本实验对汕优63、J85、老来青等三种不同基因型水稻品种衰老过程中的生理生化指标进行了分析。结果表明,随着叶片衰老进程,剑叶中的叶绿素含量逐渐下降,丙二醛含量逐渐上升,而根系活力在抽穗后一直呈下降趋势。进一步比较三种不同类型水稻叶片的衰老进程,发现:(1)在外界环境一致的情况下,三种水稻在这三个指标上变化明显不同,其中粳稻老来青抗衰老程度最高,而最易衰老的是杂交稻汕优63;(2)汕优63在抽穗后5～10d内衰老最为迅速;(3)根系活力的下降早于叶片中与衰老有关的指标变化。这表明在衰老过程中,根系的衰老先于叶片。     

不同基因型水稻抽穗后衰老进程的比较研究

本实验对汕优63、J85、老来青等三种不同基因型水稻品种衰老过程中的生理生化指标进行了分析。结果表明，随着叶片衰老进程，剑叶中的叶绿素含量逐渐下降，丙二醛含量逐渐上升，而根系活力在抽穗后一直呈下降趋势。进一步比较三种不同类型水稻叶片的衰老进程，发现：(1)在外界环境一致的情况下，三种水稻在这三个指标上变化明显不同，其中粳稻老来青抗衰老程度最高，而最易衰老的是杂交稻汕优63；(2)汕优63在抽穗后5～10d内衰老最为迅速；(3)根系活力的下降早于叶片中与衰老有关的指标变化。这表明在衰老过程中，根系的衰老先于叶片。     

水稻叶片衰老研究进展

叶片的衰老是植物生长发育的最后阶段,是一种基因编码的程序性事件。但在农业生产上,水稻叶片的过早衰老已经成为制约水稻发展的重要因素,研究水稻叶片的早衰对水稻育种改良具有重要意义。本文综述了近年来国内外有关水稻叶片衰老研究结果,主要包括水稻叶片衰老的机理、特征、影响叶片衰老的因子、常用的水稻叶片衰老的生理生化指标等,同时介绍了近年来有关水稻叶片衰老遗传研究进展。     

杂交粳稻屉优418及其亲本灌浆期叶片内源激素含量变化与衰老之间的关系

以杂交水稻屉优418及其亲本屉锦和C418为材料,研究了生育后期叶片内源激素含量变化与衰老之间的关系。结果表明杂交水稻及其亲本剑叶中ABA、GA1/3、ZR和IAA含量变化趋势基本一致。3个材料的（GA1/3+ZR+IAA）/ABA的比值变化总的趋势也基本相同,前期较平稳,后期比值迅速下降。在生育后期,屉优418和C418的（GA1/3+ZR+IAA）/ABA比值下降速率较慢,与衰老相对较慢一致;而屉锦的（GA1/3+ZR+IAA）/ABA比值下降速率较快,与其衰老较快相一致。     

Relationship Between Changes in Leaf Endogenous Hormone Contents and Senescence During Grain Filling Stage of a Rice Hybrid and its Parents

A major problem in hybrid rice production is the occurrence of leaf senescence during the grain filling stage that can result in reduction of yield. Changes in contents of several endogenous hormones are related to leaf senescence. The relationship between endogenous hormones and leaf senescence in the rice hybrid Tiyou 418 and its parents Tijin and C418, was undertaken for investigation. Indicators of leaf senescence, including superoxide dismutase (SOD) activity, malondialdehyde (MDA) accumulation and chlorophyll content, as well as the contents of abscisic acid (ABA), zeatin riboside (ZR), gibberellin (GA1/3) and auxin (IAA) in the leaves were determined. Different rates of leaf senescence were observed in the three materials. Senescence occurred earliest and fastest in Tijin, followed by Tiyou 418 and then C418. A similar trend was recorded in ABA, ZR, and IAA contents during the grain filling stage in the three materials. Changes in (GA1/3 ZR IAA)/ABA ratios were also similar, being quite stable during the early stage of leaf senescence, and decreasing markedly during the late stage. The ratio declined more dramatically in Tijin, in accordance with its faster leaf senescence. The results suggest that the ratio of (GA1/3 ZR IAA)/ABA regulates chlorophyll content, SOD activity, MDA content and membrane lipid peroxidation. It is postulated that endogenous hormones may play a role in the regulation of leaf senescence in a systematic way.     

Relationship Between Changes in Leaf Endogenous Hormone Contents and Senescence During Grain Filling Stage of a Rice Hybrid and its Parents

A major problem in hybrid rice production is the occurrence of leaf senescence during the grain filling stage that can result in reduction of yield. Changes in contents of several endogenous hormones are related to leaf senescence. The relationship between endogenous hormones and leaf senescence in the rice hybrid Tiyou 418 and its parents Tijin and C418, was undertaken for investigation. Indicators of leaf senescence, including superoxide dismutase (SOD) activity, malondialdehyde (MDA) accumulation and chlorophyll content, as well as the contents of abscisic acid (ABA), zeatin riboside (ZR), gibberellin (GA1/3) and auxin (IAA) in the leaves were determined. Different rates of leaf senescence were observed in the three materials. Senescence occurred earliest and fastest in Tijin, followed by Tiyou 418 and then C418. A similar trend was recorded in ABA, ZR, and IAA contents during the grain filling stage in the three materials. Changes in (GA1/3 ZR IAA)/ABA ratios were also similar, being quite stable during the early stage of leaf senescence, and decreasing markedly during the late stage. The ratio declined more dramatically in Tijin, in accordance with its faster leaf senescence. The results suggest that the ratio of (GA1/3 ZR IAA)/ABA regulates chlorophyll content, SOD activity, MDA content and membrane lipid peroxidation. It is postulated that endogenous hormones may play a role in the regulation of leaf senescence in a systematic way.     

稻田冬季保护性耕作条件下的土壤酶活性与水稻成熟期叶片衰老和籽粒产量之间的关系

 为研究稻田冬季保护性耕作不同种植模式对土壤酶活性以及水稻成熟期叶片衰老的影响, 设计了5种种植模式：免耕直播稻草全程覆盖马铃薯（T1）、免耕直播黑麦草（T2）、免耕直播油菜（T3）、免耕直播燕麦草（T4）和空闲（CK）,供试品种为杂交组合国稻6号。与对照（CK）相比,T1、T2和T3处理均可显著提高土壤蛋白酶、中性磷酸酶、过氧化氢酶以及转化酶活性,尤其是T1处理,其土壤酶活性明显高于其他处理,而T4处理与对照相比,差异不明显。T1、T2和T3处理均可有效延缓水稻成熟期剑叶的衰老,主要表现为成熟期剑叶中的超氧化物歧化酶、过氧化物酶活性、可溶性蛋白质和叶绿素含量下降缓慢,丙二醛含量上升的幅度较小,相对于对照（CK）,差异均达显著水平。在本试验条件下,土壤酶对水稻叶片衰老的影响程度依次为蛋白酶>过氧化氢酶>中性磷酸酶>转化酶。叶片衰老对水稻产量的影响主要表现在灌浆中后期。     

Relationship between grain filling duration and leaf senescence of temperate rice under high temperature

High temperature during grain filling period has been reported to decrease the grain filling duration, leading to the lower grain weight and yield of rice. Two experiments in the phytotron and field were carried out to test the hypothesis that the leaf senescence of rice plants may determine the grain filling duration under high temperature. In the phytotron experiment in 2008, rice plants of a japonica cultivar “Ilpumbyeo” were subjected to three minimum/maximum (mean) temperature regimes of 11/19 (15), 17/25 (21), and 23/31 °C (27 °C). In the field experiment, rice seedlings of the same rice cultivar were transplanted on May 6th and June 19th in 2009 and the mean temperatures during the grain filling period were 24.4 and 21.9 °C, respectively. Both experiments revealed consistently that high temperature increased the rates of grain filling and leaf senescence while it reduced the durations of them. However, grain filling was terminated earlier than complete leaf senescence, the time gap being greater at higher temperature. In addition, the fraction of dry matter partitioning to the leaf sheath + culm resumed to increase following the termination of grain filling under high temperature, indicating that leaves were still maintaining photosynthetic capacity and supplying assimilates into the other plant tissues except grain even after the termination of grain filling. These findings suggest that an early termination of grain filling in temperate rice under high temperature was not resulted from the lack of assimilate owing to the early leaf senescence but from the loss of sink activity owing to the earlier senescence of panicle.     

水稻结实期间叶片衰老与膜脂过氧化的关系

 以结实期间的稻为材料，在抽穗开花期通过剪去穗部部分枝梗和全穗等处理后，测定作为叶片衰老指标的叶绿素和蛋白质含量及作为膜脂过氧化指标的丙二醛含量的变化，探讨了水稻结实期间叶片衰老与膜脂过氧化之间的关系。试验结果指出，在水稻抽穗开花期间，剪去穗部枝梗或整个穗子，叶片中的叶绿素和蛋白质含量下降速率在处理后的7 d内就开始明显减慢，而MDA含量增加速率只有在处理7 d后才开始减缓，SOD活性并不因剪穗处理而有所提高。表明因剪穗(或穗部枝梗)而导致的叶绿素和蛋白质含量的变化早于MDA含量的变化，与SOD活性的变化无直接关系。说明在结实期间，水稻叶片在衰老过程中表现出的叶绿素和蛋白质含量的下降，并不主要是由于SOD的活性降低而导致膜脂过氧化作用增强所造成的结果，SOD活性下降和MDA含量的增加可能与叶绿素及蛋白质含量减少一样，只是叶片衰老开始后的一种结果，而非引起叶片衰老的原因。     

水稻水作与旱作对抽穗后剑叶光合特性、衰老及根系活性的影响

对水稻水作与旱作栽培模式下水稻衰老和后期一些生理活性的影响进行了研究。结果表明,由于水分的影响,水稻旱作不同程度加速了水稻生理机能的衰退,使籽粒干物质积累受阻,粒重下降而减产。水稻水作不仅改善了水稻剑叶光合因子,提高了光合效率,而且剑叶呼吸速率下降缓慢,相对电导率上升缓慢,SOD酶活性较高,MDA含量低,表明剑叶后期衰老进程慢,促进籽粒增重,产量高。此外,旱作水稻生育末期根系干物质积累、活力高及代谢旺盛的优势对籽粒增重也是一种负向影响;而水稻水作开花-灌浆期根系活力高,生育末期根系生理活性迅速衰退,保证了籽粒的充实