东乡野生稻抗旱鉴定指标的筛选和QTL定位研究
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摘要
干旱是导致水稻减产的重要非生物因素之一,因而培育和筛选抗旱性强的水稻品种是保障粮食供给和水稻生产可持续发展的关键所在。为此,本论文以协青早B//协青早B/东乡野生稻BC1F1o回交重组自交系群体为研究对象进行抗旱性研究,分别测定干旱胁迫和对照条件下苗期和孕穗期的形态、生理和产量等性状指标,并结合隶属函数、主成分分析和逐步回归分析等方法对群体抗旱性进行综合分析,建立水稻苗期和孕穗期抗旱评价体系,同时利用已构建的遗传图谱对东乡野生稻苗期和孕穗期两个生育期抗旱相关性状进行QTL定位分析,深入探讨东乡野生稻在干旱环境下的抗旱分子作用机理。论文获得的主要结果如下：
1.采用PEG6000模拟干旱法,在苗期对旱稻、群体及亲本进行抗旱鉴定及综合指标筛选,测定了最大根长、根基粗、根数、茎长、根鲜重、根干重、萎蔫率等10个性状,并利用主成分分析和逐步回归分析法对各株系抗旱性进行综合评价和抗旱鉴定指标筛选。研究结果表明,与抗旱性密切相关的3个指标分别为最大根长、根基粗和萎蔫率,筛选出的这三个指标可作为水稻苗期抗旱鉴定综合指标。
2.根据苗期抗旱性鉴定指标的筛选结果,利用该群体已建立的遗传图谱,对包括最大根长、根基粗和萎蔫率等共10个性状进行QTL定位分析。试验发现在干旱环境下,8个性状共定位到16个QTLs,最大根长、根基粗和萎蔫率这3个性状分别检测到3个、1个和4个QTLs。最大根长的qMRL-4和qMRL-7的加性效应为负值,说明这2个QTLs均来自于东乡野生稻,其总贡献率为11.87%。影响根基粗的QTL(qBRT-7)位于第7染色体的RM11～RM18标记区域内,其加性效应为负值,贡献率为24.11%,表明该位点来自于“东野”并且可能是个主效QTL。萎蔫率的4个QTLs,分别位于第1、7和11染色体上,其加性效应均为正值,其他苗期性状的大部分QTLs加性效应值也为正值,说明这些位点的增效等位基因来自于亲本协青早B,可能是由其自身遗传控制的。
3.采用传统干旱棚田间鉴定法评价了上述群体孕穗期的抗旱性,并测定了参试材料在正常水分和干旱胁迫下的株高、单株有效穗数、穗长、每穗总粒数、实粒数、千粒重、结实率、单株产量等性状值。结果表明,相比于非干旱胁迫环境,水稻的性状值在干旱胁迫条件下均有所降低,其中穗长对干旱胁迫表现不敏感,而千粒重对水分胁迫表现为敏感,结实率和单株产量均对干旱胁迫表现为极敏感。此外,实验采用隶属函数值和主成分分析法获得各株系的抗旱综合值,进一步以抗旱综合值为因变量,9个指标性状值为自变量,通过逐步回归分析法建立了评价孕穗期抗旱性的最优回归方程,筛选出结实率、单株产量和千粒重3个抗旱性鉴定指标。
4.试验对群体孕穗期的结实率、单株产量和千粒重等多个性状进行QTL定位,共检测到32个QTLs位点,分别位于水稻第1、2、3、7、8、9、10、11染色体上。结实率、单株产量和千粒重3个性状分别定位到了4个、2个和8个QTLs,其中qSR-1和qSR-10的贡献率较大,分别为41.30%和43.48%,加性效应值分别为-12.04、-11.97,可能是两个主效QTL。qGWP-1、qGWP-1a的加性效应值分别为-6.17、-5.16,贡献率分别为33.93%和23.35%,也可能是两个主效QTLs,千粒重的QTLs的加性效应均为负值。由此可见,3个性状的大部分QTLs的增效等位基因均来自“东野”,可见是东乡野生稻等位基因的表达结果,说明东乡野生稻含有抗旱基因。
5.对水稻苗期和孕穗期的抗旱综合值(D值)进行相关性分析后发现,二者的相关系数为0.935,达到极显著水平,这说明了苗期与孕穗期抗旱性关系密切,并获得了10个在两个时期都表现强抗旱的株系。
6.比较分析苗期与孕穗期的QTL定位结果,发现在苗期与孕穗期都存在一因多效或紧密连锁现象,而且控制孕穗期相关性状的QTL热点区也是控制苗期相关性状的QTL区域,再次说明了苗期和孕穗期的抗旱性紧密相关。
7.初步探讨了强抗旱株系和弱抗旱株系孕穗期水分胁迫下的的生理生化特性。研究结果表明,在水分胁迫下,各株系的可溶性糖和脯氨酸含量随着干旱加剧而增加,而且抗旱性强的株系增加的幅度大于抗旱性弱的株系。丙二醛含量也随着干旱加剧而增加,但抗旱性强的株系增加幅度小于抗旱性弱的株系。3个生理指标相对值与株系抗旱系数的相关性分析表明二者显著相关。
Drought is one of important abiotic factors for the decrease of rice yield. Screening of rice varieties with drought-resistance therefore is the key to the sustainable development of rice production and food supply. In this thesis, a backcross inbred lines (BIL) population including234lines was chosen as the research object, which was derived from an interspecies cross XieqingzaoB(Oryza sativa)//XieqingzaoB/Dongxiang wild rice (O.rufipogon). The morphological, physiological and yield characteristics were investigated in detail under both normal and drought stress conditions at seedling and booting stages. Through subordinate funcation analysis, principal component analysis and step wise regression analysis, the drought-resistance of population was analyzed comprehensively. The assessment index system of drought-resistance was established at both seedling and booting stages. Meanwhile, by using constructed molecular linkage map, we mapped the drought-resistance QTL locus from Dongxiang wild rice at both seedling and booting stages. These results will contribute to futher explore the molecular mechanism of the drought resistance of Dongxiang wild rice.The main results were summarized as follows:
1. Upland rice, BILs and their parents were employed to evaluate drought-resistance at seedling stage under PEG6000simulated drought stress. Ten traits were recorded for drought-resistance, such as MRL, NR, SL, DRW, FRW, DSW, BRT, WR,DR. Comprehensive assessment of drought-resistance based on the principal component analysis (PCA) was applied. The result of the stepwise regression analysis demonstrated that MRL、BRT and WR significantly were influenced under drought condition. Therefore, the three index could be used to assess the drought-resistance at seedling stage.
2. According to the results of index screening of drought-resistance, we used the established genetic linkage map to analyze the drought-resistance related QTLs for ten traits at seedling stage, including MRL, BRT, WL, etc. A total of16QTLs were detected in eight traits under drought condition. Among them, there were3QTLs for MRL,1QTL for BRT and4QTLs for WR. The addictive genetic effect of qMRL-4, qMRL-7and qBRT-7was negative, indicating the alleles from DongXiang common wild rice. The general contributions of qMRL-4and qMRL-7were11.87%and the contribution of qBRT-7was24.11%, which could be a large effect QTL.Four QTLs detected for WR were located on1,7and11chromosomes,which addictive effect was positive, indicating the alleles from Xieqingzao B and decreasing the drought-resistance ability of population. Meanwhile, the additive genetic effect of QTLs in other traits were all positive, indicating these locus from parent Xieqingzao B. The other traits variations between normal condition and drought condition might be caused by their own genetics.
3. Drought-resistance of the BILs population was appraised by the traditional method at booting stage in rainless shed. Nine traits related with drought-resistance were analyzed under drought stress condition and normal condition, including EIPN, SP, FSP, PL, PD, TGW, SR and GWP. The results showed, to different extent, the nine traits were all influenced under drought condition, among which the lines with weak drought-resistance were more seriously influenced than that with strong drought-resistance. PL was not sensitive to drought stress, otherwise TGW expressed sensitive to drought stress.Other yield traits were very sensitive. In addition, by applying subordinate funcation analysis, principal component analysis, and step wise regression analysis, we screened three index, including SR, GWP and TGW, which could be used as evaluation index at booting stage.
4. A total of32QTLs for yield traits were detected at booting stage,which located on chromosome1,2,3,7,8,10,11, respectively. Among these traits, SR, GWP and TGW were respectively identified4,2and8QTLs.The contribution of four main effect QTLs including qSR-1, qSR-10, qGWP-1and qGWP-1a was41.30%、43.48%、33.93%and23.35%,respectively.Their addictive effect of QTLs for SR、GWP and TGW were all negative, indicating the alleles from DXWR. These results showed that the most QTLs of the three traits with enhancing alleles came from DXWR, suggesting DXWR had drought-resistance genes.
5. We analyzed the correlation between comprehensive index (D value) at seedling and booting stages. The correlation coefficient was0.935, reaching a extremely remarkable level. The lines with strong drought-resistance at booting stage were identical with the lines at seedling stage. Meanwhile, the lines with weak drought-resistance at seedling stage were identical with the lines at booting stage. It indicated that the drought resistance at seedling stage was closely related to that at booting stage. Moreover, ten lines with strong drought-resistance at two stages had been obtained.
6. There were pleiotropic or tightly linked QTL regions respectively at two stages by comparing the QTL mapping results of seedling and booting stage. And some QTL regions at booting stage were found to be the co-localized regions at seedling, indicating the seedling was associated with booting stage closely again.
7. The physiology and biochemistry properties were studied between strong and weak drought-resistance lines at booting stage under drought condition. The results indicated that soluble sugar and proline content accelerated more rapidly in stronger drought-resistance lines than that in weaker ones with water stress aggravated. The contents of MDA increased in better drought-resistance lines obviously less than that in weaker ones. The above results indicated that the three index were remarkably related to drought-resistance.
1.采用PEG6000模拟干旱法,在苗期对旱稻、群体及亲本进行抗旱鉴定及综合指标筛选,测定了最大根长、根基粗、根数、茎长、根鲜重、根干重、萎蔫率等10个性状,并利用主成分分析和逐步回归分析法对各株系抗旱性进行综合评价和抗旱鉴定指标筛选。研究结果表明,与抗旱性密切相关的3个指标分别为最大根长、根基粗和萎蔫率,筛选出的这三个指标可作为水稻苗期抗旱鉴定综合指标。
2.根据苗期抗旱性鉴定指标的筛选结果,利用该群体已建立的遗传图谱,对包括最大根长、根基粗和萎蔫率等共10个性状进行QTL定位分析。试验发现在干旱环境下,8个性状共定位到16个QTLs,最大根长、根基粗和萎蔫率这3个性状分别检测到3个、1个和4个QTLs。最大根长的qMRL-4和qMRL-7的加性效应为负值,说明这2个QTLs均来自于东乡野生稻,其总贡献率为11.87%。影响根基粗的QTL(qBRT-7)位于第7染色体的RM11～RM18标记区域内,其加性效应为负值,贡献率为24.11%,表明该位点来自于“东野”并且可能是个主效QTL。萎蔫率的4个QTLs,分别位于第1、7和11染色体上,其加性效应均为正值,其他苗期性状的大部分QTLs加性效应值也为正值,说明这些位点的增效等位基因来自于亲本协青早B,可能是由其自身遗传控制的。
3.采用传统干旱棚田间鉴定法评价了上述群体孕穗期的抗旱性,并测定了参试材料在正常水分和干旱胁迫下的株高、单株有效穗数、穗长、每穗总粒数、实粒数、千粒重、结实率、单株产量等性状值。结果表明,相比于非干旱胁迫环境,水稻的性状值在干旱胁迫条件下均有所降低,其中穗长对干旱胁迫表现不敏感,而千粒重对水分胁迫表现为敏感,结实率和单株产量均对干旱胁迫表现为极敏感。此外,实验采用隶属函数值和主成分分析法获得各株系的抗旱综合值,进一步以抗旱综合值为因变量,9个指标性状值为自变量,通过逐步回归分析法建立了评价孕穗期抗旱性的最优回归方程,筛选出结实率、单株产量和千粒重3个抗旱性鉴定指标。
4.试验对群体孕穗期的结实率、单株产量和千粒重等多个性状进行QTL定位,共检测到32个QTLs位点,分别位于水稻第1、2、3、7、8、9、10、11染色体上。结实率、单株产量和千粒重3个性状分别定位到了4个、2个和8个QTLs,其中qSR-1和qSR-10的贡献率较大,分别为41.30%和43.48%,加性效应值分别为-12.04、-11.97,可能是两个主效QTL。qGWP-1、qGWP-1a的加性效应值分别为-6.17、-5.16,贡献率分别为33.93%和23.35%,也可能是两个主效QTLs,千粒重的QTLs的加性效应均为负值。由此可见,3个性状的大部分QTLs的增效等位基因均来自“东野”,可见是东乡野生稻等位基因的表达结果,说明东乡野生稻含有抗旱基因。
5.对水稻苗期和孕穗期的抗旱综合值(D值)进行相关性分析后发现,二者的相关系数为0.935,达到极显著水平,这说明了苗期与孕穗期抗旱性关系密切,并获得了10个在两个时期都表现强抗旱的株系。
6.比较分析苗期与孕穗期的QTL定位结果,发现在苗期与孕穗期都存在一因多效或紧密连锁现象,而且控制孕穗期相关性状的QTL热点区也是控制苗期相关性状的QTL区域,再次说明了苗期和孕穗期的抗旱性紧密相关。
7.初步探讨了强抗旱株系和弱抗旱株系孕穗期水分胁迫下的的生理生化特性。研究结果表明,在水分胁迫下,各株系的可溶性糖和脯氨酸含量随着干旱加剧而增加,而且抗旱性强的株系增加的幅度大于抗旱性弱的株系。丙二醛含量也随着干旱加剧而增加,但抗旱性强的株系增加幅度小于抗旱性弱的株系。3个生理指标相对值与株系抗旱系数的相关性分析表明二者显著相关。
Drought is one of important abiotic factors for the decrease of rice yield. Screening of rice varieties with drought-resistance therefore is the key to the sustainable development of rice production and food supply. In this thesis, a backcross inbred lines (BIL) population including234lines was chosen as the research object, which was derived from an interspecies cross XieqingzaoB(Oryza sativa)//XieqingzaoB/Dongxiang wild rice (O.rufipogon). The morphological, physiological and yield characteristics were investigated in detail under both normal and drought stress conditions at seedling and booting stages. Through subordinate funcation analysis, principal component analysis and step wise regression analysis, the drought-resistance of population was analyzed comprehensively. The assessment index system of drought-resistance was established at both seedling and booting stages. Meanwhile, by using constructed molecular linkage map, we mapped the drought-resistance QTL locus from Dongxiang wild rice at both seedling and booting stages. These results will contribute to futher explore the molecular mechanism of the drought resistance of Dongxiang wild rice.The main results were summarized as follows:
1. Upland rice, BILs and their parents were employed to evaluate drought-resistance at seedling stage under PEG6000simulated drought stress. Ten traits were recorded for drought-resistance, such as MRL, NR, SL, DRW, FRW, DSW, BRT, WR,DR. Comprehensive assessment of drought-resistance based on the principal component analysis (PCA) was applied. The result of the stepwise regression analysis demonstrated that MRL、BRT and WR significantly were influenced under drought condition. Therefore, the three index could be used to assess the drought-resistance at seedling stage.
2. According to the results of index screening of drought-resistance, we used the established genetic linkage map to analyze the drought-resistance related QTLs for ten traits at seedling stage, including MRL, BRT, WL, etc. A total of16QTLs were detected in eight traits under drought condition. Among them, there were3QTLs for MRL,1QTL for BRT and4QTLs for WR. The addictive genetic effect of qMRL-4, qMRL-7and qBRT-7was negative, indicating the alleles from DongXiang common wild rice. The general contributions of qMRL-4and qMRL-7were11.87%and the contribution of qBRT-7was24.11%, which could be a large effect QTL.Four QTLs detected for WR were located on1,7and11chromosomes,which addictive effect was positive, indicating the alleles from Xieqingzao B and decreasing the drought-resistance ability of population. Meanwhile, the additive genetic effect of QTLs in other traits were all positive, indicating these locus from parent Xieqingzao B. The other traits variations between normal condition and drought condition might be caused by their own genetics.
3. Drought-resistance of the BILs population was appraised by the traditional method at booting stage in rainless shed. Nine traits related with drought-resistance were analyzed under drought stress condition and normal condition, including EIPN, SP, FSP, PL, PD, TGW, SR and GWP. The results showed, to different extent, the nine traits were all influenced under drought condition, among which the lines with weak drought-resistance were more seriously influenced than that with strong drought-resistance. PL was not sensitive to drought stress, otherwise TGW expressed sensitive to drought stress.Other yield traits were very sensitive. In addition, by applying subordinate funcation analysis, principal component analysis, and step wise regression analysis, we screened three index, including SR, GWP and TGW, which could be used as evaluation index at booting stage.
4. A total of32QTLs for yield traits were detected at booting stage,which located on chromosome1,2,3,7,8,10,11, respectively. Among these traits, SR, GWP and TGW were respectively identified4,2and8QTLs.The contribution of four main effect QTLs including qSR-1, qSR-10, qGWP-1and qGWP-1a was41.30%、43.48%、33.93%and23.35%,respectively.Their addictive effect of QTLs for SR、GWP and TGW were all negative, indicating the alleles from DXWR. These results showed that the most QTLs of the three traits with enhancing alleles came from DXWR, suggesting DXWR had drought-resistance genes.
5. We analyzed the correlation between comprehensive index (D value) at seedling and booting stages. The correlation coefficient was0.935, reaching a extremely remarkable level. The lines with strong drought-resistance at booting stage were identical with the lines at seedling stage. Meanwhile, the lines with weak drought-resistance at seedling stage were identical with the lines at booting stage. It indicated that the drought resistance at seedling stage was closely related to that at booting stage. Moreover, ten lines with strong drought-resistance at two stages had been obtained.
6. There were pleiotropic or tightly linked QTL regions respectively at two stages by comparing the QTL mapping results of seedling and booting stage. And some QTL regions at booting stage were found to be the co-localized regions at seedling, indicating the seedling was associated with booting stage closely again.
7. The physiology and biochemistry properties were studied between strong and weak drought-resistance lines at booting stage under drought condition. The results indicated that soluble sugar and proline content accelerated more rapidly in stronger drought-resistance lines than that in weaker ones with water stress aggravated. The contents of MDA increased in better drought-resistance lines obviously less than that in weaker ones. The above results indicated that the three index were remarkably related to drought-resistance.
引文
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