田间冬小麦叶片光合午休过程中的非气孔限制

1　引　　言作物的光合午休是近几年作物栽培生理研究的热点之一 ,研究大都从生态、生理的各个方面阐述光合午休的成因与机理 ,如土壤水分降低 ,ＣＯ2 浓度下降 ,ＶＰＤ过高 ,气孔限制与非气孔限制 ,羧化效率降低等[4 ,6,10 ] .其中有关作物光合午休过程中的非气孔限制问题 ,多数研究仅根据Ｆａｒｑｕｈａｒ判据[2 ] 判断发生了非气孔限制即止 ,至于非气孔限制的具体内容 ,多见于讨论 ,缺乏直接的实验证据 .本文对大田生态条件下 ,冬小麦生育后期旗叶光合午休过程中的非气孔限制问题进行了较深入的研究 .2　材料与方法2 1　供试材料选用冬…     

小麦拔节期淹水对叶片光合特性的影响

为研究拔节期涝害对小麦叶片光合特性的影响, 以小麦品种烟农19 为试验材料, 采用盆栽试验, 于小麦拔节 期进行淹水(涝害)处理, 分析小麦叶片光合气体交换参数的变化。结果表明, 拔节期淹水6 d 后小麦叶片SPAD 值较对照显著下降。在淹水3–9 d 内, 随着小麦叶片净光合速率逐渐下降, 叶片气孔导度和胞间CO2 浓度亦显著下降, 说明烟农19 光合速率下降主要是叶片气孔限制引起的。而在淹水后期(9 d 之后), 净光合速率持续下降, 而叶片胞间CO2浓度却升高, 说明在涝害发生后期叶片光合速率下降主要由非气孔因素引起。在小麦拔节期涝害前期叶片光合速率下降主要是由叶片气孔限制引起的, 而在涝害发生后期叶片光合速率下降主要由非气孔因素引起。     

不同供水条件下冬小麦叶与非叶绿色 器官光合日变化特征

为揭示小麦叶片与非叶绿色器官的光合活性在一日中的变化特性及其在器官间的差异性,探讨群体及不同器官光合日变化对不同供水条件的响应特征,在田间设置生育期不灌水(I0)、灌2水(I2,拔节水＋开花水)和灌4水(I4,起身水+孕穗水+开花水+灌浆水)3个处理,于灌浆期测定了群体光合与呼吸速率的日变化, 旗叶片、叶鞘、穗、穗下节间各器官光合速率、蒸腾速率、气孔导度及叶绿素荧光参数的日变化。结果表明,灌浆期小麦穗和穗下节间光合速率日变化呈单峰曲线,而旗叶叶片与旗叶鞘光合速率均呈双峰型,表现出不同程度的午休。随着灌水次数减少,各器官光合速率降低,叶片对严重水分亏缺的反应大于各非叶器官。器官光合速率的日变化与F_v/F_m变化相一致,而与气孔导度日变化有较大差异。各器官上午的累积光合量均高于下午,上午光合量占日总光合量的比例为51％-62％,随着灌水次数减少而增大。不同灌水处理群体光合速率、呼吸速率日变化均未出现午休现象。春季浇2水处理与春浇4水处理相比,灌浆期群体光合速率及日光合积累量没有显著差异。综合研究认为,小麦叶与非叶器官光合性能及其日变化特征有较大不同,非叶光合对水分亏缺的敏感性低于叶片,生育期浇2水可以获得与浇4水相似的群体日光合积累量。     

两种高产小麦旗叶光合功能衰退特性的比较

以高产小麦宁麦8号和9号为材料,研究了自开花期旗叶光合功能衰退和品种间的差异。结果表明:开花期后,随着穗重的增加,旗叶的比叶重、相对含水量、叶绿素含量、饱和光强下的净光合速率P_nmax、光饱和点LSP、表观量子产量AQY以及PSI、PS II和全电子传递链活性不断降低,黄熟期后下降均较显著,而叶绿素a/b、光补偿点LCP和呼吸速率R_d开花期至黄熟期间缓慢升高。和宁麦8号相比,宁麦9号叶绿素含量开始时较低,衰老后期含量较高。结论:黄熟期前,旗叶对光的利用范围较光,光合功能降低缓慢;黄熟期后,利用强光和弱光能力均下降,光合功能的衰退显著。和宁麦8号相比,高产小麦宁麦9号旗叶光合功能早衰。     

水肥运筹对小麦旗叶光合特性及产量的影响

研究了不同灌水次数及等氮量不同施氮方式对优质强筋小麦品种郑麦9023旗叶净光合速率(Pn)、叶绿素荧光参数等光合性能及产量的影响。结果表明,旗叶Pn、PSⅡ的潜在活性(Fv／Fo)、PSⅡ的光化学最大效率(Fv／Fm)、光化学猝灭系数(qP)、非光化学猝灭系数(qN)及产量在不同灌水次数间的差异均达到显著或极显著水平,其中不灌水处理与灌水处理间的差异较大;在不同施氮处理问,氮肥全部基施与返青期追施(占40％)处理间植株旗叶的Pn、Fv／Fo、Fv／Fm、qP和qN差异不大,但两处理的上述光合参数及产量均明显低于拔节期或孕穗期追施(40％)处理,表明拔节期或孕穗期追施氮肥有助于减缓旗叶光合功能的衰退,延长叶片功能期,提高植株的光合产物积累。研究结果还表明,水、氮运筹对Pn、叶绿素荧光参数及产量性状的调控存在显著的互作效应,其中以灌拔节水 孕穗水并于孕穗期追施氮肥(W2N4)处理组合表现最好。     

不同发育时期追施氮肥对冬小麦旗叶中光合酶活性的影响

小麦开花后,随着旗叶的衰老,旗叶中1,5-二磷酸核酮糖羧化酶（RuBPC）、磷酸烯醇式丙酮酸羧化酶（PEPC）和乙醇酸氧化酶（GO）活性呈下降趋势。随着追施氮肥时期的推迟,光合酶活性呈增加趋势,这意味着氮肥追施时间后移有利于提高小麦光合速率。在旗叶衰老后期,大穗型品种小麦旗叶中光合酶活性略高于多穗型品种小麦。     

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。     

不同品质类型小麦非叶光合器官光合荧光特性研究

以强筋(郑麦9023)、中筋(新麦13)和弱筋(豫麦50)型小麦品种为试材,测定了它们非叶光合器官的叶绿素含量、荧光动力学参数、净光合速率及对籽粒的贡献率.结果显示,不同品质类型小麦非叶光合器官叶绿素含量表现为旗叶鞘>穗下节间>芒>穗部颖片,各非叶光合器官开花18d后叶绿素含量均表现为弱筋小麦>中筋小麦>强筋小麦;弱筋小麦的非叶光合器官在灌浆前期具有较高的Fv/Fm、ΦPSⅡ值,在灌浆后期其Fv/Fm和ΦPSⅡ值下降幅度显著小于强筋小麦并有较高的净光合速率;非叶光合器官对籽粒的相对贡献率依次为穗部>旗叶>节鞘,且对弱筋小麦籽粒的贡献率高于中筋和强筋小麦,其中节鞘和穗对弱筋小麦粒重的相对贡献率分别为15.7%和35.4%.研究表明,弱筋小麦豫麦50的非叶光合器官能维持较较强的光合能力,生产更多的光合产物,花后形成的光合产物对其粒重的增加起重要作用.     

小麦灌水时期与灌水量对花后果聚糖积累与转运及水分利用效率的影响

于2004-2005年和2005-2006年冬小麦生长季,在山东泰安和兖州进行田间试验,研究不同灌水时期和灌水量处理对冬小麦开花后倒二节间果聚糖积累与转运和水分利用效率的影响.结果表明：全生育期不灌水促进了灌浆后期倒二节间贮藏果聚糖向籽粒的转运.在拔节期和开花期各灌水60 mm,可提高开花后旗叶的光合速率和同化物输入籽粒量及其对籽粒的贡献率,拔节期、开花期和灌浆期各灌水60 mm或拔节期和开花期各灌水90 mm,灌浆后期旗叶的光合速率显著降低,营养器官花前贮藏同化物转运量及其对籽粒的贡献率升高,花后同化物输入籽粒量及其对籽粒的贡献率降低,灌浆后期倒二节间的聚合度（DP）≥4、DP=3果聚糖滞留量增加,不利于果聚糖向籽粒的转运.两个生长季中,拔节期和开花期各灌水60mm处理的小麦籽粒产量较高,水分利用效率最高.拔节期、开花期和灌浆期各灌水60 mm或拔节期和开花期各灌水90 mm,小麦籽粒产量无显著变化,水分利用效率降低.     

追施氮肥时期对冬小麦旗叶叶绿素荧光特性的影响

在大田条件下,研究了不同追氮时期对小麦旗叶叶绿素荧光特性、光合速率及籽粒产量的影响.结果表明,拔节期追肥较起身期或挑旗期追肥,改善了小麦旗叶PSⅡ的活性(F_v/F_o)、光化学最大效率(F_v/F_m)、光化学猝灭系数(qP)、实际量子产量(ΦPSⅡ)及光合速率,降低了籽粒灌浆中前期非辐射能量耗散,有利于叶片所吸收的光能较充分地用于光合作用,提高了籽粒灌浆后期非辐射能量的耗散,减缓了叶片光抑制程度和衰老进程.拔节期追肥可显著增加穗粒数和千粒重,提高产量.     

The temporal and species dynamics of photosynthetic acclimation in flag leaves of rice (Oryza sativa) and wheat (Triticum aestivum) under elevated carbon dioxide

In this study, we tested for the temporal occurrence of photosynthetic acclimation to elevated [CO₂] in the flag leaf of two important cereal crops, rice and wheat. In order to characterize the temporal onset of acclimation and the basis for any observed decline in photosynthetic rate, we characterized net photosynthesis, g_s, g_m, C_i/C_a, C_i/C_c, V_cmax, J_max, cell wall thickness, content of Rubisco, cytochrome (Cyt) f, N, chlorophyll and carbohydrate, mRNA expression for rbcL and petA, activity for Rubisco, sucrose phosphate synthase (SPS) and sucrose synthase (SS) at full flag expansion, mid‐anthesis and the late grain‐filling stage. No acclimation was observed for either crop at full flag leaf expansion. However, at the mid‐anthesis stage, photosynthetic acclimation in rice was associated with RuBP carboxylation and regeneration limitations, while wheat only had the carboxylation limitation. By grain maturation, the decline of Rubisco content and activity had contributed to RuBP carboxylation limitation of photosynthesis in both crops at elevated [CO₂]; however, the sharp decrease of Rubisco enzyme activity played a more important role in wheat. Although an increase in non‐structural carbohydrates did occur during these later stages, it was not consistently associated with changes in SPS and SS or photosynthetic acclimation. Rather, over time elevated [CO₂] appeared to enhance the rate of N degradation and senescence so that by late‐grain fill, photosynthetic acclimation to elevated [CO₂] in the flag leaf of either species was complete. These data suggest that the basis for photosynthetic acclimation with elevated [CO₂] may be more closely associated with enhanced rates of senescence, and, as a consequence, may be temporally dynamic, with significant species variation.     

不同有机肥量对旱地玉米光合特性和产量的影响

在渭北旱塬合阳试验基地进行了4年的旱地有机培肥试验, 研究不同有机肥施用量对旱地玉米光合特性和产量的影响.结果表明： 与单施化肥相比,施有机肥处理的玉米各生育时期的叶片光合速率和气孔导度显著增大,胞间CO2浓度显著减小;随着有机肥施用量的增加,玉米叶片各生育时期的叶片光合速率和气孔导度逐渐增大,胞间CO2浓度逐渐减小.玉米各生育时期的光合作用主要受非气孔因素限制,施用有机肥显著降低了非气孔因素对光合的限制.连续4年施用有机肥,改善了土壤养分状况,使养分不再是玉米光合速率和产量的主要限制因子.     

Severe water deficit-induced ethylene production decreases photosynthesis and photochemical efficiency in flag leaves of wheat

Wheat (Triticum aestivum L.) cv. Jimai22 was used to evaluate the effect of ethylene evolution rate (EER) and 1-aminocyclopropane-1-carboxylic acid (ACC) and their relations with photosynthesis and photochemical efficiency in plants well-watered (WW) and under a severe water deficit (SWD). SWD caused a noticeable reduction in the grain mass. The marked increases in both EER and the ACC concentration were observed under SWD; it was reversed effectively by exogenous spermidine (Spd) or amino-ethoxyvinylglycine (AVG). Thermal images indicated that SWD increased obviously the temperature of flag leaves, mainly due to the decrease in transpiration rate under SWD. Exogenous Spd or AVG decreased to some extent the temperature of the flag leaves. The strong decline in photosynthetic rate (P _N) and stomatal conductance as well as the photodamage of PSII were also observed under SWD after 14 and 21 days after anthesis (DAA). Intercellular CO₂ concentration was reduced at 7 DAA, but slightly increased at 14 and 21 DAA under SWD, indicating that the decreased P _N at 7 DAA might result from stomatal limitations, while the decline after 14 and 21 DAA might be attributed to nonstomatal limitations. Correlation analysis suggested that EER and ACC showed negative relations to photosynthesis and photochemical efficiency. Data obtained suggested that the effects of SWD were mediated predominantly by the increase in EER and ACC concentration, which greatly decreased the leaf photosynthesis and photochemical efficiency, and, therefore, reduced the grain mass. Application of Spd or AVG reduced the EER and ACC, and thus positively influenced photosynthesis and photochemical efficiency under SWD.     

小麦根系活力变化与叶片衰老的研究

对小麦根系活力、叶片叶绿素含量和光合速率变化的田问实验结果表明,根系活力的衰退早且快于叶片叶绿素含量和光合速率的衰退;根系活力和叶片光合速率呈正相关关系(r=0．8006)．拔节期追施氮肥,改善根际营养条件后,叶片的光合速率、根系活力明显提高,叶片中ABA含量降低,ZRs升高,SOD活性增加,MDA的高峰期比对照推迟8～10d,说明根系活力变化与地上部叶片的衰老密切相关,改善根际营养,提高生育前期的根系活力有助于延缓地上部分的衰老．     

水分胁迫下烯效唑对百日草幼苗光合特性及叶解剖结构的影响

以百日草‘芳菲1号’为试材,研究不同水分胁迫下烯效唑(S3307)对其幼苗生长、光合特性及叶解剖结构的影响,以明确S3307对百日草的抗旱作用及其机理。结果显示:(1)在水分胁迫下,百日草的生长均受到不同程度的抑制,叶绿素含量显著降低,光合作用受到抑制,叶解剖结构有所变化。(2)S3307处理后,均能够显著降低所对应的不同程度水分胁迫下百日草的株高,显著增加茎粗、叶面积、叶片厚度、栅栏组织厚度和根冠比,显著增加叶绿素含量,提高百日草的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和水分利用效率(WUE)。研究表明,S3307能够提高百日草的抗旱性,而且在轻度和中度水分胁迫下Pn的下降主要是由气孔因素引起,而在重度水分胁迫下光合速率的下降是由非气孔因素引起的。     

Cultivar differences in leaf photosynthesis of rice bred in Japan

The grain yield of rice (Oryza sativa L.), as well as of other cereal crops, is limited to a large extent, by the supply of photosynthates produced during grain filling period. In this study, flag leaf photosynthesis (LPS) after heading was compared among 32 cultivars bred during the past century in Japan, to determine if the improvement of LPS has occurred with the breeding advance of high yielding cultivars. Measurement of LPS was made for 5 consecutive years in the paddy field, on the flag leaf of the main stem, at heading (LPS-0), and 2 weeks (LPS-2) and 4 weeks (LPS-4) after heading. LPS decreased with advance of leaf senescence from LPS-0 to LPS-2, and then to LPS-4. However, if nitrogen was top-dressed at the heading time, high LPS-2 was maintained, particularly in the newer cultivars. A significant positive correlation between LPS and the released year of cultivar was found at LPS-2, especially in the nitrogen top-dressed plot, but not at LPS-0 or LPS-4. Cultivar difference in LPS of the senescing leaves were not stable through the different years, whereas LPS-0 was stable over years, suggesting that the LPS in the senescent leaf is susceptible to the environmental variation due to the effects on leaf senescence. Cultivar difference in LPS at any stage was closely associated with mesophyll conductance to CO₂, and stomatal conductance was also associated with cultivar difference in such a high LPS as LPS-0 and nitrogen top-dressed LPS-2. Significant correlation between LPS and specific leaf weight was not observed at any stage of the flag leaf.Abbreviations CV coefficient of variation - g_m mesophyll conductance - g_s stomatal conductance - LPS apparent photosynthetic rate per unit leaf area (leaf photosynthesis) - LPS-0 LPS at heading - LPS-2 LPS at active grain filling - LPS-4 LPS at maturity of grain - NT non-top dressed plot - PPFD photosynthetic photon flux density - r_m mesophyll resistance - r_s stomatal diffusion resistance against CO₂ - r_s(H₂O) stomatal diffusion resistance against H₂O - RuBisCO ribulose-1,5-bisphosphate carboxylase/oxygenase - SLW specific leaf weight - TD nitrogen top-dressed plot     

Phosphoglycerate dehydrogenase from soybean nodules : partial purification and some kinetic properties

The relationship between single leaf photosynthesis and conductance was examined in cotton (Gossypium hirsutum L.) across a range of environmental conditions. The purpose of this research was to separate and define the degree of stomatal and nonstomatal limitations in the photosynthetic process of field-grown cotton.

Photosynthetic rates were related to leaf conductance of upper canopy leaves in a curvilinear manner. Increases in leaf conductance of CO₂ in excess of 0.3 to 0.4 mole per square meter per second did not result in significant increases in gross or net photosynthetic rates. No tight coupling between environmental influences on photosynthetic rates and those affecting conductance levels was evident, since photosynthesis per unit leaf conductance did not remain constant. Slowly developing water stress caused greater reductions in photosynthesis than in leaf conductance, indicating nonstomatal limitations of photosynthesis.

Increases in external CO₂ concentration to levels above ambient did not produce proportional increases in photosynthesis even though substomatal or intercellular CO₂ concentration increased. The lack of a linear increase in photosynthetic rate in response to increases in leaf conductance and in response to increases in external CO₂ concentration demonstrated that nonstomatal factors are major photosynthetic rate determinants of cotton under field conditions.

     

增强UV-B辐射和NaCl复合胁迫下绿豆光合作用的气孔和非气孔限制

研究了0.35 W/m2的UV-B辐射、0.4%NACl及其复合胁迫下绿豆(Phaseolus radiatus L.)幼苗光合作用的气孔和非气孔限制.发现各胁迫处理下,幼苗净光合速率、气孔导度、光合能力、羧化效率和Rubisco含量均明显降低;细胞间隙CO2浓度在各胁迫处理前期低于对照,后期高于对照;气孔限制值除复合处理第5天外,其余均高于对照;复合处理下上述指标的变化程度均大于两胁迫因子单独处理.表明各胁迫下光合速率的降低既有气孔因素也有非气孔因素,但前期以气孔限制为主,后期以非气孔限制为主;Rubisco含量的降低是各胁迫下光合速率降低的非气孔因素.