甘蓝型油菜不同氮素籽粒生产效率类型品种的基本特征
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摘要
以甘蓝型常规油菜品种(2006-2007年度73个品种,2007-2008年度98个品种并设置NO和N1两个氮肥处理)为研究对象,通过测定初花期不同器官(叶片和茎枝)干重和氮素含量,成熟期测定植株农艺性状(包括株高、一次分枝数和角果数)以及不同器官(茎枝、果壳和籽粒)的干重和氮素含量。采用组内最小平方和的动态聚类方法,按氮素籽粒生产效率为指标将不同品种从低到高依次分为A、B、C、D、E和F六种类型。从农艺性状、干物质和氮素的积累与分配等方面研究不同氮素籽粒生产效率类型品种的基本特征。对2007-2008年度98个品种分别在N0和N1条件下按氮素籽粒生产效率分别进行聚类(分低效、中效和高效三类),结合不同品种从低氮到高氮条件下类型的转换将供试品种分成六种类型低到低(L→L)、中到低(M→L)、中到中(M→M)、中到高(M→H)、高到中(H→M)和高到高(H→H)6种类型,研究了不同氮素籽粒生产效率类型品种的基本特征及其对氮肥的响应。2007-2008年度测定了28个油菜品种苗期的SPAD值、光合参数等,研究了不同氮素籽粒生产效率品种苗期光合特性的差异。2008-2009年度在两个氮肥水平下以6个亲本(浙双3号、扬油7号、ZJ1、史力佳、宁油14号和沪油16)及其完全双列杂交组合为研究对象,研究氮素籽粒生产效率的杂种优势以及亲本的一般配合力和组合的特殊配合力,并分析遗传力大小。希望通过本研究为生产上高氮素籽粒生产效率品种的筛选和培育以及合理栽培措施的制定提供参考依据。本文主要研究结果如下：
1.衡量作物对已吸收氮素利用效率的高低的指标主要有氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数。本研究结果表明,甘蓝型油菜的氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数的变化趋势基本一致,其中以氮素籽粒生产效率对产量的影响最大,并且不同试验条件下不同品种的氮素籽粒生产效率的变幅也较大。所以用氮素籽粒生产效率作为反映油菜对氮素利用效率高低的指标更为合适。
2.供试品种(系)氮素籽粒生产效率差异较大,2007试验供试73个品种的变幅为15.73～21.20g.g-1,平均值18.66 g.g-1；2008N0和2008N1处理98个供试品种的变幅分别13.61～19.16 g.g-1和12.28～16.54g.g-1,平均值分别为16.76 g.g-1和14.51 g.g-1。随着土壤肥力或施氮量增加,氮素籽粒生产效率减小。研究结果显示HY04、SY01、SY03、SY07、YN06、YN07、YN14、红油3号、沪油15、宁油18号、湘05483、湘05484、湘05487、扬油7号等14个品种(系)氮素籽粒生产效率在不同的试验条件下都属于F和E类型。
3.苗期叶片PSⅡ最大光化学量子产量(Fv／Fm)与氮素籽粒生产效率呈显著正相关,低氮条件下苗期叶片SPAD值与氮素籽粒生产效率也呈显著正相关。增施氮肥,叶片SPAD值和净光合速率增加；PSⅡ最大光化学量子产量(Fv／Fm)和光化学淬灭系数(qP)增加,非光化学淬灭系数(qN)减小。
4.随着氮素籽粒生产效率的增加,不同类型品种的株高、一次分枝数和单位面积角果数呈增加的趋势。库容量(总籽粒数)与氮素籽粒生产效率呈极显著正相关(r2007=0.4057**,r2008N0=0.5945**,r2008N1=0.5412**)。通径分析表明,提高氮素籽粒生产效率应重点提高单位面积上的角果数,增加每角粒数也有一定的作用。
5.初花期干重与氮素籽粒生产效率关系不显著,而花后干物质积累量以及成熟期干重与氮素籽粒生产效率呈极显著正相关。随着氮素籽粒生产效率增加,不同类型品种果壳重、籽粒重和籽粒干重比例增加,茎枝干重比例下降。进一步表明,成熟期果壳重和籽粒重对氮素籽粒生产效率有显著影响,其中籽粒重对氮素籽粒生产效率影响最大,其次是果壳重。提高氮素籽粒生产效率重点是增加花后干物质积累量,促进茎枝干物质向籽粒输送,降低茎枝重所占的比例,提高角果重,尤其是籽粒重。
6.初花期各器官氮素积累量以及成熟期全株氮素积累量与氮素籽粒生产效率之间关系不密切,但成熟期不同器官氮素积累量以及不同器官的氮素分配比例与氮素籽粒生产效率都达显著相关,其中茎枝和果壳氮素积累量及其分配比例与氮素籽粒生产效率呈显著负相关,籽粒氮素积累量及其分配比例与氮素籽粒生产效率呈显著正相关。回归分析表明,成熟期不同器官氮素积累量对氮素籽粒生产效率都有显著影响。提高氮素籽粒生产效率应增加籽粒中的氮素积累量,降低茎枝和果壳中的氮素积累量。
7.氮肥对不同品种氮素籽粒生产效率的影响存在差异,本研究结果显示低到低、中到低、中到中、中到高、高到中和高到高类型品种数分别为14、8、21、6、19和30。其中施用氮肥变高类型(中到高)品种主要特征表现为每角粒数、籽粒重增加值比较大,库容量增加幅度大。茎枝和果壳氮素积累量增加少,茎枝和果壳氮素积累比例增加值较小,籽粒氮素分配比例下降较小。施用氮肥变低类型(中到低,高到中)变化趋势与由低到高类型相反。
8.本文研究结果表明,油菜氮素籽粒生产效率的杂种优势比较明显。配合力方差分析显示GCA、SCA和反交效应的方差均达到极显著水平,说明氮素籽粒生产效率受加性效应、非加性效应和细胞质效应的共同作用。通过亲本与杂交种的相关分析表明,在氮素籽粒生产效率的组配过程中要注重母本性状的选择。在遗传力方面研究结果显示广义遗传力都比较高,在低氮条件下氮素籽粒生产效率的基因加性作用和非加性作用都比较重要,而高氮条件下以基因的非加性效应为主。
Field experiments were carried out to study basic characteristics of rapeseed varieties (Brassica napus L.) with different types of nitrogen use efficiency for grain production.73 conventional rapeseed varieties were cultured in 2006-2007 (2007 treatment), and 98 conventional rapeseed varieties were cultured in 2007-2008 under two nitrogen fertilizer levels (2008N0 treatment and 2008N1 treatment). At beginning flowering stage, dry matter weight (DMW)and the nitrogen content of different organs (leave and stem) were tested, and at maturing stage agronomic characteristics, DMW and the nitrogen content of different organs (stem, shell and seed) were tested. The tested rapeseed varieties were classified into 6 types (A, B, C, D, E, F) based on their nitrogen use efficiency for grain production level by the MinSSw method, agronomic characteristics and nitrogen accumulation and distribution of different types were studied. According to 2008N0 and 2008N1 treatments, the tested rapeseed varieties were classified into 3 types (low type, middle type and high type) under different nitrogen fertilizer levels, then these varieties were divided into six types (L→L, M→L, M→M, M→H, H→M, H→H) baded on change of different types from low nitrogen fertilizer to high nitrogen fertilizer, responses of NUEg on nitrogen fertilizer were studied. In order to study relationships between photosynthetic characteristic of leaf and NUEg, SPAD value and photosynthetic parameters were tested at seeding stage in 28 cultivars. To study heterosis, GCA, SCA and heritability of NUEg, six parents (Zheshuang 3,Yangyou7, ZJ1, Shilijia, Ningyou 14, Huyou 16) and their F1 combinations from 6×6 complete double allele crosses were tested under two levels of nitrogen application in 2008-2009. The purposes of these studies were to investigate the basic characteristics of rapeseed with higher nitrogen use efficiency for grain production and to provide fundamental information for genetic improvement of nitrogen use efficiency in rapeseed breeding and feasible cultivation measures designment. Results were as following:
1. There are there indexes of nitrogen use effiency from plant itself including nitrogen use efficiency for grain production (NUEg), nitrogen use efficiency for plant biomass(NUEp) and nitrogen harvest index (NHI). Change trends of these indexes were consistent.The study of path analysis indicated that NUEg on yield was biggest. Change ranges of NUEg were more bigger. So NUEg was a more suitable index to indicate nitrogen efficiency than NUEp and NHI.
2. Nitrogen use efficiency for grain production differed greatly among rapeseed varieties used in this study. Range of NUEg was from 15.73 g.g-1 to 21.20g.g-1 in 2007 treatment, from 13.61 to 19.16 g.g-1 in 2008N0 treatment,from 12.28 g.g-1 to 16.54g.g-1, and mean value was 18.66 g.g-1,16.76g.g-1 and 4.51 g.g-1, respectively. With the increasing of soil fertilizer and nitrogen fertilization, NUEg decreased.There were fourteen varieties in E or F types in different treatments, and they were HY04, SY01, SY03, SY07, YN06, YN07, YN14, Hongyou3, Huyou 15,Ningyou18, Xiang 05483, Xiang 05484, Xiang 05487 and Yangyou 7。
3. There were significant positive correlations between Fv/Fm and NUEg in 2008N0 and 2008N1 treatments. SPAD values were significantly positive correlated with NUEg in 2008N0 treatment. With the increasing of nitrogen fertilization, SPAD value, Pn, and qP increased, and qN decreased.
4. With increasing of NUEg, plant height, number of primary banches and number of pods per area with different type varieties increased. Sink capacity (total number of seeds) increased with increasing of NUEg, and there were significantly positive correlations between total number of seeds and NUEg (r2007=0.4057**, r2008=0.5945**, r2009=0.5412**). The study of path analysis indicated that the indirect influence of number of pods on NUEg was biggest, and secondary was number of seeds per pod.
5. In the aspect of dry matter accumulation and distribution, the effect of DMW at beginning flowering on NUEg was slight.There were significantly positive correlations between dry matter accumulation after flowering and NUEg, biomass and NUEg. With increasing of NUEg, DMW of shell, yield and the ratio of DMW of seed to biomass with different type varieties increased, the ratio of DMW of stem to biomass decreased. The study of multiple regression analysis indicated that yield and DMW of shell significantly effected NUEg. The study of path analysis indicated that the indirect influence of yield on NUEg was most important, and secondary was DMW of shell. To increasing NUEg it was needed to increase dry matter accumulation from beginning flowering stage to maturing stage, and promote photosynthate to transport from stem to pods, so as to increase dry matter accumulation in pods, especially increase yield.
6. In the aspect of nitrogen accumulation and distribution, correlations between the amount of nitrogen accumulation at beginning flowering stage and NUEg, the total amount of nitrogen accumulation at maturing stage were not significant. But there were significantly negative correlations between nitrogen accumulation and distribution of stem and shell and NUEg, and there were significantly positive correlations between the amount of nitrogen of seed, NHI and NUEg. The studies of multiple regression analysis indicated that amounts of nitrogen of different organs at maturing stage significantly effected NUEg. To increase NUEg, increasing nitrogen accumulation of seeds was important, and at the same time decreasing nitrogen accumulation of stem and shell was also important.
7. There were different effects of nitrogen fertilizer on NUEg. Results showed that the number of different types (L→L, M→L, M→M, M→H, H→M, H→H) was 14,8,21,6,19 and 30, respectively. Though studying variety characteristics of different types, results showed that in the varieties from low type to high type by nitrogen application (M→H), added values of the number of seeds per pod, yield and the total number of seeds were larger, arid added values of nitrogen accumulation of stem and shell, ratio of nitrogen accumulation of stem and shell to total nitrogen accumulation were smaller. Decreasing values of ratio of nitrogen of seed to total nitrogen accumulation were smaller. The trends of the varieties from high type to low type by nitrogen application (M→L, H→M) were opposite.
8. Through testing NUEg of six parents and their F1 combinations from 6×6 complete double allele crosses, results showed that positive heterosis of NUEg was significant. NUEg were likely affected by additive effects, dominance effects and cytoplasm effects by variance analysis of combining ability. The correlation analysis of parents and F1 combinations showed that famale selection was important. In the aspect of heritability, results showed that broad heritability were higher in different treatments. Additive effects and dominance effects were all significant in low nitrogen fertilizer and dominance effects were significant in high nitrogen fertilizer.
1.衡量作物对已吸收氮素利用效率的高低的指标主要有氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数。本研究结果表明,甘蓝型油菜的氮素籽粒生产效率、氮素干物质生产效率和氮素收获指数的变化趋势基本一致,其中以氮素籽粒生产效率对产量的影响最大,并且不同试验条件下不同品种的氮素籽粒生产效率的变幅也较大。所以用氮素籽粒生产效率作为反映油菜对氮素利用效率高低的指标更为合适。
2.供试品种(系)氮素籽粒生产效率差异较大,2007试验供试73个品种的变幅为15.73～21.20g.g-1,平均值18.66 g.g-1；2008N0和2008N1处理98个供试品种的变幅分别13.61～19.16 g.g-1和12.28～16.54g.g-1,平均值分别为16.76 g.g-1和14.51 g.g-1。随着土壤肥力或施氮量增加,氮素籽粒生产效率减小。研究结果显示HY04、SY01、SY03、SY07、YN06、YN07、YN14、红油3号、沪油15、宁油18号、湘05483、湘05484、湘05487、扬油7号等14个品种(系)氮素籽粒生产效率在不同的试验条件下都属于F和E类型。
3.苗期叶片PSⅡ最大光化学量子产量(Fv／Fm)与氮素籽粒生产效率呈显著正相关,低氮条件下苗期叶片SPAD值与氮素籽粒生产效率也呈显著正相关。增施氮肥,叶片SPAD值和净光合速率增加；PSⅡ最大光化学量子产量(Fv／Fm)和光化学淬灭系数(qP)增加,非光化学淬灭系数(qN)减小。
4.随着氮素籽粒生产效率的增加,不同类型品种的株高、一次分枝数和单位面积角果数呈增加的趋势。库容量(总籽粒数)与氮素籽粒生产效率呈极显著正相关(r2007=0.4057**,r2008N0=0.5945**,r2008N1=0.5412**)。通径分析表明,提高氮素籽粒生产效率应重点提高单位面积上的角果数,增加每角粒数也有一定的作用。
5.初花期干重与氮素籽粒生产效率关系不显著,而花后干物质积累量以及成熟期干重与氮素籽粒生产效率呈极显著正相关。随着氮素籽粒生产效率增加,不同类型品种果壳重、籽粒重和籽粒干重比例增加,茎枝干重比例下降。进一步表明,成熟期果壳重和籽粒重对氮素籽粒生产效率有显著影响,其中籽粒重对氮素籽粒生产效率影响最大,其次是果壳重。提高氮素籽粒生产效率重点是增加花后干物质积累量,促进茎枝干物质向籽粒输送,降低茎枝重所占的比例,提高角果重,尤其是籽粒重。
6.初花期各器官氮素积累量以及成熟期全株氮素积累量与氮素籽粒生产效率之间关系不密切,但成熟期不同器官氮素积累量以及不同器官的氮素分配比例与氮素籽粒生产效率都达显著相关,其中茎枝和果壳氮素积累量及其分配比例与氮素籽粒生产效率呈显著负相关,籽粒氮素积累量及其分配比例与氮素籽粒生产效率呈显著正相关。回归分析表明,成熟期不同器官氮素积累量对氮素籽粒生产效率都有显著影响。提高氮素籽粒生产效率应增加籽粒中的氮素积累量,降低茎枝和果壳中的氮素积累量。
7.氮肥对不同品种氮素籽粒生产效率的影响存在差异,本研究结果显示低到低、中到低、中到中、中到高、高到中和高到高类型品种数分别为14、8、21、6、19和30。其中施用氮肥变高类型(中到高)品种主要特征表现为每角粒数、籽粒重增加值比较大,库容量增加幅度大。茎枝和果壳氮素积累量增加少,茎枝和果壳氮素积累比例增加值较小,籽粒氮素分配比例下降较小。施用氮肥变低类型(中到低,高到中)变化趋势与由低到高类型相反。
8.本文研究结果表明,油菜氮素籽粒生产效率的杂种优势比较明显。配合力方差分析显示GCA、SCA和反交效应的方差均达到极显著水平,说明氮素籽粒生产效率受加性效应、非加性效应和细胞质效应的共同作用。通过亲本与杂交种的相关分析表明,在氮素籽粒生产效率的组配过程中要注重母本性状的选择。在遗传力方面研究结果显示广义遗传力都比较高,在低氮条件下氮素籽粒生产效率的基因加性作用和非加性作用都比较重要,而高氮条件下以基因的非加性效应为主。
Field experiments were carried out to study basic characteristics of rapeseed varieties (Brassica napus L.) with different types of nitrogen use efficiency for grain production.73 conventional rapeseed varieties were cultured in 2006-2007 (2007 treatment), and 98 conventional rapeseed varieties were cultured in 2007-2008 under two nitrogen fertilizer levels (2008N0 treatment and 2008N1 treatment). At beginning flowering stage, dry matter weight (DMW)and the nitrogen content of different organs (leave and stem) were tested, and at maturing stage agronomic characteristics, DMW and the nitrogen content of different organs (stem, shell and seed) were tested. The tested rapeseed varieties were classified into 6 types (A, B, C, D, E, F) based on their nitrogen use efficiency for grain production level by the MinSSw method, agronomic characteristics and nitrogen accumulation and distribution of different types were studied. According to 2008N0 and 2008N1 treatments, the tested rapeseed varieties were classified into 3 types (low type, middle type and high type) under different nitrogen fertilizer levels, then these varieties were divided into six types (L→L, M→L, M→M, M→H, H→M, H→H) baded on change of different types from low nitrogen fertilizer to high nitrogen fertilizer, responses of NUEg on nitrogen fertilizer were studied. In order to study relationships between photosynthetic characteristic of leaf and NUEg, SPAD value and photosynthetic parameters were tested at seeding stage in 28 cultivars. To study heterosis, GCA, SCA and heritability of NUEg, six parents (Zheshuang 3,Yangyou7, ZJ1, Shilijia, Ningyou 14, Huyou 16) and their F1 combinations from 6×6 complete double allele crosses were tested under two levels of nitrogen application in 2008-2009. The purposes of these studies were to investigate the basic characteristics of rapeseed with higher nitrogen use efficiency for grain production and to provide fundamental information for genetic improvement of nitrogen use efficiency in rapeseed breeding and feasible cultivation measures designment. Results were as following:
1. There are there indexes of nitrogen use effiency from plant itself including nitrogen use efficiency for grain production (NUEg), nitrogen use efficiency for plant biomass(NUEp) and nitrogen harvest index (NHI). Change trends of these indexes were consistent.The study of path analysis indicated that NUEg on yield was biggest. Change ranges of NUEg were more bigger. So NUEg was a more suitable index to indicate nitrogen efficiency than NUEp and NHI.
2. Nitrogen use efficiency for grain production differed greatly among rapeseed varieties used in this study. Range of NUEg was from 15.73 g.g-1 to 21.20g.g-1 in 2007 treatment, from 13.61 to 19.16 g.g-1 in 2008N0 treatment,from 12.28 g.g-1 to 16.54g.g-1, and mean value was 18.66 g.g-1,16.76g.g-1 and 4.51 g.g-1, respectively. With the increasing of soil fertilizer and nitrogen fertilization, NUEg decreased.There were fourteen varieties in E or F types in different treatments, and they were HY04, SY01, SY03, SY07, YN06, YN07, YN14, Hongyou3, Huyou 15,Ningyou18, Xiang 05483, Xiang 05484, Xiang 05487 and Yangyou 7。
3. There were significant positive correlations between Fv/Fm and NUEg in 2008N0 and 2008N1 treatments. SPAD values were significantly positive correlated with NUEg in 2008N0 treatment. With the increasing of nitrogen fertilization, SPAD value, Pn, and qP increased, and qN decreased.
4. With increasing of NUEg, plant height, number of primary banches and number of pods per area with different type varieties increased. Sink capacity (total number of seeds) increased with increasing of NUEg, and there were significantly positive correlations between total number of seeds and NUEg (r2007=0.4057**, r2008=0.5945**, r2009=0.5412**). The study of path analysis indicated that the indirect influence of number of pods on NUEg was biggest, and secondary was number of seeds per pod.
5. In the aspect of dry matter accumulation and distribution, the effect of DMW at beginning flowering on NUEg was slight.There were significantly positive correlations between dry matter accumulation after flowering and NUEg, biomass and NUEg. With increasing of NUEg, DMW of shell, yield and the ratio of DMW of seed to biomass with different type varieties increased, the ratio of DMW of stem to biomass decreased. The study of multiple regression analysis indicated that yield and DMW of shell significantly effected NUEg. The study of path analysis indicated that the indirect influence of yield on NUEg was most important, and secondary was DMW of shell. To increasing NUEg it was needed to increase dry matter accumulation from beginning flowering stage to maturing stage, and promote photosynthate to transport from stem to pods, so as to increase dry matter accumulation in pods, especially increase yield.
6. In the aspect of nitrogen accumulation and distribution, correlations between the amount of nitrogen accumulation at beginning flowering stage and NUEg, the total amount of nitrogen accumulation at maturing stage were not significant. But there were significantly negative correlations between nitrogen accumulation and distribution of stem and shell and NUEg, and there were significantly positive correlations between the amount of nitrogen of seed, NHI and NUEg. The studies of multiple regression analysis indicated that amounts of nitrogen of different organs at maturing stage significantly effected NUEg. To increase NUEg, increasing nitrogen accumulation of seeds was important, and at the same time decreasing nitrogen accumulation of stem and shell was also important.
7. There were different effects of nitrogen fertilizer on NUEg. Results showed that the number of different types (L→L, M→L, M→M, M→H, H→M, H→H) was 14,8,21,6,19 and 30, respectively. Though studying variety characteristics of different types, results showed that in the varieties from low type to high type by nitrogen application (M→H), added values of the number of seeds per pod, yield and the total number of seeds were larger, arid added values of nitrogen accumulation of stem and shell, ratio of nitrogen accumulation of stem and shell to total nitrogen accumulation were smaller. Decreasing values of ratio of nitrogen of seed to total nitrogen accumulation were smaller. The trends of the varieties from high type to low type by nitrogen application (M→L, H→M) were opposite.
8. Through testing NUEg of six parents and their F1 combinations from 6×6 complete double allele crosses, results showed that positive heterosis of NUEg was significant. NUEg were likely affected by additive effects, dominance effects and cytoplasm effects by variance analysis of combining ability. The correlation analysis of parents and F1 combinations showed that famale selection was important. In the aspect of heritability, results showed that broad heritability were higher in different treatments. Additive effects and dominance effects were all significant in low nitrogen fertilizer and dominance effects were significant in high nitrogen fertilizer.
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