水稻超高产株型模式的生理生态基础研究
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摘要
本文以直立和弯曲穗型水稻品种杂交组合(晚轮42/沈农265；泸恢99/沈农265；晚轮422/辽粳5号)的不同世代群体为试材,分别在华南稻区广东、西南稻区四川、东北稻区辽宁和长江中下游稻区上海四个生态区同年种植,比较了多生态环境条件下不同世代株型特性、产量构成、亚种属性、穗部性状的差异,并对不同株型类型的差异进行了分析,同时研究了生态环境对其相互关系的影响,为不同地区确定最佳超高产株型模式并发挥其潜力提供科学依据。主要结果如下：
(1)生态环境对杂交F2代个体株型特性和产量构成因素具有较为明显的影响。从四川到辽宁,株高和颈穗弯曲度显著提高,而穗长、剑叶长、剑叶宽及剑叶弯曲度则表现为降低趋势。辽宁的有效穗数、千粒重、结实率以及产量均显著高于四川,穗粒数则表现为显著低于四川。千粒重、结实率和产量表现为Cw(长剑叶、弯穗)>Cz(长剑叶、直穗)>Dw(短剑叶、弯穗)>Dz(短剑叶、直穗),穗数和穗粒数以Dz最低,Cz和Cw较高。剑叶长与产量构成因素间的相关性均未达到显著水平,而剑叶宽与穗粒数均呈显著或极显著正相关,辽宁剑叶宽与结实率呈显著负相关。颈穗弯曲度只在四川分别与结实率及产量显著或极显著正相关,辽宁则表现为与大多产量构成因素相关不显著。
(2)不同生态环境下F2代植株籼粳属性均呈接近正态的连续变异,相对于四川,辽宁呈现明显的偏籼分布,生态环境对程氏指数六性状的影响并不完全一致。生态环境对株型特性也有较明显的影响,从四川到辽宁,株高和颈穗弯曲度显著提高,而穗长、剑叶长、剑叶宽及剑叶张角则表现降低趋势。不同地区株高的类型间差异没有明显规律性,剑叶长表现为籼型显著大于其他类型的趋势,剑叶弯曲度地区和类型间大多无显著差异,不同地区剑叶宽的类型间差异因组合而异。总之不同地区从籼型到粳型,均有穗长递减而颈穗弯曲度递增的趋势。籼粳稻杂交F2代植株个体籼粳属性与株型性状表现出一定的相关性,但相关的方向和程度在不同地区和组合间不完全一致。
(3)不同生态区间杂交后代产量自南向北递增,由低海拔向高海拔递增；株型特性在不同区域间存在较大差异,株高表现为四川、辽宁显著高于上海和广东；穗长区域间呈上海>四川>广东>辽宁的趋势,剑叶长表现为上海>广东>四川>辽宁,差异达显著水平；剑叶宽呈四川>辽宁>上海>广东,剑叶角度辽宁、广东高于上海和四川地区,劲穗弯曲度受遗传因素影响更为显著。在杂交后代所划分的4类株型类型中,具超级杂交稻株型模式的Cw在广东、四川和上海三个生态区表现出明显的优势性；具直立大穗型株型模式的Dz更加适应水稻生育期较长,天气条件较好,穗数水平较高的辽宁稻区。株型性状与产量表现出一定的相关性,但相关的方向和程度在不同地区和组合间表现不完全一致。在水稻理想株型育种实践中,必须适应当地生态条件和生产实际的要求,不仅要求具有“空间”的特点,也要表现出特定的生态型。
(4)糙米率、精米率、整精米率、直链淀粉含量和食味值在辽宁均显著高于四川；垩白粒率和垩白度在辽宁极显著低于四川。蛋白质含量区域间差异因组合而异,受遗传因素影响更为显著。在杂交后代所划分的4类株型类型中,具直立穗型模式的Dz(短剑叶、直立穗)精米率和整精米率在四川表现较好；辽宁糙米率、精米率和整精米率不同株型类型间差异不明显。不同生态区间垩白度、垩白粒率、蛋白质含量、直链淀粉含量以及食味值株型类型间差异不明显。品质与株型性状的关系受生态环境影响,遗传因素对其相关也存在显著影响。籼粳属性在四川地区对品质性状无明显选择压力,形态学上的分化对四川地区的影响不是很大；辽宁地区籼粳属性的分化对稻米品质具有一定的影响,糙米率、精米率、整精米率以及食味值随粳型血缘的增加而提高,特别是整精米率受籼粳属性影响最为明显；辽宁地区粳型血缘的提高外观品质有改善的趋势。
(5)不同生态区域间穗部性状发生了显著的变化,但变化的幅度和方向不尽一致。辽宁一次枝梗数显著高于其他生态区；二次枝梗数广东最低；穗粒数、次枝梗结实率和着粒密度辽宁>四川>广东和上海；千粒重四川地区最高,广东最低；总结实率呈辽宁>四川>广东>上海；穗重在不同生态区间表现为四川>辽宁>上海>广东。生态环境是引起穗部性状区域间差异的原因之一。穗部性状与株型性状表现出一定的相关性,但相关的方向和程度在不同地区和组合间表现不完全一致。辽宁稻区在育种实践中可适当引入部分籼性血缘,从而将枝梗数、粒数与结实率、穗重合理地统一起来。四川和上海地区随粳型血缘的增加,结实率也有增加的趋势,在广东地区表现在粒重和穗重的增加。在保证一定穗粒数的前提下,粳性血缘的加入可改善四川和上海籼稻结实性差的缺点,同时提高广东地区的穗重。
The different filial generations of cross between curvature panicle and erect panicle rice varieties were used as the experimental materials and were grown in in south China, Guangdong province; in southwest China, Sichuan province; in northeast China, Liaoning and in middle and lowed Reaches of Yangtze River, Shanghai. It was designed to compare the differences in plant types and rice yields, to reveal the effect of ecological environments on plant type traits, yields, subspecies characteristics and panicle traits of different filial generations of cross between curvature panicle and erect panicle rice varieties; the differences of different plant types were analyzed and to identify the correlation in different ecological environments. The purpose of this study was to provide theoretical basis in identifying the best plant type mode of super high-yeilding rice in different regions. The main results are as follow:
(1) The ecological conditions had a significant effect on plant type traits and yield traits. From Sichuan to Liaoning, the plant high and panicle curvature had increased significantly, the panicle length, flag leaf length, flag leaf width and Leaf angle decreased. Number of panicles,1000-grains weight, seed setting rate and yields in Liaoning were significantly higher than Sichuan, grains per panicle in Sichuan was significantly greater than the Liaoning. In the four patterns of plant type yield components in the comparison, the no. of panicles1000-grains weight, seed setting rate and yields had demonstrated Cw> Cz> Dw> Dz, the number of panicles and grains per panicle of Dz minimum, Cz and Cw higher. The relationship between flag leaf length and yield traits was not to reached significant, the flag leaf weight and no. of panicles achieved significant level under different environment. in Liaoning, the seed setting rates indicated a significant negative correlation with flag leaf weight. The panicle curvature showed a significantly positive correlation with the Seed setting rate and yield traits in Sichuan, but it's not significant correlation with the most of yield traits in Liaoning.
(2) The subspecies characters in F2generations were continuously varied in different environments. Two populations were generally normal distribution in Sichuan, but obviously indica-like in Liaoning. The effects of environment on six traits of Cheng's index were incompletely consistent. The ecological conditions had a significant effect on plant type traits. The plant height and panicle curvature increased significantly, but the panicle length, flag leaf length, flag leaf width and leaf angle decreased in Sichuan. Plant height in different subspecies had no obvious regularity in different regions. Flag leaf length was longer in indica(H) than in other types, but flag leaf angle had no significant difference in different subspecies and environments. Flag leaf width was changed with the cross combinations. Generally, panicle length reduced, but panicle curvature increased progressively from indica(H) to japonica(K) type. The subspecies characteristics were certainly related with plant type traits, but the correlation between them was not consistent in different cross combinations and environments.
(3) the grain yield showed a downward trend from high latitude to low latitude. There existed a large difference in plant type traits across regions. The plant high was significantly higher at Sichuan and Liaoning than that at Shanghai and Guangdong. The panicle length in the four regions showed trend of Shanghai>Sichuan>Guangdong>Liaoning. There was a significant difference in flag leaf length, ie. Shanghai>Guangdong>Sichuan>Liaoning. The flag leaf width was Sichuan>Liaoning>Shanghai>Guangdong. The flag leaf angle in Liaoning and Guangdong was larger than that in Shanghai and Sichuan. Panicle curvature was significantly influenced by genetic factors.(The difference in panicle curvature was due to the different populations in different environment). Hereditary factor played a major role in the selection of panicle type. Cw with characteristics of super hybrid plant type model showed significant vigor in Guangdong, Sichuan and Shanghai. However, erect panicle plant type (Dz) was more adaptable to the environmental conditions in Liaoning, where rice grows longer with relatively fine weather condition and with more spike numbers. There existed a closely correlation between the plant type traits and yield components. Yet, the correlative direction and degree were not completely consistent due to different populations in different ecological environments. In the ideotype breeding practice, the criteria should be regionalized based on local climatic and cultivation conditions. Such breeding should not only have the traits of "space", but also have specific geoecotype.
(4) The brown rice rate, milled rice rate, head rice rate, amylase content were significantly higher at Liaoning than that at Sichuan. The Chalky grains and Chalkiness were significantly lower at Liaoning than that at Sichuan. The difference in protein content was due to the different populations in different environment and was significantly influenced by genetic factors. In the four patterns of plant type, the milled rice rate and head rice rate of erect panicle plant type (Dz) showed significant vigor in Sichuan, the brown rice rate, milled rice rate and head rice rate at Liaoning had no obvious differences in different plant type model. The Chalky grains, Chalkiness, protent content and amylase content showed no obvious difference among different plant type model. The relationship between grain quality and plant type traits was not only significantly influenced by ecological environments, but by hereditary factor. Subspecies characterictics in Sichuan has no significantly selection pressure on grain quality, the effect of morphological classification on grain quality at Sichuan had not significant. There had a certain effect of Subspecies characterictics on grain quality at Liaoning. With the increasement of japonica ingredient, brown rice rate, milled rice rate, head rice rate and score showed increased trend, especially the head rice rate affected by subspecies characteristics was most significantly. The appearance quality at Liaoning showed an improved trend with the increasement of japonica ingredient.
(5) Panicle traits had changed significantly under different ecological environments. Yet, the varied direction and degree were not completely consistent. The number of primary branch was significantly higher at Liaoning than that at other regions. The number of Secondary branch at Guangdong showed lowest. The Grains number per panicle, Seed setting rate of primary branch and spikelet density in the four regions showed trend of Liaoning>Sichuan>Guangdong and Shanghai. Thousand grains weight was highest at Sichuan and lowest at Guangdong. The seed setting rate showed trend of Liaoning>Sichuan> Guangdong> Shanghai and the panicle weigth showed Sichuan>Liaoning> Shanghai> Guangdong. The ecological environment was one of the most important causes of the difference of panicle traits under different regions. There existed a closely correlation between the plant type traits and panicle traits. Yet, the correlative direction and degree were not completely consistent due to different populations in different ecological environments. In the Liaoning breeding practice, the partial indica pedigree should be appropriately introduced, thus the branch number, seed number and seed setting rate, grains weight could reasonably integratation. With the increasement of japonica ingredient, seed setting rate showed increased trend at Sichan and Shanghai, yet, grains weight and panicle weight showed trend of increasement at Guangdong. On the premise of guarantee of grains number per panicle, with the join of japonica ingredient could improve the defect of inferior seed setting rate at Sichuan and Shanghai, while could raise the panicle weight at Guangdong.
(1)生态环境对杂交F2代个体株型特性和产量构成因素具有较为明显的影响。从四川到辽宁,株高和颈穗弯曲度显著提高,而穗长、剑叶长、剑叶宽及剑叶弯曲度则表现为降低趋势。辽宁的有效穗数、千粒重、结实率以及产量均显著高于四川,穗粒数则表现为显著低于四川。千粒重、结实率和产量表现为Cw(长剑叶、弯穗)>Cz(长剑叶、直穗)>Dw(短剑叶、弯穗)>Dz(短剑叶、直穗),穗数和穗粒数以Dz最低,Cz和Cw较高。剑叶长与产量构成因素间的相关性均未达到显著水平,而剑叶宽与穗粒数均呈显著或极显著正相关,辽宁剑叶宽与结实率呈显著负相关。颈穗弯曲度只在四川分别与结实率及产量显著或极显著正相关,辽宁则表现为与大多产量构成因素相关不显著。
(2)不同生态环境下F2代植株籼粳属性均呈接近正态的连续变异,相对于四川,辽宁呈现明显的偏籼分布,生态环境对程氏指数六性状的影响并不完全一致。生态环境对株型特性也有较明显的影响,从四川到辽宁,株高和颈穗弯曲度显著提高,而穗长、剑叶长、剑叶宽及剑叶张角则表现降低趋势。不同地区株高的类型间差异没有明显规律性,剑叶长表现为籼型显著大于其他类型的趋势,剑叶弯曲度地区和类型间大多无显著差异,不同地区剑叶宽的类型间差异因组合而异。总之不同地区从籼型到粳型,均有穗长递减而颈穗弯曲度递增的趋势。籼粳稻杂交F2代植株个体籼粳属性与株型性状表现出一定的相关性,但相关的方向和程度在不同地区和组合间不完全一致。
(3)不同生态区间杂交后代产量自南向北递增,由低海拔向高海拔递增；株型特性在不同区域间存在较大差异,株高表现为四川、辽宁显著高于上海和广东；穗长区域间呈上海>四川>广东>辽宁的趋势,剑叶长表现为上海>广东>四川>辽宁,差异达显著水平；剑叶宽呈四川>辽宁>上海>广东,剑叶角度辽宁、广东高于上海和四川地区,劲穗弯曲度受遗传因素影响更为显著。在杂交后代所划分的4类株型类型中,具超级杂交稻株型模式的Cw在广东、四川和上海三个生态区表现出明显的优势性；具直立大穗型株型模式的Dz更加适应水稻生育期较长,天气条件较好,穗数水平较高的辽宁稻区。株型性状与产量表现出一定的相关性,但相关的方向和程度在不同地区和组合间表现不完全一致。在水稻理想株型育种实践中,必须适应当地生态条件和生产实际的要求,不仅要求具有“空间”的特点,也要表现出特定的生态型。
(4)糙米率、精米率、整精米率、直链淀粉含量和食味值在辽宁均显著高于四川；垩白粒率和垩白度在辽宁极显著低于四川。蛋白质含量区域间差异因组合而异,受遗传因素影响更为显著。在杂交后代所划分的4类株型类型中,具直立穗型模式的Dz(短剑叶、直立穗)精米率和整精米率在四川表现较好；辽宁糙米率、精米率和整精米率不同株型类型间差异不明显。不同生态区间垩白度、垩白粒率、蛋白质含量、直链淀粉含量以及食味值株型类型间差异不明显。品质与株型性状的关系受生态环境影响,遗传因素对其相关也存在显著影响。籼粳属性在四川地区对品质性状无明显选择压力,形态学上的分化对四川地区的影响不是很大；辽宁地区籼粳属性的分化对稻米品质具有一定的影响,糙米率、精米率、整精米率以及食味值随粳型血缘的增加而提高,特别是整精米率受籼粳属性影响最为明显；辽宁地区粳型血缘的提高外观品质有改善的趋势。
(5)不同生态区域间穗部性状发生了显著的变化,但变化的幅度和方向不尽一致。辽宁一次枝梗数显著高于其他生态区；二次枝梗数广东最低；穗粒数、次枝梗结实率和着粒密度辽宁>四川>广东和上海；千粒重四川地区最高,广东最低；总结实率呈辽宁>四川>广东>上海；穗重在不同生态区间表现为四川>辽宁>上海>广东。生态环境是引起穗部性状区域间差异的原因之一。穗部性状与株型性状表现出一定的相关性,但相关的方向和程度在不同地区和组合间表现不完全一致。辽宁稻区在育种实践中可适当引入部分籼性血缘,从而将枝梗数、粒数与结实率、穗重合理地统一起来。四川和上海地区随粳型血缘的增加,结实率也有增加的趋势,在广东地区表现在粒重和穗重的增加。在保证一定穗粒数的前提下,粳性血缘的加入可改善四川和上海籼稻结实性差的缺点,同时提高广东地区的穗重。
The different filial generations of cross between curvature panicle and erect panicle rice varieties were used as the experimental materials and were grown in in south China, Guangdong province; in southwest China, Sichuan province; in northeast China, Liaoning and in middle and lowed Reaches of Yangtze River, Shanghai. It was designed to compare the differences in plant types and rice yields, to reveal the effect of ecological environments on plant type traits, yields, subspecies characteristics and panicle traits of different filial generations of cross between curvature panicle and erect panicle rice varieties; the differences of different plant types were analyzed and to identify the correlation in different ecological environments. The purpose of this study was to provide theoretical basis in identifying the best plant type mode of super high-yeilding rice in different regions. The main results are as follow:
(1) The ecological conditions had a significant effect on plant type traits and yield traits. From Sichuan to Liaoning, the plant high and panicle curvature had increased significantly, the panicle length, flag leaf length, flag leaf width and Leaf angle decreased. Number of panicles,1000-grains weight, seed setting rate and yields in Liaoning were significantly higher than Sichuan, grains per panicle in Sichuan was significantly greater than the Liaoning. In the four patterns of plant type yield components in the comparison, the no. of panicles1000-grains weight, seed setting rate and yields had demonstrated Cw> Cz> Dw> Dz, the number of panicles and grains per panicle of Dz minimum, Cz and Cw higher. The relationship between flag leaf length and yield traits was not to reached significant, the flag leaf weight and no. of panicles achieved significant level under different environment. in Liaoning, the seed setting rates indicated a significant negative correlation with flag leaf weight. The panicle curvature showed a significantly positive correlation with the Seed setting rate and yield traits in Sichuan, but it's not significant correlation with the most of yield traits in Liaoning.
(2) The subspecies characters in F2generations were continuously varied in different environments. Two populations were generally normal distribution in Sichuan, but obviously indica-like in Liaoning. The effects of environment on six traits of Cheng's index were incompletely consistent. The ecological conditions had a significant effect on plant type traits. The plant height and panicle curvature increased significantly, but the panicle length, flag leaf length, flag leaf width and leaf angle decreased in Sichuan. Plant height in different subspecies had no obvious regularity in different regions. Flag leaf length was longer in indica(H) than in other types, but flag leaf angle had no significant difference in different subspecies and environments. Flag leaf width was changed with the cross combinations. Generally, panicle length reduced, but panicle curvature increased progressively from indica(H) to japonica(K) type. The subspecies characteristics were certainly related with plant type traits, but the correlation between them was not consistent in different cross combinations and environments.
(3) the grain yield showed a downward trend from high latitude to low latitude. There existed a large difference in plant type traits across regions. The plant high was significantly higher at Sichuan and Liaoning than that at Shanghai and Guangdong. The panicle length in the four regions showed trend of Shanghai>Sichuan>Guangdong>Liaoning. There was a significant difference in flag leaf length, ie. Shanghai>Guangdong>Sichuan>Liaoning. The flag leaf width was Sichuan>Liaoning>Shanghai>Guangdong. The flag leaf angle in Liaoning and Guangdong was larger than that in Shanghai and Sichuan. Panicle curvature was significantly influenced by genetic factors.(The difference in panicle curvature was due to the different populations in different environment). Hereditary factor played a major role in the selection of panicle type. Cw with characteristics of super hybrid plant type model showed significant vigor in Guangdong, Sichuan and Shanghai. However, erect panicle plant type (Dz) was more adaptable to the environmental conditions in Liaoning, where rice grows longer with relatively fine weather condition and with more spike numbers. There existed a closely correlation between the plant type traits and yield components. Yet, the correlative direction and degree were not completely consistent due to different populations in different ecological environments. In the ideotype breeding practice, the criteria should be regionalized based on local climatic and cultivation conditions. Such breeding should not only have the traits of "space", but also have specific geoecotype.
(4) The brown rice rate, milled rice rate, head rice rate, amylase content were significantly higher at Liaoning than that at Sichuan. The Chalky grains and Chalkiness were significantly lower at Liaoning than that at Sichuan. The difference in protein content was due to the different populations in different environment and was significantly influenced by genetic factors. In the four patterns of plant type, the milled rice rate and head rice rate of erect panicle plant type (Dz) showed significant vigor in Sichuan, the brown rice rate, milled rice rate and head rice rate at Liaoning had no obvious differences in different plant type model. The Chalky grains, Chalkiness, protent content and amylase content showed no obvious difference among different plant type model. The relationship between grain quality and plant type traits was not only significantly influenced by ecological environments, but by hereditary factor. Subspecies characterictics in Sichuan has no significantly selection pressure on grain quality, the effect of morphological classification on grain quality at Sichuan had not significant. There had a certain effect of Subspecies characterictics on grain quality at Liaoning. With the increasement of japonica ingredient, brown rice rate, milled rice rate, head rice rate and score showed increased trend, especially the head rice rate affected by subspecies characteristics was most significantly. The appearance quality at Liaoning showed an improved trend with the increasement of japonica ingredient.
(5) Panicle traits had changed significantly under different ecological environments. Yet, the varied direction and degree were not completely consistent. The number of primary branch was significantly higher at Liaoning than that at other regions. The number of Secondary branch at Guangdong showed lowest. The Grains number per panicle, Seed setting rate of primary branch and spikelet density in the four regions showed trend of Liaoning>Sichuan>Guangdong and Shanghai. Thousand grains weight was highest at Sichuan and lowest at Guangdong. The seed setting rate showed trend of Liaoning>Sichuan> Guangdong> Shanghai and the panicle weigth showed Sichuan>Liaoning> Shanghai> Guangdong. The ecological environment was one of the most important causes of the difference of panicle traits under different regions. There existed a closely correlation between the plant type traits and panicle traits. Yet, the correlative direction and degree were not completely consistent due to different populations in different ecological environments. In the Liaoning breeding practice, the partial indica pedigree should be appropriately introduced, thus the branch number, seed number and seed setting rate, grains weight could reasonably integratation. With the increasement of japonica ingredient, seed setting rate showed increased trend at Sichan and Shanghai, yet, grains weight and panicle weight showed trend of increasement at Guangdong. On the premise of guarantee of grains number per panicle, with the join of japonica ingredient could improve the defect of inferior seed setting rate at Sichuan and Shanghai, while could raise the panicle weight at Guangdong.
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