小麦抗叶锈病基因Lr1的分子进化
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摘要
小麦作为人类重要的粮食作物受到多种病害的侵袭,叶锈病是其生产上的主要病害之一,传播范围广,发生区域大。它的流行将导致小麦严重的减产和品质下降。利用植物本身的抗病基因控制叶锈病的流行,是一种最经济、有效和最安全的措施,然而自然环境与人工栽培的双重选择压力加速了植物抗病基因与病原体无毒基因间的共进化过程,常常导致原有抗病基因的快速进化或丢失,因此,探索植物抗病基因进化的分子机制显得尤为重要。
本研究利用与小麦抗叶锈病基因Lrl共分离的分子标记WR003筛选了来自世界各地的128份山羊草品系和283份六倍体小麦品种(包括262份六倍体小麦微核心种质),并通过PCR克隆测序获得了42个Lrl等位基因和6个Lrl同源基因的全长序列,随后利用DNAMAN和Clustal W进行Lrl等位基因和同源基因间核酸水平和蛋白水平的序列比对,并利用程序MEGA4.0.1采用neighbor-joining(NJ)方法构建系统发生树,对小麦抗叶锈病基因Lrl进行分子进化生物学方面的研究,结果表明:
1.六倍体小麦抗叶锈病基因Lrl起源于二倍体粗山羊草,并且在其进化为六倍体小麦之前就已分化出抗病和感病两种基因型,所以六倍体小麦的Lrl至少有两个起源。
2.Lrl感病基因型和抗病基因型的分离主要体现在一个570 bp的变异区内(在第2479-3048碱基之间,对应于第827-1017氨基酸),在该变异区内除两处点突变外具有相同抗性的山羊草品系和六倍体小麦品种的核酸序列完全相同,说明在山羊草进化为六倍体小麦的过程中该区域几乎没有发生过遗传重组和基因转换。
3.在此变异区两侧的相对保守区共有21个点突变,其中有9个同义碱基突变,12个非同义碱基突变,这些点突变并非发生在抗感基因型之间,而是发生在具有相同抗性基因型的山羊草与山羊草或山羊草与六倍体小麦之间。
4.发生在感病粗山羊草与六倍体小麦之间的点突变只有一个(在2460bp处,为同义碱基突变);发生在抗病粗山羊草与六倍体小麦之间的点突变为9个(2个同义碱基突变,7个非同义碱基突变),其中1个点突变发生在570bp的变异区内,8个点突变发生在该变异区两侧的相对保守区。
5.Lrl等位基因序列中的LRR编码区内串联排列着14个同义碱基突变和51个非同义碱基突变,Ka(非同义碱基突变率)╱Ks(同义碱基突变率)＞1,说明该区域经受着岐化选择的作用,这也与前人报道的该区域为配基结合部位、控制着抗病基因的特异性相吻合。
Wheat,as one of the important crops,suffers from the infection of various diseases, wheat leaf rust is one of the major diseases in wheat production.It spreads wildly and can lead to severe yield loss and poor seed quality.The use of resistance genes is the most effective,economical and ecological method to control epidemics of leaf rust disease. However,natural environment and artificial domestication accelerated the co-evolution of host-pathogen interaction,which moreover resulted in the rapid evolution and disappearance of resistance genes.Therefore,it is necessary to study the evolutionary mechanism of plant resistance genes.
In our research,we screened 128 Aegilops.Tauschii(goat grass) accessions and 283 hexaploid wheat varieties(including 262 wheat core germplasms) with marker WR003 co-segregated with Lr1,and obtained the full-length sequences of 42 Lr1 alleles and 6 Lr1 homologous genes.Afterwards,we aligned DNA and protein sequences of Lr1 allelic or homologous genes by DNAMAN and Clustal W,and analyzed the molecular evolution mechanism for the wheat leaf rust resistance genes according to the phylogenetic tree established by neighbor-joining(NJ) method by MEGA 4.0.1.The results showed that:
1.Wheat leaf rust resistance gene Lr1 was originated from Ae.Tauschii,and differentiated into resistant and susceptible genotypes prior to its evolution into hexaploid wheat. Therefore,wheat leaf rust resistance gene Lr1 has two ancestral genes at least.
2.The allelic sequences alignment of Lr1 from resistant and susceptible lines revealed a divergent DNA sequence block of 570 bp,where Lr1 nucleotide sequences were mostly similar between hexaploid wheat and Ae.Tauschii with the same resistance except for two point mutations.So it is concluded that no genetic recombination and gene conversion happened during the evolution from Ae.Tauschii.into hexaploid wheat.
3.The rest of the sequences were mostly similar,where only 21 point mutations existed(9 synonymous mutations,and 12 nonsynonymous mutations).Interestingly,these 21 point mutations occur between lines with identical resistance rather than different resistance.
4.There was only one point mutation between resistant hexaploid wheat and Ae.Tauschii (at 2400bp down stream of the start codon ATG,a synonymous mutation);9 point mutations took place between susceptible hexaploid wheat and Ae.Tauschii(2 synonymous mutations and 7 nonsynonymous mutations),1 in the 570bp block and 8 in the rest of the sequence.
5.Fourteen synonymous mutations and 51 nonsynonymous mutations lined in tandem on the LRR coding region of the allele sequences,Ka(rate of nonsynonymous mutation)/Ks (rate of synonymous mutation)＞1.These results indicated that LRR domain suffered from the pressure of strong selection,consistent with the former reports that LRR domains were involved in the interaction with effector proteins and were the major determinant of resistance specificity.
本研究利用与小麦抗叶锈病基因Lrl共分离的分子标记WR003筛选了来自世界各地的128份山羊草品系和283份六倍体小麦品种(包括262份六倍体小麦微核心种质),并通过PCR克隆测序获得了42个Lrl等位基因和6个Lrl同源基因的全长序列,随后利用DNAMAN和Clustal W进行Lrl等位基因和同源基因间核酸水平和蛋白水平的序列比对,并利用程序MEGA4.0.1采用neighbor-joining(NJ)方法构建系统发生树,对小麦抗叶锈病基因Lrl进行分子进化生物学方面的研究,结果表明:
1.六倍体小麦抗叶锈病基因Lrl起源于二倍体粗山羊草,并且在其进化为六倍体小麦之前就已分化出抗病和感病两种基因型,所以六倍体小麦的Lrl至少有两个起源。
2.Lrl感病基因型和抗病基因型的分离主要体现在一个570 bp的变异区内(在第2479-3048碱基之间,对应于第827-1017氨基酸),在该变异区内除两处点突变外具有相同抗性的山羊草品系和六倍体小麦品种的核酸序列完全相同,说明在山羊草进化为六倍体小麦的过程中该区域几乎没有发生过遗传重组和基因转换。
3.在此变异区两侧的相对保守区共有21个点突变,其中有9个同义碱基突变,12个非同义碱基突变,这些点突变并非发生在抗感基因型之间,而是发生在具有相同抗性基因型的山羊草与山羊草或山羊草与六倍体小麦之间。
4.发生在感病粗山羊草与六倍体小麦之间的点突变只有一个(在2460bp处,为同义碱基突变);发生在抗病粗山羊草与六倍体小麦之间的点突变为9个(2个同义碱基突变,7个非同义碱基突变),其中1个点突变发生在570bp的变异区内,8个点突变发生在该变异区两侧的相对保守区。
5.Lrl等位基因序列中的LRR编码区内串联排列着14个同义碱基突变和51个非同义碱基突变,Ka(非同义碱基突变率)╱Ks(同义碱基突变率)＞1,说明该区域经受着岐化选择的作用,这也与前人报道的该区域为配基结合部位、控制着抗病基因的特异性相吻合。
Wheat,as one of the important crops,suffers from the infection of various diseases, wheat leaf rust is one of the major diseases in wheat production.It spreads wildly and can lead to severe yield loss and poor seed quality.The use of resistance genes is the most effective,economical and ecological method to control epidemics of leaf rust disease. However,natural environment and artificial domestication accelerated the co-evolution of host-pathogen interaction,which moreover resulted in the rapid evolution and disappearance of resistance genes.Therefore,it is necessary to study the evolutionary mechanism of plant resistance genes.
In our research,we screened 128 Aegilops.Tauschii(goat grass) accessions and 283 hexaploid wheat varieties(including 262 wheat core germplasms) with marker WR003 co-segregated with Lr1,and obtained the full-length sequences of 42 Lr1 alleles and 6 Lr1 homologous genes.Afterwards,we aligned DNA and protein sequences of Lr1 allelic or homologous genes by DNAMAN and Clustal W,and analyzed the molecular evolution mechanism for the wheat leaf rust resistance genes according to the phylogenetic tree established by neighbor-joining(NJ) method by MEGA 4.0.1.The results showed that:
1.Wheat leaf rust resistance gene Lr1 was originated from Ae.Tauschii,and differentiated into resistant and susceptible genotypes prior to its evolution into hexaploid wheat. Therefore,wheat leaf rust resistance gene Lr1 has two ancestral genes at least.
2.The allelic sequences alignment of Lr1 from resistant and susceptible lines revealed a divergent DNA sequence block of 570 bp,where Lr1 nucleotide sequences were mostly similar between hexaploid wheat and Ae.Tauschii with the same resistance except for two point mutations.So it is concluded that no genetic recombination and gene conversion happened during the evolution from Ae.Tauschii.into hexaploid wheat.
3.The rest of the sequences were mostly similar,where only 21 point mutations existed(9 synonymous mutations,and 12 nonsynonymous mutations).Interestingly,these 21 point mutations occur between lines with identical resistance rather than different resistance.
4.There was only one point mutation between resistant hexaploid wheat and Ae.Tauschii (at 2400bp down stream of the start codon ATG,a synonymous mutation);9 point mutations took place between susceptible hexaploid wheat and Ae.Tauschii(2 synonymous mutations and 7 nonsynonymous mutations),1 in the 570bp block and 8 in the rest of the sequence.
5.Fourteen synonymous mutations and 51 nonsynonymous mutations lined in tandem on the LRR coding region of the allele sequences,Ka(rate of nonsynonymous mutation)/Ks (rate of synonymous mutation)＞1.These results indicated that LRR domain suffered from the pressure of strong selection,consistent with the former reports that LRR domains were involved in the interaction with effector proteins and were the major determinant of resistance specificity.
引文
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