遮光对北方粳稻产量和品质的形成及其生理机制的研究
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摘要
试验于2011-2012年在沈阳农业大学水稻研究所试验田进行。2011年选用粳稻品种秋光为试料,分别在分蘖期、拔节期、抽穗期和抽穗后10d四个关键生育时期研究了不同遮光处理对北方粳稻品种产量和品质的影响,探究了光照不足影响北方粳稻产量和品质的关键时期及其变化规律。2012年,在上一年试验结果基础上,选用沈农9741(直立穗型)、盐丰47(半直立穗型)和辽盐16(弯穗型)3种不同穗型粳稻品种,·对大田条件下抽穗期遮光对水稻光合生理特性、碳氮代谢及籽粒产量的影响及其生理机制进行了研究。主要研究结果如下：
1.不同时期遮光导致了水稻品种单株叶面积和地上部各器官干重的降低；分蘖期遮光导致了单株叶片重的降低幅度最大,为26.77%,而拔节期遮光主要导致了单株茎鞘重的下降,达46.02%,抽穗后遮光主要导致了穗重的降低,最大降幅约为45%左右。对各器官干物质分配比率的研究表明,分蘖期和拔节期遮光相对提高了干物质向叶重分配的比率,而抽穗后遮光降低了穗部干物质分配率,不同时期遮光并未显著改变水稻品种植株干物质向各器官分配比例,而只是延缓了各时期关键器官形态建成的速率。不同时期遮光对北方粳稻产量的影响有所不同,分蘖期和拔节期遮光后产量分别下降至8t·hm-2和7t·hm-+2左右,与对照相比最大降幅达10%以上,抽穗后遮光产量显著降低至6t·hm-2左右,最大降幅达40%以上。遮光对有效穗数的影响未达到显著水平,抽穗期以前遮光导致了每穗粒数的减少,而抽穗后遮光导致了结实率和千粒重的显著降低,这可能是遮光导致水稻产量显著下降的原因所在。抽穗期遮光导致了籽粒产量贡献最大的穗中上部产量显著降低。
2.不同时期遮光对北方粳稻品质的影响表明,不同时期和不同遮光强度下稻米的碾磨、外观和营养品质均有所下降,而遮光10d内遮光天数对稻米品质的影响不大。遮光导致了稻米外观品质显著降低；随着遮光时期的后移和遮光强度的增加,垩白粒率和垩白度呈显著的增加趋势。对不同时期遮光后稻米品质变化的研究表明,抽穗前遮光对稻米品质的影响不大,遮光对稻米品质的影响主要在抽穗后籽粒结实阶段,这也进一步说明弱光胁迫导致稻米品质下降的原因可能是籽粒灌浆充实不良所致。
3.遮光胁迫后不同穗型粳稻产量变化的结果表明,遮光后不同穗型水稻品种的产量均随遮光天数的增加而显著降低,其中以沈农9741产量的降幅最大,达45.59%,而辽盐16和盐丰47降幅基本一致,分别为40.07%和39.29%。对产量结构的分析表明,抽穗期遮光对不同穗型水稻品种每穗粒数的影响不大,未达到显著水平,但均显著降低了不同穗型品种的千粒重和结实率。总体来看,遮光对千粒重和结实率的影响以沈农9741受遮光胁迫的程度最大。
4.遮光胁迫后不同穗型粳稻叶片光合特性的结果表明,遮光导致了粳稻叶片叶绿素含量的增加,其中以对沈农9741的影响最大；同时,遮光导致了叶片净光合速率(Pn)的降低,以沈农9741的降幅最大,为49.87%,辽盐16降幅最小,为31.78%；同时,遮光也导致了粳稻品种剑叶叶片蒸腾速率与表观叶肉导度的降低,但增加了叶片胞间CO2浓度,且不同穗型粳稻品种之间差异并不显著；遮光后不同穗型粳稻叶片的原初光能转化效率(Fv/Fm)有所增加,但主要显著降低了抽穗后6d内的实际光化学效率(Yield),这说明遮光主要导致了粳稻叶片光能用于光化学反应比例的降低,而提高了光能转化为热能耗散的比例。50%遮光导致了不同穗型粳稻品种SOD、POD和CAT活性的降低,但未达到显著水平,同时遮光增加了品种MDA的含量。这说明50%遮光严重的抑制了粳稻品种叶片抗氧化酶活性,活性氧的产生超过了抗氧化酶的清除能力,进而导致膜系统被伤害,这可能也是不同穗型粳稻品种叶片光合能力下降的原因所在。遮光处理对不同穗型粳稻叶绿体结构均有较大的影响,具体表现在叶绿体外形肿胀,与对照相比出现了较多的嗜锇颗粒,类囊体垛叠程度增高且紧密,淀粉粒数目有所减少。遮光后叶绿体内细胞中胞质出现了一定程度的浓缩和周缘化,但类囊体与基粒片层结构尚为完整。遮光后不同穗型粳稻品种都呈现了一定的耐阴性,抽穗期遮光对直立穗型的影响大于对半直立穗型和弯穗型的影响。直立穗型遮光后叶片叶绿素含量的增加虽提高了吸收光能的能力,但其实际光化学效率显著降低,大部分光能转化为热能耗散。
5.遮光导致了水稻品种穗部干物质的积累降低,对单株叶重和单株茎鞘重的影响并不是一味的降低,而是因品种穗型的不同而不同。总体来讲,遮光对沈农9741的单株叶重和单株茎鞘重的影响较小,而对盐丰47和辽盐16的单株叶重和茎鞘干重的影响较大；遮光对盐丰47单株穗重的降低最小,连续遮光8d以上可显著降低沈农9741与辽盐16的单株穗重。进一步分析抽穗10d内遮光不同穗型各器官干物质分配比率的变化表明,遮光均降低了不同穗型水稻品种干物质向穗部分配的比例,随着穗部干物质分配的降低,沈农9741和盐丰47的叶片干物质分配率有所上升,而辽盐16的茎鞘干物质分配比率有所增加。
6.遮光后粳稻碳氮代谢酶活性的变化表明,遮光后水稻叶片和籽粒的SS和SBS活性均降低,导致蔗糖含量的降低,限制了叶片光合产物的输出,间接的延缓了穗部籽粒淀粉的合成。遮光对品种叶片内氮代谢酶活性的影响要大于籽粒中的酶活性,且品种之间的比较表明,遮光对沈农9741的氮代谢酶活性影响较大。遮光均导致了不同穗型水稻品种GS和GOGAT活性的降低,而增加了品种GDH的活性。
The experiment was carried out in Rice Research Institute of Shenyang Agricultural University from2011to2012. In the first year, the japonica rice variety, Akihikari, was used to research the effects of shading on the yield and quality of japonica rice within different growth stages such as tillering stage (TS), jointing stage (JS), heading stage (HS) and10days after heading stage (10DAH) which are the critical stages of that, and find out the regularity of effects of shading on the yield and quality of japonica rice in northern China. Based on the results of2011, three different panicle types japonica rice cultivars including erect panicle rice Shennong9741, half-erect panicle rice Yanfeng47and curved panicle rice Liaoyanl6were used in2012, and further some major development of japonica rice were investigated with shading at heading stage under field conditions, such as photosynthetic physiological characteristics, carbon and nitrogen metabolism and grain yield and its physiological mechanism. The mainly results were as follows:
1. The leaf area per plant and dry matter weight of each part aboveground were reduced by shading at different stages; the largest reduction of leaves weight per plant,26.77%, was got at tillering stage under shading treatment, while shading treatment mainly decreased the weight of stem per plant that there was a46.02%decrease at jointing stage. The weight of panicle was mainly decreased by shading after heading stage and the biggest drop of that was about45%. Studying on dry matter distribution of each part, it showed that there has been increased in the dry matter partition of leaf weight per plant under shading treatment at tillering stage and jointing stage, but decreased in dry matter partition of panicle per plant under shading treatment after heading stage. It was no obvious changes in dry matter partition of each part with shading treatment at different growth stages, however, shading just retarded the development of leaf or panicle (sink) at each growth stage. The influence of shading on yield of japonica rice was different at different stages, which indicated that the yield after shading within TS and JS was reduced from about9t·hm-2to8t·hm-2and7t·hm-2respectively, and the drop of yield was more than40%in HS and10DAH which was significant reduced from about9t·hm-2to approximately6t·hm-2under shading stress. The effects of shading on spikelets per panicle, seed setting rate,1000-grains weight and grain yield were significant (P<0.05), however there was no obvious effects of shading on effective panicles. Shading mainly led to the decrease of spikelets per panicle in the TS and JS and decrease of seed setting rate and1000-grains weight in HS and10DAH. That might be the reason for the marked reduction of grain of yield of rice under shading treatment. The yield of the top and middle of panicle which had the most contribution to the grain of yield was reduced evidently with shading treatment at heading stage.
2. Studying on the effects of shading on quality of japonica rice at different stages, it showed that there were no obvious changes in quality of rice under shading treatment during TS and JS. With the increase of shading degree, the milling quality, appearance quality and nutrient quality were decreased in different degrees after HS. Shading mainly resulted in significant increase of chalkiness rate and chalkiness degree (P<0.05), however there was no significant influence on milled rice rate (P>0.05). With the increase of the days for shading, the quality of rice was decreased slightly but was not significant. This study also indicated that the quality of rice wasn't changed evidently before heading. The changes in the quality of rice mainly happened at grain-filling stage after heading, which was a more advance explanation that the reason why the quality of rice was reduced by shading treatment was the poor grain plumpness.
3. Studying on the changes in yield of japonica rice with different panicle types under shading treatment, it showed that the yield was decreased with increase of shading days significantly. The reduction in the yield of Shennong9741was the most, which was up to45.49%, while the drop of Liaoyan16and Yanfeng47were almost the same, which was40.07%and39.29%, respectively. The analysis of yield components indicated that there was no obvious influence on spikelets per panicle of rice with different panicle types under shading treatment at heading stage, and it was not up to a significant level. But both1000-grains weight and the rate of fructification were decreased markedly. Overall, the changes in Shennong9741from the aspect of1000-grains weight and the rate of fructification were the biggest under shading treatment.
4. The chlorophyll content of flag-leaf was increased, and the increase of chlorophyll content in Shennong9741was most, which were resulted from studying on the photosynthetic characteristics of flag-leaf in japonica rice with different panicle types with shading treatment. Meanwhile, the net photosynthesis rate (Pn) of flag-leaf was decreased that the drop of Shennong9741was the biggest (49.87%) and the drop of Liaoyanl6was the smallest (31.78%). The transpiration rate (Tr) and apparent mesophyll conductance (Gs) of flag-leaf were also reduced with shading treatment, but CO2concentration intercellular (Ci) of flag-leaf was increased, and the difference among the photosynthetic characteristics of japonica rice with different panicle types was not significant. It was increased in Fv/Fm of flag-leaf in japonica rice with different panicle types under shading, but the Yield of flag-leaf was significantly decreased under shading within6days after heading. That suggested that shading led mainly to reduce the proportion of visible light for photochemical reaction, while increased the proportion of light energy conversion for heat dissipation. The results aslo indicated that shading could result in decrease of SOD、POD and CAT activity of flag-leaf in japonica rice, which did not reach a significant level, but it increased the MDA contents in cultivars at the same time. That suggests that50%shading constrained the activity of antioxidant enzymes in flag-leaf severely, which was resulted in the generation of active oxygen exceeding the scavenging ability of antioxidant enzymes, and then the membrane system was damaged eventually. It also might be one of the main reasons for the reduction of leaf photosynthetic capacity in japonica rice cultivars with different panicle types. The Chloroplast structure of japonica rice with different panicle types was strongly influenced under shading, and specific performance in the shape of chloroplast was swelled slightly. Meanwhile, there has more numbers of osmphore, fewer numbers of starch granule and higher of thylakoids stacking in chloroplast. There was a certain degree of concentration and periphery in cytoplasmic of chloroplast cell under shading, but thylakoid and lamella structure of grana were almost complete. Our studies indicated that there had different degrees of shading tolerance in japonica rice with different panicle types under shading stress, but the influences of shading on Shennong9741were greater than that of Yanfeng47and Liaoyanl6. The ability to absorb light energy was improved increasing chlorophyll content in Shennong9741, but the most of the light energy was converted into heat and dissipated, which was resulted in the actual photochemical efficiency decreasing significantly.
5. The accumulation of dry weight was decreased under shading, while the effects on leaf weight per plant and the weight of stem and sheath per plant were complex, which was different with different panicle types. Generally, the effects of shading on the weight of leaf and stem per plant of Shennong9741were smaller than those of Yanfeng47and Liaoyanl6. The reduction of panicle weight per plant of Yanfeng47was the smallest under shading, and panicle weight per plant of Shennong9741and Liaoyanl6were decreased significantly within8days after shading. Further analysis on the change of dry matter distribution of each part within shading after heading10days showed that, dry matter distribution of panicle were all reduced. As dry matter distribution of panicle was decreased, dry matter distribution of leaves in Shennong9741and Yanfeng47was increased slightly, while those of stem in Liaoyan16were increased slimly.
6. Analyzing the changes in enzymes of carbon and nitrogen metabolism in leaf and grain of rice with different panicle types, it indicated that the activity of SS and SBS was declined under shading treatment, which was resulted in the decrease of sucrose contents. That had limited output of photosynthetic products and delayed starch synthesis in panicle indirectly. The influence on enzymes of nitrogen metabolism in leaf was larger than those in grain with shading treatment. Meanwhile, the comparison among rice with different panicle types showed that the effects of shading on enzymes of nitrogen metabolism in Shennong9741were larger than others. The activity of GS and GOGAT of different panicle types was decreased by shading, but the activity of GDH was increased.
1.不同时期遮光导致了水稻品种单株叶面积和地上部各器官干重的降低；分蘖期遮光导致了单株叶片重的降低幅度最大,为26.77%,而拔节期遮光主要导致了单株茎鞘重的下降,达46.02%,抽穗后遮光主要导致了穗重的降低,最大降幅约为45%左右。对各器官干物质分配比率的研究表明,分蘖期和拔节期遮光相对提高了干物质向叶重分配的比率,而抽穗后遮光降低了穗部干物质分配率,不同时期遮光并未显著改变水稻品种植株干物质向各器官分配比例,而只是延缓了各时期关键器官形态建成的速率。不同时期遮光对北方粳稻产量的影响有所不同,分蘖期和拔节期遮光后产量分别下降至8t·hm-2和7t·hm-+2左右,与对照相比最大降幅达10%以上,抽穗后遮光产量显著降低至6t·hm-2左右,最大降幅达40%以上。遮光对有效穗数的影响未达到显著水平,抽穗期以前遮光导致了每穗粒数的减少,而抽穗后遮光导致了结实率和千粒重的显著降低,这可能是遮光导致水稻产量显著下降的原因所在。抽穗期遮光导致了籽粒产量贡献最大的穗中上部产量显著降低。
2.不同时期遮光对北方粳稻品质的影响表明,不同时期和不同遮光强度下稻米的碾磨、外观和营养品质均有所下降,而遮光10d内遮光天数对稻米品质的影响不大。遮光导致了稻米外观品质显著降低；随着遮光时期的后移和遮光强度的增加,垩白粒率和垩白度呈显著的增加趋势。对不同时期遮光后稻米品质变化的研究表明,抽穗前遮光对稻米品质的影响不大,遮光对稻米品质的影响主要在抽穗后籽粒结实阶段,这也进一步说明弱光胁迫导致稻米品质下降的原因可能是籽粒灌浆充实不良所致。
3.遮光胁迫后不同穗型粳稻产量变化的结果表明,遮光后不同穗型水稻品种的产量均随遮光天数的增加而显著降低,其中以沈农9741产量的降幅最大,达45.59%,而辽盐16和盐丰47降幅基本一致,分别为40.07%和39.29%。对产量结构的分析表明,抽穗期遮光对不同穗型水稻品种每穗粒数的影响不大,未达到显著水平,但均显著降低了不同穗型品种的千粒重和结实率。总体来看,遮光对千粒重和结实率的影响以沈农9741受遮光胁迫的程度最大。
4.遮光胁迫后不同穗型粳稻叶片光合特性的结果表明,遮光导致了粳稻叶片叶绿素含量的增加,其中以对沈农9741的影响最大；同时,遮光导致了叶片净光合速率(Pn)的降低,以沈农9741的降幅最大,为49.87%,辽盐16降幅最小,为31.78%；同时,遮光也导致了粳稻品种剑叶叶片蒸腾速率与表观叶肉导度的降低,但增加了叶片胞间CO2浓度,且不同穗型粳稻品种之间差异并不显著；遮光后不同穗型粳稻叶片的原初光能转化效率(Fv/Fm)有所增加,但主要显著降低了抽穗后6d内的实际光化学效率(Yield),这说明遮光主要导致了粳稻叶片光能用于光化学反应比例的降低,而提高了光能转化为热能耗散的比例。50%遮光导致了不同穗型粳稻品种SOD、POD和CAT活性的降低,但未达到显著水平,同时遮光增加了品种MDA的含量。这说明50%遮光严重的抑制了粳稻品种叶片抗氧化酶活性,活性氧的产生超过了抗氧化酶的清除能力,进而导致膜系统被伤害,这可能也是不同穗型粳稻品种叶片光合能力下降的原因所在。遮光处理对不同穗型粳稻叶绿体结构均有较大的影响,具体表现在叶绿体外形肿胀,与对照相比出现了较多的嗜锇颗粒,类囊体垛叠程度增高且紧密,淀粉粒数目有所减少。遮光后叶绿体内细胞中胞质出现了一定程度的浓缩和周缘化,但类囊体与基粒片层结构尚为完整。遮光后不同穗型粳稻品种都呈现了一定的耐阴性,抽穗期遮光对直立穗型的影响大于对半直立穗型和弯穗型的影响。直立穗型遮光后叶片叶绿素含量的增加虽提高了吸收光能的能力,但其实际光化学效率显著降低,大部分光能转化为热能耗散。
5.遮光导致了水稻品种穗部干物质的积累降低,对单株叶重和单株茎鞘重的影响并不是一味的降低,而是因品种穗型的不同而不同。总体来讲,遮光对沈农9741的单株叶重和单株茎鞘重的影响较小,而对盐丰47和辽盐16的单株叶重和茎鞘干重的影响较大；遮光对盐丰47单株穗重的降低最小,连续遮光8d以上可显著降低沈农9741与辽盐16的单株穗重。进一步分析抽穗10d内遮光不同穗型各器官干物质分配比率的变化表明,遮光均降低了不同穗型水稻品种干物质向穗部分配的比例,随着穗部干物质分配的降低,沈农9741和盐丰47的叶片干物质分配率有所上升,而辽盐16的茎鞘干物质分配比率有所增加。
6.遮光后粳稻碳氮代谢酶活性的变化表明,遮光后水稻叶片和籽粒的SS和SBS活性均降低,导致蔗糖含量的降低,限制了叶片光合产物的输出,间接的延缓了穗部籽粒淀粉的合成。遮光对品种叶片内氮代谢酶活性的影响要大于籽粒中的酶活性,且品种之间的比较表明,遮光对沈农9741的氮代谢酶活性影响较大。遮光均导致了不同穗型水稻品种GS和GOGAT活性的降低,而增加了品种GDH的活性。
The experiment was carried out in Rice Research Institute of Shenyang Agricultural University from2011to2012. In the first year, the japonica rice variety, Akihikari, was used to research the effects of shading on the yield and quality of japonica rice within different growth stages such as tillering stage (TS), jointing stage (JS), heading stage (HS) and10days after heading stage (10DAH) which are the critical stages of that, and find out the regularity of effects of shading on the yield and quality of japonica rice in northern China. Based on the results of2011, three different panicle types japonica rice cultivars including erect panicle rice Shennong9741, half-erect panicle rice Yanfeng47and curved panicle rice Liaoyanl6were used in2012, and further some major development of japonica rice were investigated with shading at heading stage under field conditions, such as photosynthetic physiological characteristics, carbon and nitrogen metabolism and grain yield and its physiological mechanism. The mainly results were as follows:
1. The leaf area per plant and dry matter weight of each part aboveground were reduced by shading at different stages; the largest reduction of leaves weight per plant,26.77%, was got at tillering stage under shading treatment, while shading treatment mainly decreased the weight of stem per plant that there was a46.02%decrease at jointing stage. The weight of panicle was mainly decreased by shading after heading stage and the biggest drop of that was about45%. Studying on dry matter distribution of each part, it showed that there has been increased in the dry matter partition of leaf weight per plant under shading treatment at tillering stage and jointing stage, but decreased in dry matter partition of panicle per plant under shading treatment after heading stage. It was no obvious changes in dry matter partition of each part with shading treatment at different growth stages, however, shading just retarded the development of leaf or panicle (sink) at each growth stage. The influence of shading on yield of japonica rice was different at different stages, which indicated that the yield after shading within TS and JS was reduced from about9t·hm-2to8t·hm-2and7t·hm-2respectively, and the drop of yield was more than40%in HS and10DAH which was significant reduced from about9t·hm-2to approximately6t·hm-2under shading stress. The effects of shading on spikelets per panicle, seed setting rate,1000-grains weight and grain yield were significant (P<0.05), however there was no obvious effects of shading on effective panicles. Shading mainly led to the decrease of spikelets per panicle in the TS and JS and decrease of seed setting rate and1000-grains weight in HS and10DAH. That might be the reason for the marked reduction of grain of yield of rice under shading treatment. The yield of the top and middle of panicle which had the most contribution to the grain of yield was reduced evidently with shading treatment at heading stage.
2. Studying on the effects of shading on quality of japonica rice at different stages, it showed that there were no obvious changes in quality of rice under shading treatment during TS and JS. With the increase of shading degree, the milling quality, appearance quality and nutrient quality were decreased in different degrees after HS. Shading mainly resulted in significant increase of chalkiness rate and chalkiness degree (P<0.05), however there was no significant influence on milled rice rate (P>0.05). With the increase of the days for shading, the quality of rice was decreased slightly but was not significant. This study also indicated that the quality of rice wasn't changed evidently before heading. The changes in the quality of rice mainly happened at grain-filling stage after heading, which was a more advance explanation that the reason why the quality of rice was reduced by shading treatment was the poor grain plumpness.
3. Studying on the changes in yield of japonica rice with different panicle types under shading treatment, it showed that the yield was decreased with increase of shading days significantly. The reduction in the yield of Shennong9741was the most, which was up to45.49%, while the drop of Liaoyan16and Yanfeng47were almost the same, which was40.07%and39.29%, respectively. The analysis of yield components indicated that there was no obvious influence on spikelets per panicle of rice with different panicle types under shading treatment at heading stage, and it was not up to a significant level. But both1000-grains weight and the rate of fructification were decreased markedly. Overall, the changes in Shennong9741from the aspect of1000-grains weight and the rate of fructification were the biggest under shading treatment.
4. The chlorophyll content of flag-leaf was increased, and the increase of chlorophyll content in Shennong9741was most, which were resulted from studying on the photosynthetic characteristics of flag-leaf in japonica rice with different panicle types with shading treatment. Meanwhile, the net photosynthesis rate (Pn) of flag-leaf was decreased that the drop of Shennong9741was the biggest (49.87%) and the drop of Liaoyanl6was the smallest (31.78%). The transpiration rate (Tr) and apparent mesophyll conductance (Gs) of flag-leaf were also reduced with shading treatment, but CO2concentration intercellular (Ci) of flag-leaf was increased, and the difference among the photosynthetic characteristics of japonica rice with different panicle types was not significant. It was increased in Fv/Fm of flag-leaf in japonica rice with different panicle types under shading, but the Yield of flag-leaf was significantly decreased under shading within6days after heading. That suggested that shading led mainly to reduce the proportion of visible light for photochemical reaction, while increased the proportion of light energy conversion for heat dissipation. The results aslo indicated that shading could result in decrease of SOD、POD and CAT activity of flag-leaf in japonica rice, which did not reach a significant level, but it increased the MDA contents in cultivars at the same time. That suggests that50%shading constrained the activity of antioxidant enzymes in flag-leaf severely, which was resulted in the generation of active oxygen exceeding the scavenging ability of antioxidant enzymes, and then the membrane system was damaged eventually. It also might be one of the main reasons for the reduction of leaf photosynthetic capacity in japonica rice cultivars with different panicle types. The Chloroplast structure of japonica rice with different panicle types was strongly influenced under shading, and specific performance in the shape of chloroplast was swelled slightly. Meanwhile, there has more numbers of osmphore, fewer numbers of starch granule and higher of thylakoids stacking in chloroplast. There was a certain degree of concentration and periphery in cytoplasmic of chloroplast cell under shading, but thylakoid and lamella structure of grana were almost complete. Our studies indicated that there had different degrees of shading tolerance in japonica rice with different panicle types under shading stress, but the influences of shading on Shennong9741were greater than that of Yanfeng47and Liaoyanl6. The ability to absorb light energy was improved increasing chlorophyll content in Shennong9741, but the most of the light energy was converted into heat and dissipated, which was resulted in the actual photochemical efficiency decreasing significantly.
5. The accumulation of dry weight was decreased under shading, while the effects on leaf weight per plant and the weight of stem and sheath per plant were complex, which was different with different panicle types. Generally, the effects of shading on the weight of leaf and stem per plant of Shennong9741were smaller than those of Yanfeng47and Liaoyanl6. The reduction of panicle weight per plant of Yanfeng47was the smallest under shading, and panicle weight per plant of Shennong9741and Liaoyanl6were decreased significantly within8days after shading. Further analysis on the change of dry matter distribution of each part within shading after heading10days showed that, dry matter distribution of panicle were all reduced. As dry matter distribution of panicle was decreased, dry matter distribution of leaves in Shennong9741and Yanfeng47was increased slightly, while those of stem in Liaoyan16were increased slimly.
6. Analyzing the changes in enzymes of carbon and nitrogen metabolism in leaf and grain of rice with different panicle types, it indicated that the activity of SS and SBS was declined under shading treatment, which was resulted in the decrease of sucrose contents. That had limited output of photosynthetic products and delayed starch synthesis in panicle indirectly. The influence on enzymes of nitrogen metabolism in leaf was larger than those in grain with shading treatment. Meanwhile, the comparison among rice with different panicle types showed that the effects of shading on enzymes of nitrogen metabolism in Shennong9741were larger than others. The activity of GS and GOGAT of different panicle types was decreased by shading, but the activity of GDH was increased.
引文
1.艾希珍,郭延奎,马兴庄,等.2004.弱光条件下日光温室黄瓜需光特性及叶绿体超微结构[J].中国农业科学,2：268-273.
2.本庄一雄.1971.稻米蛋白质含量的研究Ⅰ不同施肥条件对糙米蛋白质含有率和蛋白质总量的影响[J].日本作物学会纪事,40：190-197.
3.蔡昆争,骆世明.1999.不同生育期遮光对水稻生长发育和产量形成的影响[J].应用生态学报,10(2)：193-196.
4.曾韶西,王以柔,刘鸿先.1991.低温光照下与黄瓜子叶叶绿素低有关的酶促反应[J].植物生理学报,17(2)：177-182.
5.曾勇军,石庆华,潘晓华,等.2008.施氮量对高产早稻氮素利用特征及产量形成的影响[J].作物学报,34(8)：1409-1416.
6.柴小清,印莉萍,刘祥林,等.1996.不同浓度的N03-和NH4+对小麦根谷氨酰胺合成酶及其相关酶的影响[J].植物学报,38(10)：803-808.
7.陈二影.2012.花后弱光对小麦产量和蛋白质品质的影响及氮素调控研究[D].山东农业大学
8.陈根云,缪有刚,李立人.1993.光合蛋白合成抑制剂对水稻RubisCo大、小亚基和RobisCo亚基结合蛋白基因表达的影响[J].植物生理学报,19(3)：243-249
9.陈温福,徐正进.2007.水稻超高产育种理论与方法[M].北京：科学出版社
10.陈以峰.1997.耐阴水稻种质资源的快速筛选技术研究[J].生态农业研究,5(1)：37-41.
11.陈裕,林坤瑞.1996.金线莲生长发育与光照强度关系[J].福建热作科技,(4)：22-23.
12.程方民,张篙午.1999.水稻籽粒灌浆过程中稻米品质动态变化及温度影响效应[J].浙江人学学报,25(4)：347-350
13.程方民,朱碧岩.1998.气象生态因子对稻米品质影响的研究进展[J].中国农业气象,10(5)：39-45.
14.崔秀明,王朝梁,刘丹.1993.三七荫棚透光度初步研究[J].中药材,16(3)：3-6.
15.邓飞,王丽,姚雄,等.2009.不同生育阶段遮阴对水稻籽粒充实和产量的影响[J].四川农业大学学报,27(3)：265-269.
16.董明辉,惠锋,顾俊荣,等.2013.灌浆期不同光强对水稻不同粒位籽粒品质的影响[J].中国生态农业学报,21(2)：164-170.
17.杜春丽,沈新勇,陈渭民,等.2008.43年来我国城市气候和太阳辐射的变化特征[J].南京气象学院学报,(2)：200-207.
18.杜彦修,徐国强,季新,等.2012.扬花灌浆期抗耐弱光水稻品种评价及筛选[J].中国稻米,18(6)：35-39.
19.段若溪,姜会飞.2002.农业气象学[M].北京：气象出版社
20.冯仕喜,文云书.2006.水稻施用生长调节剂对产量和结实率的影响[J].耕作与栽培,(4)：29.
21.符冠富,李华,陶龙兴,等.灌浆期遮光对水稻籽粒生长和Q酶活性的影响[J].生态学杂志,2009,28(3)：438-444.
22.傅春霞,顾形影,聂毓琦,等.1989.光和温度逆境对水稻光合抑制的影响[J].江苏农业学报,5(1)：1-10
23.傅春霞,谭秀云,邵红宁,等.1995.广幅光强适应种质的筛选[J].江苏农业学报,11(2)：16-21.
24.高士杰,陈温福,徐正进,等.2000.直立穗型水稻的研究Ⅱ.外界环境对直立穗型水稻品种粒重的影响[J].吉林农业科学,(4)：3-6.
25.高廷东.2003.光和不同形态氮互作对小麦幼苗碳、氮同化的影响[D].山东农业大学
26.葛体达,隋方功,白莉萍,等.2005.水分胁迫下夏玉米根叶保护酶活性变化及其对膜脂过氧化作用的影响[J].中国农业科学,(5)：922-928.
27.关义新,林葆,凌碧莹.2000.光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J].作物学报,(06)：806-812.
28.郭翠花,高志强,苗果园.2010.花后遮阴对小麦旗叶光合特性及籽粒产量和品质的影响[J].作物学报,36(4)：673-679.
29.郭凤鸣.1990.弱光条件下黄瓜的生长解析[J].吉林农业大学学报,12(1)：32-35.
30.何康,刘瑞龙,卢良恕,等编著.1996.中国农业百科全书(农业化学卷)[M].北京出版社
31.侯守贵,隋国民,马兴全,等.2012.辽宁省水稻产业发展现状及展望[J].北方水稻,42(5)：70-73.
32.胡文海,师恺,曹玉林,等.2008.低温弱光对黄瓜幼苗地上部和根系生长与呼吸作用的 影响[J].江西农业大学学报,30(1)：16-19
33.黄发松,孙宗修,胡培松,等.1998.食用稻米品质形成研究的现状与展望[J].中国水稻科学,(3)：172-176.
34.黄俊,郭世荣,蒋芳玲,等.2008遮荫处理及恢复光照对白菜生长及活性氧代谢的影响[J].园艺学报,(5)：753-756.
35.黄俊,郭世荣,吴震,等.2007.弱光对不结球白菜光合特性与叶绿体超微结构的影响[J].应用生态学报,(2)：352-358.
36.黄勤妮,印莉萍,柴小清,等.1995.不同氮源对小麦幼苗谷氨酰胺合成酶的影响[J].植物学报,37(11)：856-862.
37.黄正来,韦朝领,刘敏华,等.1999.水稻抽穗灌浆期模拟阴天的光质环境分析及其对产量形成的影响[J].安徽农业大学学报,26(4)：403-409.
38.纪瑞鹏,张玉书,冯锐,等.2007.辽宁省农业气候资源变化特征分析[J].资源科学,2：74-82.
39.贾志宽,高如嵩,张嵩午.1992.稻米垩白形成的气象生态基础研究[J].应用生态学报,3(4)：321-326.
40.江苏省杂交水稻气象问题研究协作组.1983.温光条件与水稻叶片生长和干物质增长的关系[J].作物学报,9(2)：99-106.
41.焦德茂,戴秋杰,聂毓琦.1985.在水稻F4代分离群体中筛选鉴定高光合耐阴的变异体[J].江苏农业学报,1(3)：10-16.
42.焦德茂,高亮之,金之庆,等.1995.水稻耐光氧化和耐阴特性的鉴定及其生理基础[J].中国水稻科学,9(4)：245-248.
43.金丽晨,耿志明,李金州,等.2011.稻米淀粉组成及分子结构与食味品质的关系[J].江苏农业学报,27(1)：13-18.
44.巨关升,武菊英,赵军锋,等.2005.观赏狼尾草光合特性的研究[J].核农学报,19(6)：451-455.
45.孔海江,吴胜安,王蕊,等.2011.2009年秋季河南一次连阴雨天气成因分析[J].气象与环境学报,27(2)：39-44.
46.匡廷云.1979.叶绿体膜的结构与功能Ⅰ组成与PSⅡ功能的关系[J].作物生理学报,(2)：99-107.
47.李合生.2006.现代植物生理学[M].北京：高等教育出版社
48.李林,张更生,嵇福建,等.1990.水稻分蘖期的阴害研究[J].中国农业气象,11(3)：12-17.
49.李林,姚永康,刘敏华,等.1998.开花灌浆期模拟阴天水稻群体内光质环境的调控研究[J].中国农业气象,19()2：1-4
50.李林,张更生,姚永康.1994.水稻开花灌浆期间阴害形成的光谱辐射能特征初步研究[J].中国水稻科学,8(2)：91-96.
51.李林,张更生.1994.阴害影响水稻产量的机制及其调控技术Ⅱ灌浆期模拟阴害影响水稻产量的机制[J].中国农业气象,15(3)：5-9.
52.李林,张更生.1991.阴雨害对江苏水稻生产的影响及其防御对策[J].江苏农业科学,(5)：17-19.
53.李太贵,沈波,陈能,等.1997.Q酶在水稻籽粒至白形成中作用的研究[J].作物学报,5,338-344.
54.李天,大杉立,山岸,等.2005.灌浆结实期弱光对水稻籽粒淀粉积累及相关酶活性的影响[J].中国水稻科学,19(6)：545-550.
55.李天,大杉立,山岸,等.2006.灌浆结实期弱光对水稻籽粒蔗糖及其降解酶活性的影响[J].作物学报,32(6)：943-945.
56.李天.2005.温光对水稻籽粒碳水化合物代谢及品质的影响[D].四川农业大学
57.李霞,刘友良,焦德茂,等.2002.不同高产水稻品种叶片的荧光参数的日变化和光适应特性的关系[J].作物学报,28(2)：145-153.
58.李霞,严建民,季本华,等.1999.光氧化和遮荫条件下水稻的光合生理特性的品种差异[J].作物学报,25(3)：301-308.
59.李想.2003.番茄株型与耐弱光性研究[D].沈阳：沈阳农业大学
60.李晓光,金正勋,刘海英,等.2011.水稻杂种后代籽粒直链淀粉和蛋白质含量选择对产量和品质性状的影响[J].东北农业大学学报,42(4)：13-17.
61.李长缨,朱其杰.1997.光强对黄瓜光合特性及亚适温下生长的影响[J].园艺学报,(1)：98-100.
62.梁建生,曹显祖,徐生,等.1994.水稻籽粒库强与其淀粉积累之间关系的研究[J].作物学报,(6)：685-691.
63.林洪鑫,肖运萍,袁展汽,等.2011.水稻合理密植及其优质高产机理研究进展[J.].中国农学通报,27(9)：1-4.
64.林振武,孙惠珍,陈敬祥.1985.硝酸还原酶活力的体外测定[J].植物生理学通讯,(3)：33-35.
65.刘博,韩勇,谢文孝,等.2008.灌浆结实期弱光对水稻产量、生理及品质的影响[J].中国稻米,(5)：36-40.
66.刘奇华,蔡建,李天,等.2007.水稻籽粒灌浆特性及品质对孕穗期弱光胁迫的响应[J].江西农业大学学报,29(2)：172-175.
67.刘奇华,李天,张建军.2006.生育前期遮光对水稻后期功能叶生长及稻米品质的影响[J].生态学杂志,25(10)：1167-1172.
68.刘奇华,周学标,杨连群,等.2009.生育前期遮光对水稻灌浆期剑叶生理特性及籽粒生长的影响[J].应用生态学报,20(9)：2135-2141.
69.刘永华,朱祝军,魏国强.2004.不同光强下氮素形态对番茄谷氨酰胺合成酶和光呼吸的影响[J].植物生理学通讯,(6)：680-682.
7.吕军,王伯伦,孟维韧,等.2007.不同穗型粳稻的光合作用与物质生产特性[J].中国农业科学,(5)：902-908.
71.吕宁,刘荣高,刘纪远.2009,1998-2002年中国地表太阳辐射的时空变化分析[J].地球信息科学学报,(5)：5623-5630.
7.孟军,陈温福.2005.不同穗型水稻群体垂直入射光分布及其对光合作用的影响[J].沈阳农业大学学报,(1)：9-13.
73.孟雷,陈温福,李磊鑫,等.2002.减弱光照强度对水稻叶片气孔性状的影响[J].沈阳农业大学学报,33(2)：87-89.
74.孟亚利,高如嵩,张嵩午.1994.影响稻米品质的主要气候生态因子研究[J].西北农业大学学报,22(1)：40-45.
75.牟会荣,姜东,戴廷波,等.2008.遮荫对小麦旗叶光合及叶绿素荧光特性的影响[J].中国农业科学,2：599-606.
76.彭建宗,陈兆平.1999.遮阴对多年生花生生长的影响[J].华南师范大学学报,21(2)：94-96.
77.秦建权,唐启源,李迪秦,等.2010.抽穗后光照强度对超级杂交稻干物质生产及氮素吸收与分配的影响[J].四川农业大学学报,(1)：28-34.
78.任久江.1991.气候波动对水稻产量的影响[J].西南农业大学学报,(3)：43-47.
79.任万军,杨文钰,徐精文,等.2003.弱光对水稻籽粒生长及品质的影响[J].作物学报,29(5)：785-790
80.任万军,杨文钰,徐精文,等.2002.始穗后弱光对不同基因型水稻叶片特性的影响[J].四川农业大学学报,3-205-208,278.
81.任万军,杨文钰,张国珍,等.2003.弱光对杂交稻氮素积累、分配与子粒蛋白质含量的影响[J].植物营养与肥料学报,9(3)：288-293.
82.任万军.2001.始穗后弱光对水稻籽粒生长及稻米品质的影响[D].雅安：四川农业大学.
83.邵红宁,傅春霞,曹显祖,等.1998.水稻叶片光氧化敏感性与活性氧清除系统的关系[J].作物学报,24(5)：577-583.
84.申彦波,赵宗慈,石广玉.2008.地面太阳辐射的变化、影响因子及其可能的气候效应最新研究进展[J].地球科学进展,(9)：915-923.
86.石广玉.2007.大气辐射学[M].北京：科学出版社
87.松岛省三.1978.水稻栽培新技术[M].肖连成,译.长春：吉林人民出版社
88.宋桂云,徐正进,王云,等.2005.氮素水平对不同穗型水稻品种光合作用、生产特性的影响[J].作物杂志,(2)：23-25.
89.苏丽英,吴勇,於新建,等.1989.水稻叶片蔗糖磷酸合成酶的一些特性[J].植物生理学报,(2)：117-123.
90.孙金华,贺逸秋,宋鸿遇.1985.酰胺态氮瞬间调节荚膜红假单孢菌光合固氮活性的GS传感机制[J].植物生理学报,(3)：268-278.
91.孙艳,徐伟君,范爱丽.2006.高温强光下水杨酸对黄瓜叶片叶绿素荧光和叶黄素循环的影响[J].应用生态学报,17(3)：399-402.
92.孙艳,徐伟君,范爱丽.2006.高温强光下水杨酸对黄瓜叶片叶绿素荧光和叶黄素循环的影响[J].应用生态学报,17(3)：399-402.
93.孙园园,孙永健,陈林,等.2012.不同播期和抽穗期弱光胁迫对杂交稻生理性状及产量的影响[J].应用生态学报,23(10)：2737-2744.
94.谭周兹,周光恰.1989.水稻结实期光照强度对米质影响的研究[J].杂交水稻,4(1)：39-43.
95.唐焕伟,曲彦婷,张兴.2011.遮光对郁金香叶片叶绿素含量及叶绿体超微结构的影响[J]. 东北农业大学学报,(4)：77-82.
96.田鹏波,赵凤,白静,等.2010.沈阳东陵区近58年农业气候资源趋势变化[J].安徽农业科学,38(21)：11302-11304,11332.
97.童平,杨世民,马均,等.2008.不同水稻品种在不同光照条件下的光合特性及干物质积累[J].应用生态学报,19(3)：505-511.
98.王丹英,章秀福,邵国胜,等.2008.高土壤肥力环境下不同类型粳稻品种产量对氮肥用量的响应[J].作物学报,34(9)：1623-1628.
99.王丹英,章秀福,邵国胜,等.2008.不同叶色水稻叶片的衰老及对光强的响应[J].中国水稻科学,22(1)：77-81.
100.王国民,丁元树.1989.水稻抽穗和结实期的生态因子研究Ⅰ土壤水分对早稻结实和籽粒品质的影响[J].浙江农业大学学报,(1)：17-23.
101.王丽,邓飞,郑军,等.2012.水稻根系生长对弱光胁迫的响应[J].浙江大学学报(农业与生命科学版),38(6)：700-708.
102.王志敏,王树安,苏宝林.1997.小麦穗粒数的调节Ⅱ开花前遮光对穗碳水化合物代谢和内源激素水平的影响[J].华北农学报,12(4)：42-47.
103.王忠.2000.植物生理学[M].中国农业出版社,130-131
104.韦朝领,刘敏华,李林,等.2001.化学调控剂对减轻水稻抽穗灌浆期阴害效应的综合评判[J].应用生态学报,12(1)：55-58.
105.魏珉,邢禹贤,王秀峰,等.2002.C02加富对黄瓜叶片显微和亚显微结构的影响[J].园艺学报,29(1)：30-34
106.谢立勇,徐正进,刘宏光.2004.不同穗型水稻生育后期群体光合有效辐射的分布状况[J].吉林农业大学学报,(5)：476-479,487.
107.熊杰,符冠富,宋建,等.2010.植物生长调节物质对灌浆期遮光水稻生长及结实的影响[J].中国稻米,16(6)：9-13.
108.徐彩龙,尹燕枰,蔡瑞国,等.2012.弱光条件下不同穗型小麦品种旗叶光合特性和抗氧化代谢[J].作物学报,38(7)：1295-1306.
109.徐富贤,郑家奎,朱永川,等.2004.川东南高温伏旱区杂交中稻品种库源结构对稻米整精米率与垩白粒率的影响[J].作物学报,(5)：432-437.
110.徐正进,陈温福,黄瑞冬,等.2007.水稻穗型改良的生理与遗传基础研究进展[J].自然科 学进展,(9)：1161-1167.
111.徐正进,陈温福,张龙步.1990.直立和弯曲穗型水稻冠层光分布比较初报[J].辽宁农业科学,(3)：8-12.
112.徐正进,邵国军,韩勇,等.2006.东北三省水稻产量和品质及其与穗部性状关系的初步研究[J].作物学报,32(12)：1878-1883.
113.徐正进,张龙步,陈温福.1988.不同穗型水稻抽穗后物质生产特性比较[J].辽宁农业科学,(4)：38-40.
114.许人全.2002.光合作用效率[M].上海：上海科学技术出版社,164-167.
115.许正刚,史正军,谢良生,等.2009.遮荫处理下两种园林植物叶绿素含量及荧光参数的研究[J].甘肃科技,25(3)：158-160.
116.严建民,翟虎渠,万建民,等.2002.几种重穗型杂交稻的耐阴性差异[J].江苏农业学报,18(4)：193-198.
117.杨东,段留生,谢华安,等.2011.花前光照亏缺对水稻物质积累及生理特性的影响[J].中国生态农业学报,(2)：347-352.
118.杨建昌.2011.水稻根系形态生理与产量,品质形成及养分吸收利用的关系[J].中国农业科学,44(1)：36-46.
119.杨俊霞,郭宝林,鲁韧强,等.2005.遮阴对美国黑蓦生长及光和特性的影响[J].园艺学报,5(2)：108-110.
120.杨丽敏.1999.寒地水稻低直链淀粉材料的筛选及影响因素的研究[J].黑龙江农业科学,(1)：12-14.
121.杨联松,孙明,张培江,等.1998.温度、光照对杂交中粳80优121稻米品质的影响[J].杂交水稻,13(6)：23-28.
122.杨守仁,张龙步,王进民.1984.水稻理想株形育种的理论和方法初论[J].中国农业科学,(3)：6-13.
123.姚允聪,王绍辉,孔云.2007.弱光条件下桃叶片结构及光合特性与叶绿体超微结构变化[J].中国农业科学,04：855-863.
124.义鸣放,Berghoef J.1994.小苍兰生长发育与光照强度关系的研究[J].园艺学报,21(4)：377-350.
125.殷延勃,朱美静,马洪文,等.2002.环境因子对宁夏水稻品质性状的影响[J].宁夏农林科技,(2)：17-19.
126.殷正华.2004.始穗后不同形式的遮光对稻米品质的影响[D].雅安：四川农业大学
127.于振文.2003.作物栽培学各论：北方本[M].北京：中国农业出版社,155-157.
128.袁继超,丁志勇,赵超,等.2005.高海拔地区水稻遮光、剪叶和疏花对米质影响的研究[J].作物学报,11:429-436.
129.袁刘正.2009.玉米耐阴性的QTL分析[D].郑州：河南农业大学
130.岳寿松,余松烈,于振文.1998.小麦旗叶衰老期间光合作用与叶肉细胞超微结构变化[J].沈阳农业大学学报,1：1-5.
131.张春红,李金州,田孟祥,等.2010.不同食昧粳稻品种稻米蛋白质相关性状与食味的关系[J].江苏农业学报,26(6)：1126-1132.
132.张大鹏.1989.水稻叶片气孔的研究Ⅱ不同生态条件下气孔的动态[J].福建农学院学报,18(3)：302-307.
133.张更生,李林.1996.阴害影响水稻产量的机制及其调控技术Ⅳ植物生长调节剂防御水稻阴害的效果[J].中国农业气象,17(1)：1-5.
134.张国发,丁艳锋.2004.温光因子对稻米品质影响的研究进展[J].中国稻米,(1)：11-14.
135.张吉旺,董树亭,王空军,等.2007.大田遮荫对夏玉米光合特性的影响[J].作物学报,2：216-222.
136.张吉旺.2005.光温胁迫对玉米产量和品质及其生理特性的影响[D].山东农业大学
137.张建军.2005.播期与不同生育时期遮光对水稻生理特性及品质的影响[D].四川农业大学
138.张丽丽,张文忠,韩亚东,等.2007.不同穗型水稻群体遮光对品质的影响[J].辽宁农业科学,2：18-21.
139.张文香,王成缓,王伯伦,等.2006.寒冷地区温度!光照对水稻产量及品质的影响[J].吉林农业科学,31(1),19-20.
140.张文忠,徐正进,陈温福,等.2002.直立穗型水稻研究进展[J].沈阳农业大学学报,(6)：471-475.
141.张玉烛,张桂和,朱国奇,等.1995.阴雨对早稻开花及受精结实的影响[J].中国水稻科学,9(3)：173-178.
142.张振贤,郭延奎,艾希珍,等.2003.日光温室光温因子对黄瓜叶绿体超微结构及其功能的影响[J].应用生态学报,(8)：1287-1290.
143.长户一雄.1972.谷粒蛋白质含量研究[J].日作记,(41)：472-479.
144.赵春雨,任国玉,张运福,等.2009.近50年东北地区的气候变化事实检测分析[J].干旱区资源与环境,23(7)：25-30.
145.赵弢,高志奎,徐广辉,等.2006.非调制式荧光仪测定叶绿素荧光参数的研究[J].生物物理学报,22(1)：34-38.
146.赵晓玲.2009.植物中与光敏色素相互作用的因子PIFs[J].植物生理学通讯,45(6)：531-536.
147.甄伟,张福墁.2000.弱光对黄瓜功能叶片光合特性及超微结构的影响[J].园艺学报,27(4)：290-292.
148.郑红星,刘静.2011.东北地区近40年干燥指数变化趋势及其气候敏感性[J].地理研究,30(10)：1765-1774.
149.郑旭川.2009.灌浆结实期高温弱光对水稻籽粒蛋白质代谢关键酶活性及氨基酸组分的影响[D].四川农业大学
150.郑志广.2003.光温条件对水稻结实及干物质生产的影响[J].北京农学院学报,(1)：13-16.
151.朱萍,杨世民,马均,等.2008.遮光对杂交水稻组合生育后期光合特性和产量的影响[J].作物学报,34(11)：2003-2009.
152.朱萍.2009.孕穗期和抽穗开花期弱光胁迫对杂交水稻生理特性的影响[D].雅安：四川农业大学
153.朱庆森,杜永,王志琴,等.2001.杂交稻米的直链淀粉含量与米饭口感粘度硬度关系的研究[J].作物学报,27(3)：377-382.
154.朱旭东,何洪林,刘敏,等.2010.近50年中国光合有效辐射的时空变化特征[J].地理学报,(3)：270-280.
155.邹琦.1995.植物生理生化实验指导[M].北京：中国农业出版社
156.邹应斌.2006.籼型超级杂交水稻高产栽培研究进展[J].耕作与栽培,(5)：1-5.
157. Akita K, Tanaka N.1990. Effects of limited space on foliage development in rice plants [J]. Science Reports of Faculty of Agriculture, Kobe University,20(1):71-78.
158. Awang Y B, Atherton J G, Taylor A J.1995.Growth and fruiting responses of strawberry plant grown on rockwool to shading and salinity[J].Scientia Hort,62(1):25-31.
159. Baig M J, Anand A, Mandal P K, et al.2005.Irradiance influences contents of photosynthetic pigments and proteins in tropical grasses and legumes [J]. Photosynthetica, 43(1):47-53.
160. Century K, Reuber T L, Ratcliffe O J.2008.Regulating the regulators:the future prospects for transcription-factor-based agricultural biotechnology products [J].Plant physiology,147(1):20-29.
161. Chance B, Maehly A C.1955.Assay of catalases and peroxidases [M]. Methods in Enzymology.
162. Chang X, Alderson P G, Wright C J.2008.Solar irradiance level alters the growth of basil (Ocimum basilicum L.) and its content of volatile oils [J].Environmental and Experimental Botany,63(1):216-223.
163. Chaturvedi G S, Ingram K T.1989.Growth and yield of lowland rice in response to shade and drainage[J].Philippine Journal of Crop Science,14(2):61-67.
164. Dela Torre W R, Burkey K O.1990.Acclimation of barley to change in light intensity: chlorophyll organization [J]. Photosynth Res,24:117-125.
165. Douglas C D,Tsung M K, Ferderiek C F.1988.Enzymes of sucrose and hexose metabolism in developing kernels of two inbreds of maize[J].Plant Physiology, 86:1013-1019.
166. Fischer R A, Stockman Y M.1980. Kernel number per spike in wheat (Triticum aestivum L.):Respones to preanthesis shading. Aust J Plant Physiol,7:169-180.
167. Fisher R A, Rees D, Sayre K D et al.1998. Wheat yield progress associated with higher stomatal conductance and pHotosynthetic rate, and cooler canopiesog [J]. Crop sci, 38:1467-1475
168. Galstyan A, Martinez-Garcia J F.2010.Light signalling in plant developmental regulation [J].Plant Developmental Biology,3:255-274.
169. Garg A K, Sawers R J H, Wang H, et al.2006.Light-regulated overexpression of an Arabidopsis phytochrome A gene in rice alters plant architecture and increases grain yield[J].Planta,223(4):627-636.
170. Guenni O, Seiter S, Figueroa R.2008.Growth responses of three Brachiaria species to light intensity and nitrogen supply [J].Tropical Grasslands,42(2):75-87.
171. Lichtenthaler H K, Buschmann C, Doll M, et al.1981.Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves [J]. Photosynthesis Research,2(2):115-141.
172. Hashemi D A, Herbert S J.1992. Intensifying plant density response of corn with artificial shade [J]. Agron J,84:547-551.
173. Hashemi Dezfouli A, Herbert S J.1992.Effect of leaf orientation and density on yield of corn[J].Iran Agric. Res,11:89-104.
174. Horton P.1999.Prospects for improvement through the genetic manipulation of photosynthesis:morphological and biochemical aspects of light capture [J].Journa of Experimental Botany,51(3):475-485.
175. Jacovides C P, Timvios F S, Papaioannou G, et al.2004. Ratio of PAR to broadband solar radiation measured in Cyprus [J]. Agricultural and Forest Meteorology,121(3/4): 135-140.
176. Janardhan K V, Murty K S, Dash. N B.1980.Effect of low light during ripening period on grain yield and translocation of assimilates in rice varieties[J].Indian Journal of Plant pHysiology, ⅩⅩⅢ (2):163-168.
177. Jiao Y, Lau O S, Deng X W.2007.Light-regulated transcriptional networks in higher plants[J].Nature Reviews Genetics,8(3):217-230.
178. Jun H, Amane M, Yasuko K, et al.1992.Changes in the levels of chlorophyll and light-harvesting chlorophyll a/b protein of PSⅡ in rice leaves aged under different irradiances from full expansion through Senescenee [J].Plant Cell Physiol,33(8): 1209-1214.
179. Keeling P L, Wood J R, Tyson R H, et al.1988.Starch biosynthesis in developing wheat grain [J]. Plant Physiol,87:311-319.
180. Kennedy D.2003. Break through of the Year. Science.302:2033.
181. Kim K S, Kim S K, Huh B L, et al.1991.Effects of shading at heading stage on yield components in rice[J]. K o. J. Crop Sci,36(2):127-133.
182. Kobata T, Sugawara S,Takatu S.2000.Shading during the early grain filling period does not affect potential grain dry matter increase in rice[J].Agron,92:411-417.
183. Kong S G, Lee D S, Kwak S N, et al.2004.Characterization of sunlight-grown transgenic rice plants expressing Arabidopsis pHytochrome A[J].Molecular Breeding, 14(1):35-45.
184. Kumari B P, Londhe A L, Danie S, et al.2007. Observational evidence of solar dimming: Offsetting surface warming over India[J]. GeopHysical Research Letters,34, L21810, do:i 10.1029/2007GL031133.
185. Lea P J and Mifiin B J.1974.Altenrative route for nitrogen assimilation in higher plants.Nature,251:614-616.
186. Lin C C, Kao C H.1996.Disturbed ammonium associated with growth inhibition of roots assimilation is in rice seedlings caused by NaCI[J]. Plant Growth Regulation,18: 233-238.
187. Mawaki M, Morita S, Suga T, et al.1990.Effect of shading on root system morphology and grain yield of rice plants(Oryza sativa L.). I. An analysis on root length density. Japanese Journal of Crop Science,59:89-94.
188. Murty K S, Dey S K, Swain P, et al.1992. Low light adapted restorers of different maturity durations for hybrid rice breeding[J]. Int Rice Res Newsletter.17(6):6-7.
189. Oaks A, Stulen I, Jones K.1980. Enzymes of nitrogen assimilation in maize roots [J]. Planta,148:477-484.
190. Osaki M, Tyoda M, Yamada S, et al.1995.Effect of mutual shading on carbon distribution in rice plant [J]. Soil Sci Plant Nutr,41(2):235-244.
191. Philippe M, Claudio P, Matthew P.2003.Limitations to pHotosynthesis under light and heat stress in three high-yielding wheat genotypes [J]. J Plant Physiol,160:657-666.
192. Pinker R T, Zhang B, Dutton E G.2005.Do satellites detect trends in surface solar radiation?[J]. Science,308:850-854.
193. Power H C, Mills D M.2005.Solar radiation climate change over South Africa and an assessment of the radiative impact of volcanic eruptions [J]. International Journal of Climatology,25:295-318.
194. Ruberti I, Sessa G, Ciolfi A, et al.2012.Plant adaptation to dynamically changing environment:the shade avoidance response [J].Biotechnology Advances, 30(5):1047-1058.
195. Saha A and Dasgupta D K.1989.Effect of low light on yield stability of elite rice cultivars[J]. Indian J Plant Physiol.32(4):372-373.
196. Sasidharan R, Chinnappa C C, Voesenek L A C J, et al.2008.The regulation of cell wall extensibility during shade avoidance:a study using two contrasting ecotypes of Stellaria longipes[J].Plant pHysiology,148(3):1557-1569.
197. Sawers R J H, Sheehan M J, Brutnell T P.2005.Cereal phytochromes:targets of selection, targets for manipulation?[J].Trends in Plant Science,10(3):138-143.
198. Seemarm J R.1989.Light adaptation acclimation of photosynthesis and there gelation of ribulos-1,5-bisphosphate carboxylase activity in sun and shade plants[J].Plant Physiol, 91(5):379-386.
199. Shingnori O, Amane M, Tadahiko M.2003.Effect of irradiance on the partitioning of assimilated carbon during the early phase of grain filling in rice[J].Annals of Botany, 92:357-364.
200. Singh R D and Srivastava H S.1986.Increase in glutamate synthase (NADH) activity in maize seedings in response to nitrate and ammonium nitrogen [J]. Physiologia Plantarum, 66 (3):413-416.
201. Singh S.1994.Physiological response of different crop species to low light stress[J]. Indian J Plant Physiol,37(3):147-151.
202. Stamm P and Kumar P P.2010.The phytohormone signal network regulating elongation growth during shade avoidance [J].Journal of Experimental Botany,61(11):2889-2903.
203. Stuefer J F and Heidrun H.1998.Differential effects of light quantity and spectral light quality on growth, morpHology and development of two stoloniferous Potentilla species [J].Oecologia,117(1):1-8.
204. Tanaka A.1958.Studies on the charaeteristics of the physiological function of leaf of definite position on stem of the rice plant [J].Science Soil Manure,29:327-331.
205. Thangaraj M and Sivasubramanian V.1990.Effects of low light intensity on growth and productivity of irrigated rice (Oryza sativa L.). Madras Agricultural Journal,77:220-224.
206. Tsai Y Z and Lai K L.1990.The effects of temperature and light intensity on tiller development of rice [J]. Memoirs of the College of Agriculture, National Taiwan University,30(2):22-30.
207. Tsai-Mei O L and Setter T L.1985.Enzyme activities of starch and sucrose pathways and growth of apical and based maize kernels[J]. Plant Physiology,79:848-851.
208. Tsuneo Kato.1986.Effects of the shading and rachis branch clipping on the grain filling proeess of rice cultivars differing in the grain [J].Japan Jour Crop Sci,55(2):252-260.
209. Udo S O and Aro T O.1999.Global PAR related to global solar radiation for central Nigeria [J]. Agricultural and Forest Meteorology,97(1):21-31.
210. Vityakon P, Sae Lee S, Seripong S.1993.Effects of tree leaf litter and shading on growth and yield of paddy rice in northeast Thailand [J]. Kasetsart J Nat Sci,27(2):219-222.
211. Wang Z, Yin Y, He M, et al.2003. Allocation of pHotosynthates and grain growth of two wheat cultivars with different potential grain growth in response to pre- and post-anthesis shading[J]. Crop Sci,189:280-285.
212. Wardlaw I F.1994.The early stages of grain development in wheat:response to water stress in a single variety[J].Aust J Biol Sci,44:1047-1055.
213. Wherley B G, Gardner D S, Metzger J D.2005.Tall fescue photomorphogenesis as influenced by changes in the spectral composition and light intensity [J].Crop Science, 45(2):562-568.
214. Winston Raul De la Torre and Kent Oliver Burkey.1990.Acclimation of barley to changes in light intensity:chlorophyll organization [J].Photosynthesis Research, 24(2):117-125.
215. Awang Y B and Atherton J G.1995.Growth and fruiting responses of strawberry plants grown on rockwool to shading and salinity [J]. Scientia Horticulturae,62(1-2):25-31.
216. Yamazaki J.2010.Is light quality involved in the regulation of the photosynthetic apparatus in attached rice leaves? [J].Photosynthesis Research,105(1):63-67.
217. Yang J, Lin R, Sullivan J, et al.2005.Light regulates COP1-mediated degradation of HFR1, a transcription factor essential for light signaling in Arabidopsis [J].The Plant Cell, 17(3):804-821.
218. Zhang C F, Peng S B, Peng X X, et al.1997.Response of glutamine synthetase isoforms to nitrogen sources in rice (Oryza sativa L.) roots [J]. PIant Sci,125:163-170.
2.本庄一雄.1971.稻米蛋白质含量的研究Ⅰ不同施肥条件对糙米蛋白质含有率和蛋白质总量的影响[J].日本作物学会纪事,40：190-197.
3.蔡昆争,骆世明.1999.不同生育期遮光对水稻生长发育和产量形成的影响[J].应用生态学报,10(2)：193-196.
4.曾韶西,王以柔,刘鸿先.1991.低温光照下与黄瓜子叶叶绿素低有关的酶促反应[J].植物生理学报,17(2)：177-182.
5.曾勇军,石庆华,潘晓华,等.2008.施氮量对高产早稻氮素利用特征及产量形成的影响[J].作物学报,34(8)：1409-1416.
6.柴小清,印莉萍,刘祥林,等.1996.不同浓度的N03-和NH4+对小麦根谷氨酰胺合成酶及其相关酶的影响[J].植物学报,38(10)：803-808.
7.陈二影.2012.花后弱光对小麦产量和蛋白质品质的影响及氮素调控研究[D].山东农业大学
8.陈根云,缪有刚,李立人.1993.光合蛋白合成抑制剂对水稻RubisCo大、小亚基和RobisCo亚基结合蛋白基因表达的影响[J].植物生理学报,19(3)：243-249
9.陈温福,徐正进.2007.水稻超高产育种理论与方法[M].北京：科学出版社
10.陈以峰.1997.耐阴水稻种质资源的快速筛选技术研究[J].生态农业研究,5(1)：37-41.
11.陈裕,林坤瑞.1996.金线莲生长发育与光照强度关系[J].福建热作科技,(4)：22-23.
12.程方民,张篙午.1999.水稻籽粒灌浆过程中稻米品质动态变化及温度影响效应[J].浙江人学学报,25(4)：347-350
13.程方民,朱碧岩.1998.气象生态因子对稻米品质影响的研究进展[J].中国农业气象,10(5)：39-45.
14.崔秀明,王朝梁,刘丹.1993.三七荫棚透光度初步研究[J].中药材,16(3)：3-6.
15.邓飞,王丽,姚雄,等.2009.不同生育阶段遮阴对水稻籽粒充实和产量的影响[J].四川农业大学学报,27(3)：265-269.
16.董明辉,惠锋,顾俊荣,等.2013.灌浆期不同光强对水稻不同粒位籽粒品质的影响[J].中国生态农业学报,21(2)：164-170.
17.杜春丽,沈新勇,陈渭民,等.2008.43年来我国城市气候和太阳辐射的变化特征[J].南京气象学院学报,(2)：200-207.
18.杜彦修,徐国强,季新,等.2012.扬花灌浆期抗耐弱光水稻品种评价及筛选[J].中国稻米,18(6)：35-39.
19.段若溪,姜会飞.2002.农业气象学[M].北京：气象出版社
20.冯仕喜,文云书.2006.水稻施用生长调节剂对产量和结实率的影响[J].耕作与栽培,(4)：29.
21.符冠富,李华,陶龙兴,等.灌浆期遮光对水稻籽粒生长和Q酶活性的影响[J].生态学杂志,2009,28(3)：438-444.
22.傅春霞,顾形影,聂毓琦,等.1989.光和温度逆境对水稻光合抑制的影响[J].江苏农业学报,5(1)：1-10
23.傅春霞,谭秀云,邵红宁,等.1995.广幅光强适应种质的筛选[J].江苏农业学报,11(2)：16-21.
24.高士杰,陈温福,徐正进,等.2000.直立穗型水稻的研究Ⅱ.外界环境对直立穗型水稻品种粒重的影响[J].吉林农业科学,(4)：3-6.
25.高廷东.2003.光和不同形态氮互作对小麦幼苗碳、氮同化的影响[D].山东农业大学
26.葛体达,隋方功,白莉萍,等.2005.水分胁迫下夏玉米根叶保护酶活性变化及其对膜脂过氧化作用的影响[J].中国农业科学,(5)：922-928.
27.关义新,林葆,凌碧莹.2000.光、氮及其互作对玉米幼苗叶片光合和碳、氮代谢的影响[J].作物学报,(06)：806-812.
28.郭翠花,高志强,苗果园.2010.花后遮阴对小麦旗叶光合特性及籽粒产量和品质的影响[J].作物学报,36(4)：673-679.
29.郭凤鸣.1990.弱光条件下黄瓜的生长解析[J].吉林农业大学学报,12(1)：32-35.
30.何康,刘瑞龙,卢良恕,等编著.1996.中国农业百科全书(农业化学卷)[M].北京出版社
31.侯守贵,隋国民,马兴全,等.2012.辽宁省水稻产业发展现状及展望[J].北方水稻,42(5)：70-73.
32.胡文海,师恺,曹玉林,等.2008.低温弱光对黄瓜幼苗地上部和根系生长与呼吸作用的 影响[J].江西农业大学学报,30(1)：16-19
33.黄发松,孙宗修,胡培松,等.1998.食用稻米品质形成研究的现状与展望[J].中国水稻科学,(3)：172-176.
34.黄俊,郭世荣,蒋芳玲,等.2008遮荫处理及恢复光照对白菜生长及活性氧代谢的影响[J].园艺学报,(5)：753-756.
35.黄俊,郭世荣,吴震,等.2007.弱光对不结球白菜光合特性与叶绿体超微结构的影响[J].应用生态学报,(2)：352-358.
36.黄勤妮,印莉萍,柴小清,等.1995.不同氮源对小麦幼苗谷氨酰胺合成酶的影响[J].植物学报,37(11)：856-862.
37.黄正来,韦朝领,刘敏华,等.1999.水稻抽穗灌浆期模拟阴天的光质环境分析及其对产量形成的影响[J].安徽农业大学学报,26(4)：403-409.
38.纪瑞鹏,张玉书,冯锐,等.2007.辽宁省农业气候资源变化特征分析[J].资源科学,2：74-82.
39.贾志宽,高如嵩,张嵩午.1992.稻米垩白形成的气象生态基础研究[J].应用生态学报,3(4)：321-326.
40.江苏省杂交水稻气象问题研究协作组.1983.温光条件与水稻叶片生长和干物质增长的关系[J].作物学报,9(2)：99-106.
41.焦德茂,戴秋杰,聂毓琦.1985.在水稻F4代分离群体中筛选鉴定高光合耐阴的变异体[J].江苏农业学报,1(3)：10-16.
42.焦德茂,高亮之,金之庆,等.1995.水稻耐光氧化和耐阴特性的鉴定及其生理基础[J].中国水稻科学,9(4)：245-248.
43.金丽晨,耿志明,李金州,等.2011.稻米淀粉组成及分子结构与食味品质的关系[J].江苏农业学报,27(1)：13-18.
44.巨关升,武菊英,赵军锋,等.2005.观赏狼尾草光合特性的研究[J].核农学报,19(6)：451-455.
45.孔海江,吴胜安,王蕊,等.2011.2009年秋季河南一次连阴雨天气成因分析[J].气象与环境学报,27(2)：39-44.
46.匡廷云.1979.叶绿体膜的结构与功能Ⅰ组成与PSⅡ功能的关系[J].作物生理学报,(2)：99-107.
47.李合生.2006.现代植物生理学[M].北京：高等教育出版社
48.李林,张更生,嵇福建,等.1990.水稻分蘖期的阴害研究[J].中国农业气象,11(3)：12-17.
49.李林,姚永康,刘敏华,等.1998.开花灌浆期模拟阴天水稻群体内光质环境的调控研究[J].中国农业气象,19()2：1-4
50.李林,张更生,姚永康.1994.水稻开花灌浆期间阴害形成的光谱辐射能特征初步研究[J].中国水稻科学,8(2)：91-96.
51.李林,张更生.1994.阴害影响水稻产量的机制及其调控技术Ⅱ灌浆期模拟阴害影响水稻产量的机制[J].中国农业气象,15(3)：5-9.
52.李林,张更生.1991.阴雨害对江苏水稻生产的影响及其防御对策[J].江苏农业科学,(5)：17-19.
53.李太贵,沈波,陈能,等.1997.Q酶在水稻籽粒至白形成中作用的研究[J].作物学报,5,338-344.
54.李天,大杉立,山岸,等.2005.灌浆结实期弱光对水稻籽粒淀粉积累及相关酶活性的影响[J].中国水稻科学,19(6)：545-550.
55.李天,大杉立,山岸,等.2006.灌浆结实期弱光对水稻籽粒蔗糖及其降解酶活性的影响[J].作物学报,32(6)：943-945.
56.李天.2005.温光对水稻籽粒碳水化合物代谢及品质的影响[D].四川农业大学
57.李霞,刘友良,焦德茂,等.2002.不同高产水稻品种叶片的荧光参数的日变化和光适应特性的关系[J].作物学报,28(2)：145-153.
58.李霞,严建民,季本华,等.1999.光氧化和遮荫条件下水稻的光合生理特性的品种差异[J].作物学报,25(3)：301-308.
59.李想.2003.番茄株型与耐弱光性研究[D].沈阳：沈阳农业大学
60.李晓光,金正勋,刘海英,等.2011.水稻杂种后代籽粒直链淀粉和蛋白质含量选择对产量和品质性状的影响[J].东北农业大学学报,42(4)：13-17.
61.李长缨,朱其杰.1997.光强对黄瓜光合特性及亚适温下生长的影响[J].园艺学报,(1)：98-100.
62.梁建生,曹显祖,徐生,等.1994.水稻籽粒库强与其淀粉积累之间关系的研究[J].作物学报,(6)：685-691.
63.林洪鑫,肖运萍,袁展汽,等.2011.水稻合理密植及其优质高产机理研究进展[J.].中国农学通报,27(9)：1-4.
64.林振武,孙惠珍,陈敬祥.1985.硝酸还原酶活力的体外测定[J].植物生理学通讯,(3)：33-35.
65.刘博,韩勇,谢文孝,等.2008.灌浆结实期弱光对水稻产量、生理及品质的影响[J].中国稻米,(5)：36-40.
66.刘奇华,蔡建,李天,等.2007.水稻籽粒灌浆特性及品质对孕穗期弱光胁迫的响应[J].江西农业大学学报,29(2)：172-175.
67.刘奇华,李天,张建军.2006.生育前期遮光对水稻后期功能叶生长及稻米品质的影响[J].生态学杂志,25(10)：1167-1172.
68.刘奇华,周学标,杨连群,等.2009.生育前期遮光对水稻灌浆期剑叶生理特性及籽粒生长的影响[J].应用生态学报,20(9)：2135-2141.
69.刘永华,朱祝军,魏国强.2004.不同光强下氮素形态对番茄谷氨酰胺合成酶和光呼吸的影响[J].植物生理学通讯,(6)：680-682.
7.吕军,王伯伦,孟维韧,等.2007.不同穗型粳稻的光合作用与物质生产特性[J].中国农业科学,(5)：902-908.
71.吕宁,刘荣高,刘纪远.2009,1998-2002年中国地表太阳辐射的时空变化分析[J].地球信息科学学报,(5)：5623-5630.
7.孟军,陈温福.2005.不同穗型水稻群体垂直入射光分布及其对光合作用的影响[J].沈阳农业大学学报,(1)：9-13.
73.孟雷,陈温福,李磊鑫,等.2002.减弱光照强度对水稻叶片气孔性状的影响[J].沈阳农业大学学报,33(2)：87-89.
74.孟亚利,高如嵩,张嵩午.1994.影响稻米品质的主要气候生态因子研究[J].西北农业大学学报,22(1)：40-45.
75.牟会荣,姜东,戴廷波,等.2008.遮荫对小麦旗叶光合及叶绿素荧光特性的影响[J].中国农业科学,2：599-606.
76.彭建宗,陈兆平.1999.遮阴对多年生花生生长的影响[J].华南师范大学学报,21(2)：94-96.
77.秦建权,唐启源,李迪秦,等.2010.抽穗后光照强度对超级杂交稻干物质生产及氮素吸收与分配的影响[J].四川农业大学学报,(1)：28-34.
78.任久江.1991.气候波动对水稻产量的影响[J].西南农业大学学报,(3)：43-47.
79.任万军,杨文钰,徐精文,等.2003.弱光对水稻籽粒生长及品质的影响[J].作物学报,29(5)：785-790
80.任万军,杨文钰,徐精文,等.2002.始穗后弱光对不同基因型水稻叶片特性的影响[J].四川农业大学学报,3-205-208,278.
81.任万军,杨文钰,张国珍,等.2003.弱光对杂交稻氮素积累、分配与子粒蛋白质含量的影响[J].植物营养与肥料学报,9(3)：288-293.
82.任万军.2001.始穗后弱光对水稻籽粒生长及稻米品质的影响[D].雅安：四川农业大学.
83.邵红宁,傅春霞,曹显祖,等.1998.水稻叶片光氧化敏感性与活性氧清除系统的关系[J].作物学报,24(5)：577-583.
84.申彦波,赵宗慈,石广玉.2008.地面太阳辐射的变化、影响因子及其可能的气候效应最新研究进展[J].地球科学进展,(9)：915-923.
86.石广玉.2007.大气辐射学[M].北京：科学出版社
87.松岛省三.1978.水稻栽培新技术[M].肖连成,译.长春：吉林人民出版社
88.宋桂云,徐正进,王云,等.2005.氮素水平对不同穗型水稻品种光合作用、生产特性的影响[J].作物杂志,(2)：23-25.
89.苏丽英,吴勇,於新建,等.1989.水稻叶片蔗糖磷酸合成酶的一些特性[J].植物生理学报,(2)：117-123.
90.孙金华,贺逸秋,宋鸿遇.1985.酰胺态氮瞬间调节荚膜红假单孢菌光合固氮活性的GS传感机制[J].植物生理学报,(3)：268-278.
91.孙艳,徐伟君,范爱丽.2006.高温强光下水杨酸对黄瓜叶片叶绿素荧光和叶黄素循环的影响[J].应用生态学报,17(3)：399-402.
92.孙艳,徐伟君,范爱丽.2006.高温强光下水杨酸对黄瓜叶片叶绿素荧光和叶黄素循环的影响[J].应用生态学报,17(3)：399-402.
93.孙园园,孙永健,陈林,等.2012.不同播期和抽穗期弱光胁迫对杂交稻生理性状及产量的影响[J].应用生态学报,23(10)：2737-2744.
94.谭周兹,周光恰.1989.水稻结实期光照强度对米质影响的研究[J].杂交水稻,4(1)：39-43.
95.唐焕伟,曲彦婷,张兴.2011.遮光对郁金香叶片叶绿素含量及叶绿体超微结构的影响[J]. 东北农业大学学报,(4)：77-82.
96.田鹏波,赵凤,白静,等.2010.沈阳东陵区近58年农业气候资源趋势变化[J].安徽农业科学,38(21)：11302-11304,11332.
97.童平,杨世民,马均,等.2008.不同水稻品种在不同光照条件下的光合特性及干物质积累[J].应用生态学报,19(3)：505-511.
98.王丹英,章秀福,邵国胜,等.2008.高土壤肥力环境下不同类型粳稻品种产量对氮肥用量的响应[J].作物学报,34(9)：1623-1628.
99.王丹英,章秀福,邵国胜,等.2008.不同叶色水稻叶片的衰老及对光强的响应[J].中国水稻科学,22(1)：77-81.
100.王国民,丁元树.1989.水稻抽穗和结实期的生态因子研究Ⅰ土壤水分对早稻结实和籽粒品质的影响[J].浙江农业大学学报,(1)：17-23.
101.王丽,邓飞,郑军,等.2012.水稻根系生长对弱光胁迫的响应[J].浙江大学学报(农业与生命科学版),38(6)：700-708.
102.王志敏,王树安,苏宝林.1997.小麦穗粒数的调节Ⅱ开花前遮光对穗碳水化合物代谢和内源激素水平的影响[J].华北农学报,12(4)：42-47.
103.王忠.2000.植物生理学[M].中国农业出版社,130-131
104.韦朝领,刘敏华,李林,等.2001.化学调控剂对减轻水稻抽穗灌浆期阴害效应的综合评判[J].应用生态学报,12(1)：55-58.
105.魏珉,邢禹贤,王秀峰,等.2002.C02加富对黄瓜叶片显微和亚显微结构的影响[J].园艺学报,29(1)：30-34
106.谢立勇,徐正进,刘宏光.2004.不同穗型水稻生育后期群体光合有效辐射的分布状况[J].吉林农业大学学报,(5)：476-479,487.
107.熊杰,符冠富,宋建,等.2010.植物生长调节物质对灌浆期遮光水稻生长及结实的影响[J].中国稻米,16(6)：9-13.
108.徐彩龙,尹燕枰,蔡瑞国,等.2012.弱光条件下不同穗型小麦品种旗叶光合特性和抗氧化代谢[J].作物学报,38(7)：1295-1306.
109.徐富贤,郑家奎,朱永川,等.2004.川东南高温伏旱区杂交中稻品种库源结构对稻米整精米率与垩白粒率的影响[J].作物学报,(5)：432-437.
110.徐正进,陈温福,黄瑞冬,等.2007.水稻穗型改良的生理与遗传基础研究进展[J].自然科 学进展,(9)：1161-1167.
111.徐正进,陈温福,张龙步.1990.直立和弯曲穗型水稻冠层光分布比较初报[J].辽宁农业科学,(3)：8-12.
112.徐正进,邵国军,韩勇,等.2006.东北三省水稻产量和品质及其与穗部性状关系的初步研究[J].作物学报,32(12)：1878-1883.
113.徐正进,张龙步,陈温福.1988.不同穗型水稻抽穗后物质生产特性比较[J].辽宁农业科学,(4)：38-40.
114.许人全.2002.光合作用效率[M].上海：上海科学技术出版社,164-167.
115.许正刚,史正军,谢良生,等.2009.遮荫处理下两种园林植物叶绿素含量及荧光参数的研究[J].甘肃科技,25(3)：158-160.
116.严建民,翟虎渠,万建民,等.2002.几种重穗型杂交稻的耐阴性差异[J].江苏农业学报,18(4)：193-198.
117.杨东,段留生,谢华安,等.2011.花前光照亏缺对水稻物质积累及生理特性的影响[J].中国生态农业学报,(2)：347-352.
118.杨建昌.2011.水稻根系形态生理与产量,品质形成及养分吸收利用的关系[J].中国农业科学,44(1)：36-46.
119.杨俊霞,郭宝林,鲁韧强,等.2005.遮阴对美国黑蓦生长及光和特性的影响[J].园艺学报,5(2)：108-110.
120.杨丽敏.1999.寒地水稻低直链淀粉材料的筛选及影响因素的研究[J].黑龙江农业科学,(1)：12-14.
121.杨联松,孙明,张培江,等.1998.温度、光照对杂交中粳80优121稻米品质的影响[J].杂交水稻,13(6)：23-28.
122.杨守仁,张龙步,王进民.1984.水稻理想株形育种的理论和方法初论[J].中国农业科学,(3)：6-13.
123.姚允聪,王绍辉,孔云.2007.弱光条件下桃叶片结构及光合特性与叶绿体超微结构变化[J].中国农业科学,04：855-863.
124.义鸣放,Berghoef J.1994.小苍兰生长发育与光照强度关系的研究[J].园艺学报,21(4)：377-350.
125.殷延勃,朱美静,马洪文,等.2002.环境因子对宁夏水稻品质性状的影响[J].宁夏农林科技,(2)：17-19.
126.殷正华.2004.始穗后不同形式的遮光对稻米品质的影响[D].雅安：四川农业大学
127.于振文.2003.作物栽培学各论：北方本[M].北京：中国农业出版社,155-157.
128.袁继超,丁志勇,赵超,等.2005.高海拔地区水稻遮光、剪叶和疏花对米质影响的研究[J].作物学报,11:429-436.
129.袁刘正.2009.玉米耐阴性的QTL分析[D].郑州：河南农业大学
130.岳寿松,余松烈,于振文.1998.小麦旗叶衰老期间光合作用与叶肉细胞超微结构变化[J].沈阳农业大学学报,1：1-5.
131.张春红,李金州,田孟祥,等.2010.不同食昧粳稻品种稻米蛋白质相关性状与食味的关系[J].江苏农业学报,26(6)：1126-1132.
132.张大鹏.1989.水稻叶片气孔的研究Ⅱ不同生态条件下气孔的动态[J].福建农学院学报,18(3)：302-307.
133.张更生,李林.1996.阴害影响水稻产量的机制及其调控技术Ⅳ植物生长调节剂防御水稻阴害的效果[J].中国农业气象,17(1)：1-5.
134.张国发,丁艳锋.2004.温光因子对稻米品质影响的研究进展[J].中国稻米,(1)：11-14.
135.张吉旺,董树亭,王空军,等.2007.大田遮荫对夏玉米光合特性的影响[J].作物学报,2：216-222.
136.张吉旺.2005.光温胁迫对玉米产量和品质及其生理特性的影响[D].山东农业大学
137.张建军.2005.播期与不同生育时期遮光对水稻生理特性及品质的影响[D].四川农业大学
138.张丽丽,张文忠,韩亚东,等.2007.不同穗型水稻群体遮光对品质的影响[J].辽宁农业科学,2：18-21.
139.张文香,王成缓,王伯伦,等.2006.寒冷地区温度!光照对水稻产量及品质的影响[J].吉林农业科学,31(1),19-20.
140.张文忠,徐正进,陈温福,等.2002.直立穗型水稻研究进展[J].沈阳农业大学学报,(6)：471-475.
141.张玉烛,张桂和,朱国奇,等.1995.阴雨对早稻开花及受精结实的影响[J].中国水稻科学,9(3)：173-178.
142.张振贤,郭延奎,艾希珍,等.2003.日光温室光温因子对黄瓜叶绿体超微结构及其功能的影响[J].应用生态学报,(8)：1287-1290.
143.长户一雄.1972.谷粒蛋白质含量研究[J].日作记,(41)：472-479.
144.赵春雨,任国玉,张运福,等.2009.近50年东北地区的气候变化事实检测分析[J].干旱区资源与环境,23(7)：25-30.
145.赵弢,高志奎,徐广辉,等.2006.非调制式荧光仪测定叶绿素荧光参数的研究[J].生物物理学报,22(1)：34-38.
146.赵晓玲.2009.植物中与光敏色素相互作用的因子PIFs[J].植物生理学通讯,45(6)：531-536.
147.甄伟,张福墁.2000.弱光对黄瓜功能叶片光合特性及超微结构的影响[J].园艺学报,27(4)：290-292.
148.郑红星,刘静.2011.东北地区近40年干燥指数变化趋势及其气候敏感性[J].地理研究,30(10)：1765-1774.
149.郑旭川.2009.灌浆结实期高温弱光对水稻籽粒蛋白质代谢关键酶活性及氨基酸组分的影响[D].四川农业大学
150.郑志广.2003.光温条件对水稻结实及干物质生产的影响[J].北京农学院学报,(1)：13-16.
151.朱萍,杨世民,马均,等.2008.遮光对杂交水稻组合生育后期光合特性和产量的影响[J].作物学报,34(11)：2003-2009.
152.朱萍.2009.孕穗期和抽穗开花期弱光胁迫对杂交水稻生理特性的影响[D].雅安：四川农业大学
153.朱庆森,杜永,王志琴,等.2001.杂交稻米的直链淀粉含量与米饭口感粘度硬度关系的研究[J].作物学报,27(3)：377-382.
154.朱旭东,何洪林,刘敏,等.2010.近50年中国光合有效辐射的时空变化特征[J].地理学报,(3)：270-280.
155.邹琦.1995.植物生理生化实验指导[M].北京：中国农业出版社
156.邹应斌.2006.籼型超级杂交水稻高产栽培研究进展[J].耕作与栽培,(5)：1-5.
157. Akita K, Tanaka N.1990. Effects of limited space on foliage development in rice plants [J]. Science Reports of Faculty of Agriculture, Kobe University,20(1):71-78.
158. Awang Y B, Atherton J G, Taylor A J.1995.Growth and fruiting responses of strawberry plant grown on rockwool to shading and salinity[J].Scientia Hort,62(1):25-31.
159. Baig M J, Anand A, Mandal P K, et al.2005.Irradiance influences contents of photosynthetic pigments and proteins in tropical grasses and legumes [J]. Photosynthetica, 43(1):47-53.
160. Century K, Reuber T L, Ratcliffe O J.2008.Regulating the regulators:the future prospects for transcription-factor-based agricultural biotechnology products [J].Plant physiology,147(1):20-29.
161. Chance B, Maehly A C.1955.Assay of catalases and peroxidases [M]. Methods in Enzymology.
162. Chang X, Alderson P G, Wright C J.2008.Solar irradiance level alters the growth of basil (Ocimum basilicum L.) and its content of volatile oils [J].Environmental and Experimental Botany,63(1):216-223.
163. Chaturvedi G S, Ingram K T.1989.Growth and yield of lowland rice in response to shade and drainage[J].Philippine Journal of Crop Science,14(2):61-67.
164. Dela Torre W R, Burkey K O.1990.Acclimation of barley to change in light intensity: chlorophyll organization [J]. Photosynth Res,24:117-125.
165. Douglas C D,Tsung M K, Ferderiek C F.1988.Enzymes of sucrose and hexose metabolism in developing kernels of two inbreds of maize[J].Plant Physiology, 86:1013-1019.
166. Fischer R A, Stockman Y M.1980. Kernel number per spike in wheat (Triticum aestivum L.):Respones to preanthesis shading. Aust J Plant Physiol,7:169-180.
167. Fisher R A, Rees D, Sayre K D et al.1998. Wheat yield progress associated with higher stomatal conductance and pHotosynthetic rate, and cooler canopiesog [J]. Crop sci, 38:1467-1475
168. Galstyan A, Martinez-Garcia J F.2010.Light signalling in plant developmental regulation [J].Plant Developmental Biology,3:255-274.
169. Garg A K, Sawers R J H, Wang H, et al.2006.Light-regulated overexpression of an Arabidopsis phytochrome A gene in rice alters plant architecture and increases grain yield[J].Planta,223(4):627-636.
170. Guenni O, Seiter S, Figueroa R.2008.Growth responses of three Brachiaria species to light intensity and nitrogen supply [J].Tropical Grasslands,42(2):75-87.
171. Lichtenthaler H K, Buschmann C, Doll M, et al.1981.Photosynthetic activity, chloroplast ultrastructure, and leaf characteristics of high-light and low-light plants and of sun and shade leaves [J]. Photosynthesis Research,2(2):115-141.
172. Hashemi D A, Herbert S J.1992. Intensifying plant density response of corn with artificial shade [J]. Agron J,84:547-551.
173. Hashemi Dezfouli A, Herbert S J.1992.Effect of leaf orientation and density on yield of corn[J].Iran Agric. Res,11:89-104.
174. Horton P.1999.Prospects for improvement through the genetic manipulation of photosynthesis:morphological and biochemical aspects of light capture [J].Journa of Experimental Botany,51(3):475-485.
175. Jacovides C P, Timvios F S, Papaioannou G, et al.2004. Ratio of PAR to broadband solar radiation measured in Cyprus [J]. Agricultural and Forest Meteorology,121(3/4): 135-140.
176. Janardhan K V, Murty K S, Dash. N B.1980.Effect of low light during ripening period on grain yield and translocation of assimilates in rice varieties[J].Indian Journal of Plant pHysiology, ⅩⅩⅢ (2):163-168.
177. Jiao Y, Lau O S, Deng X W.2007.Light-regulated transcriptional networks in higher plants[J].Nature Reviews Genetics,8(3):217-230.
178. Jun H, Amane M, Yasuko K, et al.1992.Changes in the levels of chlorophyll and light-harvesting chlorophyll a/b protein of PSⅡ in rice leaves aged under different irradiances from full expansion through Senescenee [J].Plant Cell Physiol,33(8): 1209-1214.
179. Keeling P L, Wood J R, Tyson R H, et al.1988.Starch biosynthesis in developing wheat grain [J]. Plant Physiol,87:311-319.
180. Kennedy D.2003. Break through of the Year. Science.302:2033.
181. Kim K S, Kim S K, Huh B L, et al.1991.Effects of shading at heading stage on yield components in rice[J]. K o. J. Crop Sci,36(2):127-133.
182. Kobata T, Sugawara S,Takatu S.2000.Shading during the early grain filling period does not affect potential grain dry matter increase in rice[J].Agron,92:411-417.
183. Kong S G, Lee D S, Kwak S N, et al.2004.Characterization of sunlight-grown transgenic rice plants expressing Arabidopsis pHytochrome A[J].Molecular Breeding, 14(1):35-45.
184. Kumari B P, Londhe A L, Danie S, et al.2007. Observational evidence of solar dimming: Offsetting surface warming over India[J]. GeopHysical Research Letters,34, L21810, do:i 10.1029/2007GL031133.
185. Lea P J and Mifiin B J.1974.Altenrative route for nitrogen assimilation in higher plants.Nature,251:614-616.
186. Lin C C, Kao C H.1996.Disturbed ammonium associated with growth inhibition of roots assimilation is in rice seedlings caused by NaCI[J]. Plant Growth Regulation,18: 233-238.
187. Mawaki M, Morita S, Suga T, et al.1990.Effect of shading on root system morphology and grain yield of rice plants(Oryza sativa L.). I. An analysis on root length density. Japanese Journal of Crop Science,59:89-94.
188. Murty K S, Dey S K, Swain P, et al.1992. Low light adapted restorers of different maturity durations for hybrid rice breeding[J]. Int Rice Res Newsletter.17(6):6-7.
189. Oaks A, Stulen I, Jones K.1980. Enzymes of nitrogen assimilation in maize roots [J]. Planta,148:477-484.
190. Osaki M, Tyoda M, Yamada S, et al.1995.Effect of mutual shading on carbon distribution in rice plant [J]. Soil Sci Plant Nutr,41(2):235-244.
191. Philippe M, Claudio P, Matthew P.2003.Limitations to pHotosynthesis under light and heat stress in three high-yielding wheat genotypes [J]. J Plant Physiol,160:657-666.
192. Pinker R T, Zhang B, Dutton E G.2005.Do satellites detect trends in surface solar radiation?[J]. Science,308:850-854.
193. Power H C, Mills D M.2005.Solar radiation climate change over South Africa and an assessment of the radiative impact of volcanic eruptions [J]. International Journal of Climatology,25:295-318.
194. Ruberti I, Sessa G, Ciolfi A, et al.2012.Plant adaptation to dynamically changing environment:the shade avoidance response [J].Biotechnology Advances, 30(5):1047-1058.
195. Saha A and Dasgupta D K.1989.Effect of low light on yield stability of elite rice cultivars[J]. Indian J Plant Physiol.32(4):372-373.
196. Sasidharan R, Chinnappa C C, Voesenek L A C J, et al.2008.The regulation of cell wall extensibility during shade avoidance:a study using two contrasting ecotypes of Stellaria longipes[J].Plant pHysiology,148(3):1557-1569.
197. Sawers R J H, Sheehan M J, Brutnell T P.2005.Cereal phytochromes:targets of selection, targets for manipulation?[J].Trends in Plant Science,10(3):138-143.
198. Seemarm J R.1989.Light adaptation acclimation of photosynthesis and there gelation of ribulos-1,5-bisphosphate carboxylase activity in sun and shade plants[J].Plant Physiol, 91(5):379-386.
199. Shingnori O, Amane M, Tadahiko M.2003.Effect of irradiance on the partitioning of assimilated carbon during the early phase of grain filling in rice[J].Annals of Botany, 92:357-364.
200. Singh R D and Srivastava H S.1986.Increase in glutamate synthase (NADH) activity in maize seedings in response to nitrate and ammonium nitrogen [J]. Physiologia Plantarum, 66 (3):413-416.
201. Singh S.1994.Physiological response of different crop species to low light stress[J]. Indian J Plant Physiol,37(3):147-151.
202. Stamm P and Kumar P P.2010.The phytohormone signal network regulating elongation growth during shade avoidance [J].Journal of Experimental Botany,61(11):2889-2903.
203. Stuefer J F and Heidrun H.1998.Differential effects of light quantity and spectral light quality on growth, morpHology and development of two stoloniferous Potentilla species [J].Oecologia,117(1):1-8.
204. Tanaka A.1958.Studies on the charaeteristics of the physiological function of leaf of definite position on stem of the rice plant [J].Science Soil Manure,29:327-331.
205. Thangaraj M and Sivasubramanian V.1990.Effects of low light intensity on growth and productivity of irrigated rice (Oryza sativa L.). Madras Agricultural Journal,77:220-224.
206. Tsai Y Z and Lai K L.1990.The effects of temperature and light intensity on tiller development of rice [J]. Memoirs of the College of Agriculture, National Taiwan University,30(2):22-30.
207. Tsai-Mei O L and Setter T L.1985.Enzyme activities of starch and sucrose pathways and growth of apical and based maize kernels[J]. Plant Physiology,79:848-851.
208. Tsuneo Kato.1986.Effects of the shading and rachis branch clipping on the grain filling proeess of rice cultivars differing in the grain [J].Japan Jour Crop Sci,55(2):252-260.
209. Udo S O and Aro T O.1999.Global PAR related to global solar radiation for central Nigeria [J]. Agricultural and Forest Meteorology,97(1):21-31.
210. Vityakon P, Sae Lee S, Seripong S.1993.Effects of tree leaf litter and shading on growth and yield of paddy rice in northeast Thailand [J]. Kasetsart J Nat Sci,27(2):219-222.
211. Wang Z, Yin Y, He M, et al.2003. Allocation of pHotosynthates and grain growth of two wheat cultivars with different potential grain growth in response to pre- and post-anthesis shading[J]. Crop Sci,189:280-285.
212. Wardlaw I F.1994.The early stages of grain development in wheat:response to water stress in a single variety[J].Aust J Biol Sci,44:1047-1055.
213. Wherley B G, Gardner D S, Metzger J D.2005.Tall fescue photomorphogenesis as influenced by changes in the spectral composition and light intensity [J].Crop Science, 45(2):562-568.
214. Winston Raul De la Torre and Kent Oliver Burkey.1990.Acclimation of barley to changes in light intensity:chlorophyll organization [J].Photosynthesis Research, 24(2):117-125.
215. Awang Y B and Atherton J G.1995.Growth and fruiting responses of strawberry plants grown on rockwool to shading and salinity [J]. Scientia Horticulturae,62(1-2):25-31.
216. Yamazaki J.2010.Is light quality involved in the regulation of the photosynthetic apparatus in attached rice leaves? [J].Photosynthesis Research,105(1):63-67.
217. Yang J, Lin R, Sullivan J, et al.2005.Light regulates COP1-mediated degradation of HFR1, a transcription factor essential for light signaling in Arabidopsis [J].The Plant Cell, 17(3):804-821.
218. Zhang C F, Peng S B, Peng X X, et al.1997.Response of glutamine synthetase isoforms to nitrogen sources in rice (Oryza sativa L.) roots [J]. PIant Sci,125:163-170.