小麦苗期耐旱耐低磷胁迫相关性状的QTL定位及遗传图谱的构建
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摘要
小麦（Triticum aestivum L.）是我国主要的粮食作物之一，干旱和土壤缺磷都是限制小麦获得高产、稳产的重要因素，而耐旱、耐低磷胁迫都是由微效多基因控制的复杂数量性状，利用传统的细胞遗传学和数量遗传学方法难以对其进行深入研究。本研究以3个关联RIL群体为材料，对小麦苗期耐旱和耐低磷胁迫的相关性状进行条件和非条件QTL分析，以期明确与苗期耐旱和耐低磷胁迫相关的位点在染色体上的位置和效应，在QTL水平揭示耐旱和耐低磷胁迫与苗期相关性状之间的遗传关系，此研究对于小麦耐旱、耐低磷胁迫相关性状的遗传改良具有重要的意义。同时以潍麦8号和冀8901为亲本杂交获得的RIL群体为材料，采用基于PCR原理的分子标记和单核苷酸多态性（SNP）标记构建新的连锁图谱，以期将其与本实验室已构建的3个遗传图谱进行整合，为小麦农艺性状的多群体联合定位奠定基础。
本研究获得以下主要结果：
（1）以WL（179个株系）、WY（172个株系）、WJ（175个株系）三个关联的重组自交系群体为材料，对小麦苗期在正常水分和干旱胁迫条件下的胚芽鞘长、苗高、最大根长、根数、苗干重、茎叶干重、根干重、根冠干重比等8个表型性状进行非条件和条件QTL分析，共检测到121个位点，分布在小麦的21条染色体上，可分别解释表型变异的2.12～28.56%，71个位点为贡献率大于10%的主效QTL。其中，通过非条件QTL分析检测到87个QTL，12个QTL在两个或两个以上的群体内同时检测到，18个QTL在两种水分条件下同时检测到，38个QTL只在正常水分条件下检测到，31个QTL只在干旱胁迫条件下检测到。通过条件QTL分析共检测到34个与苗期耐旱性相关的QTL，其中8个QTL在干旱胁迫下苗期性状的非条件QTL分析中也被检测到，说明条件QTL可以有效地排除性状正常表现效应的影响，而检测到真正与耐旱性有关的位点。发现了11个与小麦幼苗性状和耐旱性表达有关的重要的QTL簇，分别位于1BL、1D、2A、2B、2D、4A、6B、7B染色体上，9个QTL簇包含两个以上的群体检测到的位点，8个QTL簇包含通过条件QTL分析检测到的与耐旱性相关的位点，说明这些QTL簇所在的染色体区段可能为调控苗期耐旱相关性状表达的重要区域。
（2）以WL、WY、WJ三个关联重组自交系群体为材料，对小麦苗期在正常供磷和低磷胁迫条件下的苗高、最大根长、主根数、苗干重、茎叶干重、根干重、茎叶磷利用效率、根磷利用效率、苗磷利用效率、茎叶磷含量、根磷含量、苗磷含量等12个表型性状进行了非条件和条件QTL分析，共检测到159个位点，分布在小麦的21条染色体上，可分别解释表型变异的4.04～71.12%，其中，105个位点为贡献率大于10%的主效QTL。通过条件QTL分析检测到110个QTL，10个QTL在同一群体的两种磷素水平下被检测到，38个QTL只在正常供磷条件下检测到，48个QTL只在低磷胁迫条件下检测到，19个QTL被两个群体同时检测到。条件QTL分析检测到54个QTL，10个QTL在低磷胁迫的非条件QTL分析中也被检测到。说明低磷胁迫下通过性状的非条件QTL分析检测到的大部分QTL只与性状的表达有关，而与耐低磷胁迫没有直接的关系。检测到16个与小麦苗期耐低磷胁迫性状相关的重要的QTL簇，分别位于1D、2A、2B、3A、3B、4B、4D、5A、5D、6A、6B染色体上，其中4个QTL簇包含的位点来自WL、WY、WJ三个群体，10个QTL簇的位点来自两个群体，10个QTL簇既包括非条件分析检测到的位点，又包括条件分析检测到的耐低磷胁迫位点。说明这些QTL簇所在的区段可能为调控苗期耐低磷胁迫相关性状表达的重要区域。
（3）利用包含163个株系的潍冀RIL群体进行基于PCR原理的分子标记和单核苷酸多态性（SNP）标记的遗传图谱的构建，获得了包括3916个标记，1257个位点的分子标记遗传连锁图谱，其中，基于PCR原理的分子标记有143个，SNP标记有3773个，图谱总长度2291.6cM，位点间平均遗传距离为1.82cM，由于相邻标记间距离大于50cM，在2A染色体上形成了一个连锁断点。A基组含有541个位点，全长为925.9cM，位点间平均遗传距离为1.71cM；B基组含有574个位点，全长为899.1cM，位点间平均遗传距离为1.57cM；D基组位点数最少，含有142个位点，全长为466.6cM，位点间平均遗传距离为3.29cM。
（4）3个重要的QTL簇在小麦苗期耐旱和耐低磷胁迫这两个试验中被同时检测到，第一个QTL簇位于1D染色体的wpt67199-Xwmc429.3标记区间内，与耐旱、耐低磷胁迫、根长、根数、根磷利用效率、苗和茎叶磷含量等性状的表达有关。第二个QTL簇位于2B染色体的Xbarc55.1-wpt-3561标记区间内，与耐旱、耐低磷胁迫、根数、苗和茎叶磷利用效率、磷积累量、苗干重、茎叶干重等性状的表达有关。第三个QTL簇位于7B染色体wpt669158-wpt-666615标记区间内，与耐旱、耐低磷胁迫、茎叶、苗干重、苗磷含量、苗磷利用效率等性状的表达有关。能在不同群体，不同环境，不同生长发育阶段，不同性状的QTL分析中被检测到，说明这三个QTL簇所在的染色体区段可能对小麦的生长发育具有重要的影响，有必要对其进一步深入研究。
Wheat(Triticum aestivum L.）is one of the major food crops. Drought and phosphorusdeficiency in soil are major factors limiting high and stable yield of wheat. Drought andlow-phosphorus tolerance are all complex quantitative traits controlled by polygenes withminor effects, and it is difficult to conduct intensive study by traditional cytogenetics andquantitative genetics. Using three related F9recombinant inbred line (RIL) populations asmaterials, unconditional and conditional QTL analysis of seedling traits related to drought andlow-phosphorus tolerance in wheat were conducted in this paper. Aim was to explicit theposition and the effect of loci concerned with drought and low-phosphorus tolerance in wheat,and reveal the genetic relationship between drought and low-phosphorus tolerance andseedling traits at the QTL level，respectively. It would be important for the geneticimprovement of wheat for increasing drought tolerance and efficient utilization of phosphorus.At the same time, a new genetic map based on PCR-based molecular marker and singlenucleotide polymorphism (SNP) was constructed using another RIL population derived fromcrossing between Weimai8and Ji8901as material. Aim was to make it integrate with otherthree genetic maps constructed by our lab and take foundation for multi-population jointlyQTL analysis of agricultural traits in wheat. The main results were as follows:
(1) Conditional and unconditional analysis of8traits of wheat seedling—coleoptilelength (CL), plant height (PH), the longest root length (RL), root number (RN), seedling dryweight (SDW), stem-and-leaves dry weight (SLDW), root dry weight (RDW), root-to-shootdry weight ratio (RSDWR)—were conducted under two water conditions using three F9RILpopulations.Total of121QTLs located on all of the21chromosomes of wheat were detected,accounting for2.12-28.56%of phenotypic variance respectively. All of them,71were majorQTLs with a contribution rate greater than10%. Eighty seven QTLs were detected byunconditional QTL analysis. All of them,12were detected in more than one population,18were simultaneously detected under the two water conditions,38were detected under onlynormal water condition, and31were detected only under drought stress condition. Total34 QTL related to drought tolerance of seedling were detected. Only8QTL were also detectedby unconditional analyses of the seedling traits under drought stress condition. It indicatedthat conditional QTL analysis could exclude effectively the influence of normal expressioneffect of trait to detect QTL really related to drought tolerance. Eleven important QTL clusterslocated on chromosome1BL,1D,2A,2B,2D,4A,6B, and7B were discoved. All of them,9included QTLs detected in more than one population, and8included QTLs related to droughttolerance by conditional QTL analysis. It indicated that these chromosomal intervals withQTL clusters should be important areas controlling the expression of seedling traits related todrought tolerance.
(2) Conditional and unconditional QTL analyses of the12traits of wheatseedlings—plant height (PH), the longest root length (RL), root number (RN), seedling dryweight (SDW), stem-and-leave dry weight (SLDW), root dry weight (RDW), stem and leavephosphorus utilization efficiency (SLPU), root phosphorus utilization efficiency (RPU), shootphosphorus utilization efficiency (SPU), stem and leave phosphorus content (SLPC), rootphosphorus content (RPC), shoot phosphorus content (SPC)—were conducted under normalphosphorus and low-phosphorus conditions using three F9recombinant inbred line (RIL)populations. Total of159QTLs located on the21chromosomes of wheat were detected,accounting for4.04-71.12%of phenotypic variance respectively. All of them,105were majorQTLs with a contribution rate greater than10%. One hundred and ten QTLs were detected byunconditional QTL analysis. Among of these QTLs,19were detected simultaneously in twopopulations,10were detected simultaneously under two phosphorus conditions,38weredetected only under normal phosphorus condition, and48were detected underlow-phosphorus condition. Total of54QTLs related to low-phosphorus tolerance of seedlingwere detected. Only10QTLs were also detected by unconditional analysis of seedling traitsunder low-phosphorus condition. It indicated that most of the QTLs detected by unconditionalQTL analysis of seedling traits under low-phosphorus condition were only concerned withexpression of these traits, but not concerned directly with low-phosphorus tolerance. Sixteenimportant QTL clusters located on chromosome1D,2A,2B,3A,3B,4B,4D,5A,5D,6A, and6B were discovered. All of them,4included loci detected in WL, WY, and WJ threepopulations,10included loci detected in two populations, and10included loci related tolow-phosphorus tolerance. It indicated that these chromosome intervals with these QTLclusters should be important areas controlling the expression of seedling traits related tolow-phosphorus tolerance.
(3) A genetic map comprising3916marker and1257loci were constructed based on PCR-based and SNP molecular markers using the recombinant inbred line (RIL) populationderived from crossing of Weimai8and Ji8901. The genetic map spaned2291.6cM, with anaverage density of one marker per1.82cM. Due to the linkage distance>50cM between theadjacent loci, there was a linkage gaps in chromosome2A. A genome included541loci andspaned925.9cM, with an average density of one locus per1.71cM. B genome included574loci and spaned899.1cM, with an average density of one locus per1.57cM. D genomeincluded142loci and spaned466.6cM, with an average density of one locus per3.29cM.
(4) Three important QTL clusters were detected simultaneously in both drought toleranceand low-phosphorus tolerance trials of wheat seedling. The first QTL cluster located onchromosome1D between wpt-67199and Xwmc429.3, related to drought tolerance,low-phosphorus tolerance, RL, RN, RPUE, SPC, and SLPC. The second QTL cluster locatedon chromosome2B between Xbarc55.1and wpt-3561, related to drought tolerance,low-phosphorus tolerance, RN, SPUE, SLPUE, SPC, SLPC, SDW, and SLDW. The thirdQTL cluster located on chromosome7B between wpt-669158and wpt-666615, related todrought tolerance, low-phosphorus tolerance, SPUE, SPC, SDW, and SLDW. These threeQTL clusters could be detected by QTL analyses of different population, differentenvironment, different growth and development stage, and different trait, which indicated thatthese three QTL clusters could have significant influence in the growth and development ofwheat and it is necessary to conduct more intensive study.
本研究获得以下主要结果：
（1）以WL（179个株系）、WY（172个株系）、WJ（175个株系）三个关联的重组自交系群体为材料，对小麦苗期在正常水分和干旱胁迫条件下的胚芽鞘长、苗高、最大根长、根数、苗干重、茎叶干重、根干重、根冠干重比等8个表型性状进行非条件和条件QTL分析，共检测到121个位点，分布在小麦的21条染色体上，可分别解释表型变异的2.12～28.56%，71个位点为贡献率大于10%的主效QTL。其中，通过非条件QTL分析检测到87个QTL，12个QTL在两个或两个以上的群体内同时检测到，18个QTL在两种水分条件下同时检测到，38个QTL只在正常水分条件下检测到，31个QTL只在干旱胁迫条件下检测到。通过条件QTL分析共检测到34个与苗期耐旱性相关的QTL，其中8个QTL在干旱胁迫下苗期性状的非条件QTL分析中也被检测到，说明条件QTL可以有效地排除性状正常表现效应的影响，而检测到真正与耐旱性有关的位点。发现了11个与小麦幼苗性状和耐旱性表达有关的重要的QTL簇，分别位于1BL、1D、2A、2B、2D、4A、6B、7B染色体上，9个QTL簇包含两个以上的群体检测到的位点，8个QTL簇包含通过条件QTL分析检测到的与耐旱性相关的位点，说明这些QTL簇所在的染色体区段可能为调控苗期耐旱相关性状表达的重要区域。
（2）以WL、WY、WJ三个关联重组自交系群体为材料，对小麦苗期在正常供磷和低磷胁迫条件下的苗高、最大根长、主根数、苗干重、茎叶干重、根干重、茎叶磷利用效率、根磷利用效率、苗磷利用效率、茎叶磷含量、根磷含量、苗磷含量等12个表型性状进行了非条件和条件QTL分析，共检测到159个位点，分布在小麦的21条染色体上，可分别解释表型变异的4.04～71.12%，其中，105个位点为贡献率大于10%的主效QTL。通过条件QTL分析检测到110个QTL，10个QTL在同一群体的两种磷素水平下被检测到，38个QTL只在正常供磷条件下检测到，48个QTL只在低磷胁迫条件下检测到，19个QTL被两个群体同时检测到。条件QTL分析检测到54个QTL，10个QTL在低磷胁迫的非条件QTL分析中也被检测到。说明低磷胁迫下通过性状的非条件QTL分析检测到的大部分QTL只与性状的表达有关，而与耐低磷胁迫没有直接的关系。检测到16个与小麦苗期耐低磷胁迫性状相关的重要的QTL簇，分别位于1D、2A、2B、3A、3B、4B、4D、5A、5D、6A、6B染色体上，其中4个QTL簇包含的位点来自WL、WY、WJ三个群体，10个QTL簇的位点来自两个群体，10个QTL簇既包括非条件分析检测到的位点，又包括条件分析检测到的耐低磷胁迫位点。说明这些QTL簇所在的区段可能为调控苗期耐低磷胁迫相关性状表达的重要区域。
（3）利用包含163个株系的潍冀RIL群体进行基于PCR原理的分子标记和单核苷酸多态性（SNP）标记的遗传图谱的构建，获得了包括3916个标记，1257个位点的分子标记遗传连锁图谱，其中，基于PCR原理的分子标记有143个，SNP标记有3773个，图谱总长度2291.6cM，位点间平均遗传距离为1.82cM，由于相邻标记间距离大于50cM，在2A染色体上形成了一个连锁断点。A基组含有541个位点，全长为925.9cM，位点间平均遗传距离为1.71cM；B基组含有574个位点，全长为899.1cM，位点间平均遗传距离为1.57cM；D基组位点数最少，含有142个位点，全长为466.6cM，位点间平均遗传距离为3.29cM。
（4）3个重要的QTL簇在小麦苗期耐旱和耐低磷胁迫这两个试验中被同时检测到，第一个QTL簇位于1D染色体的wpt67199-Xwmc429.3标记区间内，与耐旱、耐低磷胁迫、根长、根数、根磷利用效率、苗和茎叶磷含量等性状的表达有关。第二个QTL簇位于2B染色体的Xbarc55.1-wpt-3561标记区间内，与耐旱、耐低磷胁迫、根数、苗和茎叶磷利用效率、磷积累量、苗干重、茎叶干重等性状的表达有关。第三个QTL簇位于7B染色体wpt669158-wpt-666615标记区间内，与耐旱、耐低磷胁迫、茎叶、苗干重、苗磷含量、苗磷利用效率等性状的表达有关。能在不同群体，不同环境，不同生长发育阶段，不同性状的QTL分析中被检测到，说明这三个QTL簇所在的染色体区段可能对小麦的生长发育具有重要的影响，有必要对其进一步深入研究。
Wheat(Triticum aestivum L.）is one of the major food crops. Drought and phosphorusdeficiency in soil are major factors limiting high and stable yield of wheat. Drought andlow-phosphorus tolerance are all complex quantitative traits controlled by polygenes withminor effects, and it is difficult to conduct intensive study by traditional cytogenetics andquantitative genetics. Using three related F9recombinant inbred line (RIL) populations asmaterials, unconditional and conditional QTL analysis of seedling traits related to drought andlow-phosphorus tolerance in wheat were conducted in this paper. Aim was to explicit theposition and the effect of loci concerned with drought and low-phosphorus tolerance in wheat,and reveal the genetic relationship between drought and low-phosphorus tolerance andseedling traits at the QTL level，respectively. It would be important for the geneticimprovement of wheat for increasing drought tolerance and efficient utilization of phosphorus.At the same time, a new genetic map based on PCR-based molecular marker and singlenucleotide polymorphism (SNP) was constructed using another RIL population derived fromcrossing between Weimai8and Ji8901as material. Aim was to make it integrate with otherthree genetic maps constructed by our lab and take foundation for multi-population jointlyQTL analysis of agricultural traits in wheat. The main results were as follows:
(1) Conditional and unconditional analysis of8traits of wheat seedling—coleoptilelength (CL), plant height (PH), the longest root length (RL), root number (RN), seedling dryweight (SDW), stem-and-leaves dry weight (SLDW), root dry weight (RDW), root-to-shootdry weight ratio (RSDWR)—were conducted under two water conditions using three F9RILpopulations.Total of121QTLs located on all of the21chromosomes of wheat were detected,accounting for2.12-28.56%of phenotypic variance respectively. All of them,71were majorQTLs with a contribution rate greater than10%. Eighty seven QTLs were detected byunconditional QTL analysis. All of them,12were detected in more than one population,18were simultaneously detected under the two water conditions,38were detected under onlynormal water condition, and31were detected only under drought stress condition. Total34 QTL related to drought tolerance of seedling were detected. Only8QTL were also detectedby unconditional analyses of the seedling traits under drought stress condition. It indicatedthat conditional QTL analysis could exclude effectively the influence of normal expressioneffect of trait to detect QTL really related to drought tolerance. Eleven important QTL clusterslocated on chromosome1BL,1D,2A,2B,2D,4A,6B, and7B were discoved. All of them,9included QTLs detected in more than one population, and8included QTLs related to droughttolerance by conditional QTL analysis. It indicated that these chromosomal intervals withQTL clusters should be important areas controlling the expression of seedling traits related todrought tolerance.
(2) Conditional and unconditional QTL analyses of the12traits of wheatseedlings—plant height (PH), the longest root length (RL), root number (RN), seedling dryweight (SDW), stem-and-leave dry weight (SLDW), root dry weight (RDW), stem and leavephosphorus utilization efficiency (SLPU), root phosphorus utilization efficiency (RPU), shootphosphorus utilization efficiency (SPU), stem and leave phosphorus content (SLPC), rootphosphorus content (RPC), shoot phosphorus content (SPC)—were conducted under normalphosphorus and low-phosphorus conditions using three F9recombinant inbred line (RIL)populations. Total of159QTLs located on the21chromosomes of wheat were detected,accounting for4.04-71.12%of phenotypic variance respectively. All of them,105were majorQTLs with a contribution rate greater than10%. One hundred and ten QTLs were detected byunconditional QTL analysis. Among of these QTLs,19were detected simultaneously in twopopulations,10were detected simultaneously under two phosphorus conditions,38weredetected only under normal phosphorus condition, and48were detected underlow-phosphorus condition. Total of54QTLs related to low-phosphorus tolerance of seedlingwere detected. Only10QTLs were also detected by unconditional analysis of seedling traitsunder low-phosphorus condition. It indicated that most of the QTLs detected by unconditionalQTL analysis of seedling traits under low-phosphorus condition were only concerned withexpression of these traits, but not concerned directly with low-phosphorus tolerance. Sixteenimportant QTL clusters located on chromosome1D,2A,2B,3A,3B,4B,4D,5A,5D,6A, and6B were discovered. All of them,4included loci detected in WL, WY, and WJ threepopulations,10included loci detected in two populations, and10included loci related tolow-phosphorus tolerance. It indicated that these chromosome intervals with these QTLclusters should be important areas controlling the expression of seedling traits related tolow-phosphorus tolerance.
(3) A genetic map comprising3916marker and1257loci were constructed based on PCR-based and SNP molecular markers using the recombinant inbred line (RIL) populationderived from crossing of Weimai8and Ji8901. The genetic map spaned2291.6cM, with anaverage density of one marker per1.82cM. Due to the linkage distance>50cM between theadjacent loci, there was a linkage gaps in chromosome2A. A genome included541loci andspaned925.9cM, with an average density of one locus per1.71cM. B genome included574loci and spaned899.1cM, with an average density of one locus per1.57cM. D genomeincluded142loci and spaned466.6cM, with an average density of one locus per3.29cM.
(4) Three important QTL clusters were detected simultaneously in both drought toleranceand low-phosphorus tolerance trials of wheat seedling. The first QTL cluster located onchromosome1D between wpt-67199and Xwmc429.3, related to drought tolerance,low-phosphorus tolerance, RL, RN, RPUE, SPC, and SLPC. The second QTL cluster locatedon chromosome2B between Xbarc55.1and wpt-3561, related to drought tolerance,low-phosphorus tolerance, RN, SPUE, SLPUE, SPC, SLPC, SDW, and SLDW. The thirdQTL cluster located on chromosome7B between wpt-669158and wpt-666615, related todrought tolerance, low-phosphorus tolerance, SPUE, SPC, SDW, and SLDW. These threeQTL clusters could be detected by QTL analyses of different population, differentenvironment, different growth and development stage, and different trait, which indicated thatthese three QTL clusters could have significant influence in the growth and development ofwheat and it is necessary to conduct more intensive study.
引文
曹卫东,贾继增,金继运.小麦磷素利用效率的基因位点及其交互作用[J].植物营养与肥料学报.2001,7(3):285-292
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