水稻源库结构、稻米品质以及对氮响应的基因型差异
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摘要
水稻源库结构与稻米品质的关系是水稻品种选育和栽培必须考虑的两大方面,扩库强源是实现水稻高产的必由之路,而在提高产量的同时如何保证优质,已成为水稻育种和栽培的热点和难点。本研究以27个早稻品种和30个晚稻品种为材料,采用系统聚类法将早、晚稻品种分别分为大库容、中库容和小库容3种类型,从源、氮素吸收与利用、稻米品质等方面分析了不同库容类型品种之间的差异,以及氮素穗肥对早、晚稻不同库容类型品种源库结构和品质的调控效应,并探讨了源库结构与稻米品质的关系,探明与品质性状密切相关的源库指标,以期为早、晚稻品种的遗传改良和调优栽培提供参考依据。主要研究结果如下：
(1)研究了不同早、晚稻品种库容及其构成因子的差异,并对早稻和晚稻品种的库容类型进行了划分。结果表明,供试品种间库容量的差异较大,早稻品种间变异系数达到了40.5%、晚稻达到了35.0%；库容构成因子对库容量的贡献早、晚稻品种表现一致,穗数>每穗粒数>千粒重。聚类结果,27个早稻品种和30个晚稻品种均分为大库容、中库容和小库容3种类型,类型间库容量特征值分别为1211.9、1073.2、921.6gm-2(早稻)和1424.7、1201.8、1007.1gm-2(晚稻)。其结果为下一步开展不同库容类型品种源特性、氮素吸收利用特性和品质形成特性等研究奠定了基础。
(2)研究了早、晚稻不同库容类型品种叶光合、干物质生产与转运、籽粒灌浆和源库比的差异。结果显示,大库容品种叶量、叶光合能力较小库容品种大,具有齐穗期和成熟期叶面积大、叶重高,灌浆期间叶面积消亡慢、净同化率和净光合速率高,顶部3张叶的叶基角小等特性；大库容品种干物质积累和再利用能力强,齐穗期和成熟期干物质积累量显著高于小库容品种,灌浆期茎鞘的干物质转运量、转运率也显著大于小库容品种；库容量对籽粒灌浆影响明显,随品种库容量的增大,灌浆历期明显增长,灌浆速率减小。研究认为大库容品种的叶库比、茎库比和净库比小于小库容品种,这可能是影响籽粒灌浆的主因。
(3)研究了早、晚稻不同库容类型品种氮素积累和转运的差异,以及库容量与氮素吸收利用的关系。结果表明,大库容品种的氮素吸收与转运能力较强,齐穗期和成熟期的氮素积累量显著高于小库容品种,氮素转运量、贡献率也显著大于小库容品种；库容量对氮素利用效率的影响明显,随库容量的增大,早稻品种的氮素干物质生产效率略有减小,籽粒生产效率和氮肥利用率提高或显著提高,而晚稻品种的氮素干物质生产效率、氮素籽粒生产效率、氮肥利用率则显著提高。
(4)研究了早、晚稻不同库容类型品种加工品质、外观品质、蒸煮食味品质和营养品质的差异,以及库容量与稻米品质的关系。结果显示,不同库容类型品种的糙米率、精米率、粒型差异较小,而整精米率、垩白差异较大,早稻整精米率表现为中库容品种>小库容品种>大库容品种,晚稻整精米率表现为小库容品种>中库容品种>大库容品种,垩白粒率、垩白度表现出与整精米率相反的趋势；库容量对稻米理化特性的影响明显,随库容量的增大,胶稠度变短、直链淀粉含量降低,蛋白质及其组分的含量增加。
(5)研究了早、晚稻不同库容类型品种对氮素穗肥响应的差异。结果表明,适量增施氮素穗肥,提高了水稻库容量,增强叶片的光合能力,增加干物质积累量,促进茎鞘中贮存干物质向穗部的转运,并使源库比降低。氮素穗肥对垩白的影响较大,随施氮量增加早稻小库容品种的垩白粒率和垩白度先降后升,而早稻大、中库容品种和晚稻3种库容品种的垩白粒率、垩白度则随氮素穗肥水平提高而增加；氮素穗肥对稻米理化特性的影响显著,随施氮量提高,不同库容类型品种的蛋白质及其组分含量均增加、直链淀粉含量下降、胶稠度变短。源库结构、稻米品质对穗肥施氮量响应的敏感程度与品种库容量有关,表现出大库容品种比小库容品种敏感的特性。在本试验条件下,大库容品种获得较高产量和较优品质的适宜穗肥施氮量为135kg hm-2(早稻)、180kg hm-2(晚稻)；小库容品种获得较高产量和较优品质的适宜穗肥施氮量为90kg hm-2(早稻)、120kg hm-2(晚稻)。
(6)采用组相关分析法探究了早、晚稻源库结构与品质性状之间的关系。结果表明,库容构成因子组、叶光合因子组、物质生产因子组、氮素积累因子组、源库比因子组与品质性状组之间存在显著或极显著的相关,而株型因子组与品质性状组之间的相关不显著。其中,每穗粒数、齐穗期LAI、叶面积消亡率、茎鞘干物质转运量、灌浆期物质生产量、灌浆期吸氮量、叶茎鞘氮素转运量、叶库比、茎库比等性状或指标在各自的因子组中起主导作用,并与垩白粒率、垩白度、蛋白质含量等性状密切关联,进而对稻米品质产生显著影响。
The relationship between source-sink structure and grain quality is one of the most important aspects should be considered in rice breeding and cultivation. Expanding the sink and enhancing the source of rice is the only way to achieve high yield. How to increase yield and keep good grain quality at the same time became one of the hot and difficulty point in rice study. In this study, a field experiment was carried out with27early rice and30late rice cultivars. The cultivars of early rice and late rice were classified into3types respectively based on their sink potential level using the system clustering method. Then we analyzed the differences of source, nitrogen uptake and utilization and rice quality among the large, medium and small sink-potential types of early and late rice, investigated the effects of panicle nitrogen fertilizer on source-sink structure and rice quality. And moreover, we discussed the relationship between source-sink structure and rice quality, in order to provide a reference for the genetic improvement of rice quality and optimized cultivate. The main results of the study were as follows:
(1)Sink potential and its components of early and late rice cultivars were studied, and sink-potential of different cultivars were classified. The sink-potential of the cultivars used in study showed a huge difference. The coefficient of variation of early rice and late rice were40.5%and35.0%respectively. The same as late rice, contribution of components to sink potential of early rice was manifested as panicles>grains per panicle>1000-grain weight.And Cluster analysis results showed that27early rice and30late rice cultivars could both be divided into three types:large, medium, and small. The averages of sink potential of the3tyes were1211.9,1073.2,921.6gm-2of early rice and1424.7,1201.8,1007.1g m-2of late rice. These results will lay a foundation for the further research on the source, nitrogen uptake and utilization and rice quality characteristics.
(2) The difference of leaf photosynthesis characteristics, dry matter production and transportation, grain filling and source-sink ratio among the large, medium and small sink-potential types of early and late rice were studied. The results showed that leaf mass and photosynthetic capacity of large and medium sink-potential types were higher than that of small sink-potential type. Large sink-potential type rice had larger LAI and leaf weight at heading and maturing stage, lower extinction rate of leaf area and higher net assimilation rate photosynthetic rate and chlorophyll content, smaller leaf angle and leaf drop angle of the top three leaves during grain filling period. Dry matter accumulation and reuse capacity of large sink-potential type rice was better than that of small sink-potential type. Large sink-potential type rice had higher dry matter accumulation especially at heading and maturing stage, dry matter of stem and sheath translocation and translocation rate than those of small sink-potential type. Sink potential obviously influented the grain filling, while with its increasing, the grain filling days significantly increased and the grain filling rate decreased, which was due to the less leaf area/sink potential, stem and sheath weight/sink potential and net assimilation rate/sink potential of large sink-potential type rice.
(3) The difference of nitrogen accumulation and translocation among different sink-potential types of early and late rice were analyzed. The relationship between sink potential and nitrogen uptake and utilization was also studied. The results showed that the large sink-potential type had higher nitrogen accumulation at heading and maturity stage, nitrogen translocation and contribution than that of the small sink-potential type rice. Nitrogen utilization efficiency was significantly influenced by sink potential. With the increasing of sink potential, nitrogen use efficiency of the plant of the early rice cultivars was slightly decreased, while nitrogen use efficiency for grain output and nitrogen use efficiency were increased or significantly increased. But nitrogen use efficiency for the plant, nitrogen use efficiency for grain output and nitrogen use efficiency of the late rice cultivars were significantly increased.
(4) The difference of processing quality, appearance quality, cooking and eating quality and nutrient quality among large, medium and small sink-potential types of early and late rice were analyzed. The relationship between sink potential and grain quality were studied. The results showed that there is a great difference in head rice rate and grain chalky whereas brown rice rate, milled rice rate and grain type was relatively small in term of different sink potential types. The head rice rate of early rice manifested as medium>small>large sink-potential type, and late rice manifested as small>medium>large sink-potential type, while the grain chalky was opposite with head rice rate. The effects of sink potential on physical and chemical characteristics were obvious. With the sink potential increases, the gel consistency and amylose content were decreased, protein content and its components was increased.
(5) The differences of response to panicle nitrogen in rice cultivars with different sink-potential were studied. The results showed that with nitrogen fertilizer increased to a proper extent, sink potential increased, the photosynthetic capacity of leaves and dry matter accumulation were enhanced, dry matter transport from stem and sheath to panicle was promoted and source-sink ratio decreased. Top dressing nitrogen fertilizer had great impact on the chalky, which showed some differences because of sink potential. Chalky grain rate and chalkiness degree of early rice with small sink-potential were decreased and then increased with the nitrogen increasing, but that of early rice with large and medium sink-potential and all three sink-potential types of late rice were increased. The effects of panicle nitrogen fertilizer on physical and chemical characteristics were significant. With the nitrogen level increased, protein and its components content were increased and amylose content and gel consistency were decreased. The responses of source-sink structure and rice quality to top dressing nitrogen fertilizer was manifested as large> medium> small sink-potential type. In this experiment,135kg hm-2N (early rice) and180kg hm"2N (late rice) were optimum level for large sink-potential type cultivars to gain higher yield and better quality;90kg hm-2N (early rice) and120kg hm-2N (late rice) were optimum level for small sink-potential type cultivars.
(6) The relationship between source-sink structure and rice quality were studied by the canonical analysis. The results showed that the rice quality factor group was significantly correlated with the sink component, leaf photosynthetic, material production, nitrogen accumulation and source-sink ratio factor group. But the quality factor group was not significantly correlated with the plant type factor group. The number of grains per panicle, LAI at heading, extinction rate of leaf area, dry matter translocation, dry matter production during grain filling, nitrogen uptake during grain filling, nitrogen translocation, leaf area/sink potential and stem and sheath weight/sink potential played an important role in each group. These indexes had great effect on chalky grain rate, chalkiness degree and protein content.
(1)研究了不同早、晚稻品种库容及其构成因子的差异,并对早稻和晚稻品种的库容类型进行了划分。结果表明,供试品种间库容量的差异较大,早稻品种间变异系数达到了40.5%、晚稻达到了35.0%；库容构成因子对库容量的贡献早、晚稻品种表现一致,穗数>每穗粒数>千粒重。聚类结果,27个早稻品种和30个晚稻品种均分为大库容、中库容和小库容3种类型,类型间库容量特征值分别为1211.9、1073.2、921.6gm-2(早稻)和1424.7、1201.8、1007.1gm-2(晚稻)。其结果为下一步开展不同库容类型品种源特性、氮素吸收利用特性和品质形成特性等研究奠定了基础。
(2)研究了早、晚稻不同库容类型品种叶光合、干物质生产与转运、籽粒灌浆和源库比的差异。结果显示,大库容品种叶量、叶光合能力较小库容品种大,具有齐穗期和成熟期叶面积大、叶重高,灌浆期间叶面积消亡慢、净同化率和净光合速率高,顶部3张叶的叶基角小等特性；大库容品种干物质积累和再利用能力强,齐穗期和成熟期干物质积累量显著高于小库容品种,灌浆期茎鞘的干物质转运量、转运率也显著大于小库容品种；库容量对籽粒灌浆影响明显,随品种库容量的增大,灌浆历期明显增长,灌浆速率减小。研究认为大库容品种的叶库比、茎库比和净库比小于小库容品种,这可能是影响籽粒灌浆的主因。
(3)研究了早、晚稻不同库容类型品种氮素积累和转运的差异,以及库容量与氮素吸收利用的关系。结果表明,大库容品种的氮素吸收与转运能力较强,齐穗期和成熟期的氮素积累量显著高于小库容品种,氮素转运量、贡献率也显著大于小库容品种；库容量对氮素利用效率的影响明显,随库容量的增大,早稻品种的氮素干物质生产效率略有减小,籽粒生产效率和氮肥利用率提高或显著提高,而晚稻品种的氮素干物质生产效率、氮素籽粒生产效率、氮肥利用率则显著提高。
(4)研究了早、晚稻不同库容类型品种加工品质、外观品质、蒸煮食味品质和营养品质的差异,以及库容量与稻米品质的关系。结果显示,不同库容类型品种的糙米率、精米率、粒型差异较小,而整精米率、垩白差异较大,早稻整精米率表现为中库容品种>小库容品种>大库容品种,晚稻整精米率表现为小库容品种>中库容品种>大库容品种,垩白粒率、垩白度表现出与整精米率相反的趋势；库容量对稻米理化特性的影响明显,随库容量的增大,胶稠度变短、直链淀粉含量降低,蛋白质及其组分的含量增加。
(5)研究了早、晚稻不同库容类型品种对氮素穗肥响应的差异。结果表明,适量增施氮素穗肥,提高了水稻库容量,增强叶片的光合能力,增加干物质积累量,促进茎鞘中贮存干物质向穗部的转运,并使源库比降低。氮素穗肥对垩白的影响较大,随施氮量增加早稻小库容品种的垩白粒率和垩白度先降后升,而早稻大、中库容品种和晚稻3种库容品种的垩白粒率、垩白度则随氮素穗肥水平提高而增加；氮素穗肥对稻米理化特性的影响显著,随施氮量提高,不同库容类型品种的蛋白质及其组分含量均增加、直链淀粉含量下降、胶稠度变短。源库结构、稻米品质对穗肥施氮量响应的敏感程度与品种库容量有关,表现出大库容品种比小库容品种敏感的特性。在本试验条件下,大库容品种获得较高产量和较优品质的适宜穗肥施氮量为135kg hm-2(早稻)、180kg hm-2(晚稻)；小库容品种获得较高产量和较优品质的适宜穗肥施氮量为90kg hm-2(早稻)、120kg hm-2(晚稻)。
(6)采用组相关分析法探究了早、晚稻源库结构与品质性状之间的关系。结果表明,库容构成因子组、叶光合因子组、物质生产因子组、氮素积累因子组、源库比因子组与品质性状组之间存在显著或极显著的相关,而株型因子组与品质性状组之间的相关不显著。其中,每穗粒数、齐穗期LAI、叶面积消亡率、茎鞘干物质转运量、灌浆期物质生产量、灌浆期吸氮量、叶茎鞘氮素转运量、叶库比、茎库比等性状或指标在各自的因子组中起主导作用,并与垩白粒率、垩白度、蛋白质含量等性状密切关联,进而对稻米品质产生显著影响。
The relationship between source-sink structure and grain quality is one of the most important aspects should be considered in rice breeding and cultivation. Expanding the sink and enhancing the source of rice is the only way to achieve high yield. How to increase yield and keep good grain quality at the same time became one of the hot and difficulty point in rice study. In this study, a field experiment was carried out with27early rice and30late rice cultivars. The cultivars of early rice and late rice were classified into3types respectively based on their sink potential level using the system clustering method. Then we analyzed the differences of source, nitrogen uptake and utilization and rice quality among the large, medium and small sink-potential types of early and late rice, investigated the effects of panicle nitrogen fertilizer on source-sink structure and rice quality. And moreover, we discussed the relationship between source-sink structure and rice quality, in order to provide a reference for the genetic improvement of rice quality and optimized cultivate. The main results of the study were as follows:
(1)Sink potential and its components of early and late rice cultivars were studied, and sink-potential of different cultivars were classified. The sink-potential of the cultivars used in study showed a huge difference. The coefficient of variation of early rice and late rice were40.5%and35.0%respectively. The same as late rice, contribution of components to sink potential of early rice was manifested as panicles>grains per panicle>1000-grain weight.And Cluster analysis results showed that27early rice and30late rice cultivars could both be divided into three types:large, medium, and small. The averages of sink potential of the3tyes were1211.9,1073.2,921.6gm-2of early rice and1424.7,1201.8,1007.1g m-2of late rice. These results will lay a foundation for the further research on the source, nitrogen uptake and utilization and rice quality characteristics.
(2) The difference of leaf photosynthesis characteristics, dry matter production and transportation, grain filling and source-sink ratio among the large, medium and small sink-potential types of early and late rice were studied. The results showed that leaf mass and photosynthetic capacity of large and medium sink-potential types were higher than that of small sink-potential type. Large sink-potential type rice had larger LAI and leaf weight at heading and maturing stage, lower extinction rate of leaf area and higher net assimilation rate photosynthetic rate and chlorophyll content, smaller leaf angle and leaf drop angle of the top three leaves during grain filling period. Dry matter accumulation and reuse capacity of large sink-potential type rice was better than that of small sink-potential type. Large sink-potential type rice had higher dry matter accumulation especially at heading and maturing stage, dry matter of stem and sheath translocation and translocation rate than those of small sink-potential type. Sink potential obviously influented the grain filling, while with its increasing, the grain filling days significantly increased and the grain filling rate decreased, which was due to the less leaf area/sink potential, stem and sheath weight/sink potential and net assimilation rate/sink potential of large sink-potential type rice.
(3) The difference of nitrogen accumulation and translocation among different sink-potential types of early and late rice were analyzed. The relationship between sink potential and nitrogen uptake and utilization was also studied. The results showed that the large sink-potential type had higher nitrogen accumulation at heading and maturity stage, nitrogen translocation and contribution than that of the small sink-potential type rice. Nitrogen utilization efficiency was significantly influenced by sink potential. With the increasing of sink potential, nitrogen use efficiency of the plant of the early rice cultivars was slightly decreased, while nitrogen use efficiency for grain output and nitrogen use efficiency were increased or significantly increased. But nitrogen use efficiency for the plant, nitrogen use efficiency for grain output and nitrogen use efficiency of the late rice cultivars were significantly increased.
(4) The difference of processing quality, appearance quality, cooking and eating quality and nutrient quality among large, medium and small sink-potential types of early and late rice were analyzed. The relationship between sink potential and grain quality were studied. The results showed that there is a great difference in head rice rate and grain chalky whereas brown rice rate, milled rice rate and grain type was relatively small in term of different sink potential types. The head rice rate of early rice manifested as medium>small>large sink-potential type, and late rice manifested as small>medium>large sink-potential type, while the grain chalky was opposite with head rice rate. The effects of sink potential on physical and chemical characteristics were obvious. With the sink potential increases, the gel consistency and amylose content were decreased, protein content and its components was increased.
(5) The differences of response to panicle nitrogen in rice cultivars with different sink-potential were studied. The results showed that with nitrogen fertilizer increased to a proper extent, sink potential increased, the photosynthetic capacity of leaves and dry matter accumulation were enhanced, dry matter transport from stem and sheath to panicle was promoted and source-sink ratio decreased. Top dressing nitrogen fertilizer had great impact on the chalky, which showed some differences because of sink potential. Chalky grain rate and chalkiness degree of early rice with small sink-potential were decreased and then increased with the nitrogen increasing, but that of early rice with large and medium sink-potential and all three sink-potential types of late rice were increased. The effects of panicle nitrogen fertilizer on physical and chemical characteristics were significant. With the nitrogen level increased, protein and its components content were increased and amylose content and gel consistency were decreased. The responses of source-sink structure and rice quality to top dressing nitrogen fertilizer was manifested as large> medium> small sink-potential type. In this experiment,135kg hm-2N (early rice) and180kg hm"2N (late rice) were optimum level for large sink-potential type cultivars to gain higher yield and better quality;90kg hm-2N (early rice) and120kg hm-2N (late rice) were optimum level for small sink-potential type cultivars.
(6) The relationship between source-sink structure and rice quality were studied by the canonical analysis. The results showed that the rice quality factor group was significantly correlated with the sink component, leaf photosynthetic, material production, nitrogen accumulation and source-sink ratio factor group. But the quality factor group was not significantly correlated with the plant type factor group. The number of grains per panicle, LAI at heading, extinction rate of leaf area, dry matter translocation, dry matter production during grain filling, nitrogen uptake during grain filling, nitrogen translocation, leaf area/sink potential and stem and sheath weight/sink potential played an important role in each group. These indexes had great effect on chalky grain rate, chalkiness degree and protein content.
引文
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