甘蓝型油菜硼营养及离子组遗传学研究
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摘要
作物的生长因为土壤中矿质营养元素的缺乏或过量,会出现生长的紊乱,并发各种缺素综合症,最终导致作物减产和品质下降。在我国长江中下游油菜主产区微量元素硼缺乏比较严重,该地区曾出现过因为硼的缺乏导致油菜和棉花大量的减产或绝收。尽管现代农业在作物栽培过程中以增施硼肥的措施能保证油菜正常生长,但肥料的施用无疑给农户增添了经济上的负担,而且肥料随着农田地表水的流失,不仅仅带来肥料的浪费,也给江河湖泊水资源带来了严重的污染。发展大田绿色作物,培育肥料高效利用的作物品种,保持高产稳产的同时尽量降低肥料用量是现在农业研究中的重要课题。对于油菜硼高效的生理基础,前人作了很多研究,也提出了很多的假设,但都没有从根本上解释硼高效的生理和遗传机理。本研究为探明硼高效遗传规律,调查了缺硼环境下油菜籽粒产量和硼效率系数两个性状,检测到其对应的遗传位点。并调查油菜苗期地上部和根中硼和其他六种矿质离子的含量,将离子组和数量遗传学结合,寻找油菜基因组中控制离子含量的遗传位点。借助油菜和拟南芥基因组同源关系,用比较定位的办法找到同源候选基因。
1.大田试验的油菜产量相关性状硼效率遗传分析
TN群体202个株系,于2004和2005两年栽种在蕲春缺硼大田土壤中,苗期分别施用硼1.5 kg/ha和15 kg/ha以设置两组硼水平,三个重复,每个重复内的株系采取随机区组分布种植。营养生长阶段,低硼环境下的油菜长势明显弱于正常硼环境下的油菜;高效亲本宁油7和高效株系生长正常,而低效亲本Tapidor和低效株系出现叶缘紫红,茎纵裂,生长点坏死,多头等典型的缺硼症状。油菜蕾薹期对硼的缺乏最为敏感,缺硼油菜部分株系花期明显延迟,出现“返花”等症状,花蕾很多枯败,没有结实,出现典型的“花而不实”症状。
为调查缺硼影响的产量性状,以每株系缺硼条件下的籽粒产量和硼效率系数作为指标,两年试验共检测到10个硼高效QTL和10对相关的互作位点。其中A2和A6连锁群上的QTL位点,在两年的试验中能重复被检测到,而且其中A2上的位点与以前BQ群体中定位的位点吻合,是一个较稳定的低硼环境下的产量相关QTL。而该位点同时还和基因组中其他位点发生互作,可能涉及更复杂的分子遗传机理。
2.油菜苗期地上部和根在两种硼环境下的七种离子含量分析
设置两个硼营养液培养水平,低硼水平为0.25μM,硼正常水平为50μM。分别测量TN亲本和群体地上部和根中7种离子的含量,而受胁迫影响的B含量变化最为明显,其次为Cu,Fe和Zn。而且根中和地上部的元素受影响程度不一致。
3.七种离子含量的遗传分析
两种B水平下,共检测到离子含量相关QTL107个和互作对154对。其中发现地上部的Ca和Mg的QTL存在三处重叠,另外位于Al连锁群上的地上部主效P-QTL稳定存在,不受环境B含量高低的影响。也有多处根中离子含量的QTL位点,特别是Ca,Mg,Cu,Fe,和Zn这些阳离子之间,有多达7处重叠位点。说明根中很多位点可能存在这些二价阳离子的一些共同的非选择性吸收机制。
4.同源转运基因在TN图谱上的in silico定位及于QTL的比对
根据拟南芥和油菜之间的同源共线关系,我们借助TN遗传图上含有序列信息的分子标记,将拟南芥的基因组区段比对到TN遗传图上,用以辅助分析QTL的位置以及和拟南芥基因组的同源区段。根据拟南芥中已发现的矿质元素转运基因和控制离子自稳态的转录因子等基因位置信息,在TN遗传图谱上,我们共电子定位了352个同源基因,发现其中46个基因位于先前定位的矿质元素QTL中。这种利用比较基因组的方法确定不同物种的同源基因和QTL关系的方法,为下一步定位克隆这些基因提供了丰富的信息。
5.根据硼转运子发展特异标记定位到TN遗传图上
根据已发现的拟南芥的硼转运基因NIP5;1,BORl及其家族的同源基因设计引物,在TN遗传图谱上定位5个硼转运子同源序列的SSCP标记,其中一个位于先前定位的A5连锁群低硼环境下根长QTL区间内。并用分离集团的策略将两个转运子起源的TRAP标记定位到A2连锁群的低硼产量QTL区间中。
6.AFLP+BSA策略加密QTL区间
用AFLP分离集团策略加密A2上的低硼产量高效QTL区间分子标记密度,在原本标记稀疏的30 cM区段内,增加了9个AFLP标记,为精细定位该QTL区间作了前期准备。
The growth of crops often occur various disorders because of the lack or excessive of elements in the soil.Middle and lower reaches of the Yangtze River in China's is main producing areas of rapeseed.In the past,a large number of rapeseed and cotton decreased production or total destruction for more serious lack of boron(B) in the region.Although fertilizer in the process of cultivation could guarantee the normal growth of crop,but the application of fertilizers to farmers undoubtedly added economic burden;and the loss of fertilizer from farmland by rain,resulted in not only waste of fertilizer,also caused serious water pollution.The development of green crops,and cultivating the efficient use of fertilizer varieties,in maintaining high and stable yield at the same time reducing the amount of fertilizer is an important subject in agricultural research.A lot of studies were concerning about the physiological basis of B efficient for oilseed.Although a lot of assumptions were proposed,but there still were not fundamentally explains for the mechanism of B efficient.
Detection of B efficent loci in rapeseed genome is our research objection by investigating characters of mature stage traits combining with TN genetic resourses.A further inquiry was firstly performed about preliminary ionomics exploration in the seedling stage.
1.Genetic analysis of rapeseed B efficiency in field experiments
TN population of 202 lines was planted for two years in B deficiency soil,Qichun of Hubei.1.5 kg/ha of boron and 15 kg/ha of boron were fertilized to design two levels of controlled experiment.The experiment included three repeats,each repeat of random block design of 202 lines with the parents as control.In vegetative growth stage,rapeseed under low B environment grew significantly weaker than under normal B environment.In low B environment,efficient parent(Ningyou7) and efficient DH lines grew normally, but inefficient parent(Tapidor) and inefficient DH lines appeared with typical symptoms of B deficiency:purplish leaf edge,cracked stem,the SAM necrosis,many-branch.In bolting stage,rapeseed is most sensitive of the lack of B.Flowering time was significantly delayed for B deficiency with pod lost.It's the typical symptoms with "flowering not seeding".
Two traits(the seed production under the low B environment and B efficient coefficient) were tested.Ten QTL and ten interaction pairs were detected.The two QTL on A2 and A6 lingkage groups could be validated by two years results.The major QTL on A2 linkage group of TN pupulation could be identical with QTL in LG9 of BQ population,The result found that a stable production-related QTL under low B conditions exist across two genetic populations.At the same time,we found the QTL on A2 linkage group interacted with other genetic locus in genome.It suggest complicated epistatic effect exists in the major B QTL.
2.Concentrations of seven mineral elements of rapeseed seedling in the two B conditions
Two B levels of hydroponical culture were low B levels for 0.25μM,B normal level for 50μM,respectively.The concentrations of seven mineral elements in shoots and roots were measured.B concentration has been most intensely decreased because of the stress, followed increased with concentrations of Cu,Fe and Zn.But fluctuation of elements concentrations in roots is comparative with elements in shoots.
3.Genetic analysis for the concentrations of mineral elements
In the two B conditions,total 107 QTL and 154 interation pairs were detected for the concentrations of 7 mineral elements.In shoots,wefound three overlapping QTL for Ca/Mg and the biggest effect P-QTL across two B conditions.In roots,QTL for cations (Ca,Mg,Cu,Fe and Zn) has 7 overlapping QTL intervals.It means the cations may exist the common non-selective absorption mechanism.
4.Small regions of the Arabidopsis genome have already been targeted by comparative mapping between B.napus and Arabidopsis.
The genes that were related to mineral absorption and allocation in each syntenic block of Arabidopsis were located according to their physical positions in the genome of Arabidopsis.The positions of putative genes were then aligned with the TN linkage map according to the position of close anchor marker(s) in the same syntenic block.If the position of an aligned gene(s) was located in the confidence interval of a QTL,the orthologous gene(s) was considered to be associated with the target QTL.By the method of in silico mapping,hundreds of mineral transporting genes in Arabidopsis have been aligned with the linkage map of B.napus.
In our study,alignment results were used to construct blocks of syntenic or insertion fragment islands between the Arabidopsis chromosomes and TN linkage groups. According to homologue relationship between Arabidopsis and rapeseed,the genetic map of TN population with Arabidopsis thaliana section,used to support the position of QTL and there may be some candidate genes.We have found in the various ion transporter gene through this comparative genomic methods to determine their genes in the genome, to pave the way for map-based cloning of these genes.
5.According to the sequence of the transporter genes of NIP5;1,BORl family in Arabidopsis,the primers were designed and amplified in TN population.Five single-strand conformation polymorphism(SSCP) markers were got and single marker was located in QTL intervcal for root length in low B condition.
By using target region polymorphism(TRAP) and bulked segregate analysis(BSA), we identified 2 TRAP markers linked to the B efficient locus on A2 lingkage group.The markers will be useful for marker-assisted selection(MAS) in breeding programs and provide the bases for map-based cloning of this gene.
6.AFLP markers were used to densify the target QTL interval on A2 linkage group. The original interval covers 30 cM with sparse markers.By BSA method,9 AFLP markers were added in the QTL interval.
1.大田试验的油菜产量相关性状硼效率遗传分析
TN群体202个株系,于2004和2005两年栽种在蕲春缺硼大田土壤中,苗期分别施用硼1.5 kg/ha和15 kg/ha以设置两组硼水平,三个重复,每个重复内的株系采取随机区组分布种植。营养生长阶段,低硼环境下的油菜长势明显弱于正常硼环境下的油菜;高效亲本宁油7和高效株系生长正常,而低效亲本Tapidor和低效株系出现叶缘紫红,茎纵裂,生长点坏死,多头等典型的缺硼症状。油菜蕾薹期对硼的缺乏最为敏感,缺硼油菜部分株系花期明显延迟,出现“返花”等症状,花蕾很多枯败,没有结实,出现典型的“花而不实”症状。
为调查缺硼影响的产量性状,以每株系缺硼条件下的籽粒产量和硼效率系数作为指标,两年试验共检测到10个硼高效QTL和10对相关的互作位点。其中A2和A6连锁群上的QTL位点,在两年的试验中能重复被检测到,而且其中A2上的位点与以前BQ群体中定位的位点吻合,是一个较稳定的低硼环境下的产量相关QTL。而该位点同时还和基因组中其他位点发生互作,可能涉及更复杂的分子遗传机理。
2.油菜苗期地上部和根在两种硼环境下的七种离子含量分析
设置两个硼营养液培养水平,低硼水平为0.25μM,硼正常水平为50μM。分别测量TN亲本和群体地上部和根中7种离子的含量,而受胁迫影响的B含量变化最为明显,其次为Cu,Fe和Zn。而且根中和地上部的元素受影响程度不一致。
3.七种离子含量的遗传分析
两种B水平下,共检测到离子含量相关QTL107个和互作对154对。其中发现地上部的Ca和Mg的QTL存在三处重叠,另外位于Al连锁群上的地上部主效P-QTL稳定存在,不受环境B含量高低的影响。也有多处根中离子含量的QTL位点,特别是Ca,Mg,Cu,Fe,和Zn这些阳离子之间,有多达7处重叠位点。说明根中很多位点可能存在这些二价阳离子的一些共同的非选择性吸收机制。
4.同源转运基因在TN图谱上的in silico定位及于QTL的比对
根据拟南芥和油菜之间的同源共线关系,我们借助TN遗传图上含有序列信息的分子标记,将拟南芥的基因组区段比对到TN遗传图上,用以辅助分析QTL的位置以及和拟南芥基因组的同源区段。根据拟南芥中已发现的矿质元素转运基因和控制离子自稳态的转录因子等基因位置信息,在TN遗传图谱上,我们共电子定位了352个同源基因,发现其中46个基因位于先前定位的矿质元素QTL中。这种利用比较基因组的方法确定不同物种的同源基因和QTL关系的方法,为下一步定位克隆这些基因提供了丰富的信息。
5.根据硼转运子发展特异标记定位到TN遗传图上
根据已发现的拟南芥的硼转运基因NIP5;1,BORl及其家族的同源基因设计引物,在TN遗传图谱上定位5个硼转运子同源序列的SSCP标记,其中一个位于先前定位的A5连锁群低硼环境下根长QTL区间内。并用分离集团的策略将两个转运子起源的TRAP标记定位到A2连锁群的低硼产量QTL区间中。
6.AFLP+BSA策略加密QTL区间
用AFLP分离集团策略加密A2上的低硼产量高效QTL区间分子标记密度,在原本标记稀疏的30 cM区段内,增加了9个AFLP标记,为精细定位该QTL区间作了前期准备。
The growth of crops often occur various disorders because of the lack or excessive of elements in the soil.Middle and lower reaches of the Yangtze River in China's is main producing areas of rapeseed.In the past,a large number of rapeseed and cotton decreased production or total destruction for more serious lack of boron(B) in the region.Although fertilizer in the process of cultivation could guarantee the normal growth of crop,but the application of fertilizers to farmers undoubtedly added economic burden;and the loss of fertilizer from farmland by rain,resulted in not only waste of fertilizer,also caused serious water pollution.The development of green crops,and cultivating the efficient use of fertilizer varieties,in maintaining high and stable yield at the same time reducing the amount of fertilizer is an important subject in agricultural research.A lot of studies were concerning about the physiological basis of B efficient for oilseed.Although a lot of assumptions were proposed,but there still were not fundamentally explains for the mechanism of B efficient.
Detection of B efficent loci in rapeseed genome is our research objection by investigating characters of mature stage traits combining with TN genetic resourses.A further inquiry was firstly performed about preliminary ionomics exploration in the seedling stage.
1.Genetic analysis of rapeseed B efficiency in field experiments
TN population of 202 lines was planted for two years in B deficiency soil,Qichun of Hubei.1.5 kg/ha of boron and 15 kg/ha of boron were fertilized to design two levels of controlled experiment.The experiment included three repeats,each repeat of random block design of 202 lines with the parents as control.In vegetative growth stage,rapeseed under low B environment grew significantly weaker than under normal B environment.In low B environment,efficient parent(Ningyou7) and efficient DH lines grew normally, but inefficient parent(Tapidor) and inefficient DH lines appeared with typical symptoms of B deficiency:purplish leaf edge,cracked stem,the SAM necrosis,many-branch.In bolting stage,rapeseed is most sensitive of the lack of B.Flowering time was significantly delayed for B deficiency with pod lost.It's the typical symptoms with "flowering not seeding".
Two traits(the seed production under the low B environment and B efficient coefficient) were tested.Ten QTL and ten interaction pairs were detected.The two QTL on A2 and A6 lingkage groups could be validated by two years results.The major QTL on A2 linkage group of TN pupulation could be identical with QTL in LG9 of BQ population,The result found that a stable production-related QTL under low B conditions exist across two genetic populations.At the same time,we found the QTL on A2 linkage group interacted with other genetic locus in genome.It suggest complicated epistatic effect exists in the major B QTL.
2.Concentrations of seven mineral elements of rapeseed seedling in the two B conditions
Two B levels of hydroponical culture were low B levels for 0.25μM,B normal level for 50μM,respectively.The concentrations of seven mineral elements in shoots and roots were measured.B concentration has been most intensely decreased because of the stress, followed increased with concentrations of Cu,Fe and Zn.But fluctuation of elements concentrations in roots is comparative with elements in shoots.
3.Genetic analysis for the concentrations of mineral elements
In the two B conditions,total 107 QTL and 154 interation pairs were detected for the concentrations of 7 mineral elements.In shoots,wefound three overlapping QTL for Ca/Mg and the biggest effect P-QTL across two B conditions.In roots,QTL for cations (Ca,Mg,Cu,Fe and Zn) has 7 overlapping QTL intervals.It means the cations may exist the common non-selective absorption mechanism.
4.Small regions of the Arabidopsis genome have already been targeted by comparative mapping between B.napus and Arabidopsis.
The genes that were related to mineral absorption and allocation in each syntenic block of Arabidopsis were located according to their physical positions in the genome of Arabidopsis.The positions of putative genes were then aligned with the TN linkage map according to the position of close anchor marker(s) in the same syntenic block.If the position of an aligned gene(s) was located in the confidence interval of a QTL,the orthologous gene(s) was considered to be associated with the target QTL.By the method of in silico mapping,hundreds of mineral transporting genes in Arabidopsis have been aligned with the linkage map of B.napus.
In our study,alignment results were used to construct blocks of syntenic or insertion fragment islands between the Arabidopsis chromosomes and TN linkage groups. According to homologue relationship between Arabidopsis and rapeseed,the genetic map of TN population with Arabidopsis thaliana section,used to support the position of QTL and there may be some candidate genes.We have found in the various ion transporter gene through this comparative genomic methods to determine their genes in the genome, to pave the way for map-based cloning of these genes.
5.According to the sequence of the transporter genes of NIP5;1,BORl family in Arabidopsis,the primers were designed and amplified in TN population.Five single-strand conformation polymorphism(SSCP) markers were got and single marker was located in QTL intervcal for root length in low B condition.
By using target region polymorphism(TRAP) and bulked segregate analysis(BSA), we identified 2 TRAP markers linked to the B efficient locus on A2 lingkage group.The markers will be useful for marker-assisted selection(MAS) in breeding programs and provide the bases for map-based cloning of this gene.
6.AFLP markers were used to densify the target QTL interval on A2 linkage group. The original interval covers 30 cM with sparse markers.By BSA method,9 AFLP markers were added in the QTL interval.
引文
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