高温诱导水稻颖花不育特性研究
摘要
针对当前高温热害频发严重影响我国水稻生产这一问题，以Ⅱ优7954（热敏感品种）和Ⅱ优7号（耐热品种）为主要供试材料研究高温诱导水稻颖花不育特性。本研究首先明确水稻高温敏感期和敏感时段。其次研究高温对开花习性的影响，分析高温下颖花受精率下降的原因，探讨高温致害机理。再次研究开花前高温和开花时高温对花药开裂、花粉可染性、花粉粒直径、花粉粒萌发和花粉管延伸的影响。最后研究土壤含水量和空气湿度对高温热害的影响，探讨品种间耐热性差异产生原因以及高温胁迫条件的设置对品种耐热性筛选的影响。主要研究结果如下：
颖花育性高温敏感期的确定。在水稻生殖生长期，以开花期最为敏感；在开花期内，以开花当天最为敏感；而在开花当天，以正在开花的时刻最为敏感。人工气候箱高温时段的设置会影响开花期内高温敏感期，当高温后移时（高温时段为11:00-16:00），敏感期为开花前一天。
高温对水稻开花习性的影响。首先，高温下籼稻花时提前，粳稻花时后延，耐热品种比热敏感品种开花集中。其次，高温下耐热品种较热敏感品种花期缩短，开颖率高。再次，高温下颖花的开放时刻直接决定其育性。早开花有利于籼稻、粳稻提高受精率，晚开花即高温结束后开花有利于籼稻而不利于粳稻提高受精率。再次，高温时段设置直接影响开花习性和颖花受精率。高温发生晚(高温时段为11:00-16:00)比高温发生早(高温时段为9:00-14:00)时，颖花开放高峰出现在高温前，育性主要受开花前期高温影响，颖花育性敏感温度较高。最后，高温下闭颖现象对避热没有作用。
高温对花药开裂影响。常温条件开颖花药即裂的品种，开花前高温对花药开裂没有显著影响，但是散粉性变差。花药开裂较晚的品种，开花前高温对花药开裂影响较大。开颖时高温对花药开裂影响最大，其次是开颖前高温。
高温对花粉粒影响。高温对花粉可染性影响与高温发生时花粉正处的发育阶段紧密相关；开花前遭遇高温时花粉粒内淀粉降解较对照提前，开颖时花粉可染率显著变小，花药开裂越差淀粉降解越严重；开花时高温直接破坏花粉粒内部结构，花粉破裂，可染性降低；高温处理下花粉粒直径开花前一天开始增大，开颖前直径达到最大，开颖后花粉粒皱缩直径变小；开花时高温则是直接引起花粉粒皱缩，直径减小。
高温对花粉粒萌发和花粉管延伸的影响。开颖前和开颖时高温影响散粉、花粉粒萌发和花粉管延伸；开颖后高温主要影响花粉管延伸；开颖后高温处理时间越早，颖花受精率受到影响越大，并且开颖40min花粉管达到珠孔时，颖花受精率不会受到高温的显著影响。除此之外，开花前期高温严重影响了花粉活力，花粉内淀粉降解严重，花粉管延伸短。
水分胁迫和空气湿度对高温热害的影响。高温高湿下，饱和蒸汽压差（VPD）小，气孔导度小，蒸腾速率小，叶温高，花器官受害严重，散粉差，柱头萌发花粉数目少，花粉延伸差。高温干旱条件下，由于水分胁迫气孔关闭，气孔导度小也导致叶温高，伤害颖花。除此之外干旱引起的抽穗不畅，花时晚，开颖率低，花丝伸长而不开颖，高温炙烤下花丝花药萎焉，直接导致颖花不育。
品种间耐热性差异原因。首先是耐热品种常温下开颖即裂药，开颖早散粉好；其次开花习性方面，耐热品种开花早、开花集中、开花峰值高、开花前期日开花量大、开颖率高；再次从颖花不育时段来看，以花药开裂到受精过程为顺序，耐热品种颖花不育发生阶段晚；最后从父母本来讲，品种之间的抗逆性与父母本双方都相关。
Heat stress during reproductive stage was one of the prominent factors to reduce rice yield in China.Ⅱyou7954(heat sensitive variety) and Ⅱyou7(heat tolerant variety) were used to study sterilityinduced by high temperature at reproductive stage. The sensitive stage and time from booting toanthesis to high temperature were determined. The effect of high temperature on floweringcharacteristics was studied, and the cause of spikelet sterility was clear. Then effects of hightemperature in the pre-blooming and blooming periods on anther dehiscence, pollen dyeability, pollengrains diameter, pollen germination and tube growth were studied. Effects of water stress and humidityon heat stress were studied, and heat resistance among varieties and the effect of high temperaturesetting mode on evaluation of heat resisitance in rice were discussed. The main results in this study wereas follows:
The sensitive stage of spikelets sterility to high temperure at reproductive stage was anthesis stage.The sensitive stage of spikelets sterility to high temperure at anthesis stage was the day at blooming, andthe sensitive time of spikelets sterility to high temperure at blooming was glume opening. Hightemperature setting mode affected the sensitive stage of spikelets sterility to high temperure at anthesisstage, which was the day before blooming when exposing to11:00-16:00heat treatment.
High temperature advanced the flowering clock of Ⅱyou7andⅡyou7954, delayed that oflianjing6. Ⅱyou7had obvious flowering peak, yet Ⅱyou7954and lianjing6had low or indistinctflowering peaks. The flowering period of heat tolerant sensitive variety was shortened and the dailynumber of spikelet flowering was more than that of heat sensitive variety. Glume opening time affectedspikelet fertility. Early flowering raised the spikelet fertility both in indica and japonica rice, andflowering after the end of heat stress raised the spikelet fertility of indica rice, not japonica rice. Hightemperature setting mode affected flowering characteristics and spikelet fertility. In contrast with hightemperature during9:00-14:00, the flowering peak of high temperature during11:00-16:00occurredbefore heat stress, then sterility was induced by high temperature before blooming not by hightemperature occurring at blooming, and the sensitive temperature was higher. High temperatureinhibited glume opening for good at anthesis stage, which didn’t take a part in avoiding heat injury.
High temperature before blooming didn’t affected the anther dehiscence of varieties whose antherstarting splitting up just after glume opening under moderate temperature, while the pollen sheddingwas still bad, and affected anther dehiscence of varieties whose anther dehiscence was late. Hightemperature at glume opening affected anther dehiscence most seriously, and the temperature beforeglume opening affected that less.
Pollen dyeability closely related to the developmental stages. The starch in pollen grains subjectedto high temperature before blooming began to degcompose, and became very few when glume opening.The worse the anther dehiscence, the worse the pollen dyeabilitity was. Temperature at bloomingdestroyed the internal structure of pollen grains, the pollen grains were broke, and the pollen dyeability was bad. Under high temperature, the pollen diameter at one day before blooming began to expand,became largest just before glume opening, then the pollen grains shrived after glum opening. Hightemperature at blooming made pollen shrived, then the diameter of which reduced.
High temperature before and at glume opening affected pollen shedding, pollen germination andtube growth. High temperature after glume opening affected tube growth. The earlier the hightemperature came, the lower the spikelet fertility was, and when pollen tube entered into micropyl40min after glume opening, high temperature had no significant effect on spikelet fertility. Hightemperature before blooming affected pollen viability greatly, the starch in pollen grains degraded, andthe length of pollen tube was short.
When relative humidity and temperature was high, the VPD was low, stomatal conductancedecreased, transpiration rate decreased, the leaf temperature was high, floral organ suffered severely,pollen shedding was bad, the germinated pollen grains on the stigma were few, and the extension ofpollen tube was bad. When blooming under water and heat stress, stomatal conductance decreased, leaftemperature was high, and floral organ suffered severely. Besides, water stress induced poor heading,late flowering clock, poor proportion of spikelets anthesing, then when the filament elongated the glumedidn’t open, which induced anther shriveled and sterility.
The anther of heat tolerant varieties began to split up just after glume opening under moderatetemperature, then the early dehiscence led to good pollen shedding. Early flowering, obvious floweringpeak, high flowering peak, high proportion of spikelets anthesing at the earlier stages during floweringperiod, high proportion of spikelets anthesing under high temperature were found in heat tolerantvarieties. The sterility of heat tolerant varieties occurred at later stages Heat tolerance of varieties wasrelated with their male parents and female parents.
颖花育性高温敏感期的确定。在水稻生殖生长期，以开花期最为敏感；在开花期内，以开花当天最为敏感；而在开花当天，以正在开花的时刻最为敏感。人工气候箱高温时段的设置会影响开花期内高温敏感期，当高温后移时（高温时段为11:00-16:00），敏感期为开花前一天。
高温对水稻开花习性的影响。首先，高温下籼稻花时提前，粳稻花时后延，耐热品种比热敏感品种开花集中。其次，高温下耐热品种较热敏感品种花期缩短，开颖率高。再次，高温下颖花的开放时刻直接决定其育性。早开花有利于籼稻、粳稻提高受精率，晚开花即高温结束后开花有利于籼稻而不利于粳稻提高受精率。再次，高温时段设置直接影响开花习性和颖花受精率。高温发生晚(高温时段为11:00-16:00)比高温发生早(高温时段为9:00-14:00)时，颖花开放高峰出现在高温前，育性主要受开花前期高温影响，颖花育性敏感温度较高。最后，高温下闭颖现象对避热没有作用。
高温对花药开裂影响。常温条件开颖花药即裂的品种，开花前高温对花药开裂没有显著影响，但是散粉性变差。花药开裂较晚的品种，开花前高温对花药开裂影响较大。开颖时高温对花药开裂影响最大，其次是开颖前高温。
高温对花粉粒影响。高温对花粉可染性影响与高温发生时花粉正处的发育阶段紧密相关；开花前遭遇高温时花粉粒内淀粉降解较对照提前，开颖时花粉可染率显著变小，花药开裂越差淀粉降解越严重；开花时高温直接破坏花粉粒内部结构，花粉破裂，可染性降低；高温处理下花粉粒直径开花前一天开始增大，开颖前直径达到最大，开颖后花粉粒皱缩直径变小；开花时高温则是直接引起花粉粒皱缩，直径减小。
高温对花粉粒萌发和花粉管延伸的影响。开颖前和开颖时高温影响散粉、花粉粒萌发和花粉管延伸；开颖后高温主要影响花粉管延伸；开颖后高温处理时间越早，颖花受精率受到影响越大，并且开颖40min花粉管达到珠孔时，颖花受精率不会受到高温的显著影响。除此之外，开花前期高温严重影响了花粉活力，花粉内淀粉降解严重，花粉管延伸短。
水分胁迫和空气湿度对高温热害的影响。高温高湿下，饱和蒸汽压差（VPD）小，气孔导度小，蒸腾速率小，叶温高，花器官受害严重，散粉差，柱头萌发花粉数目少，花粉延伸差。高温干旱条件下，由于水分胁迫气孔关闭，气孔导度小也导致叶温高，伤害颖花。除此之外干旱引起的抽穗不畅，花时晚，开颖率低，花丝伸长而不开颖，高温炙烤下花丝花药萎焉，直接导致颖花不育。
品种间耐热性差异原因。首先是耐热品种常温下开颖即裂药，开颖早散粉好；其次开花习性方面，耐热品种开花早、开花集中、开花峰值高、开花前期日开花量大、开颖率高；再次从颖花不育时段来看，以花药开裂到受精过程为顺序，耐热品种颖花不育发生阶段晚；最后从父母本来讲，品种之间的抗逆性与父母本双方都相关。
Heat stress during reproductive stage was one of the prominent factors to reduce rice yield in China.Ⅱyou7954(heat sensitive variety) and Ⅱyou7(heat tolerant variety) were used to study sterilityinduced by high temperature at reproductive stage. The sensitive stage and time from booting toanthesis to high temperature were determined. The effect of high temperature on floweringcharacteristics was studied, and the cause of spikelet sterility was clear. Then effects of hightemperature in the pre-blooming and blooming periods on anther dehiscence, pollen dyeability, pollengrains diameter, pollen germination and tube growth were studied. Effects of water stress and humidityon heat stress were studied, and heat resistance among varieties and the effect of high temperaturesetting mode on evaluation of heat resisitance in rice were discussed. The main results in this study wereas follows:
The sensitive stage of spikelets sterility to high temperure at reproductive stage was anthesis stage.The sensitive stage of spikelets sterility to high temperure at anthesis stage was the day at blooming, andthe sensitive time of spikelets sterility to high temperure at blooming was glume opening. Hightemperature setting mode affected the sensitive stage of spikelets sterility to high temperure at anthesisstage, which was the day before blooming when exposing to11:00-16:00heat treatment.
High temperature advanced the flowering clock of Ⅱyou7andⅡyou7954, delayed that oflianjing6. Ⅱyou7had obvious flowering peak, yet Ⅱyou7954and lianjing6had low or indistinctflowering peaks. The flowering period of heat tolerant sensitive variety was shortened and the dailynumber of spikelet flowering was more than that of heat sensitive variety. Glume opening time affectedspikelet fertility. Early flowering raised the spikelet fertility both in indica and japonica rice, andflowering after the end of heat stress raised the spikelet fertility of indica rice, not japonica rice. Hightemperature setting mode affected flowering characteristics and spikelet fertility. In contrast with hightemperature during9:00-14:00, the flowering peak of high temperature during11:00-16:00occurredbefore heat stress, then sterility was induced by high temperature before blooming not by hightemperature occurring at blooming, and the sensitive temperature was higher. High temperatureinhibited glume opening for good at anthesis stage, which didn’t take a part in avoiding heat injury.
High temperature before blooming didn’t affected the anther dehiscence of varieties whose antherstarting splitting up just after glume opening under moderate temperature, while the pollen sheddingwas still bad, and affected anther dehiscence of varieties whose anther dehiscence was late. Hightemperature at glume opening affected anther dehiscence most seriously, and the temperature beforeglume opening affected that less.
Pollen dyeability closely related to the developmental stages. The starch in pollen grains subjectedto high temperature before blooming began to degcompose, and became very few when glume opening.The worse the anther dehiscence, the worse the pollen dyeabilitity was. Temperature at bloomingdestroyed the internal structure of pollen grains, the pollen grains were broke, and the pollen dyeability was bad. Under high temperature, the pollen diameter at one day before blooming began to expand,became largest just before glume opening, then the pollen grains shrived after glum opening. Hightemperature at blooming made pollen shrived, then the diameter of which reduced.
High temperature before and at glume opening affected pollen shedding, pollen germination andtube growth. High temperature after glume opening affected tube growth. The earlier the hightemperature came, the lower the spikelet fertility was, and when pollen tube entered into micropyl40min after glume opening, high temperature had no significant effect on spikelet fertility. Hightemperature before blooming affected pollen viability greatly, the starch in pollen grains degraded, andthe length of pollen tube was short.
When relative humidity and temperature was high, the VPD was low, stomatal conductancedecreased, transpiration rate decreased, the leaf temperature was high, floral organ suffered severely,pollen shedding was bad, the germinated pollen grains on the stigma were few, and the extension ofpollen tube was bad. When blooming under water and heat stress, stomatal conductance decreased, leaftemperature was high, and floral organ suffered severely. Besides, water stress induced poor heading,late flowering clock, poor proportion of spikelets anthesing, then when the filament elongated the glumedidn’t open, which induced anther shriveled and sterility.
The anther of heat tolerant varieties began to split up just after glume opening under moderatetemperature, then the early dehiscence led to good pollen shedding. Early flowering, obvious floweringpeak, high flowering peak, high proportion of spikelets anthesing at the earlier stages during floweringperiod, high proportion of spikelets anthesing under high temperature were found in heat tolerantvarieties. The sterility of heat tolerant varieties occurred at later stages Heat tolerance of varieties wasrelated with their male parents and female parents.
引文
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