水稻抗旱相关重要农艺生理特性及其与耐盐性遗传重叠
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摘要
在亚洲许多地区,干旱和盐胁迫是限制水稻生产的最主要非生物因素。培育高产和抗逆的水稻品种或许是解决这一问题的最好方法。由于水稻抗旱和耐盐性是受多基因或多位点控制的数量性状,培育高产抗旱又耐盐的水稻品种十分困难。近10年来,我们在国际水稻研究所,通过分子标记辅助选育了大量的抗旱和耐盐的导入系和聚合系。为了解高产,抗旱以及耐盐之间的关系及其它们内在机理,本研究利用抗旱QTL聚合系从表型和生理的角度阐述了抗旱机理,并利用导入系阐述了抗旱和耐盐的遗传机制。
第一套试验材料包括48个聚合系。在严重大田干旱胁迫和灌溉条件下,以轮回亲本IR64为对照,连续三年分别在营养期和生殖期,对48个IR64背景的二轮抗旱QTL聚合系得抗旱性以及产量潜力,进行了鉴定评估。研究表明,相对于轮回亲本IR64,在生殖期旱胁迫条件下所有48个聚合系,在营养期胁迫条件下36个株系的抗旱性均显著提高(p＜0.05)。另外,在正常灌溉条件下23个株系产量显著高于IR64(p<0.05),其余25株系产量与IR64相近。通过对产量以及次级性状的比较分析发现,3种重要的抗旱机理相互协调提高聚合系的抗旱能力和产量潜力,最主要的是避旱性(drought avoidance),具体表现为在生殖期胁迫条件下,通过减少分蘖数和茎叶干物质量,提高水分利用效率,增加了有效穗数和育性,从而大大提高了收获指数和产量,这表明在干旱胁迫条件下,这些聚合系能够更有效利用土壤水分,确保重要功能的正常进行。而在正常灌溉条件下这些聚合系比IR64没有减少甚至更多的分蘖数和穗数。第二种机理是一种特殊的抗早性(drought tolerance),表现为在生殖期干旱胁迫下加速了“源”的有效转运和利用,显著提高收获指数,表现为所有的聚合系在三种条件下(正常,营养期胁迫,以及生殖期胁迫),均比IR64表现出了显著高的千粒重或者显著高的育性(P<0.05)。逃旱性(drought escape)是第三种机理,在生殖期,通过加速生长避免花期遭遇更严重干旱伤害；在营养期,加速生长为复水后后期库的积累提供了更多“源的供应”。同时我们发现,在具有相似抗旱性和产量潜力的株系间,次级性状表现差异较大,这为通过分子标记聚合这些有利性状提供了信息,从而可以进一步提高植株抗旱能力和产量潜力。因此胁迫条件下通过选择产量和优良次级性状相结合可以大大提高培育高产抗旱品种的效率。
为进一步了解这些聚合系的抗旱机理,尤其是在干旱胁迫条件下动态反应,选择12个或生殖期或营养期或兼而有之表现较好的株系,以IR64为对照,在温室条件下进行盆栽实验。研究发现,在营养期胁迫条件下,比较对照,拥有发达根系的株系29和较低单位面积蒸腾量(较厚叶片)的株系30,在土壤水分含量较低的情况下,均能保持较高蒸腾速率和光合作用,这样有效积累了地上地下部生物量,从而提高了水分利用效率。ABA的积累是其主要的信号调节机理。同时发现非ABA代谢途径也同样调节植株对水分匮缺的反应。在生殖期间歇性水分胁迫下,土壤水分的过快汲取导致了株系间表现相似,但复水后差异明显(p<0.01),株系30复水前水分利用率高,生物量积累多,恢复能力最强,而具有发达根系的line29并没有表现出强的恢复能力。根系解剖结构表明,株系29具有厚的木质部层和更多导管,这有利于土壤水分的吸收,但是在严重土壤水分胁迫下易造成导管“气穴”现象,这将极大降低根系对土壤水分的汲取能力。同时在田间旱胁迫下,我们尝试观察了植株的动态变化,结果再次证明,适时的开花期,高效水分利用或者发达根系,以及碳水化合物的有效转运等,协同调节是聚合系抗旱的机理所在。
在第二套实验中,利用两个以高产稻黄花占为背景BC1F3群体(HHZ/OM1723和HHZ/Teqing)3000个单株,在中国广东和国际水稻研究所进行了抗旱,耐盐和高产选择,选择到24个高产,36个极端抗旱和30个极端耐盐的导入系进行了卡方和多位点关联概率测验,以检测供体导入片段中与可能与高产、抗旱以及耐盐相关联的功能遗传单位(FGU)。根据连锁不平衡分析和最新数量遗传理论,共检测到53个FGU构成了抗旱和耐盐的遗传网络。在群体HHZ/OM1723中检测到28个FGU,其中9个在三种选择群体中检测到,14个在两种选择群体中,5个只在单个选择群体中检测到,而且有三个具有高导入的位点总是处于抗旱耐盐遗传网络的最上游,其中3个在三种遗传网络的最上游具有重叠。同样地,在群体HHZ／Teqing中,检测到25个FGU,其中19个在三种选择群体中检测到,4个只在两种选择群体中检测到,2个只在单个选择群体中检测到。同样地,总有两个位点处于高产、抗旱和耐盐遗传网络的最上游。而且同一个途径的位点间表现出高度正相关,不同途径间表现出独立或者负相关关系。但是比较发现,很少FGU在两个群体发生重叠,这表明大多数检测到的FGU来自于不同的供体。这与当前对抗旱和耐盐的研究结论一致,但这些遗传信息为我们通过聚合高度正相关或者独立的不同的功能遗传位点,进一步提高水稻的高产、抗旱以及耐盐水平,提供了有益的帮助,也为通过分子标记高通量检测抗旱耐盐QTL位点提供了重要的参考。
Abiotic stresses such as drought and salinity are very important factor limiting rice productivity in the rainfed and costal areas of Asia. Development and adoption of high-yielding and stress tolerant rice varieties are the best solution for solving the problems. However, it is well known that rice tolerances to both drought and salinity are controlled by large numbers of genes or quantitative trait loci (QTLs), making it difficult to breed superior rice varieties with good tolerances to both stresses. In past 10 years, many sets of introgression lines (ILs) and pyramiding lines (PLs) with significantly improved drought tolerance (DT) and/or salt tolerance (ST) have been developed using backcross breeding and molecular pyramiding at IRRI. The objectives of this study were:(1) to characterize the phenotypic and physiological traits associated with DT PLs and (2) to understand the genetic basis of DT and ST using ILs and molecular markers.
Two series experiments were performed in this study. In the first set of experiments,48 PLs, their parental ILs and recurrent parent, IR64 (check) were evaluated for their yield performances and related traits under severe drought stress at the reproductive stage (RS) and vegetative stage (VS) and irrigated control in three years to understand the relationship between drought tolerance (DT) and yield potential (YP) in rice and their underlying mechanisms. When compared to IR64, all PLs had significantly improved DT to RS and 36 PLs also had significantly improved DT to VS (p<0.05). In addition,23 PLs had higher YP than IR64 (p<0.05) and the remaining 25 PLs had similar YP as IR64 under the irrigated control. Detailed characterization of the PLs revealed 3 mechanisms that functioned together to contribute to their improved DT and/or YP. The most important mechanism was drought avoidance (DA) characterized with significantly higher biomass and harvest index (HI) of the PLs than IR64 under stress, suggesting the PLs' better abilities to maintain higher water status and function under stress. Because most PLs did not show any reduction in biomass under control, this type of DA resulted most likely from improved dehydration resistance. The second mechanism was efficient partitioning characterized by improved HI in all PLs as compared to IR64, resulting primarily from higher GW and/or SF under control and was the major constituent of the improved YP in the 17 PLs. Drought escape (DE) by accelerated heading under drought was the third mechanism that contributed to DT of the PLs to RS. The considerable variation in the measured traits among the PLs with similar levels of DT and YP implies the complex genetic control of the mechanisms for DT/YP and offers opportunities to further improve DT and/or YP by fine-tuning of QTLs/genes among the PLs using MAS. Finally, our results indicate that selection for yield plus some secondary traits under appropriate type(s) of stress and non-stress conditions similar to target environments are critically important for improving both DT and YP in rice.
A selected set of promising PLs was further analyzed for the dynamic responses of plant growth and water use to progressive water deficit at different growth stages in lysimeters under controlled phytotron conditions and rainfed lowland condition. In vegetative stage dry-down experiment, the PLs that performed best under drought were those that had the lowest threshold of transpiration response to soil moisture decrease (P<0.01), i.e. lines that kept stomata open longer or had large root system during the soil drying, and hence extracted more soil water, which resulted in higher shoot and root biomass, and an overall improvement of transpiration efficiency. ABA-dependant and ABA-independent signal pathway co-regulated the different response to the FTSW among the genotypes. In intermittent water stress, the sharp dry-down stress resulted in the similar performance between lines in the threshold of FTSW. The lines with higher water use efficiency (WUE) and large biomass at the end of stress showed good drought recovery ability, and thus produced higher spikelet fertility and grain yield. However, those lines produced large root biomass did not show superior drought recovery, probably due to the specific root anonomy structure. Under the lowland progressive water stress, the PLs produced high yield potential as the result of large biomass production (more tillers and panicles), while under drought, they improved WUE by balancing the transpiration and tiller and leaf area production, and performed higher yield, harvest index and spikelet fertility, but lower above ground shoot production. As well, the lines with large root system also conferred to drought tolerance in lowland condition, to some extent.
In the second set of experiments, two BC1F2 populations (1500plants) in the genetic background of a high-yielding indica variety, Huang-Hua-Zan, were screened under drought and salinity conditions at IRRI and China, resulting in development of 36 DT ILs and 30 salt tolerant (ST) ILs. These ILs were analyzed using X2 and multilocus probability tests to identify functional genetic units (FGUs) located in the donor segments associated with DT and ST. Together, a total of 53 FGUs associated with HY, DT and/or ST were identified in the ILs, forming 3 genetic networks underlying HY, DT and ST in rice. The first one detected in the HHZ/OM1723 population contained 28 FGUs, including 9 FGUs contributing to HY, DT, and ST,14 FGUs affecting 2 traits, and 5 FGUs that contributed to one trait only. Of these,3 loci (RM24, RM6 and RM130) appeared to be important as they had high frequencies of introgression and were placed in the upstream of the network based on the new molecular-quantitative genetics theory. Similarly, the genetic network identified in the HHZ/Teqing population had 25 FGUs, including 19 FGUs contributing to HY, DT, and ST,4 FGUs affecting 2 traits, and 2 FGUs that contributed to one trait only. Again, these FGUs included 2 important ones in the upstream of the genetic network. In contrast to the high genetic overlap among the three traits, especially between DT and ST, however, there were few shared loci identified in the HHZ/OM1723 and those in the HHZ/Teqing population, indicating the 2 donors contributed different sets of loci to the DT and/or ST in the selected ILs. Our results were in consistent with our current knowledge that DT and ST in rice are under complex genetic control, and provided useful information and materials for further improvement of DT and ST in rice by pathway pyramiding, and for detailed molecular dissection of DT and ST in rice using high throughput genomic tools.
第一套试验材料包括48个聚合系。在严重大田干旱胁迫和灌溉条件下,以轮回亲本IR64为对照,连续三年分别在营养期和生殖期,对48个IR64背景的二轮抗旱QTL聚合系得抗旱性以及产量潜力,进行了鉴定评估。研究表明,相对于轮回亲本IR64,在生殖期旱胁迫条件下所有48个聚合系,在营养期胁迫条件下36个株系的抗旱性均显著提高(p＜0.05)。另外,在正常灌溉条件下23个株系产量显著高于IR64(p<0.05),其余25株系产量与IR64相近。通过对产量以及次级性状的比较分析发现,3种重要的抗旱机理相互协调提高聚合系的抗旱能力和产量潜力,最主要的是避旱性(drought avoidance),具体表现为在生殖期胁迫条件下,通过减少分蘖数和茎叶干物质量,提高水分利用效率,增加了有效穗数和育性,从而大大提高了收获指数和产量,这表明在干旱胁迫条件下,这些聚合系能够更有效利用土壤水分,确保重要功能的正常进行。而在正常灌溉条件下这些聚合系比IR64没有减少甚至更多的分蘖数和穗数。第二种机理是一种特殊的抗早性(drought tolerance),表现为在生殖期干旱胁迫下加速了“源”的有效转运和利用,显著提高收获指数,表现为所有的聚合系在三种条件下(正常,营养期胁迫,以及生殖期胁迫),均比IR64表现出了显著高的千粒重或者显著高的育性(P<0.05)。逃旱性(drought escape)是第三种机理,在生殖期,通过加速生长避免花期遭遇更严重干旱伤害；在营养期,加速生长为复水后后期库的积累提供了更多“源的供应”。同时我们发现,在具有相似抗旱性和产量潜力的株系间,次级性状表现差异较大,这为通过分子标记聚合这些有利性状提供了信息,从而可以进一步提高植株抗旱能力和产量潜力。因此胁迫条件下通过选择产量和优良次级性状相结合可以大大提高培育高产抗旱品种的效率。
为进一步了解这些聚合系的抗旱机理,尤其是在干旱胁迫条件下动态反应,选择12个或生殖期或营养期或兼而有之表现较好的株系,以IR64为对照,在温室条件下进行盆栽实验。研究发现,在营养期胁迫条件下,比较对照,拥有发达根系的株系29和较低单位面积蒸腾量(较厚叶片)的株系30,在土壤水分含量较低的情况下,均能保持较高蒸腾速率和光合作用,这样有效积累了地上地下部生物量,从而提高了水分利用效率。ABA的积累是其主要的信号调节机理。同时发现非ABA代谢途径也同样调节植株对水分匮缺的反应。在生殖期间歇性水分胁迫下,土壤水分的过快汲取导致了株系间表现相似,但复水后差异明显(p<0.01),株系30复水前水分利用率高,生物量积累多,恢复能力最强,而具有发达根系的line29并没有表现出强的恢复能力。根系解剖结构表明,株系29具有厚的木质部层和更多导管,这有利于土壤水分的吸收,但是在严重土壤水分胁迫下易造成导管“气穴”现象,这将极大降低根系对土壤水分的汲取能力。同时在田间旱胁迫下,我们尝试观察了植株的动态变化,结果再次证明,适时的开花期,高效水分利用或者发达根系,以及碳水化合物的有效转运等,协同调节是聚合系抗旱的机理所在。
在第二套实验中,利用两个以高产稻黄花占为背景BC1F3群体(HHZ/OM1723和HHZ/Teqing)3000个单株,在中国广东和国际水稻研究所进行了抗旱,耐盐和高产选择,选择到24个高产,36个极端抗旱和30个极端耐盐的导入系进行了卡方和多位点关联概率测验,以检测供体导入片段中与可能与高产、抗旱以及耐盐相关联的功能遗传单位(FGU)。根据连锁不平衡分析和最新数量遗传理论,共检测到53个FGU构成了抗旱和耐盐的遗传网络。在群体HHZ/OM1723中检测到28个FGU,其中9个在三种选择群体中检测到,14个在两种选择群体中,5个只在单个选择群体中检测到,而且有三个具有高导入的位点总是处于抗旱耐盐遗传网络的最上游,其中3个在三种遗传网络的最上游具有重叠。同样地,在群体HHZ／Teqing中,检测到25个FGU,其中19个在三种选择群体中检测到,4个只在两种选择群体中检测到,2个只在单个选择群体中检测到。同样地,总有两个位点处于高产、抗旱和耐盐遗传网络的最上游。而且同一个途径的位点间表现出高度正相关,不同途径间表现出独立或者负相关关系。但是比较发现,很少FGU在两个群体发生重叠,这表明大多数检测到的FGU来自于不同的供体。这与当前对抗旱和耐盐的研究结论一致,但这些遗传信息为我们通过聚合高度正相关或者独立的不同的功能遗传位点,进一步提高水稻的高产、抗旱以及耐盐水平,提供了有益的帮助,也为通过分子标记高通量检测抗旱耐盐QTL位点提供了重要的参考。
Abiotic stresses such as drought and salinity are very important factor limiting rice productivity in the rainfed and costal areas of Asia. Development and adoption of high-yielding and stress tolerant rice varieties are the best solution for solving the problems. However, it is well known that rice tolerances to both drought and salinity are controlled by large numbers of genes or quantitative trait loci (QTLs), making it difficult to breed superior rice varieties with good tolerances to both stresses. In past 10 years, many sets of introgression lines (ILs) and pyramiding lines (PLs) with significantly improved drought tolerance (DT) and/or salt tolerance (ST) have been developed using backcross breeding and molecular pyramiding at IRRI. The objectives of this study were:(1) to characterize the phenotypic and physiological traits associated with DT PLs and (2) to understand the genetic basis of DT and ST using ILs and molecular markers.
Two series experiments were performed in this study. In the first set of experiments,48 PLs, their parental ILs and recurrent parent, IR64 (check) were evaluated for their yield performances and related traits under severe drought stress at the reproductive stage (RS) and vegetative stage (VS) and irrigated control in three years to understand the relationship between drought tolerance (DT) and yield potential (YP) in rice and their underlying mechanisms. When compared to IR64, all PLs had significantly improved DT to RS and 36 PLs also had significantly improved DT to VS (p<0.05). In addition,23 PLs had higher YP than IR64 (p<0.05) and the remaining 25 PLs had similar YP as IR64 under the irrigated control. Detailed characterization of the PLs revealed 3 mechanisms that functioned together to contribute to their improved DT and/or YP. The most important mechanism was drought avoidance (DA) characterized with significantly higher biomass and harvest index (HI) of the PLs than IR64 under stress, suggesting the PLs' better abilities to maintain higher water status and function under stress. Because most PLs did not show any reduction in biomass under control, this type of DA resulted most likely from improved dehydration resistance. The second mechanism was efficient partitioning characterized by improved HI in all PLs as compared to IR64, resulting primarily from higher GW and/or SF under control and was the major constituent of the improved YP in the 17 PLs. Drought escape (DE) by accelerated heading under drought was the third mechanism that contributed to DT of the PLs to RS. The considerable variation in the measured traits among the PLs with similar levels of DT and YP implies the complex genetic control of the mechanisms for DT/YP and offers opportunities to further improve DT and/or YP by fine-tuning of QTLs/genes among the PLs using MAS. Finally, our results indicate that selection for yield plus some secondary traits under appropriate type(s) of stress and non-stress conditions similar to target environments are critically important for improving both DT and YP in rice.
A selected set of promising PLs was further analyzed for the dynamic responses of plant growth and water use to progressive water deficit at different growth stages in lysimeters under controlled phytotron conditions and rainfed lowland condition. In vegetative stage dry-down experiment, the PLs that performed best under drought were those that had the lowest threshold of transpiration response to soil moisture decrease (P<0.01), i.e. lines that kept stomata open longer or had large root system during the soil drying, and hence extracted more soil water, which resulted in higher shoot and root biomass, and an overall improvement of transpiration efficiency. ABA-dependant and ABA-independent signal pathway co-regulated the different response to the FTSW among the genotypes. In intermittent water stress, the sharp dry-down stress resulted in the similar performance between lines in the threshold of FTSW. The lines with higher water use efficiency (WUE) and large biomass at the end of stress showed good drought recovery ability, and thus produced higher spikelet fertility and grain yield. However, those lines produced large root biomass did not show superior drought recovery, probably due to the specific root anonomy structure. Under the lowland progressive water stress, the PLs produced high yield potential as the result of large biomass production (more tillers and panicles), while under drought, they improved WUE by balancing the transpiration and tiller and leaf area production, and performed higher yield, harvest index and spikelet fertility, but lower above ground shoot production. As well, the lines with large root system also conferred to drought tolerance in lowland condition, to some extent.
In the second set of experiments, two BC1F2 populations (1500plants) in the genetic background of a high-yielding indica variety, Huang-Hua-Zan, were screened under drought and salinity conditions at IRRI and China, resulting in development of 36 DT ILs and 30 salt tolerant (ST) ILs. These ILs were analyzed using X2 and multilocus probability tests to identify functional genetic units (FGUs) located in the donor segments associated with DT and ST. Together, a total of 53 FGUs associated with HY, DT and/or ST were identified in the ILs, forming 3 genetic networks underlying HY, DT and ST in rice. The first one detected in the HHZ/OM1723 population contained 28 FGUs, including 9 FGUs contributing to HY, DT, and ST,14 FGUs affecting 2 traits, and 5 FGUs that contributed to one trait only. Of these,3 loci (RM24, RM6 and RM130) appeared to be important as they had high frequencies of introgression and were placed in the upstream of the network based on the new molecular-quantitative genetics theory. Similarly, the genetic network identified in the HHZ/Teqing population had 25 FGUs, including 19 FGUs contributing to HY, DT, and ST,4 FGUs affecting 2 traits, and 2 FGUs that contributed to one trait only. Again, these FGUs included 2 important ones in the upstream of the genetic network. In contrast to the high genetic overlap among the three traits, especially between DT and ST, however, there were few shared loci identified in the HHZ/OM1723 and those in the HHZ/Teqing population, indicating the 2 donors contributed different sets of loci to the DT and/or ST in the selected ILs. Our results were in consistent with our current knowledge that DT and ST in rice are under complex genetic control, and provided useful information and materials for further improvement of DT and ST in rice by pathway pyramiding, and for detailed molecular dissection of DT and ST in rice using high throughput genomic tools.
引文
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