水稻苗期抗高温鉴定程序的建立及其基因定位
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摘要
随着全球经济的快速发展和二氧化碳排放量的逐年增加,高温热害的发生频率日趋频繁,严重时导致农作物大面积减产和产品品质下降。面对这样的栽培逆境,如何稳定和提高作物产量,确保农业经济和粮食安全,已成为农业科技领域迫切需要解决的重大科研课题。水稻是最重要的粮食农作物,全球有一半以上的人口以水稻为主食。目前,人们就水稻对高温响应及其抗高温特性开展了广泛的研究,包括水稻受高温胁迫后的蛋白质表达谱变化、生理生化反应和QTL定位等,但迄今为止,还没有通过图位克隆发掘耐热基因的报道,所报导的耐热相关基因均为同源克隆法获得,且主要为热激蛋白。
本研究以广东省农科院水稻所钟旭华研究员筛选并提供使用的水稻抗高温地方品种HT54和感高温品种HT13为材料,于不同生育期和不同栽培方式条件下对其进行不同温度处理,试图建立适宜的水稻抗高温鉴定程序和抗性分级标准,并在此基础上,对HT54中的抗高温性状进行遗传分析和基因定位以及生理生化指标测定。主要研究结果如下：
1.抗高温鉴定程序及判别标准的建立：对抗感高温水稻品种HT54和HT13分别进行水培和土培种植,待秧苗生长至两叶一心和三叶一心期移入生长箱,进行42℃、45℃和48℃的高温处理,空气相对湿度和其它栽培条件保持在相同水平上,之后,每隔一定时间观察其对高温的反应。试验结果显示：两叶一心期48℃高温处理至84h,水培的秧苗恢复5d后呈现抗性品种秧苗完全存活和感性品种秧苗一致死亡的显著差异；而土培的秧苗处理至79h恢复相同天数后即可达到相同的效果。这些结果表明,土培和水培的方法都可用于抗高温材料的筛选鉴定,但后者比前者的鉴定用时短、效率高。分蘖期土培剥蘖秧苗抗感品种间达到存活与死亡差异效果的处理温度是45℃,处理时间是72h,但秧苗培养和剥蘖较费时费工,好处是取样不受材料死亡的限制。抽穗期由于抗高温特性的考察指标通常为结实率,当以平均温度为36℃的高温处理48h时,其抗感高温品种的差异达到了显著水平,但此时抗高温品种HT54的结实率和常温下相比也显著降低,因此,不是全生育期抗高温特性鉴定的最佳处理时期。据此,本研究最终确定的水稻全生育期抗高温特性苗期鉴定程序为：两叶一心的土培或水培苗,48℃高温、相对湿度75%处理至79h或84h,恢复5d后,以秧苗能否存活作为抗感鉴定指标。
2.抗高温基因的遗传分析：以优化的抗高温鉴定程序对抗感高温亲本品种HT54和HT13、F1及F2群体进行高温处理,发现杂种F1秧苗经高温处理后的表现与抗高温品种HT54的表现一致,F2群体中抗感高温植株的分离符合3：1的分离比率。这些结果证实HT54在苗期鉴定的抗高温特性是受显性主效单基因控制,据此,我们将其命名为OsHTAS。
3.抗高温基因的定位：应用隐性极端集团群体法从F2群体中挑选131株敏感植株构建了作图群体,同时,随机从相同的F2群体中分别挑选10株抗感高温极端单株构建了用于初步连锁群分析的两个DNA池,籍此对HT54中的抗高温基因OsHTAS进行了定位。试验结果显示：位于第9染色体上的一对SSR引物RM444在两个DNA池间具有多态性,之后,通过F2群体中61株感高温,验证了RM444和抗高温基因OsHTAS司的连锁关系。再先后以61株和群体扩大后的131株感高温单株为作图群体,围绕RM444扩增片段所在的位点进行了标记加密分析,结果显示OsHTAS是位于InDel5和RM7364之间,遗传上分别相距2.5cM/3.2cM和1.7cM/1.2cM,标记间的总物理距离为420kb;根据此区间候选基因功能分析及基因芯片表达数据的比较,筛选出两个候选基因并进行测序,结果发现只有其中一个基因的序列在两亲本之间存在两处差异；根据其中一处差异开发了一个CAPS标记,该标记和抗高温基因OsHTAS表现共分离。
4.抗高温特性相关生理指标SOD和POD的测定：以已知抗高温的黄华占品种为对照,在两叶一心时期对抗感高温水稻品种HT54和HT13进行高温处理,然后与不同时间点分别取样测定高温处理过程中水稻幼苗的POD和SOD活性,结果显示：三个水稻品种黄华占、HT54和HT13在热胁迫处理过程中,各时间点测得的SOD和POD活性均受激增强,但就活性检测值大小而言,HT54在三个品种中居中。另外,该结果还显示：在热胁迫处理条件下,HT13和黄华占的POD和SOD活性大小变化的相对程度不一致,HT13的SOD活性一直是三个品种中最小的,但其POD的活性除在48℃处理24h时低于黄华占外,其它时间点的测定值均是三个品种最大的。因此,该结果说明不能简单的通过热胁迫处理后SOD和POD活性大小测定评价这三个材料对高温的抗性。
Hot disaster appears more and more frequently with the rapid development of global economy and the increasing of carbon dioxide emission year by year, leading to the decline of crop production and product quality when it becomes more serious. An urgent and significant research issue is put into place on how to stabilize and improve the yield of crop, and how to ensure agricultural economy and grain security in agricultural science domain in the face of such cultivation adversity. Rice is the most important crop in the word as it is the major source of food for more than half of the word population. At present, there are a wide range of researches on high temperature-stress responses and heat tolerance characteristics of rice, including the variation of the protein expression profile, the physiological and biochemical reactions, location of Quantitative Trait Locus (QTLs) responsible for high temperature tolerance, et al. However there is no report on map-based cloning of heat tolerance so far. The reported genes, related with heat tolerance, are homology-based cloning method to get and most are heat shock proteins.
The plant materials used in the present study include HT54, a heat tolerance cultivar, and HT13, a heat sensitive cultivar, which were provided and identified by Xuhua Zhong a research fellow from Rice Research Institute of Guangdong Academy of Agricultural Science. We attempted to establish high temperature resistance evaluation procedures as well as its assesment criteria at seedling stage, and on this basis, to further conduct inheritance analysis, gene mapping and physiological index assessment of high temperature resistance present in HT54. The main results detained are as follows:
1. The establishment of high temperature resistance evaluation procedures and its assessment criteria
HT54and HT13, both cultivated by nutrition solution or soil, were removed into growth cabinet and treated by42℃,45℃or48℃at two to three leaves or three to four leaves. The relative humidity was set at75%and the other cultural conditions were kept on the same level, and after that, the response on high temperature was observed at a certain period of time. The results showed that HT54, cultivated by nutrition solution, were all survive and HT13were all dead after84h48℃high temperature treatment at two to three leaves, but it needed only79h to achieve the same effect for soil culture. Both two culturing methods could be used in the high temperature resistance evaluation procedures, but the latter was highly efficient as compared with the former one. The obvious differences were observed after72h45℃high temperature treatment at the tillering stage. Although it solved the limits of sample for DNA isolation because of the death, more time and more work were needed to select and cultivate tillerings. At the heading stage, seed setting rate reduced vigorously after48hours under36℃high temperature and the varying degree of variance reached5%significance between HT54and HT13cultivars, but the rate of HT54after high temperature was significantly lower than that under natural condition. Therefore heading stage was not the optimal stage for high temperature resistance evaluation of HT54. According to these results, we established the high temperature procedures at seedling stage:two to three leaves, cultivated by soil or nutrition solution,84h or79h48℃high temperature treatment with75%air relative humidity, evaluation after5day recovery after treatment as the assessment criteria of high temperature resistance.
2. Genetic analysis of high temperature resistance
F2population was made by pollination of HT54with HT13. Regarding to the assessment criterion, the resistance response of F1plants was consistent with high temperature resistance cultivar HT54and the segregation of F2progeny was consistent with3:1ratio after high temperature treatment. These results thus demonstated that the high temperature resistance presence in HT54at seedling stage was controlled by complete dominance of monogenic allele, named OsHTAS (Oryza saliva heat tolerance at seedling stage).
3. Mapping of high temperature resistance gene
The mapping population was constructed with131sensitive plants randomly selected from F2generation according to the bulked recessive extreme segregats strategy. At the same time, two DNA pools for primary linkge analysis were also constructed with DNA samples extacted from10extreme sensitive and resisant plants, respectively, which were randomly selected from the same F2population. The high temperature resistance gene OsHTAS was then mapped using these populations. The results showed that the marker RM444, position on chromosome9, had the polymorphism between two DNA pools. This result was preliminarily confirmed by marker screening of61high temperature sensitive plants. Further marker density increasing analysis with61sensitive plants and an enlarged F2population containing131recessive extreme segregants demonstrated that the OsHTAS is located between markers InDel5and RM7364. The map distance between OsHTAS and the two closely linked markers is2.5cM/3.2cM and1.7cM/1.2cM, respectively. The physical distance between the two markers InDe15and RM7364is420kb. Two genes were selected and consequently sequenced according to deduced function analysis of cadidate genes within the interval of these two markers as well as previously reported chip expression data. Two single base pair (SNP) differences on the resultant sequences of of one gene between HT54and HT13were detected and one of these SNPs was developed into CAPS marker. Further analysis revealed that this CAPS marker was cosegregated with OsHTAS gene.
3. High temperature resistance related physiological index SOD and POD assay
The SOD and POD activity in the seedling samples of HT54, HT13were assayed at different time points after high temperature treatment at stage of two to three leaves and the currently well-known high temperature resistance cultivar Huanghuazhan was used as control. The result showed that the SOD and POD activity detected at different time points was stimulated to increase during heat stresses. But in case of activity detection value, the HT54was in the middl among three cultivars. In addition, these results also showed that under heat stress conditions, the relative degree of variation of the POD and SOD activity in Huang Huazhan and HT13was inconsistency. The SOD activity of HT13was the lowest but its POD activity appeared the highest among three cultivars except one case lower than Huanghuazhan at48℃. Therefore, all these results indicated that the high temperature resistance of the tested varieties could not be evaluated by simply detecting the SOD and POD activity.
本研究以广东省农科院水稻所钟旭华研究员筛选并提供使用的水稻抗高温地方品种HT54和感高温品种HT13为材料,于不同生育期和不同栽培方式条件下对其进行不同温度处理,试图建立适宜的水稻抗高温鉴定程序和抗性分级标准,并在此基础上,对HT54中的抗高温性状进行遗传分析和基因定位以及生理生化指标测定。主要研究结果如下：
1.抗高温鉴定程序及判别标准的建立：对抗感高温水稻品种HT54和HT13分别进行水培和土培种植,待秧苗生长至两叶一心和三叶一心期移入生长箱,进行42℃、45℃和48℃的高温处理,空气相对湿度和其它栽培条件保持在相同水平上,之后,每隔一定时间观察其对高温的反应。试验结果显示：两叶一心期48℃高温处理至84h,水培的秧苗恢复5d后呈现抗性品种秧苗完全存活和感性品种秧苗一致死亡的显著差异；而土培的秧苗处理至79h恢复相同天数后即可达到相同的效果。这些结果表明,土培和水培的方法都可用于抗高温材料的筛选鉴定,但后者比前者的鉴定用时短、效率高。分蘖期土培剥蘖秧苗抗感品种间达到存活与死亡差异效果的处理温度是45℃,处理时间是72h,但秧苗培养和剥蘖较费时费工,好处是取样不受材料死亡的限制。抽穗期由于抗高温特性的考察指标通常为结实率,当以平均温度为36℃的高温处理48h时,其抗感高温品种的差异达到了显著水平,但此时抗高温品种HT54的结实率和常温下相比也显著降低,因此,不是全生育期抗高温特性鉴定的最佳处理时期。据此,本研究最终确定的水稻全生育期抗高温特性苗期鉴定程序为：两叶一心的土培或水培苗,48℃高温、相对湿度75%处理至79h或84h,恢复5d后,以秧苗能否存活作为抗感鉴定指标。
2.抗高温基因的遗传分析：以优化的抗高温鉴定程序对抗感高温亲本品种HT54和HT13、F1及F2群体进行高温处理,发现杂种F1秧苗经高温处理后的表现与抗高温品种HT54的表现一致,F2群体中抗感高温植株的分离符合3：1的分离比率。这些结果证实HT54在苗期鉴定的抗高温特性是受显性主效单基因控制,据此,我们将其命名为OsHTAS。
3.抗高温基因的定位：应用隐性极端集团群体法从F2群体中挑选131株敏感植株构建了作图群体,同时,随机从相同的F2群体中分别挑选10株抗感高温极端单株构建了用于初步连锁群分析的两个DNA池,籍此对HT54中的抗高温基因OsHTAS进行了定位。试验结果显示：位于第9染色体上的一对SSR引物RM444在两个DNA池间具有多态性,之后,通过F2群体中61株感高温,验证了RM444和抗高温基因OsHTAS司的连锁关系。再先后以61株和群体扩大后的131株感高温单株为作图群体,围绕RM444扩增片段所在的位点进行了标记加密分析,结果显示OsHTAS是位于InDel5和RM7364之间,遗传上分别相距2.5cM/3.2cM和1.7cM/1.2cM,标记间的总物理距离为420kb;根据此区间候选基因功能分析及基因芯片表达数据的比较,筛选出两个候选基因并进行测序,结果发现只有其中一个基因的序列在两亲本之间存在两处差异；根据其中一处差异开发了一个CAPS标记,该标记和抗高温基因OsHTAS表现共分离。
4.抗高温特性相关生理指标SOD和POD的测定：以已知抗高温的黄华占品种为对照,在两叶一心时期对抗感高温水稻品种HT54和HT13进行高温处理,然后与不同时间点分别取样测定高温处理过程中水稻幼苗的POD和SOD活性,结果显示：三个水稻品种黄华占、HT54和HT13在热胁迫处理过程中,各时间点测得的SOD和POD活性均受激增强,但就活性检测值大小而言,HT54在三个品种中居中。另外,该结果还显示：在热胁迫处理条件下,HT13和黄华占的POD和SOD活性大小变化的相对程度不一致,HT13的SOD活性一直是三个品种中最小的,但其POD的活性除在48℃处理24h时低于黄华占外,其它时间点的测定值均是三个品种最大的。因此,该结果说明不能简单的通过热胁迫处理后SOD和POD活性大小测定评价这三个材料对高温的抗性。
Hot disaster appears more and more frequently with the rapid development of global economy and the increasing of carbon dioxide emission year by year, leading to the decline of crop production and product quality when it becomes more serious. An urgent and significant research issue is put into place on how to stabilize and improve the yield of crop, and how to ensure agricultural economy and grain security in agricultural science domain in the face of such cultivation adversity. Rice is the most important crop in the word as it is the major source of food for more than half of the word population. At present, there are a wide range of researches on high temperature-stress responses and heat tolerance characteristics of rice, including the variation of the protein expression profile, the physiological and biochemical reactions, location of Quantitative Trait Locus (QTLs) responsible for high temperature tolerance, et al. However there is no report on map-based cloning of heat tolerance so far. The reported genes, related with heat tolerance, are homology-based cloning method to get and most are heat shock proteins.
The plant materials used in the present study include HT54, a heat tolerance cultivar, and HT13, a heat sensitive cultivar, which were provided and identified by Xuhua Zhong a research fellow from Rice Research Institute of Guangdong Academy of Agricultural Science. We attempted to establish high temperature resistance evaluation procedures as well as its assesment criteria at seedling stage, and on this basis, to further conduct inheritance analysis, gene mapping and physiological index assessment of high temperature resistance present in HT54. The main results detained are as follows:
1. The establishment of high temperature resistance evaluation procedures and its assessment criteria
HT54and HT13, both cultivated by nutrition solution or soil, were removed into growth cabinet and treated by42℃,45℃or48℃at two to three leaves or three to four leaves. The relative humidity was set at75%and the other cultural conditions were kept on the same level, and after that, the response on high temperature was observed at a certain period of time. The results showed that HT54, cultivated by nutrition solution, were all survive and HT13were all dead after84h48℃high temperature treatment at two to three leaves, but it needed only79h to achieve the same effect for soil culture. Both two culturing methods could be used in the high temperature resistance evaluation procedures, but the latter was highly efficient as compared with the former one. The obvious differences were observed after72h45℃high temperature treatment at the tillering stage. Although it solved the limits of sample for DNA isolation because of the death, more time and more work were needed to select and cultivate tillerings. At the heading stage, seed setting rate reduced vigorously after48hours under36℃high temperature and the varying degree of variance reached5%significance between HT54and HT13cultivars, but the rate of HT54after high temperature was significantly lower than that under natural condition. Therefore heading stage was not the optimal stage for high temperature resistance evaluation of HT54. According to these results, we established the high temperature procedures at seedling stage:two to three leaves, cultivated by soil or nutrition solution,84h or79h48℃high temperature treatment with75%air relative humidity, evaluation after5day recovery after treatment as the assessment criteria of high temperature resistance.
2. Genetic analysis of high temperature resistance
F2population was made by pollination of HT54with HT13. Regarding to the assessment criterion, the resistance response of F1plants was consistent with high temperature resistance cultivar HT54and the segregation of F2progeny was consistent with3:1ratio after high temperature treatment. These results thus demonstated that the high temperature resistance presence in HT54at seedling stage was controlled by complete dominance of monogenic allele, named OsHTAS (Oryza saliva heat tolerance at seedling stage).
3. Mapping of high temperature resistance gene
The mapping population was constructed with131sensitive plants randomly selected from F2generation according to the bulked recessive extreme segregats strategy. At the same time, two DNA pools for primary linkge analysis were also constructed with DNA samples extacted from10extreme sensitive and resisant plants, respectively, which were randomly selected from the same F2population. The high temperature resistance gene OsHTAS was then mapped using these populations. The results showed that the marker RM444, position on chromosome9, had the polymorphism between two DNA pools. This result was preliminarily confirmed by marker screening of61high temperature sensitive plants. Further marker density increasing analysis with61sensitive plants and an enlarged F2population containing131recessive extreme segregants demonstrated that the OsHTAS is located between markers InDel5and RM7364. The map distance between OsHTAS and the two closely linked markers is2.5cM/3.2cM and1.7cM/1.2cM, respectively. The physical distance between the two markers InDe15and RM7364is420kb. Two genes were selected and consequently sequenced according to deduced function analysis of cadidate genes within the interval of these two markers as well as previously reported chip expression data. Two single base pair (SNP) differences on the resultant sequences of of one gene between HT54and HT13were detected and one of these SNPs was developed into CAPS marker. Further analysis revealed that this CAPS marker was cosegregated with OsHTAS gene.
3. High temperature resistance related physiological index SOD and POD assay
The SOD and POD activity in the seedling samples of HT54, HT13were assayed at different time points after high temperature treatment at stage of two to three leaves and the currently well-known high temperature resistance cultivar Huanghuazhan was used as control. The result showed that the SOD and POD activity detected at different time points was stimulated to increase during heat stresses. But in case of activity detection value, the HT54was in the middl among three cultivars. In addition, these results also showed that under heat stress conditions, the relative degree of variation of the POD and SOD activity in Huang Huazhan and HT13was inconsistency. The SOD activity of HT13was the lowest but its POD activity appeared the highest among three cultivars except one case lower than Huanghuazhan at48℃. Therefore, all these results indicated that the high temperature resistance of the tested varieties could not be evaluated by simply detecting the SOD and POD activity.
引文
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