两个生育时期水稻耐低磷胁迫相关性状的QTL定位
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摘要
土壤中有效磷缺乏是制约水稻生产的重要因素之一。选育磷高效水稻品种,是改善水稻磷营养状况的一种更为经济、环保、长远的途径。分子标记辅助选择结合表型鉴定是选育水稻磷高效基因型的一条途径,而QTL定位是进行分子标记辅助选择和图位克隆的基础。本研究在进行遗传连锁图谱分子标记加密的基础上,以超级杂交稻协优9308不育系的保持系协青早B和恢复系中恢9308杂交衍生的198个株系组成的重组自交系群体为材料,利用水培方法,在分蘖期和抽穗期分别对磷效率相关性状作了初步定位,同时对两个生育时期根系的酸性磷酸酶活性与磷效率之间的关系作了初步分析。主要结果如下:
1.应用328个SSR标记对亲本协青早B和中恢9308进行多态性筛选,共获得98个亲本多态性SSR分子标记,多态性频率为29.8%。在参考分析已有图谱的基础上,选取35个SSR标记对281个单株的RIL群体进行标记基因型检测。经卡方检验,35个SSR标记中,当P< 0.05时,有18个符合孟德尔1:1的单基因理论分离比,占总标记数的51.4%,其余17个为显著偏分离,占总标记数的48.6%。在所有17个偏分离标记中,7个(41.2%)偏向父本中恢9308,10个(58.8%)偏向母本协青早B。应用MAPMAKER/EXP 3.0,将35个标记全部加入到各自的连锁群。加密后的连锁图谱共有198个标记,每个连锁群上的标记数介于4~25个之间;总图距为1765.6 cM,标记间平均图距为8.96 cM,图距范围为0.2 cM-34.4 cM。
2. QTL定位结果表明,在两个生育时期共检测到控制6个性状表现的21个加性QTL,分布在第4、5、6、7、10、11染色体上。其中4个相对根干重QTL、3个相对冠干重QTL、3个相对总干重QTL、5个相对根长QTL、4个相对分蘖力QTL和2个相对根冠比QTL。除此之外,还在第1、5、6、9染色体检测到3对非加性QTL互作效应。在分蘖期于第4染色体的RM307-RM5953区间内,均定位到控制相对冠干重、相对总干重及相对根冠比的QTL;在第10染色体的RM311-RM6142区间内,各自检测到1个相对根干重和相对分蘖力的QTL。类似的现象在抽穗期也有出现。比较分蘖期和抽穗期的定位结果看,除相对根长外,其它5个性状在两个生育时期都各自在相同的区间定位到相应QTL。除此之外,在分蘖期检测到的2对上位性效应中,有1对在抽穗期也被检测到。在第4染色体的RM307-RM3317区间内检测到多个QTL,是所有染色体中聚集QTL数目最多的一个区间,在该区间可能存在控制水稻磷效率的基因。
3.在低磷条件下,两个生育时期亲本及RIL群体根系的酸性磷酸酶活性均有提高。虽然在正常供磷条件下,父本中恢9308的根系酸性磷酸酶活性高于母本协青早B,但在低磷条件下协青早B的根系酸性磷酸酶活性高于中恢9308,这暗示着根系酸性磷酸酶活性在低磷胁迫和正常处理条件下可能受不同的遗传基础调控。对比两个生育时期根系相对酸性磷酸酶活性发现,抽穗期的酶活性要低于分蘖期的酶活,这个结果也可通过相关分析得以间接佐证。相关分析结果表明,在分蘖期,根系的相对酸性磷酸酶活性与相对根干重、相对冠干重、相对总干重、相对分蘖力、根系相对磷吸收量的相关程度要高于抽穗期,这暗示着根系酸性磷酸酶活性在生育期前期对磷效率的贡献率,要高于生育期后期的贡献率。QTL定位结果表明,仅在分蘖期于第3和第7染色体分别定位到一个控制根系相对酸性磷酸酶活性的QTL,在抽穗期未定位到相关QTL。
The low level of available phosphorus in soils is a major yield-limiting factor in rice production. Breeding high phosphorus efficient cultivars may represent a more sustainable solution than sole reliance on fertilizer application. Molecular marker assisted selection (MAS) can be integrated with traditional methods of artificial selection on phenotypes to breed rice cultivars with high phosphorus efficiency. Wheares QTL mapping is the basis of the MAS and map-based cloning. The present research mainly aimed to map QTLs for traits related to phosphorus deficiency tolerance at tillering stage and heading stage in rice after increasing the density of rice genetic linkage map with a number of 35 SSR markers. In addition, the rice root acid phosphatase was evaluated and its relationship with phosphorus efficiency was studied. A RIL population consisting of 195 lines derived from maintainer line Xieqingzao B (abbreviated as Xie B) and restoring line Zhonghui 9308(abbreviated as R9308) of super hybrid rice Xieyou 9308 was used and hydrophonic method was employed in the experiment. The relative parameter, defined as a ratio of value under phosphorus deficient condition to that measured under sufficient condition was used to evaluate the tolerance ability of parents and RIL population in the experiment. The main results are outlined as follows:
1. A total of 328 SSR markers were used to screen the polymorphism between Xie B and R9308, and 98 polymorphic markers were obtained with the polymorphism rate of 29.8%. Based on the preliminary framework of the linkage map, 35 polymorphic markers were chosen to genotype the RIL population consisting of 281 lines. The chi-squares test indicated that a number of 18 markers which accounted for 51.4% were consistent with theoretical segregation ratio of 1:1 at the significance level of 5%, and the other 17 markers which accounted for 48.6% deviated significantly. Among all the deviation markers, 7 markers deviated to male parent R9308 and the other 10 deviated to female parent Xie B. By using MAPMAKER/EXP 3.0, all the 35 SSR markers were localized onto the chromosomes. The marker-newly-added linkage map consists of 198 SSR markers and covers a length of 1765.6 cM with the average distance of 8.96 cM between adjacent markers.
2. By using CIM method, a total of 21 additive QTLs for phosphorus deficiency tolerance related traits were detected at two development stages, which distributed through chromosome 4, 5, 6, 7, 10, 11. Among them, 4 was for relative root dry weight(RRDW), 3 was for relative shoot dry weight(RSDW), 3 was for relative total dry weight(RTDW), 5 was for relative root length(RRL), 4 was for relative tillering ability(RTA) and 2 was for relative root/shoot ratio(RRRS). In addition, 3 pairs of epistatic effect between QTLs which have no additive effect were also detected on chromosome 1, 5, 6, and 9. At tillering stage, QTLs for RSDW, RTDW and RRRS were detected at the same region of RM307-RM5953 on chromosome 4 and QTLs for RRDW and RTA were detected at the same region of RM311-RM6142 on chromosome 10. The situation that QTLs for different traits localized at the same region was also found at heading stage. Except for RRL, same QTLs were detected for the other 5 traits at the same region at tillering stage and heading stage. Interestingly, among three pairs of epistatic effect, one pair was detected at two development stages. A cluster of QTLs were detected at the region of RM307-RM5953 on chromosome 4 in which phosphorus efficiency related gene may lie.
3. Under phosphorus surficient condition, the root acid phosphatase activity of male parent R9308 was higher than that of female parent Xie B. Under phosphorus stress condition, however, the situation is quite reverse although the root acid phosphatase activity of the parents and RIL population lines increased at both development stages, which indicated that the genetic system for acid phosphatase under stress condition was different with that for acid phosphatase under normal culture. The simple correlation analysis showed that the contribution of AAP to phosphorus efficiency at tillering stage was greater than that of at heading stage. There were two QTLs for RRAPA at tillering stage, but no QTL was detected at heading stage.
1.应用328个SSR标记对亲本协青早B和中恢9308进行多态性筛选,共获得98个亲本多态性SSR分子标记,多态性频率为29.8%。在参考分析已有图谱的基础上,选取35个SSR标记对281个单株的RIL群体进行标记基因型检测。经卡方检验,35个SSR标记中,当P< 0.05时,有18个符合孟德尔1:1的单基因理论分离比,占总标记数的51.4%,其余17个为显著偏分离,占总标记数的48.6%。在所有17个偏分离标记中,7个(41.2%)偏向父本中恢9308,10个(58.8%)偏向母本协青早B。应用MAPMAKER/EXP 3.0,将35个标记全部加入到各自的连锁群。加密后的连锁图谱共有198个标记,每个连锁群上的标记数介于4~25个之间;总图距为1765.6 cM,标记间平均图距为8.96 cM,图距范围为0.2 cM-34.4 cM。
2. QTL定位结果表明,在两个生育时期共检测到控制6个性状表现的21个加性QTL,分布在第4、5、6、7、10、11染色体上。其中4个相对根干重QTL、3个相对冠干重QTL、3个相对总干重QTL、5个相对根长QTL、4个相对分蘖力QTL和2个相对根冠比QTL。除此之外,还在第1、5、6、9染色体检测到3对非加性QTL互作效应。在分蘖期于第4染色体的RM307-RM5953区间内,均定位到控制相对冠干重、相对总干重及相对根冠比的QTL;在第10染色体的RM311-RM6142区间内,各自检测到1个相对根干重和相对分蘖力的QTL。类似的现象在抽穗期也有出现。比较分蘖期和抽穗期的定位结果看,除相对根长外,其它5个性状在两个生育时期都各自在相同的区间定位到相应QTL。除此之外,在分蘖期检测到的2对上位性效应中,有1对在抽穗期也被检测到。在第4染色体的RM307-RM3317区间内检测到多个QTL,是所有染色体中聚集QTL数目最多的一个区间,在该区间可能存在控制水稻磷效率的基因。
3.在低磷条件下,两个生育时期亲本及RIL群体根系的酸性磷酸酶活性均有提高。虽然在正常供磷条件下,父本中恢9308的根系酸性磷酸酶活性高于母本协青早B,但在低磷条件下协青早B的根系酸性磷酸酶活性高于中恢9308,这暗示着根系酸性磷酸酶活性在低磷胁迫和正常处理条件下可能受不同的遗传基础调控。对比两个生育时期根系相对酸性磷酸酶活性发现,抽穗期的酶活性要低于分蘖期的酶活,这个结果也可通过相关分析得以间接佐证。相关分析结果表明,在分蘖期,根系的相对酸性磷酸酶活性与相对根干重、相对冠干重、相对总干重、相对分蘖力、根系相对磷吸收量的相关程度要高于抽穗期,这暗示着根系酸性磷酸酶活性在生育期前期对磷效率的贡献率,要高于生育期后期的贡献率。QTL定位结果表明,仅在分蘖期于第3和第7染色体分别定位到一个控制根系相对酸性磷酸酶活性的QTL,在抽穗期未定位到相关QTL。
The low level of available phosphorus in soils is a major yield-limiting factor in rice production. Breeding high phosphorus efficient cultivars may represent a more sustainable solution than sole reliance on fertilizer application. Molecular marker assisted selection (MAS) can be integrated with traditional methods of artificial selection on phenotypes to breed rice cultivars with high phosphorus efficiency. Wheares QTL mapping is the basis of the MAS and map-based cloning. The present research mainly aimed to map QTLs for traits related to phosphorus deficiency tolerance at tillering stage and heading stage in rice after increasing the density of rice genetic linkage map with a number of 35 SSR markers. In addition, the rice root acid phosphatase was evaluated and its relationship with phosphorus efficiency was studied. A RIL population consisting of 195 lines derived from maintainer line Xieqingzao B (abbreviated as Xie B) and restoring line Zhonghui 9308(abbreviated as R9308) of super hybrid rice Xieyou 9308 was used and hydrophonic method was employed in the experiment. The relative parameter, defined as a ratio of value under phosphorus deficient condition to that measured under sufficient condition was used to evaluate the tolerance ability of parents and RIL population in the experiment. The main results are outlined as follows:
1. A total of 328 SSR markers were used to screen the polymorphism between Xie B and R9308, and 98 polymorphic markers were obtained with the polymorphism rate of 29.8%. Based on the preliminary framework of the linkage map, 35 polymorphic markers were chosen to genotype the RIL population consisting of 281 lines. The chi-squares test indicated that a number of 18 markers which accounted for 51.4% were consistent with theoretical segregation ratio of 1:1 at the significance level of 5%, and the other 17 markers which accounted for 48.6% deviated significantly. Among all the deviation markers, 7 markers deviated to male parent R9308 and the other 10 deviated to female parent Xie B. By using MAPMAKER/EXP 3.0, all the 35 SSR markers were localized onto the chromosomes. The marker-newly-added linkage map consists of 198 SSR markers and covers a length of 1765.6 cM with the average distance of 8.96 cM between adjacent markers.
2. By using CIM method, a total of 21 additive QTLs for phosphorus deficiency tolerance related traits were detected at two development stages, which distributed through chromosome 4, 5, 6, 7, 10, 11. Among them, 4 was for relative root dry weight(RRDW), 3 was for relative shoot dry weight(RSDW), 3 was for relative total dry weight(RTDW), 5 was for relative root length(RRL), 4 was for relative tillering ability(RTA) and 2 was for relative root/shoot ratio(RRRS). In addition, 3 pairs of epistatic effect between QTLs which have no additive effect were also detected on chromosome 1, 5, 6, and 9. At tillering stage, QTLs for RSDW, RTDW and RRRS were detected at the same region of RM307-RM5953 on chromosome 4 and QTLs for RRDW and RTA were detected at the same region of RM311-RM6142 on chromosome 10. The situation that QTLs for different traits localized at the same region was also found at heading stage. Except for RRL, same QTLs were detected for the other 5 traits at the same region at tillering stage and heading stage. Interestingly, among three pairs of epistatic effect, one pair was detected at two development stages. A cluster of QTLs were detected at the region of RM307-RM5953 on chromosome 4 in which phosphorus efficiency related gene may lie.
3. Under phosphorus surficient condition, the root acid phosphatase activity of male parent R9308 was higher than that of female parent Xie B. Under phosphorus stress condition, however, the situation is quite reverse although the root acid phosphatase activity of the parents and RIL population lines increased at both development stages, which indicated that the genetic system for acid phosphatase under stress condition was different with that for acid phosphatase under normal culture. The simple correlation analysis showed that the contribution of AAP to phosphorus efficiency at tillering stage was greater than that of at heading stage. There were two QTLs for RRAPA at tillering stage, but no QTL was detected at heading stage.
引文
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