杂交稻米质与生理生化特性的遗传及相互关系研究
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摘要
本研究选用稻米RVA谱特征值差异显著的深95A、389A、中3A、T98A、五丰A、炳1A等6个籼型杂交稻不育系和R031、湘恢059、湘农恢076、优恢036、R342等5个恢复系,按不完全双列杂交设计配制了的30个杂交稻组合。以这些杂交稻组合及其亲本和天优华占(对照)为A组材料,以直链淀粉含量差异显著的62个籼稻品种(组合)为B组材料。以A组材料研究了杂交稻RVA谱特征值及光合生理生化特性的杂种优势、配合力和遗传特性及相互关系；以A、B两组材料研究了杂交稻RVA谱特征值与主要米质性状间的关系。主要结果如下：
1.杂交稻RVA谱特征值的峰值粘度存在超高亲优势,崩解值、消减值、回复值、最低粘度和最终粘度存在正向中亲优势；RVA谱特征值存在极显著的组合间遗传差异,其中变异系数最大的是消减值,最小的是糊化开始温度；基因加性效应和非加性效应对杂交稻RVA谱特征值均存在极显著的影响,以亲本基因的加性效应为主(糊化开始温度除外)：不育系GCA方差远远大于父本GCA方差和组合SCA方差,表明杂交稻RVA谱特征值主要受不育系的影响；杂交稻RVA谱特征值的竞争优势与母本GCA呈极显著正相关,且决定系数较高,表明不育系的一般配合力高,其所配杂交稻组合的竞争优势也强。
2.不同水稻品种(组合)的稻米RVA谱特征值与主要米质性状间的关系表明：精米的长/宽、精米长与峰值粘度、崩解值和糊化开始温度极显著负相关,与其它RVA谱特征值显著或极显著正相关；透明度与峰值粘度和崩解值极显著正相关,与消减值、回复值和峰值时间显著或极显著负相关。因此,优质籼型杂交稻品种(组合)选育,不宜过份追求长粒型特征,而应注重透明度的选择；蒸煮食味和营养品质与RVA谱特征值关系密切,RVA谱特征值能反应米质的优劣。直链淀粉含量越高,其崩解值越小,消减值和回复值越大。RVA谱特征值能反映中低(15%≤AC<17%)及中等AC (17%≤AC≤22%)含量相似的水稻品种间的蒸煮食味品质的差异。
3.净光合速率、气孔导度、蒸腾速率,Fm、Fv/Fm、Y(NPQ)、NPQ、qN、qP和qL存在杂种优势；光合特性和叶绿素荧光参数存在极显著的组合间差异,主要受非加性效应影响,Fm、Y(Ⅱ)、Y(NPQ)和气孔导度受不育系的影响显著,叶绿素a和叶绿素b受恢复系的影响显著；净光合速率、Fv/Fm、Y(Ⅱ)和Y(NPQ)的竞争优势与母本GCA呈显著或极显著正相关,且相关系数较大,表明不育系GCA高,其所配杂交稻组合的光合能力竞争优势也强。
4. SOD、MDA及根系活力的杂种优势并不普遍存在；SOD、MDA及根系活力存在极显著的组合间差异,亲本的遗传力低于20%,广义遗传力高,主要受非加性效应影响：SOD、根系活力的竞争优势与母本GCA分别呈显著或极显著正相关,表明杂交稻母本一般配合力高,其所配杂交稻组合的SOD和根系活力的竞争优势也强。
5.稻米RVA谱特征值与植株生理生化特性的关系表现为：峰值粘度和崩解值与气孔导度分别呈极显著正相关,糊化开始温度与蒸腾速率呈显著正相关；RVA谱特征值与叶绿素荧光参数的关系表现为：Y(Ⅱ)与峰值粘度、崩解值、糊化开始温度呈显著或极显著正相关,与其它RVA谱特征值呈显著或极显著负相关。Y(NPQ)和qN与峰值粘度和崩解值呈显著负相关,与消减值和回复值呈显著正相关。表明PS(Ⅱ)天线色素吸收的光能用于光化学电子传递的份额越多,以热的形式耗散的光能越少,光能实际利用率高,其籽粒充实度好,蒸煮食味品质优良;Fv/Fm与气孔导度、胞间CO2浓度、叶绿素含量分别呈显著或极显著正相关。表明潜在光合效率高的杂交稻组合有气孔导度大、胞间CO2浓度高、叶绿素含量高的特点。
This study was conducted via6indica hybrid rice sterile lines:SHEN95A,389A, ZHONG3A, T98A, WUFENG A and BING1A, and5indica hybrid rice restorer lines: R031, XIANGHUI059, XIANG NONGHUI076, YOUHUI036and R342, which have significant difference in starch RVA profile properties. Thirty different hybrid rice combinations were derived from above hybrid rice lines based on incomplete diallel cross. Two sets of materials were prepared for this study, these hybrid rice combinations and their parents together with Tian You Hua Zhan(Control) were set up as group A; the62indica varieties or hybrid rice combinations with significant difference in amylose contents were set up as group B. The heterosis and combining ability, inheritance traits of starch RVA profile properties, physio-biochemical characteristics and the correlations between RVA profile properties and physio-biochemical characteristics in indica hybrid rice were analyzed for samples of group A, and the correlations between RVA profile properties and the main rice qualities also were analyzed for samples of group B. The main results are listed below:
1. There were significant over high-parent heterosis in peak viscosity and over mid-parent heterosis in breakdown, setback, consistency, trough viscosity and final viscosity. Results showed that there were highly significant genetic diversities of the RVA profile properties in hybrid rice combinations, among those RVA profile properties setback had the widest range of variations while the pasting temperature had the narrowest range of variations. The starch viscosity properties in hybrid rice combinations were significantly affected by gene additive effects and non-additive effects, mainly determined by gene additive effects of parents (not covering pasting temperature). Among all the genotype variances, the GCA variance of sterile lines was far higher than that of both GCA variance in restorer lines and SCA variance in hybrid rice combinations, indicating that the starch viscosity properties in hybrid rice combinations were mainly affected by the sterile lines. Results also showed that there were significant positive correlations between the competitive advantages of hybrid rice combinations and the GCA of sterile lines in RVA profile properties with higher determination coefficients, indicating that higher general combining ability of the sterile lines had the higher competitive advantages from those hybrid rice combinations would be obtained.
2. The correlations between rice starch viscosity properties and main rice qualities from different combinations were analyzed. There were significant negative correlations between the length, the rate of length and width of polished rice and peak viscosity, breakdown and pasting temperature from different combinations, but significant positive correlations to other RVA profile properties were presented; Transparency was found to have significant positive correlations to peak viscosity and breakdown, but to have significant negative correlations to setback, consistency and peak time. In this case, the rice variety seeds with better transparency should be preferred and the ones with longer length should not be excessively focused on during any selection and breeding activity of high-quality indica hybrid rice varieties (combinations). Results also indicated that cooking/eating and nutritional qualities in different rice varieties (combinations) had close correlations with their RVA profile properties, and the superior or inferior quality of rice could be highly indicated by their RVA profile properties. The rice varieties (combinations) with higher AC content, their breakdown would be smaller while their setback and consistency would be bigger. RVA profile properties would reflect the difference in cooking and eating quality among hybrid rice varieties with similar AC content, from low-mid-level (15%≤AC<17%) to mid-level (17%≤AC≤22%).
3. The net photosynthetic rate, stomatal conductance, transpiration rate, Fm、Fv/Fm、 Y(NPQ)、NPQ、qN、qP and qL showed positive effects over mid-parent heterosis. Significant genetic diversities were found in photosynthesis traits and chlorophyll fluorescence parameters of hybrid rice combinations. Photosynthesis traits and chlorophyll fluorescence parameters were mainly affected by non-additive effects. Fm, Y (Ⅱ), Y (NPQ) and stomatal conductance were significantly affected by the sterile lines while chlorophyll a and chlorophyll b were significantly affected by the restorer lines. The net photosynthetic rate, Fv/Fm, Y (Ⅱ) and Y (NPQ) of competitive advantages in hybrid rice combinations were significantly correlated to the GCA of the sterile lines, indicating that the sterile lines with higher GCA would obtain a better competitive advantage of their combinations.
4. Heterosis in SOD, MDA and root activity of hybrid rice combinations were not universal. Significant genetic diversities were found in SOD, MDA and root activity among different hybrid rice combinations with below20%heritability from parents and higher broad heritability. SOD, MDA and root activity were mainly affected by non-additive effects. The competitive advantages of SOD and root activity in hybrid rice combinations were significantly correlated to their GCA effects respectively, indicating that the sterile lines with higher GCA effects would obtain a better competitive advantage of their combinations.
5. The correlations between RVA profile properties and Physio-biochemical characteristics indicated that there were significant positive correlations between the peak viscosity, breakdown and stomatal conductance. There were positive correlations between pasting temperature and the transpiration rate. The correlations between RVA profile properties and chlorophyll fluorescence parameters showed Y (Ⅱ) had significant positive correlations to peak viscosity, breakdown, pasting temperature but negative correlations to other RVA profile properties; Y (NPQ) and qN had significant negative correlations to peak viscosity and breakdown but significant positive correlations to setback and consistency. The analyses indicated that PS(Ⅱ) antenna pigment of hybrid rice combinations absorbed light energy that the more it transmitted by photochemical electron, the less it dissipated in form of heat, and the actual utilization rate of light energy would be higher, which would result in better grain filling and higher quality in cooking and eating etc. There were significant positive correlations between stomatal conductance, intercellular CO2concentration, chlorophyll and carotenoid content respectively. The results indicated that the hybrid rice combinations would have higher potential photosynthesis efficiency due to bigger stomatal conductance, higher intercellular CO2concentration and chlorophyll content.
1.杂交稻RVA谱特征值的峰值粘度存在超高亲优势,崩解值、消减值、回复值、最低粘度和最终粘度存在正向中亲优势；RVA谱特征值存在极显著的组合间遗传差异,其中变异系数最大的是消减值,最小的是糊化开始温度；基因加性效应和非加性效应对杂交稻RVA谱特征值均存在极显著的影响,以亲本基因的加性效应为主(糊化开始温度除外)：不育系GCA方差远远大于父本GCA方差和组合SCA方差,表明杂交稻RVA谱特征值主要受不育系的影响；杂交稻RVA谱特征值的竞争优势与母本GCA呈极显著正相关,且决定系数较高,表明不育系的一般配合力高,其所配杂交稻组合的竞争优势也强。
2.不同水稻品种(组合)的稻米RVA谱特征值与主要米质性状间的关系表明：精米的长/宽、精米长与峰值粘度、崩解值和糊化开始温度极显著负相关,与其它RVA谱特征值显著或极显著正相关；透明度与峰值粘度和崩解值极显著正相关,与消减值、回复值和峰值时间显著或极显著负相关。因此,优质籼型杂交稻品种(组合)选育,不宜过份追求长粒型特征,而应注重透明度的选择；蒸煮食味和营养品质与RVA谱特征值关系密切,RVA谱特征值能反应米质的优劣。直链淀粉含量越高,其崩解值越小,消减值和回复值越大。RVA谱特征值能反映中低(15%≤AC<17%)及中等AC (17%≤AC≤22%)含量相似的水稻品种间的蒸煮食味品质的差异。
3.净光合速率、气孔导度、蒸腾速率,Fm、Fv/Fm、Y(NPQ)、NPQ、qN、qP和qL存在杂种优势；光合特性和叶绿素荧光参数存在极显著的组合间差异,主要受非加性效应影响,Fm、Y(Ⅱ)、Y(NPQ)和气孔导度受不育系的影响显著,叶绿素a和叶绿素b受恢复系的影响显著；净光合速率、Fv/Fm、Y(Ⅱ)和Y(NPQ)的竞争优势与母本GCA呈显著或极显著正相关,且相关系数较大,表明不育系GCA高,其所配杂交稻组合的光合能力竞争优势也强。
4. SOD、MDA及根系活力的杂种优势并不普遍存在；SOD、MDA及根系活力存在极显著的组合间差异,亲本的遗传力低于20%,广义遗传力高,主要受非加性效应影响：SOD、根系活力的竞争优势与母本GCA分别呈显著或极显著正相关,表明杂交稻母本一般配合力高,其所配杂交稻组合的SOD和根系活力的竞争优势也强。
5.稻米RVA谱特征值与植株生理生化特性的关系表现为：峰值粘度和崩解值与气孔导度分别呈极显著正相关,糊化开始温度与蒸腾速率呈显著正相关；RVA谱特征值与叶绿素荧光参数的关系表现为：Y(Ⅱ)与峰值粘度、崩解值、糊化开始温度呈显著或极显著正相关,与其它RVA谱特征值呈显著或极显著负相关。Y(NPQ)和qN与峰值粘度和崩解值呈显著负相关,与消减值和回复值呈显著正相关。表明PS(Ⅱ)天线色素吸收的光能用于光化学电子传递的份额越多,以热的形式耗散的光能越少,光能实际利用率高,其籽粒充实度好,蒸煮食味品质优良;Fv/Fm与气孔导度、胞间CO2浓度、叶绿素含量分别呈显著或极显著正相关。表明潜在光合效率高的杂交稻组合有气孔导度大、胞间CO2浓度高、叶绿素含量高的特点。
This study was conducted via6indica hybrid rice sterile lines:SHEN95A,389A, ZHONG3A, T98A, WUFENG A and BING1A, and5indica hybrid rice restorer lines: R031, XIANGHUI059, XIANG NONGHUI076, YOUHUI036and R342, which have significant difference in starch RVA profile properties. Thirty different hybrid rice combinations were derived from above hybrid rice lines based on incomplete diallel cross. Two sets of materials were prepared for this study, these hybrid rice combinations and their parents together with Tian You Hua Zhan(Control) were set up as group A; the62indica varieties or hybrid rice combinations with significant difference in amylose contents were set up as group B. The heterosis and combining ability, inheritance traits of starch RVA profile properties, physio-biochemical characteristics and the correlations between RVA profile properties and physio-biochemical characteristics in indica hybrid rice were analyzed for samples of group A, and the correlations between RVA profile properties and the main rice qualities also were analyzed for samples of group B. The main results are listed below:
1. There were significant over high-parent heterosis in peak viscosity and over mid-parent heterosis in breakdown, setback, consistency, trough viscosity and final viscosity. Results showed that there were highly significant genetic diversities of the RVA profile properties in hybrid rice combinations, among those RVA profile properties setback had the widest range of variations while the pasting temperature had the narrowest range of variations. The starch viscosity properties in hybrid rice combinations were significantly affected by gene additive effects and non-additive effects, mainly determined by gene additive effects of parents (not covering pasting temperature). Among all the genotype variances, the GCA variance of sterile lines was far higher than that of both GCA variance in restorer lines and SCA variance in hybrid rice combinations, indicating that the starch viscosity properties in hybrid rice combinations were mainly affected by the sterile lines. Results also showed that there were significant positive correlations between the competitive advantages of hybrid rice combinations and the GCA of sterile lines in RVA profile properties with higher determination coefficients, indicating that higher general combining ability of the sterile lines had the higher competitive advantages from those hybrid rice combinations would be obtained.
2. The correlations between rice starch viscosity properties and main rice qualities from different combinations were analyzed. There were significant negative correlations between the length, the rate of length and width of polished rice and peak viscosity, breakdown and pasting temperature from different combinations, but significant positive correlations to other RVA profile properties were presented; Transparency was found to have significant positive correlations to peak viscosity and breakdown, but to have significant negative correlations to setback, consistency and peak time. In this case, the rice variety seeds with better transparency should be preferred and the ones with longer length should not be excessively focused on during any selection and breeding activity of high-quality indica hybrid rice varieties (combinations). Results also indicated that cooking/eating and nutritional qualities in different rice varieties (combinations) had close correlations with their RVA profile properties, and the superior or inferior quality of rice could be highly indicated by their RVA profile properties. The rice varieties (combinations) with higher AC content, their breakdown would be smaller while their setback and consistency would be bigger. RVA profile properties would reflect the difference in cooking and eating quality among hybrid rice varieties with similar AC content, from low-mid-level (15%≤AC<17%) to mid-level (17%≤AC≤22%).
3. The net photosynthetic rate, stomatal conductance, transpiration rate, Fm、Fv/Fm、 Y(NPQ)、NPQ、qN、qP and qL showed positive effects over mid-parent heterosis. Significant genetic diversities were found in photosynthesis traits and chlorophyll fluorescence parameters of hybrid rice combinations. Photosynthesis traits and chlorophyll fluorescence parameters were mainly affected by non-additive effects. Fm, Y (Ⅱ), Y (NPQ) and stomatal conductance were significantly affected by the sterile lines while chlorophyll a and chlorophyll b were significantly affected by the restorer lines. The net photosynthetic rate, Fv/Fm, Y (Ⅱ) and Y (NPQ) of competitive advantages in hybrid rice combinations were significantly correlated to the GCA of the sterile lines, indicating that the sterile lines with higher GCA would obtain a better competitive advantage of their combinations.
4. Heterosis in SOD, MDA and root activity of hybrid rice combinations were not universal. Significant genetic diversities were found in SOD, MDA and root activity among different hybrid rice combinations with below20%heritability from parents and higher broad heritability. SOD, MDA and root activity were mainly affected by non-additive effects. The competitive advantages of SOD and root activity in hybrid rice combinations were significantly correlated to their GCA effects respectively, indicating that the sterile lines with higher GCA effects would obtain a better competitive advantage of their combinations.
5. The correlations between RVA profile properties and Physio-biochemical characteristics indicated that there were significant positive correlations between the peak viscosity, breakdown and stomatal conductance. There were positive correlations between pasting temperature and the transpiration rate. The correlations between RVA profile properties and chlorophyll fluorescence parameters showed Y (Ⅱ) had significant positive correlations to peak viscosity, breakdown, pasting temperature but negative correlations to other RVA profile properties; Y (NPQ) and qN had significant negative correlations to peak viscosity and breakdown but significant positive correlations to setback and consistency. The analyses indicated that PS(Ⅱ) antenna pigment of hybrid rice combinations absorbed light energy that the more it transmitted by photochemical electron, the less it dissipated in form of heat, and the actual utilization rate of light energy would be higher, which would result in better grain filling and higher quality in cooking and eating etc. There were significant positive correlations between stomatal conductance, intercellular CO2concentration, chlorophyll and carotenoid content respectively. The results indicated that the hybrid rice combinations would have higher potential photosynthesis efficiency due to bigger stomatal conductance, higher intercellular CO2concentration and chlorophyll content.
引文
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