中国北方代表地区黏菌主要类群的个体发育及化学成分研究
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摘要
水稻品种遗传改良过程中,不同历史阶段由于育种目标的设定不同,育成品种的产量及株型、农艺性状等特征存在明显差异。关于水稻品种遗传改良的研究已有一些,但关于针对不同历史阶段的品种性状演化的研究还不多见。本研究拟分三个历史时段研究吉林省品种改良过程中农艺性状、产量和各器官特性及其与产量的关系,试图阐明吉林省三段历史时期育成品种的产量形成机制,为吉林省水稻品种改良提供可参考株型模式,同时为吉林省水稻超高产栽培实践提供量化参考依据。
本研究选择吉林省1931-2009年间育成的53个有代表性的水稻品种,分三个历史时段(1931-1969年,1970-1999年,2000年以来),采用随机区组设计,种植于同一块试验田中,研究其产量及其构成、农艺性状、各器官生育期演进规律和物质生产特性及其与产量的关系。研究结果表明：
(1)产量及组分的演化特点为,吉林省1931-2009年间育成品种产量不断增加,整个历史时期育成品种产量表现出“低-高-低-高-瓶颈”的特点。即"60年代前低,60年代高,70和80年代低,90年代高,2000年以来进入瓶颈”。1931-1969年间育成的水稻品种产量不断增加,产量的增加主要是由于对穗数和每穗粒数的改良增加引起；1970-1999年间育成的水稻品种不断增加的产量,是由于对穗数、每穗粒数、千粒重和结实率综合增加改良结果；2000年以来育成的品种产量有下滑趋势,这是由于每穗粒数增加而有效穗数逐渐减少的结果。
(2)农艺性状的演化特点为,1931-1969年间育成的水稻品种,株高显著降低,收获指数显著增加,生物量、穗重不显著增加。收获指数从40年代前的平均0.49提高到60年代的平均0.55。1970-1999年间育成的水稻品种,随育成年代增加,生物量、单穗重和穴草重显著增加,与育成年代的相关系数分别为0.49*、0.5*和0.48*,而收获指数在此期未得到改良,与各指标的相关性也不显著,但保持较高水平。此期育种策略为“保持较高的收获指数,增加生物学产量”。2000年以来育成的品种单穗重随育成年代的增加而显著增加,生物量增加不显著而收获指数有下降趋势。
(3)抽穗前的植株各器官演化特征为,1931-1969年间育成的水稻品种分蘖能力逐渐增强,倒三叶面积和鲜重显著降低。此期育成品种抽穗前叶片干重与产量呈显著正相关,相关系数为0.54**。1970-1999年间育成的水稻品种随育成年代增加倒三叶面积显著增加,相关系数为0.52*。90年代是育成品种抽穗前的分蘖和群体结构相对合理的育种时期,育成品种抽穗前的分蘖数为28.3个/穴,倒三叶面积为28.5cm2,倒三叶鲜重为0.51g,倒三叶干重为0.16g,与产量为800kg/mu的几个品种的平均值相似。2000-2009年间育成的水稻品种最大分蘖数和叶片干重出现负增长。产量与叶片干重为负相关关系,相关系数为-0.39。本试验综合倒三叶的性状与最大分蘖数的关系,提出一个可判定抽穗前群体是否合理的指标为“穴倒三叶面积”,在30cm×13cm的种植条件下,抽穗前的群体“穴倒三叶面积”应大于850cm2才是高产的合理群体结构。
(4)抽穗后的植株各器官演化特征为,1931-1969年间育成的水稻品种随着品种的改良,灌浆期植株性状各指标有降低趋势,但茎鞘干重却有增加。剑叶面积和倒三叶面积除与茎鞘为负相关外,与其它植株性状均为正相关或显著正相关。此期育成的品种产量的增加是因为倒三叶和剑叶面积的增加以及叶片干重的增加所致。1970-1999年间育成的水稻品种随育成年代增加除穗鲜重和倒三叶鲜重外,其它各指标均随育成年代增加而增加。植株各器官性状均与产量呈不显著正相关。2000-2009年间育成的水稻品种倒三叶面积、倒三叶鲜重、剑叶面积和剑叶鲜重随育成年代增加而减少,倒三叶鲜重和叶片干重与产量呈显著正相关,相关系数分别为0.61*和0.56*,这说明产量的下滑与倒三叶和叶片的改良不成功有关,可能是株型不明确的原因。
(5)吉林省超级稻适合选育以中晚熟期为主的品种,综合本试验结论与前大研究结果,提出吉林省超级稻理想株型为：植株高度为110-115cm,叶片数为14-15片,叶片直立、紧凑,半直立穗型,剑叶面积大于35cm2,根系发达,活秆成熟。每亩有效穗数30-33万穗,每穗粒数在140-160粒,千粒重为23-25g,结实率为90%以上,生育期间亩日生产量5.7-6.0kg,收获指数大于0.55,产量指标为800kg/亩以上。
综上所述,从吉林省水稻育成品种演化过程看,60年代和90年代育成品种是产量综合性状较好的时期,2000年以来育成品种的产量下滑是由于育种目标不清楚造成,吉林省超级稻育种目标应进行细致攻关研究明确,本研究提出了详细的株型及产量性状和生理性状数据,供育种者参考。
The process of genetic improvement of rice varieties in different historical stages due to the different of breeding objectives, the yield of rice varieties released and plant type, agronomic traits and characteristics such as significantly different. Genetic improvement of fice varieties have been researched, little is known about physiological and agronomical of rice varieties released in the different historical stages. In this study,53typical japonica rice cultivars applied in the production in Jilin province during the last78years were used, and classified intothree historical periods (1931-1969,1970-1999, since2000). All the tested cultivars were grown in the same field and the evolution characteristics of grain yield and its components, agronomic traits,morphology,organ characteristics and physiology were investigated and its relationship with yield. In order to clarify the formation mechanism of yield of varieties released in three historical periods, to provide the plant type mode reference for rice variety improvement, and at the same time provide a quantitative reference for the super rice cultivation practice in Jilin province.The results show that:
(1) The evolution characteristics of the yield and its composition, grain yield were progressively increased with the evolution of the cultivars from1931to2009, the entire historical period of "low-high-low-high-bottleneck" characteristics, before1960s is low,1960s is high,1970s and1980s is low,1990is high and into the bottleneck since2000. Grain yield were progressively increased from1931to1969, the increase in yield was mainly due to the increase in the panicles and grains per panicle improved; Increasing the yield of rice varieties bred between1970-1999, because of the panicles, grains per panicle, grain weight and seed setting rate increase improved the results of comprehensive; A downward trend in the yield of varieties released since2000, decrease in grain yield was attributed mainly to in the number of panicle gradually reduced.
(2) The evolution characteristics of agronomic traits, with the improvement of rice cultivars from1931to1969, plant height was significantly degraded, a significant increase in harvest index, biomass, panicle weight was not significantly increased. Harvest index increased from an average of0.49of the1940s to the1960s an average of0.55. There were extremely significant increases in biomass, single panicle weight and grass weight per hole of rice varieties released between the years1970to1999with year of release increased and the correlation coefficients were0.49*,0.5*and0.48*, and harvest index has not been improved in this period, the correlation with the index was not significant, but to maintain a high level. Breeding strategy of rice is to maintain a high harvest index. increased biomass yield in this period. Since2000, Single panicle weight were increased significantly, biomass increase was not significant, while harvest index was reduced with year of release increased.
(3) The evolution characteristics of the various organs of the plants before heading. Rice tillering ability is gradually increased, top3rd leaf area and fresh weight of rice varieties released was significantly reduced from1931to1969. Leaf dry weight of varieties released before heading and yield was significant positive correlation in this period, and correlation coefficient is0.54*. Top3rd leaf area of rice varieties released were th increased from1970to1999. the correlation coefficient was0.52*for years of released. Heading tiller and population structure of rice varieties were relativey reasonable in the1090s. the number of tillers before heading varieties bred was28.3per hole, top3rd leaf area was28.5cm2. top3rd leaf fresh weight was0.5lg. top3rd leaf dry weight was0.16g. similar to the average of several varieties of the output of800kg/mn. Since2000, tiller number and leaf dry weight were negative decreased. Yield and leaf dry weight was negatively correlated, and a correlation coefficient of-0.39. This experiment integrated top3rd leaf traits and the relationship of the maximum number of tillers, and proposed one can determine whether a reasonable population structure before heading groups as " top3rd leaf area per hole", the30cmxl3cm planting conditions before heading groups the " top3rd leaf area per hole " should be greater than850cm" is a reasonable population structure of super-high-yield.
(4) The characteristics of plant organ evolution after heading, from1931to1969, value of plant traits index of rice varieties released were decreased at the grain filling stage, but there is an increasing stem and sheath dry weight. The flag leaf area and top3rd leaf area were negatively correlated with the stem and sheath, with other plant traits were significantly positively correlated, increase in grain yield was attributed mainly to the increase in flag leaf and top3rd leaf area, leaf dry weight. From1970to1999, values of traits were increased with years of release beside panicle and top3rd leaf fresh weight; various organ traits in plants and the yield were not significantly positively correlated. Top3rd leaf are and fresh weight, flag leaf area and fresh weight of rice varieties released were decreased with years of release from2000to2009, and top3rd leaf fresh weight and leaf dry weight was significant positive relation with yield, the correlation coefficient of0.61*and0.56*, which shows the decline in yield and improvement of the top3rd leaf and leaf unsuccessful, maybe the plant type is not clear.
(5) The fundamental strategy of super high-yield breeding wound been breeding in the late-maturing of varieties in Jilin province, and the ideal plant type of super high-yield rice varieties were proposed from this study also concluded with the results of previous studies put forward in Jilin province. Plant height is110-115cm, leaves numbers is14-15piece, upright leaves, compact and semi-erect panicle type, the flag leaf area is greater than35cm2, well-developed root system, and live mature stalk. Effective panicles are30×l04-33×l04/mu. grains per panicle in the140-160grain.1000-grain weight for23-25g, setting rate above90%,85.5-90.0kg/hm2per day production of the reproductive period, harvest index is greater than0.55, yield is more than800kg/mu.
In summary, yield integrative traits of rice varieties released in the1960s and1990s, which were better than other historical periods in Jilin province from evolution of varieties bred. The yield of varieties released were declined since2000, the breeding objectives unclear cause. Jilin Province super high-yield breeding objectives detailed research should be carried out explicitly, this study proposes a detailed plant type and yield traits and physiological traits data for breeder's reference.
本研究选择吉林省1931-2009年间育成的53个有代表性的水稻品种,分三个历史时段(1931-1969年,1970-1999年,2000年以来),采用随机区组设计,种植于同一块试验田中,研究其产量及其构成、农艺性状、各器官生育期演进规律和物质生产特性及其与产量的关系。研究结果表明：
(1)产量及组分的演化特点为,吉林省1931-2009年间育成品种产量不断增加,整个历史时期育成品种产量表现出“低-高-低-高-瓶颈”的特点。即"60年代前低,60年代高,70和80年代低,90年代高,2000年以来进入瓶颈”。1931-1969年间育成的水稻品种产量不断增加,产量的增加主要是由于对穗数和每穗粒数的改良增加引起；1970-1999年间育成的水稻品种不断增加的产量,是由于对穗数、每穗粒数、千粒重和结实率综合增加改良结果；2000年以来育成的品种产量有下滑趋势,这是由于每穗粒数增加而有效穗数逐渐减少的结果。
(2)农艺性状的演化特点为,1931-1969年间育成的水稻品种,株高显著降低,收获指数显著增加,生物量、穗重不显著增加。收获指数从40年代前的平均0.49提高到60年代的平均0.55。1970-1999年间育成的水稻品种,随育成年代增加,生物量、单穗重和穴草重显著增加,与育成年代的相关系数分别为0.49*、0.5*和0.48*,而收获指数在此期未得到改良,与各指标的相关性也不显著,但保持较高水平。此期育种策略为“保持较高的收获指数,增加生物学产量”。2000年以来育成的品种单穗重随育成年代的增加而显著增加,生物量增加不显著而收获指数有下降趋势。
(3)抽穗前的植株各器官演化特征为,1931-1969年间育成的水稻品种分蘖能力逐渐增强,倒三叶面积和鲜重显著降低。此期育成品种抽穗前叶片干重与产量呈显著正相关,相关系数为0.54**。1970-1999年间育成的水稻品种随育成年代增加倒三叶面积显著增加,相关系数为0.52*。90年代是育成品种抽穗前的分蘖和群体结构相对合理的育种时期,育成品种抽穗前的分蘖数为28.3个/穴,倒三叶面积为28.5cm2,倒三叶鲜重为0.51g,倒三叶干重为0.16g,与产量为800kg/mu的几个品种的平均值相似。2000-2009年间育成的水稻品种最大分蘖数和叶片干重出现负增长。产量与叶片干重为负相关关系,相关系数为-0.39。本试验综合倒三叶的性状与最大分蘖数的关系,提出一个可判定抽穗前群体是否合理的指标为“穴倒三叶面积”,在30cm×13cm的种植条件下,抽穗前的群体“穴倒三叶面积”应大于850cm2才是高产的合理群体结构。
(4)抽穗后的植株各器官演化特征为,1931-1969年间育成的水稻品种随着品种的改良,灌浆期植株性状各指标有降低趋势,但茎鞘干重却有增加。剑叶面积和倒三叶面积除与茎鞘为负相关外,与其它植株性状均为正相关或显著正相关。此期育成的品种产量的增加是因为倒三叶和剑叶面积的增加以及叶片干重的增加所致。1970-1999年间育成的水稻品种随育成年代增加除穗鲜重和倒三叶鲜重外,其它各指标均随育成年代增加而增加。植株各器官性状均与产量呈不显著正相关。2000-2009年间育成的水稻品种倒三叶面积、倒三叶鲜重、剑叶面积和剑叶鲜重随育成年代增加而减少,倒三叶鲜重和叶片干重与产量呈显著正相关,相关系数分别为0.61*和0.56*,这说明产量的下滑与倒三叶和叶片的改良不成功有关,可能是株型不明确的原因。
(5)吉林省超级稻适合选育以中晚熟期为主的品种,综合本试验结论与前大研究结果,提出吉林省超级稻理想株型为：植株高度为110-115cm,叶片数为14-15片,叶片直立、紧凑,半直立穗型,剑叶面积大于35cm2,根系发达,活秆成熟。每亩有效穗数30-33万穗,每穗粒数在140-160粒,千粒重为23-25g,结实率为90%以上,生育期间亩日生产量5.7-6.0kg,收获指数大于0.55,产量指标为800kg/亩以上。
综上所述,从吉林省水稻育成品种演化过程看,60年代和90年代育成品种是产量综合性状较好的时期,2000年以来育成品种的产量下滑是由于育种目标不清楚造成,吉林省超级稻育种目标应进行细致攻关研究明确,本研究提出了详细的株型及产量性状和生理性状数据,供育种者参考。
The process of genetic improvement of rice varieties in different historical stages due to the different of breeding objectives, the yield of rice varieties released and plant type, agronomic traits and characteristics such as significantly different. Genetic improvement of fice varieties have been researched, little is known about physiological and agronomical of rice varieties released in the different historical stages. In this study,53typical japonica rice cultivars applied in the production in Jilin province during the last78years were used, and classified intothree historical periods (1931-1969,1970-1999, since2000). All the tested cultivars were grown in the same field and the evolution characteristics of grain yield and its components, agronomic traits,morphology,organ characteristics and physiology were investigated and its relationship with yield. In order to clarify the formation mechanism of yield of varieties released in three historical periods, to provide the plant type mode reference for rice variety improvement, and at the same time provide a quantitative reference for the super rice cultivation practice in Jilin province.The results show that:
(1) The evolution characteristics of the yield and its composition, grain yield were progressively increased with the evolution of the cultivars from1931to2009, the entire historical period of "low-high-low-high-bottleneck" characteristics, before1960s is low,1960s is high,1970s and1980s is low,1990is high and into the bottleneck since2000. Grain yield were progressively increased from1931to1969, the increase in yield was mainly due to the increase in the panicles and grains per panicle improved; Increasing the yield of rice varieties bred between1970-1999, because of the panicles, grains per panicle, grain weight and seed setting rate increase improved the results of comprehensive; A downward trend in the yield of varieties released since2000, decrease in grain yield was attributed mainly to in the number of panicle gradually reduced.
(2) The evolution characteristics of agronomic traits, with the improvement of rice cultivars from1931to1969, plant height was significantly degraded, a significant increase in harvest index, biomass, panicle weight was not significantly increased. Harvest index increased from an average of0.49of the1940s to the1960s an average of0.55. There were extremely significant increases in biomass, single panicle weight and grass weight per hole of rice varieties released between the years1970to1999with year of release increased and the correlation coefficients were0.49*,0.5*and0.48*, and harvest index has not been improved in this period, the correlation with the index was not significant, but to maintain a high level. Breeding strategy of rice is to maintain a high harvest index. increased biomass yield in this period. Since2000, Single panicle weight were increased significantly, biomass increase was not significant, while harvest index was reduced with year of release increased.
(3) The evolution characteristics of the various organs of the plants before heading. Rice tillering ability is gradually increased, top3rd leaf area and fresh weight of rice varieties released was significantly reduced from1931to1969. Leaf dry weight of varieties released before heading and yield was significant positive correlation in this period, and correlation coefficient is0.54*. Top3rd leaf area of rice varieties released were th increased from1970to1999. the correlation coefficient was0.52*for years of released. Heading tiller and population structure of rice varieties were relativey reasonable in the1090s. the number of tillers before heading varieties bred was28.3per hole, top3rd leaf area was28.5cm2. top3rd leaf fresh weight was0.5lg. top3rd leaf dry weight was0.16g. similar to the average of several varieties of the output of800kg/mn. Since2000, tiller number and leaf dry weight were negative decreased. Yield and leaf dry weight was negatively correlated, and a correlation coefficient of-0.39. This experiment integrated top3rd leaf traits and the relationship of the maximum number of tillers, and proposed one can determine whether a reasonable population structure before heading groups as " top3rd leaf area per hole", the30cmxl3cm planting conditions before heading groups the " top3rd leaf area per hole " should be greater than850cm" is a reasonable population structure of super-high-yield.
(4) The characteristics of plant organ evolution after heading, from1931to1969, value of plant traits index of rice varieties released were decreased at the grain filling stage, but there is an increasing stem and sheath dry weight. The flag leaf area and top3rd leaf area were negatively correlated with the stem and sheath, with other plant traits were significantly positively correlated, increase in grain yield was attributed mainly to the increase in flag leaf and top3rd leaf area, leaf dry weight. From1970to1999, values of traits were increased with years of release beside panicle and top3rd leaf fresh weight; various organ traits in plants and the yield were not significantly positively correlated. Top3rd leaf are and fresh weight, flag leaf area and fresh weight of rice varieties released were decreased with years of release from2000to2009, and top3rd leaf fresh weight and leaf dry weight was significant positive relation with yield, the correlation coefficient of0.61*and0.56*, which shows the decline in yield and improvement of the top3rd leaf and leaf unsuccessful, maybe the plant type is not clear.
(5) The fundamental strategy of super high-yield breeding wound been breeding in the late-maturing of varieties in Jilin province, and the ideal plant type of super high-yield rice varieties were proposed from this study also concluded with the results of previous studies put forward in Jilin province. Plant height is110-115cm, leaves numbers is14-15piece, upright leaves, compact and semi-erect panicle type, the flag leaf area is greater than35cm2, well-developed root system, and live mature stalk. Effective panicles are30×l04-33×l04/mu. grains per panicle in the140-160grain.1000-grain weight for23-25g, setting rate above90%,85.5-90.0kg/hm2per day production of the reproductive period, harvest index is greater than0.55, yield is more than800kg/mu.
In summary, yield integrative traits of rice varieties released in the1960s and1990s, which were better than other historical periods in Jilin province from evolution of varieties bred. The yield of varieties released were declined since2000, the breeding objectives unclear cause. Jilin Province super high-yield breeding objectives detailed research should be carried out explicitly, this study proposes a detailed plant type and yield traits and physiological traits data for breeder's reference.
引文
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