花后高温和水分逆境对小麦籽粒品质形成的生理影响
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摘要
温度和水分是影响小麦籽粒品质形成的重要生态因子。明确花后高温和水分逆境对小麦籽粒品质形成的生理影响对于深化小麦品质生理生态和指导专用小麦抗逆栽培具有重要的理论和实践意义。本研究以蛋白质含量差异显著的小麦品种扬麦9、徐州26和豫麦34为材料，利用人工气候室设置不同温度水平和不同昼夜温差处理，以及在高温(32℃／24℃)和适温(24℃／16℃)条件下分别设置不同水分处理(对照、干旱、渍水)，综合研究了不同温度和温差条件下以及温度与水分互作条件下，不同类型小麦籽粒品质形成、光合特性、物质运转和相关酶活性的变化规律，明确了花后高温与水分逆境对小麦籽粒品质形成的生理调控作用，为深化和拓展小麦品质生理生态，指导专用小麦调优栽培提供了理论基础和技术途径。
获得的主要研究结果如下：
1．高温显著提高了小麦籽粒蛋白质含量及清蛋白、球蛋白和醇溶蛋白含量，但降低了谷蛋白含量，导致麦谷蛋白／醇溶蛋白比值降低。高温显著降低了籽粒总淀粉和支链淀粉含量及支／直比。籽粒蛋白质和淀粉及其组分形成所需的适宜昼夜温差随小麦品质类型而异，但温度水平对籽粒蛋白质和淀粉的影响较温差大。在高温和水分逆境下，温度对籽粒蛋白质和淀粉含量的影响较水分逆境大，且存在显著的互作效应。小麦籽粒蛋白质含量均表现为干旱＞对照＞渍水，以高温干旱最高，适温渍水最低；淀粉含量为对照＞干旱＞渍水，以适温对照最高，而高温渍水最低。高温和水分逆境显著提高了籽粒醇溶蛋白含量而降低了谷蛋白含量及支链淀粉含量，使蛋白质谷／醇比和淀粉支／直比降低，以高温渍水对籽粒蛋白质和淀粉组分的影响最为显著。不同品种之间，高蛋白小麦籽粒蛋白质和组分的形成受高温和水分逆境的影响更大，而低蛋白品种籽粒淀粉形成显著受温度和水分逆境的调节。高温和水分逆境对面筋特性和沉降值的影响因品种的不同而有所差异。
2．高温提高了营养器官游离氨基酸含量，但降低了可溶性总糖含量，从而降低了糖／氮比。高温降低了营养器官花前贮藏同化物和氮素的转运量和转运率，花后同化物和氮素输入籽粒量。氨基酸含量随温差的变化因品种和温度水平不同。高温下温差对各器官可溶性总糖含量及糖／氮比影响较小；适温下温差大的处理各器官可溶性总糖含量较高，从而糖／氮比提高。干旱提高了营养器官氨基酸含量，但明显降低了营养器官和籽粒中可溶性总糖含量和糖／氮比。渍水降低了各器官游离氨基酸含量和可溶性总糖含量，但对糖／氮比的影响因品种和器官而不同。小麦营养器官花前贮藏干物质和氮素运转量和运转率在适温下表现为干旱＞对照＞渍水，高温下则表现为对照＞干旱＞渍水。适温下花后同化物积累量表现为对照＞渍水＞干旱，高温下则表现为对照＞干旱＞渍水。花后氮素积累量在适温和高温下均表现为对照＞渍水＞干旱。花前贮藏氮素再运转量和花后氮素同化量与籽粒蛋白质产量呈正相关关系。而花前贮藏干物质再运转量和花后物质同化量与籽粒重和籽粒淀粉含量均达显著相关和极显著相关。
3．水分适宜条件下，高温提高了灌浆初期扬麦9籽粒蔗糖合成酶(SS)和束缚态淀粉合成酶(GBSS)的活性，但明显降低了灌浆后期SS、GBSS和可溶性淀粉合成酶(SSS)活性。而豫麦34三种酶活性在高温下均呈下降趋势。灌浆初期高温处理显著降低了徐州26籽粒SSS活性，说明高温抑制不同类型小麦籽粒淀粉合成的酶学机制不同。此外，在不同昼夜温差处理(8℃和12℃)间，灌浆中后期，高温下SS活性随温差增大而升高，SSS和GBSS则以温差小的处理较高；适温下SS、GBSS和SSS 3种淀粉合成关键酶活性均以温差大的处理为高。干旱和渍水处理均明显降低了淀粉合成相关酶活性。高温、干旱和渍水均明显降低了旗叶谷胺酰氨合成酶(GS)和谷丙转氨酶(GPT)活性。与对照相比，高温提高了花后14天徐州26旗叶GS活性，但降低了两品种籽粒GPT活性和扬麦9旗叶GS活性。GS和GPT活性随温差的变化在不同品种间表现不同。不同温度条件下，干旱和渍水对蛋白质合成关键酶活性的影响因温度和品种的不同而异。相关分析结果表明，高温和水分逆境下SS、SSS和GBSS活性与籽粒淀粉合成关系密切，旗叶GS和籽粒GPT活性与蛋白质产量呈正相关，而与籽粒蛋白质含量呈负相关关系。
4．高温、干旱和渍水明显降低了旗叶光合速率和SPAD值。高温下，干旱和渍水对光合作用的影响加重。高温、干旱和渍水条件下，旗叶MDA含量增加，而可溶性蛋白含量降低。水分适宜条件下，与适温处理相比，高温下旗叶SOD活性先上升后迅速下降。而高温干旱和高温渍水条件下，SOD活性呈一直下降的趋势。表明随高温处理时间的延长，叶片膜脂过氧化程度加剧，细胞质膜相对透性增大，叶片衰老加速，而干旱和渍水对叶片造成了不可逆的伤害。高温下，干旱和渍水对植株衰老的影响有加重的趋势。不同温差处理间比较，高温下温差升高旗叶所受伤害加重，衰老加速。适温时，温差加大有利于延缓植株衰老。
综上所述，花后高温和水分逆境明显降低了小麦旗叶光合速率，减少了光合产物的积累，从而明显降低了花后同化物输入籽粒量；而高温和渍水处理则明显降低了花前贮藏氮素再运转量和花后氮素输入籽粒量，从而导致不同温度和水分处理下籽粒蛋白质、淀粉产量和含量的差异。调控籽粒中淀粉合成的旗叶中SPS及籽粒中SS、SSS和GBSS，调控蛋白质合成的旗叶中的GS及籽粒中的GPT等酶活性在不同温度和水分条件下的变化是影响籽粒中蛋白质和淀粉合成差异的生理基础。因此，高温和水分逆境通过对小麦物质积累和运转以及淀粉和蛋白质合成关键酶活性的互作影响而造成了籽粒品质性状在不同温度和水分条件下的差异。
Temperature and water are main ecological factors affecting formation of wheat quality. Elucidating the physiological mechanism for the effects of high temperature and water stress on grain quality formation in wheat is of great importance for understanding grain quality physiology and guiding cultural management in wheat. Two wheat cultivars differing in protein content including Yangrnai 9 and Xuzhou 26 were grown under varied temperature conditions during grain filling including 32~C/24~C, 28~C/20~C and 24~C/16 ~C in one experiment and 34~C/22~C, 32~C/24~C, 26~C/14~C and 24~C/16~C in the other experiment. Two wheat cultivars including Yangmai 9 with low protein content and Yumai 34 with high protein content were subject to temperature and water stress conditions in the third experiment. Two day/night temperature regimes of 32~C/24~C and 24~C/16~C with three soil water regimes were established as moderate water status (soil relative water content, SRWC=75%-80%), drought (SRWC=45%-50%) and waterlogging. The quality formation, photosynthetic characters, C/N assimilates translocation and regulatory enzyme activities in relation to formation of grain quality were investigated comprehensively, and the physiological regulatory mechanisms of high temperature and water stress on wheat quality formation was elucidated. The main results were as follows:
Contents of crude protein, albumin, globulin and gliadin in grains were significantly enhanced, but glutenin content was reduced by high temperatures, which resulted in decrease of the ratio of glutenin to gliadin. Moreover, contents of total starch, amylose and amylopectin were reduced, but more seriously effect of high temperature on amylopectin than amylose resulted in reducing amylopectin/amylose ratio. In addition, optimum diurnal temperature difference favoring the formations of grain protein and starch differed in wheat genotypes, but the effects of temperature level on protein and starch content were much higher than temperature differences. The second experiment showed that the effect of high temperature on grain protein and starch content were more marked than that of water stress, and the interaction effect of temperature X water also existed. Under high temperature or optimum temperature, grain protein content exhibited the pattern of drought>CK>waterlogging, the highest under drought with high temperature and the lowest under waterlogging with optimum temperature. Grain starch content showed the order CK>drought> waterlogging, the highest under CK with optimum temperature and the lowest under waterlogging with high temperature Furthermore, the content of grain gliadin was enhanced under high temperature and water stresses, while the contents of glutenin and amylopectin were reduced, which resulted in decreased tha ratios of glutenin to gliadin and amylopectin to amylose. Especially, the effect of high temperature with waterlogging was the most significant on protein and starch components. In addition, the response of Yumai 34 in grain protein formation to high temperature and water stresses were more serious than in starch, but for Yangrnai 9, starch formation was more sensitive to high temperature and water stresses. Correlation analysis revealed that protein content was positively correlated to albumin and gliadin contents, but slightly negatively to glutenin content. Starch content was positively to the contents of glutenin and amylopectin and amylopectin/amylose ratio under high temperature and water stresses. High temperature and water stress had different effects on grain gluten characters and sedimentation value in different varieties.
High temperature enhanced free amino acid content in vegetative organs, but significantly reduced soluble sugar content in vegetative organs and kernels during whole grain filling so that ratio of sugar/amino decreased. High temperature reduced remobilization amount and rate of pre-anthesis assimilates and nitrogen stored in vegetative organs and transferring into grain. Effect of diurnal temperature difference on amino acid content differed between cultivars. Under high temperature, temperature difference affected soluble sugar content slightly. Uunder optimum temperature, soluble sugar content under greater diurnal temperature difference was higher than that under lower temperature difference. Under high temperature, day/night temperature difference affected ratio of C/N slightly. Under optimum temperature ratio of C/N was higher under the day/night temperature difference of 12~C. Drought enhanced free amino acid content in vegetative organs, but reduced soluble sugar content in vegetative organs and kernels. As a result, ratio of C/N decreased under high temperature and drought. Under high and optimum temperature, waterlogging reduced contents of amino acid and soluble sugar vegetative organs and kernels. Effect of waterlogging on ratio of C/N differed among different cultivars and organs in wheat. Amount and ratio of assimilates and nitrogen translocation declined as drought>control>waterlogging under optimum temperature, while declined as control>drought>waterlogging under high temperature. The amounts of post-anthesis assimilate transferring into grain were declined as CK>waterlogging>drought under optimum temperature,while the order was CK>drought>waterlogging under high temperature.The amounts of post-anthesis accumulated nitrogen declined as CK>waterlogging>drought under both high and optimum temperature.The amounts of pre-anthesis stored and post-anthesis accumulated nitrogen transferring into grain were significantly related to protein yield.The amounts of pre-anthesis stored and post-anthesis accumulated assimilates transferring into grain were significantly related to grain weight and starch content.
Under optimum water status,high temperature enhanced the activities of sucrose synthase (SS) and granule-bound starch synthase (GBSS) at early stage of grain filling, while reduced activities of SS,GBSS and soluble starch synthase (SSS) at late grain filling stage.Activities of three enzymes were reduced by high temperature in Yumai 34.High temperature significantly reduced activity of SSS in Xuzhou 26 at early stage of grain filling.Those results indicated that the enzymatic mechanism of starch formation differed among wheat varieties.In addition,under high temperatures,SS activity was higher at 34℃/22℃during the middle and late stage of grain filling,whereas the activities of SSS and GBSS were higher under 32℃/24℃.Under optimum temperature,activities of three enzymes were higher at bigger day/night temperature difference.At different day-night temperature differences, GPT (glutamate pyruvic aminotransferase) and glutamine synthase (GS) activities changed between cultivars.Effects of drought and waterlogging on activities of three enzymes differed between different cultivars.Drought and waterlogging reduced activities of three enzymes under both high and optimum temperature.Compared with optimum temperature,high temperature reduced GPT activity in kernels for two wheat cultivars.But reduced GS activity in flag leaves of Yangmai 9 and enhanced GS activity of Xuzhou 26 on 14 DAA.At different day-night temperature differences,GPT and GS activities changed between cultivars.High temperature,drought and waterlogging reduced GS activity and GPT activities in flag leaves.The effects of drought and waterlogging on activities of enzymes involved in protein formation were different among different cultivars and temperatures.Results of correlation analyses indicated that the SPS,SSS and GBSS activities were closely correlated to the yield of starch. GS activity in grains was correlated to a lower degree to the yield and content of protein than GS in flag leaves and GPT ingrains.Under high temperature,the effects of high temperature and water stresses on grain quality and protein and starch accumulation aggravated.
High temperature,drought and watedogging reduced photosynthetic rate and chlorophyll content in flag leaves,and the consequences of drought and waterlogging were more severe under high temperature than under optimum temperature.High temperature, drought and waterlogging enhanced the content of malondiadehyde (MDA),while reduced soluble protein content in flag leaves.Under optimum water treatment,compared with optimal temperature treatment,during the early stage of high temperature treatment,activity of SOD was enhanced but dropped rapidly during late grain filling.Under optimum and high temperature, drought and waterlogging reduced SOD activity on both 15DAA and 20DAA.These results indicated that the membrane lipid peroxidation was accelerated and the permeability of plasma membrane increased as the high temperature treatment prolonged and accelerated leaf senescence.The effects of drought and waterlogging on flag leave were not reversed.The consequences of drought and waterlogging were more severe under high temperature than under optimum temperature.Higher temperature difference under high temperature condition increased senescence of flag leaf,while under optimal temperature condition,higher temperature difference delayed the plant senescence.
In summary,high temperature and water stress reduced the photosynthesis rate in flag leaves of wheat.The amount of post-anthesis assimilates transferred into grains decreased distinctly under high temperature and water stress.High temperature and waterlogging significantly reduced the pre-anthesis stored nitrogen and the translocation of post-anthesis accumulated nitrogen into grains.Thus,the differential substrates caused the differences in protein and starch contents of wheat grains under different water and temperature conditions.The further studies indicated that the activities of key enzymes for grain starch and protein formation,including SPS and GS in leaf,SSS,GBSS and GPT in grains,are the vital enzymatic reasons for affecting the yields and contents of grain starch and protein by high temperature and water stress.The differences between grain qualities under different treatment conditions were caused by the interaction of high temperature and water stress on C/N translocation and activities of key enzymes involved in starch and protein accumulation in wheat
获得的主要研究结果如下：
1．高温显著提高了小麦籽粒蛋白质含量及清蛋白、球蛋白和醇溶蛋白含量，但降低了谷蛋白含量，导致麦谷蛋白／醇溶蛋白比值降低。高温显著降低了籽粒总淀粉和支链淀粉含量及支／直比。籽粒蛋白质和淀粉及其组分形成所需的适宜昼夜温差随小麦品质类型而异，但温度水平对籽粒蛋白质和淀粉的影响较温差大。在高温和水分逆境下，温度对籽粒蛋白质和淀粉含量的影响较水分逆境大，且存在显著的互作效应。小麦籽粒蛋白质含量均表现为干旱＞对照＞渍水，以高温干旱最高，适温渍水最低；淀粉含量为对照＞干旱＞渍水，以适温对照最高，而高温渍水最低。高温和水分逆境显著提高了籽粒醇溶蛋白含量而降低了谷蛋白含量及支链淀粉含量，使蛋白质谷／醇比和淀粉支／直比降低，以高温渍水对籽粒蛋白质和淀粉组分的影响最为显著。不同品种之间，高蛋白小麦籽粒蛋白质和组分的形成受高温和水分逆境的影响更大，而低蛋白品种籽粒淀粉形成显著受温度和水分逆境的调节。高温和水分逆境对面筋特性和沉降值的影响因品种的不同而有所差异。
2．高温提高了营养器官游离氨基酸含量，但降低了可溶性总糖含量，从而降低了糖／氮比。高温降低了营养器官花前贮藏同化物和氮素的转运量和转运率，花后同化物和氮素输入籽粒量。氨基酸含量随温差的变化因品种和温度水平不同。高温下温差对各器官可溶性总糖含量及糖／氮比影响较小；适温下温差大的处理各器官可溶性总糖含量较高，从而糖／氮比提高。干旱提高了营养器官氨基酸含量，但明显降低了营养器官和籽粒中可溶性总糖含量和糖／氮比。渍水降低了各器官游离氨基酸含量和可溶性总糖含量，但对糖／氮比的影响因品种和器官而不同。小麦营养器官花前贮藏干物质和氮素运转量和运转率在适温下表现为干旱＞对照＞渍水，高温下则表现为对照＞干旱＞渍水。适温下花后同化物积累量表现为对照＞渍水＞干旱，高温下则表现为对照＞干旱＞渍水。花后氮素积累量在适温和高温下均表现为对照＞渍水＞干旱。花前贮藏氮素再运转量和花后氮素同化量与籽粒蛋白质产量呈正相关关系。而花前贮藏干物质再运转量和花后物质同化量与籽粒重和籽粒淀粉含量均达显著相关和极显著相关。
3．水分适宜条件下，高温提高了灌浆初期扬麦9籽粒蔗糖合成酶(SS)和束缚态淀粉合成酶(GBSS)的活性，但明显降低了灌浆后期SS、GBSS和可溶性淀粉合成酶(SSS)活性。而豫麦34三种酶活性在高温下均呈下降趋势。灌浆初期高温处理显著降低了徐州26籽粒SSS活性，说明高温抑制不同类型小麦籽粒淀粉合成的酶学机制不同。此外，在不同昼夜温差处理(8℃和12℃)间，灌浆中后期，高温下SS活性随温差增大而升高，SSS和GBSS则以温差小的处理较高；适温下SS、GBSS和SSS 3种淀粉合成关键酶活性均以温差大的处理为高。干旱和渍水处理均明显降低了淀粉合成相关酶活性。高温、干旱和渍水均明显降低了旗叶谷胺酰氨合成酶(GS)和谷丙转氨酶(GPT)活性。与对照相比，高温提高了花后14天徐州26旗叶GS活性，但降低了两品种籽粒GPT活性和扬麦9旗叶GS活性。GS和GPT活性随温差的变化在不同品种间表现不同。不同温度条件下，干旱和渍水对蛋白质合成关键酶活性的影响因温度和品种的不同而异。相关分析结果表明，高温和水分逆境下SS、SSS和GBSS活性与籽粒淀粉合成关系密切，旗叶GS和籽粒GPT活性与蛋白质产量呈正相关，而与籽粒蛋白质含量呈负相关关系。
4．高温、干旱和渍水明显降低了旗叶光合速率和SPAD值。高温下，干旱和渍水对光合作用的影响加重。高温、干旱和渍水条件下，旗叶MDA含量增加，而可溶性蛋白含量降低。水分适宜条件下，与适温处理相比，高温下旗叶SOD活性先上升后迅速下降。而高温干旱和高温渍水条件下，SOD活性呈一直下降的趋势。表明随高温处理时间的延长，叶片膜脂过氧化程度加剧，细胞质膜相对透性增大，叶片衰老加速，而干旱和渍水对叶片造成了不可逆的伤害。高温下，干旱和渍水对植株衰老的影响有加重的趋势。不同温差处理间比较，高温下温差升高旗叶所受伤害加重，衰老加速。适温时，温差加大有利于延缓植株衰老。
综上所述，花后高温和水分逆境明显降低了小麦旗叶光合速率，减少了光合产物的积累，从而明显降低了花后同化物输入籽粒量；而高温和渍水处理则明显降低了花前贮藏氮素再运转量和花后氮素输入籽粒量，从而导致不同温度和水分处理下籽粒蛋白质、淀粉产量和含量的差异。调控籽粒中淀粉合成的旗叶中SPS及籽粒中SS、SSS和GBSS，调控蛋白质合成的旗叶中的GS及籽粒中的GPT等酶活性在不同温度和水分条件下的变化是影响籽粒中蛋白质和淀粉合成差异的生理基础。因此，高温和水分逆境通过对小麦物质积累和运转以及淀粉和蛋白质合成关键酶活性的互作影响而造成了籽粒品质性状在不同温度和水分条件下的差异。
Temperature and water are main ecological factors affecting formation of wheat quality. Elucidating the physiological mechanism for the effects of high temperature and water stress on grain quality formation in wheat is of great importance for understanding grain quality physiology and guiding cultural management in wheat. Two wheat cultivars differing in protein content including Yangrnai 9 and Xuzhou 26 were grown under varied temperature conditions during grain filling including 32~C/24~C, 28~C/20~C and 24~C/16 ~C in one experiment and 34~C/22~C, 32~C/24~C, 26~C/14~C and 24~C/16~C in the other experiment. Two wheat cultivars including Yangmai 9 with low protein content and Yumai 34 with high protein content were subject to temperature and water stress conditions in the third experiment. Two day/night temperature regimes of 32~C/24~C and 24~C/16~C with three soil water regimes were established as moderate water status (soil relative water content, SRWC=75%-80%), drought (SRWC=45%-50%) and waterlogging. The quality formation, photosynthetic characters, C/N assimilates translocation and regulatory enzyme activities in relation to formation of grain quality were investigated comprehensively, and the physiological regulatory mechanisms of high temperature and water stress on wheat quality formation was elucidated. The main results were as follows:
Contents of crude protein, albumin, globulin and gliadin in grains were significantly enhanced, but glutenin content was reduced by high temperatures, which resulted in decrease of the ratio of glutenin to gliadin. Moreover, contents of total starch, amylose and amylopectin were reduced, but more seriously effect of high temperature on amylopectin than amylose resulted in reducing amylopectin/amylose ratio. In addition, optimum diurnal temperature difference favoring the formations of grain protein and starch differed in wheat genotypes, but the effects of temperature level on protein and starch content were much higher than temperature differences. The second experiment showed that the effect of high temperature on grain protein and starch content were more marked than that of water stress, and the interaction effect of temperature X water also existed. Under high temperature or optimum temperature, grain protein content exhibited the pattern of drought>CK>waterlogging, the highest under drought with high temperature and the lowest under waterlogging with optimum temperature. Grain starch content showed the order CK>drought> waterlogging, the highest under CK with optimum temperature and the lowest under waterlogging with high temperature Furthermore, the content of grain gliadin was enhanced under high temperature and water stresses, while the contents of glutenin and amylopectin were reduced, which resulted in decreased tha ratios of glutenin to gliadin and amylopectin to amylose. Especially, the effect of high temperature with waterlogging was the most significant on protein and starch components. In addition, the response of Yumai 34 in grain protein formation to high temperature and water stresses were more serious than in starch, but for Yangrnai 9, starch formation was more sensitive to high temperature and water stresses. Correlation analysis revealed that protein content was positively correlated to albumin and gliadin contents, but slightly negatively to glutenin content. Starch content was positively to the contents of glutenin and amylopectin and amylopectin/amylose ratio under high temperature and water stresses. High temperature and water stress had different effects on grain gluten characters and sedimentation value in different varieties.
High temperature enhanced free amino acid content in vegetative organs, but significantly reduced soluble sugar content in vegetative organs and kernels during whole grain filling so that ratio of sugar/amino decreased. High temperature reduced remobilization amount and rate of pre-anthesis assimilates and nitrogen stored in vegetative organs and transferring into grain. Effect of diurnal temperature difference on amino acid content differed between cultivars. Under high temperature, temperature difference affected soluble sugar content slightly. Uunder optimum temperature, soluble sugar content under greater diurnal temperature difference was higher than that under lower temperature difference. Under high temperature, day/night temperature difference affected ratio of C/N slightly. Under optimum temperature ratio of C/N was higher under the day/night temperature difference of 12~C. Drought enhanced free amino acid content in vegetative organs, but reduced soluble sugar content in vegetative organs and kernels. As a result, ratio of C/N decreased under high temperature and drought. Under high and optimum temperature, waterlogging reduced contents of amino acid and soluble sugar vegetative organs and kernels. Effect of waterlogging on ratio of C/N differed among different cultivars and organs in wheat. Amount and ratio of assimilates and nitrogen translocation declined as drought>control>waterlogging under optimum temperature, while declined as control>drought>waterlogging under high temperature. The amounts of post-anthesis assimilate transferring into grain were declined as CK>waterlogging>drought under optimum temperature,while the order was CK>drought>waterlogging under high temperature.The amounts of post-anthesis accumulated nitrogen declined as CK>waterlogging>drought under both high and optimum temperature.The amounts of pre-anthesis stored and post-anthesis accumulated nitrogen transferring into grain were significantly related to protein yield.The amounts of pre-anthesis stored and post-anthesis accumulated assimilates transferring into grain were significantly related to grain weight and starch content.
Under optimum water status,high temperature enhanced the activities of sucrose synthase (SS) and granule-bound starch synthase (GBSS) at early stage of grain filling, while reduced activities of SS,GBSS and soluble starch synthase (SSS) at late grain filling stage.Activities of three enzymes were reduced by high temperature in Yumai 34.High temperature significantly reduced activity of SSS in Xuzhou 26 at early stage of grain filling.Those results indicated that the enzymatic mechanism of starch formation differed among wheat varieties.In addition,under high temperatures,SS activity was higher at 34℃/22℃during the middle and late stage of grain filling,whereas the activities of SSS and GBSS were higher under 32℃/24℃.Under optimum temperature,activities of three enzymes were higher at bigger day/night temperature difference.At different day-night temperature differences, GPT (glutamate pyruvic aminotransferase) and glutamine synthase (GS) activities changed between cultivars.Effects of drought and waterlogging on activities of three enzymes differed between different cultivars.Drought and waterlogging reduced activities of three enzymes under both high and optimum temperature.Compared with optimum temperature,high temperature reduced GPT activity in kernels for two wheat cultivars.But reduced GS activity in flag leaves of Yangmai 9 and enhanced GS activity of Xuzhou 26 on 14 DAA.At different day-night temperature differences,GPT and GS activities changed between cultivars.High temperature,drought and waterlogging reduced GS activity and GPT activities in flag leaves.The effects of drought and waterlogging on activities of enzymes involved in protein formation were different among different cultivars and temperatures.Results of correlation analyses indicated that the SPS,SSS and GBSS activities were closely correlated to the yield of starch. GS activity in grains was correlated to a lower degree to the yield and content of protein than GS in flag leaves and GPT ingrains.Under high temperature,the effects of high temperature and water stresses on grain quality and protein and starch accumulation aggravated.
High temperature,drought and watedogging reduced photosynthetic rate and chlorophyll content in flag leaves,and the consequences of drought and waterlogging were more severe under high temperature than under optimum temperature.High temperature, drought and waterlogging enhanced the content of malondiadehyde (MDA),while reduced soluble protein content in flag leaves.Under optimum water treatment,compared with optimal temperature treatment,during the early stage of high temperature treatment,activity of SOD was enhanced but dropped rapidly during late grain filling.Under optimum and high temperature, drought and waterlogging reduced SOD activity on both 15DAA and 20DAA.These results indicated that the membrane lipid peroxidation was accelerated and the permeability of plasma membrane increased as the high temperature treatment prolonged and accelerated leaf senescence.The effects of drought and waterlogging on flag leave were not reversed.The consequences of drought and waterlogging were more severe under high temperature than under optimum temperature.Higher temperature difference under high temperature condition increased senescence of flag leaf,while under optimal temperature condition,higher temperature difference delayed the plant senescence.
In summary,high temperature and water stress reduced the photosynthesis rate in flag leaves of wheat.The amount of post-anthesis assimilates transferred into grains decreased distinctly under high temperature and water stress.High temperature and waterlogging significantly reduced the pre-anthesis stored nitrogen and the translocation of post-anthesis accumulated nitrogen into grains.Thus,the differential substrates caused the differences in protein and starch contents of wheat grains under different water and temperature conditions.The further studies indicated that the activities of key enzymes for grain starch and protein formation,including SPS and GS in leaf,SSS,GBSS and GPT in grains,are the vital enzymatic reasons for affecting the yields and contents of grain starch and protein by high temperature and water stress.The differences between grain qualities under different treatment conditions were caused by the interaction of high temperature and water stress on C/N translocation and activities of key enzymes involved in starch and protein accumulation in wheat
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[1] 王宪泽，李菡，张玲等．山东省推广小麦品种蛋白质和淀粉及其组分的聚类分析．麦类作物学报，2000，21(1)：19-23
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