Ribulose-1,5-Diphosphate Carboxylase Protein during Flag Leaf Senescence

Ribulose-l,5-diphosphate (RuDP) carboxylase protein and activitywere determined in relation to net photosynthetic rate duringthe senescence of intact flag leaves of wheat on the plant.Initially the decrease in RuDP carboxylase activity was greaterthan the decline in net photosynthesis. The major decrease inRuDP carboxylase activity over this period resulted from a decreasein enzyme specific activity from 11 to 2 µmol CO₂ fixedh^–1 mg^–1 protein. Loss of RuDP carboxylase proteindid not occur until late in senescence by which time chlorophyllconcentration had decreased by more than 50%. Treatment of flagleaves at weekly intervals with either 1000 parts 10^–62-chloro-ethyltrimethylammonium chloride or 100 parts 10^–6gibberellic acid with 1 part 10^–6 kinetin did not significantlyaffect net photosynthetic rate, RuDP carboxylase protein oractivity during senescence.     

Translocation from the Flag Leaf of Winter Wheat in the Field

Translocation of assimilate from the flag leaf of winter wheat(Triticum aestivum cv. Maris Huntsman) was studied in the fieldby monitoring the export of photo-assimilated [^l4C]carbon dioxidewith a Geiger-Miiller counter placed under the fed area of leaf.The resulting export curve was analysed as a sum of two exponentialterms, and interpreted as a two-pool compartmental system. Therate constant for export from the leaf increased slightly frommaximum elongation to anthesis, then declined to almost halfits peak value just before the leaf lost all visible chlorophyll.The inter-pool transfer rate constants did not change significantlyover the same period, but all rate constants varied with timeof day. Short-term changes in the environment of the flag leaf had nodiscernible influence on translocation in the field. The timeconstants of the two pools of assimilate agreed with those forother species reported in the literature. These results areconsistent with the suggestion that sucrose is stored in thevacuole of mesophyll cells. The variations in rate constants with time of day, and deviationsof the export data from the two-pool model, suggest that exportand inter-pool transport have saturation kinetics. A model withMichaelis-Menten kinetics was formulated, and simulations ofthis model showed similar deviations from a simple two-poolsystem to those seen in our data.     

小麦旗叶早衰性状的QTL定位

为小麦旗叶早衰性状的精细定位和基因克隆奠定基础,该试验以普通小麦(Triticum aestivum L.)‘宁春4号’和‘宁春27号’杂交得到的128个F10代RIL群体为研究材料,利用307对多态性SSR标记对小麦旗叶早衰性状进行了QTL定位,并通过构建整合图谱的方法进行了标记加密。结果表明,共检测到1个控制旗叶早衰性状的加性QTL,位于2A染色体长臂的gwm526和gwm382标记区间内,可解释49.88%的表型变异。经遗传图谱整合后发现,gwm526和gwm382标记之间存在124个SNP标记。     

Senescence of Detached Fern Leaves

Detached fern leaves show a loss in the content of pigments,protein and a decline in 2,6-dichlorophenol indophenol Hillreaction of isolated chloroplasts during aging. Light treatmentof leaves could considerably retard aging induced loss in 2,6-dichlorophenolindophenol photoreduction but it exhibits less effect on pigmentand protein loss. (Received February 7, 1983; Accepted July 14, 1983)     

Senescence of Flag Leaves and Ears of Wheat Hastened by Methyl Jasmonate

Treatment of flag leaves and ears of wheat plants with MJ (jasmonic acid methylester) (10⁻⁵ and 10⁻⁴ m) did not increase ethylene production, but it did accelerate senescence as indicated by the loss of chlorophyll. MJ also caused the closure of stomata, and consequently the rates of transpiration and photosynthesis decreased. Early maturity shortened the grain filling period, so the thousand grain weight was lower. Although ethylene elicited the same physiologic effects, the syndrome of senescence by MJ is independent of the former. We conclude that senescence and death in wheat are far from being elucidated; however, MJ and ethylene seem to participate in the phenomenon. Received July 10, 1997; accepted January 5, 1998     

冬小麦叶片持绿能力及其衰老特征研究

以12个冬小麦(Triticum aestivum L.)品种为供试材料,通过田间实验连续2年于开花后定期测定各品种的绿叶数目、绿叶面积、叶绿素和MDA含量以及SOD和CAT活性等指标,并以生理成熟时的保绿度、衰老启动时间为指标进行Hierarchical聚类分析,对小麦品种持绿能力进行分级.结果表明,参试冬小麦品种可分为持绿和非持绿两种类型,‘潍麦8号'(WM8)和‘豫麦66'(YM66)两年均表现为持绿型小麦.在整个灌浆期,持绿型小麦品种绿叶数目、面积、叶绿素含量明显高于非持绿型品种,叶片保护酶SOD与CAT活性也较非持绿小麦强,而其MDA含量明显低于非持绿型小麦品种.持绿型小麦叶片衰老启动时间延迟,生育后期绿叶面积较大,光合作用时间延长,具有较高的产量.本研究结果为冬小麦的品种选育、布局等相关研究奠定基础.     

The Effect of Light on the Senescence of Detached Cucumber Cotyledons

Measurements of cell constituents of cotyledons of cucumberexcised and kept in the light show differences from those excisedand kept in the dark. These differences are partly resolvedby adding glucose to the water-supply of the cotyledons in thedark suggesting that a lack of photosynthetic products is thecontrolling factor in their senescence. Under other circumstancesdifferent factors are operative suggesting that the first stepin senescence is different under different conditions.     

Hormonal Aspects of Senescence in Detached Tobacco Leaves

The objective of the present work was to describe the simultaneous changes in endogenous levels of cytokinins, abscisic acid, indoleacetic acid and ethylene in detached, senescing tobacco (Nicotiana rustica L.) leaves. These measurements were related to changes in chlorophyll contents, ¹⁴CO₂ fixation and proline contents — three parameters which have been considered to reflect senescence. Effects of exogenous hormonal treatments on these parameters, as well as on endogenous hormonal levels, provided further evidence for the interrelationships between hormones and for their roles in senescence. Starting with actively growing attached leaves and ending with well-advanced senescence in detached leaves, our data indicate a chronological sequence of three hormonal states: (a) cytokinins — high activity, abscisic acid, auxin and ethylene — low contents (actively growing, attached leaves); (b) cytokinins — low activity, abscisic acid — high, auxin and ethylene — low contents (apparent induction of senescence in detached leaves); and (c) cytokinins and abscisic acid — low, auxin and ethylene — high contents (senescence proper in detached leaves).     

The Role of Abscisic Acid in Senescence of Detached Tobacco Leaves

The role of abscisic acid in the regulation of senescence was investigated in detached tobacco leaves (Nicotiana rustica L.). Leaves senesced in darkness showed a sharp rise in abscisic acid level in the early stage of aging, followed by a rapid decline later. The same trend was found when leaves were aged in light, but the rise in abscisic acid occurred four days later than in darkness. Senescence was slower in light than in darkness, while salt stress accelerated the processes. Leaves treated with kinetin which senesced in light and darkness, did not show an increase in abscisic acid. Application of kinetin led to a transformation from free to bound ABA. These results may indicate that ABA and cytokinin are involved in a trigger mechanism which regulates senescence; the stage at which this trigger is activated determines the rate of senescence.     

小麦旗叶长、宽及面积的QTL分析

以小麦品种‘小偃81’和‘西农1376’构建的含236个家系的自交重组系(RIL)群体(F2:7、F2:8代)为研究材料,采用完全随机区组设计,连续2年在陕西杨陵、河南驻马店和山东济南于灌浆期(花后20d)随机取每个株系10株测量旗叶长、宽,并利用172个SSR标记构建了遗传连锁图谱,通过基于完备区间作图法的QTL IciMapping V3.2软件,对控制小麦旗叶长、宽和面积的数量性状位点(QTL)进行了加性效应分析。结果发现:(1)9个旗叶长QTLs位于1A、4A、3B、5D和7D染色体上,单个QTL可解释5.10%~16.44%的表型变异;10个旗叶宽QTLs位于1A、3A、5A、7A、3B和5D染色体上,单个QTL可解释4.63%~14.24%的表型变异;12个旗叶面积QTLs位于1A、4A、3B、2D和5D染色体上,单个QTL可解释4.25%~22.67%的表型变异。(2)控制小麦旗叶长、宽和面积的QTLs存在差异,同一QTL在不同性状中的遗传贡献率也不同。(3)同一性状在同一年份,不同地点和在不同年份,相同地点下检测到的QTLs有的相同,但有的差异明显。(4)有些控制不同性状的QTLs在染色体的同一标记区间,表现一因多效。研究表明:位于1A和5D染色体上的2个加性QTLs都同时控制旗叶长、宽和面积,且前者为主效基因,后者遗传贡献率也较大,可用于标记辅助育种和分子聚合育种。     

Source-Sink Relationship: Control of Photosynthesis in the Flag Leaf by the Panicle in Wheat

If the sink organs regulate the metabolic activity of the sourceorgans through the transmission of a signal and that signalis chemical in nature, then it should be present among the substancescoming out of the panicle. When flag leaf respiration and photosynthesisof wheat (Tritcum aestivum L.) cv. HP 1459 were measured withan infrared gas analyser after feeding the flag leaves withthe diffusate coming out of the panicle, it was observed thatwhile respiration was unaffected, photosynthesis was sharplyinhibited; the inhibition decreased with increasing age of thepanicle. Among the known plant growth regulators none couldsatisfy the requirements of this role with the exception ofabscisic acid which was found to inhibit photosynthesis by morethan 50% even at 10^–7 M, when respiration was affectedby only 8%. Apparently, abscisic acid inhibited photosynthesisthrough its effects on stomatal movement. (Received September 2, 1986; Accepted May 1, 1987)     

Relationship of Nitrogen Deficiency-Induced Leaf Senescence with ROS Generation and ABA Concentration in Rice Flag Leaves

Nitrogen (N) deficiency is one of the critical environmental factors that induce leaf senescence, and its occurrence may cause the shorten leaf photosynthetic period and markedly lowered grain yield. However, the physiological metabolism underlying N deficiency-induced leaf senescence and its relationship with the abscisic acid (ABA) concentration and reactive oxygen species (ROS) burst in leaf tissues are not well understood. In this paper, the effect of N supply on several senescence-related physiological parameters and its relation to the temporal patterns of ABA concentration and ROS accumulation during leaf senescence were investigated using the premature senescence of flag leaf mutant rice (psf) and its wild type under three N treatments. The results showed that N deficiency hastened the initiation and progression of leaf senescence, and this occurrence was closely associated with the upregulated expression of 9-cis-epoxycarotenoiddioxygenase genes (NCEDs) and with the downregulated expression of two ABA 8′-hydroxylase isoform genes (ABA8ox2 and ABA8ox3) under LN treatment. Contrarily, HN supply delayed the initiation and progression of leaf senescence, concurrently with the suppressed ABA biosynthesis and relatively lower level of ABA concentration in leaf tissues. Exogenous ABA incubation enhanced ROS generation and MDA accumulation in a dose-dependent manner, but it decreased the activities of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) in detached leaf. These results suggested that the participation of ABA in the regulation of ROS generation and N assimilating/remobilizing metabolism in rice leaves was strongly responsible for induction of leaf senescence by N deficiency.

     

Plant Growth Regulators and Induction of Leaf Senescence in Nitrogen-Deprived Wheat Plants

The sequence of events and the signals that regulate the remobilization of nitrogen (N) reserves during senescence induced by N starvation were studied in leaf 3, the last fully expanded leaf, in 17-day-old wheat (Triticum aestivum L.) plants. The first event observed was a rapid decrease in the isopentenyl adenosine (iPA) concentration during the first 24 h of N starvation. No differences in t-zeatin riboside and dihydrozeatin riboside concentrations were observed until the end of the assay. During the following 6 days, a decrease in soluble amino acids, chlorophyll, and protein, as well as an increase in soluble sugar concentration and endoproteolytic activity, could be observed. At day 3 of the experiment, the abscisic acid (ABA) concentration in the leaves of N-deprived plants started to increase. After 6 days of N deprivation there was a rise in oxidative stress, as indicated by the increase in malondialdehyde concentration, as well as a decrease in the activities of antioxidant enzymes catalase and ascorbate peroxidase. To analyze interactions with leaf development, the first, second, third, and fourth leaves were studied. iPA concentration decreased in all the leaf stages, including leaf 4, which was not fully expanded. A linear correlation between iPA and protein concentration was determined. These results suggest that the sharp fall in iPA could be the earliest event that induces protein degradation during the development of senescence induced by N deficiency, and that only later is ABA accumulated and oxidative stress developed.     

超级高产小麦旗叶荧光动力学研究

探明超级小麦品种的旗叶光合作用与荧光动力学特性,为超级小麦品种选育利用提供理论依据。以超级小麦临麦4号为试验材料,应用CI-301PS型便携式光合作用测定系统和FMS-2便携式荧光测定仪(英国Hansatech公司)在田间试验中测定旗叶光合作用与荧光动力学参数。结果表明,与普通高产对照品种皖麦52和烟农19相比,超级小麦临麦4号的光合作用参数光合速率、光饱和点和CO2饱和点、羧化效率高,光补偿点和CO2补偿点低;光合机构系统工作参数PSII实际的光化学效率(ΦPSII)、光化学猝灭系数(qP)、PSII反应中心的激发能捕获效率(Fv/Fm)、PSⅡ潜在活性Fv/Fo和电子传递速率(ETR)值高,非光化学猝灭系数(NPQ)值低。这表明超级小麦临麦4号的光合机构系统工作能力强和工作效率高,保证旗叶光合作用的高效运行,为子粒灌浆提供充足的能量和碳水化合物。     

Nitrogen and Photosynthesis in the Flag Leaf of Wheat (Triticum aestivum L.)

Wheat (Triticum aestivum L. cv Yecora 70) plants were grown with various concentrations of nitrate nitrogen available to the roots. Sampling of flag leaves began after they had reached full expansion and continued throughout senescence. Rates of gas exchange, ribulose-1,5-bisphosphate (RuP₂) carboxylase activity, and the amounts of chlorophyll, soluble protein, nitrogen, and phosphorus were determined for each flag leaf. Rate of CO₂ assimilation was uniquely related to total leaf nitrogen irrespective of nutrient treatment, season, and leaf age. Assimilation rate increased with leaf nitrogen, but the slope of the relationship declined markedly when leaf nitrogen exceeded 125 millimoles nitrogen per square meter. Chlorophyll content and RuP₂ carboxylase activity were approximately proportional to leaf nitrogen content. As leaves aged, RuP₂ carboxylase activity and calculated Hill activity declined in parallel. With normal ambient partial pressure of CO₂, the intercellular partial pressure of CO₂ was always such that rate of assimilation appeared colimited by RuP₂ carboxylation and RuP₂ regeneration capacity.

The initial slope of rate of CO₂ assimilation against intercellular partial pressure of CO₂ varied nonlinearly with carboxylase activity. It is suggested that this was due to a finite conductance to CO₂ diffusion in the wall and liquid phase which causes a drop in CO₂ partial pressure between the intercellular spaces and the site of carboxylation. A double reciprocal plot was used to obtain an estimate of the transfer conductance.

     

Characteristics of Chlorophyll Fluorescence and Membrane-Lipid Peroxidation During Senescence of Flag Leaf in Different Cultivars of Rice

With japonica rice 98-08, indica hybrids Shanyou 63, Gangyou 881, and X07S/Zihui 100, and sub-species hybrid Peiai 64S/9311 as materials, chlorophyll (Chl) content, Chl a fluorescence parameters, and membrane lipid peroxidation in flag leaf were measured at late developmental stages under natural conditions. F_v/F_m, q_P, _PS2, and electron transport rate gradually decreased while q_N increased conversely. Excessive photon energy led to the accumulation of active oxygen (O₂ ^–), H₂O, malonyldialdehyde, and products of membrane lipid peroxidation, and resulted in reduced Chl content and early ageing subsequent to the photooxidation during flag leaf senescence. There was obvious diversification of these parameters among rice cultivars. In comparison with japonica cv. 98-08 (tolerant to photooxidation), F_v/F_m decreased in indica cv. Shanyou 63 (susceptible to photooxidation) with greater accumulation of active oxygen and a sharp drop in Chl content, which resulted in yellowish early ageing, and affected the filling and setting of rice grains. The mechanism for premature ageing in indica rice was related to irradiance and temperature at filling stages. On a sunny day at above 25 °C, the reaction centre of photosystem 2 (PS2) exhibited a dynamic change on reversible inactivation. Under the intense irradiance at noon, PS2 function in indica rice exhibited obvious down-regulation and photoinhibition. Under intense irradiance with lowered temperatures, PS2 resulted in photo-damage and early ageing, related to the degradation of PS2-D1 protein and the inhibition of endogenous protection systems such as the xanthophyll cycle and enzymes scavenging active oxygen. Hence for high-yield breeding, based on a good plant-type and utilising heterosis and tolerance of photooxidation, the selection of japonica rice or a sterile line with the japonica genotype as female is a strategy worthy of consideration.     

水稻抽穗后剑叶衰老过程中光合关键酶的基因表达

协优 930 8、培矮 6 4s/ 9311和汕优 6 3三个超级高产水稻品种抽穗后 ,剑叶中光合速率减小、叶绿素含量下降、SOD活力降低和MDA含量上升的幅度不同 ,显示它们开花结实过程中叶片衰老的速度不同 ,以汕优 6 3衰老最快 ,协优 930 8最慢 ,培矮 6 4s/ 9311居中。Northern杂交和免疫定量研究表明 ,汕优 6 3和培矮6 4s/ 9311剑叶rbcS(Rubisco小亚基基因 )和rca(Ru bisco活化酶基因 )mRNA的表达随抽穗后天数的增加而下降 ,而协优 930 8在抽穗后第 10天才开始下降 ;Rubisco和Rubisco活化酶蛋白含量的下降趋势与rbcS和rcamRNA的变化趋势相似。因此 ,水稻抽穗后剑叶衰老过程中光合能力的下降与光合关键酶基因(rbcS和rca)表达下降密切相关     

小麦旗叶黄化转绿突变体的生理分析及细胞学研究

叶色突变体是研究植物光合作用机理的理想材料。该研究以小麦旗叶黄化转绿突变体LF2090及其野生型H_261为材料,对其主要农艺性状、光合色素含量、光合参数和叶绿体超微结构进行比较分析。结果显示:突变体在旗叶黄化期叶绿素b含量显著降低,叶绿体结构基本正常,光合速率无显著变化;在旗叶转绿期,突变体叶绿素b相对含量提高,但各色素含量均显著下降,叶绿体内基粒大部分消失,细胞间CO_2浓度及光合速率均显著下降。研究表明,叶绿素a与叶绿素b间的比例变化导致突变体旗叶叶片颜色发生由黄转绿的变化;突变体LF2090旗叶气孔部分关闭是该突变体光合速率降低和农艺性状较差的主要原因,同时色素含量降低导致叶绿体结构的改变也影响着旗叶的光合效率。     

小麦旗叶Rubisco周转与籽粒含氮量的关系

随着旗叶的衰老,Rubisco含量逐渐减少.延缓小麦旗叶的衰老进程(抽穗期施氮肥),可增加旗叶Rubisco的含量,提高籽粒的全氮含量.在小麦旗叶全展后28d内,Rubisco的15N丰度处于较高水平,表明仍有Rubisco的重新合成;而在28d以后,Rubisco的15N丰度处于低水平,表明无Rubisco的重新合成.但这时籽粒的15N丰度却上升.旗叶全展后14d内Rubisco的15N丰度高于旗叶中全氮的丰度,说明此时期Rubisco重新合成的速率高于其它蛋白质;旗叶衰老过程中Rubisco的15N丰度的净转移高于全氮,Rubi-sco净N转移也高于全氮,表明Rubisco向籽粒中转移的氮素多于其它蛋白质,对籽粒含氮量的影响最大.     

萝卜离体子叶衰老与膜脂过氧化的关系

萝卜离体子叶在光下或暗中衰老及激素调节衰老过程中,作为叶片衰老指标的叶绿素和蛋白质含量的降低,发生在MDA含量增高之前,更早于SOD活性的下降。表明由SOD活性降低所导致的膜脂过氧化的增强,并非衰老的原初反应,而是叶片衰老到一定程度的生理变化。因此,至少在萝卜离体子叶上,不能将其衰老的启动归因于受SOD控制的膜脂过氧化作用导致的膜累积性质变。