山羊草属sitopsis组物种LMW-GS及Gliadin编码基因鉴定及系统进化分析
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摘要
小麦低分子量谷蛋白和醇溶蛋白是重要的种子贮藏蛋白，与小麦的加工品质和面团特性紧密相关。前人从分子水平对小麦及其A、D染色体组二倍体原始供体物种中的这两类蛋白的编码基因进行了广泛地研究，本文以B染色体组的可能供体种，山羊草属Sitopsis组中的5个物种Aegilops longissima(2n=2x=14，S~lS~l)，Aegilopssharonensis(2n=2x=14，S~(sh)S~(sh))，Aegilops searsii(2n=2x=-14，S~sS~s)，Aegilops bicornis(2n=2x=14，S~bS~b)以及Aegilops．speltoides(2n=2x=-14，SS)为研究对象，分离其中的LMW-GS(low-molecular-weight glutenin subunits，LMW-GS)和gliadin编码基因，并进行系统进化分析，主要取得了以下结果：
1．山羊草属Sitopsis组LMW-GS基因鉴定及系统进化分析
从山羊草属Sitopsis组的5个物种中共得到69个LMW-GS基因。根据N-末端推导氨基酸序列，可将其主要划分成5种类群，其中MENSHIPGLERP-与小麦中B基因组特异的LMW-GS基因类群属于一类。而METSCIPSLER-、METSHILSLEK-和METSHIPSLEKSL-是A、B、D、G染色体组中没有的新型LMW-GS基因类型，该结果进一步丰富了对LMW-GS多基因家族的认识。系统进化表明，N-末端为METSCIPSLER-的基因与来自A和D染色体组的基因聚为一类，而N-末端为METSHILSLEK-和METSHIPSLEKSL-则与来自小麦B染色体组的LMW=GS基因类群紧密相关。这些结果支持Sitopsis组做为小麦B染色体组供体的观点。然而，这3类基因涵盖了Sitopsis组中除Ae.speltoides的其余4个物种，无法提供究竟哪一个物种更可能是B染色体组的供体种的有力证据。本研究结果显示，Ae．speltoides与其余4个物种的遗传关系可能较远。
2．山羊草属Sitopsis组α-醇溶蛋白基因鉴定及系统进化分析
从山羊草属Sitopsis组3个物种中分离了40个α-醇溶蛋白基因，并与来自Ae．speltoide和Ae．longissima的基因进行比较，发现山羊草属Sitopsis组的α-醇溶蛋白基因存在丰富的多样性，可以划分成12类，其中，Ae．speltoides中的类型最多，有6种，且4种为Ae．speltoides特有。这表明Ae．speltoides与其他4个物种的遗传差异较大。某些(或可能)位于B染色体的α-醇溶蛋白与类型group S3、group S8和group S6聚为1类，表明这些B染色体特异的基因可能来自Sitopsis组物种。此外，某些基因类型与来自A染色体组和D染色体组的基因聚为1类，类似现象也同样在LMW-GS中发现，但出现频率更高，这表明α-醇溶蛋白基因家族更加复杂。
3．山羊草属Sitopsis组γ-醇溶蛋白基因鉴定及系统进化分析
从山羊草属Sitopsis组的5个物种中分离得到55个γ-醇溶蛋白基因。序列比对显示，这些基因与已知γ-醇溶蛋白基因具有很高的序列相似性，但又确实存在差异，具有典型的γ-醇溶蛋白一级结构。根据半胱氨酸残基数目将这些基因分为两类：1类具有8个保守的半胱氨酸残基，另一类在重复区存在1个自由的半胱氨酸残基，可能参与到谷蛋白的多聚物的聚合当中。将本研究所得γ-醇溶蛋白基因与已知来自Sitopsis和A、B、D染色体组的基因进行分类比较和系统进化分析。结果显示，第1类即重复区无半胱氨酸的基因可进一步聚为3类：1类为D基因组特异类型；而定位于1B的基因与部分来自5个Sitopsis物种的基因聚为1类，表明这类基因可能为B基因组特异，且Sitopsis物种可能是其初始来源；而第3类来自AABB和AABBDD基因组物种的基因与3个来自Ae．longissima和Ae.sharonesis的基因与聚为1类，不能断定该类型所属基因组。
来自Ae．bicornis、Ae．longissima和Ae．sharonesis的第2类重复区内有半胱氨酸的基因聚为1类，而来自Ae．searsii和Ae．speltoides的基因分别单独聚为1类。同时，分别与部分来自4倍体(AABB)和(或)6倍体(AABBDD)小麦的基因聚为1类，无1A或1D染色体的基因聚于此类。这些结果可能暗示其为B染色体组特异类型，Ae．searsii或Ae．Speltoides可能是其类基因的原始供体。
The low-molecular-weight glutenin subunits and gliadins are major seed storage proteins of wheat, and closely related to the wheat quality and processing properties. Their coding genes at molecular level had been extensively conducted in wheat and its wild diploid progenitors of A and D genomes. In the present study, the characterization and phylogenetic analysis of LMW-GS (low-molecular-weight glutenin subunits, LMW-GS) and gliadin genes were conducted in the Aegilops sect. sitopsis species, Aegilops longissima (2n=2x=14, S~lS~l), Aegilops. sharonensis (2n=2x=14, S~(sh)S~(sh)), Aegilops searsii (2n=2x=14, S~sS~s), Aegilops bicornis (2n=2x=14, S~bS~b) and Aegilops speltoides (2n=2x=14, SS). The main results obtained were the following:
1. Characterization and phylogenetic analysis of the LMW-GS genes in Aegilops sect. sitopsis species
A total of 69 LMW-GS genes were obtained and could be divided into 5 groups based on the deduced amino acids of the N-terminal domain, in which the MENSHIPGLERP- could belong to a B genome specific gene group in wheat, while the others METSCIPSLER, METSHILSLEK-, and METSHIPSLEKSL were proved to be novel gene groups which had not been observed in the A, B, D, and G genomes, by PCR assaying. Phylogenetic analysis indicated the genes with N-terminal domain of METSCIPSLER were grouped with those from A and D genomes, and METSHIPSLEKSL- and METSHIPSLEKSL were highly homologous to the genes in the B genome. These results supported the Sitopsis, but none of the five species, should be assigned as the donors of B genome of wheat. It was also be speculated that the Ae. Speltoides was more divergent with the other 4 species.
2. Characterization and Phylogenetic analysis of a-gliadin genes in Aegilops sect, sitopsis species
Forty a-gliadin genes were obtained from 3 species in the sect, sitopsis. Sequence comparison with known genes from Ae. speltoide and Ae. longissima indicated that the a-gliadin genes in Aegilops sect, sitopsis could be divided into 12 groups. The genes from Ae. speltoide were involved in 6 groups, among which 4 groups were Ae. Speltoide specific, indicating that Ae. speltoide might be divergent from other 4 species. Some a-gliadin genes in B genome from wheat were clustered with groups of group S3, groupS8, and group S6, indicating that these genes might originate from the species in the sitopsis section. Further more, group S7-1 and group S7-2 were clustered with that from A and D genomes, which was similar in LMW-GS genes.
3. Characterization and Phylogenetic analysis ofγ-gliadin genes in Aegilops sect, sitopsis species
Fifty fiveγ-gliadin genes were obtained from 5 species of Aegilops sect. sitopsis. Sequence analysis showed that these genes were high similar, but not identical to the known genes. Two groups of y-gliadin genes were apparently classified based on the number of the cysteines: group 1 with 8 conserved cysteines and group 2 with an additional free cysteine in the repetitive domain which might be involved in the formation of the glutenin polymer. The phylogenetic analysis showed that the genes with 8 cysteines could be clustered into 3 sub-groups: a distinct sub-group specific to the D genome, a sub-group of B genome specific genes and that from 5 Aegilops sect, sitopsis species, indicating that sitopsis should be considered as the origin of this B-genome specific sub-group, and the 3nd sub-group of genes from species with ABD and AB genomes and sitopsis species Ae. longissima and Aesharonesis, however, it could not be concluded that which genome could be assigned to the this sub-group.
The genes of group 2 from Ae.bicornis, Ae. longissima and Ae. Sharonesis were distinctly clustered from the species, indicating it was a specific sub-group. The genes from Ae.searsii and Ae. Speltoides were clustered with some genes from species with AABBDD and AABB genomes, respectively, and no A and D genome specific genes were present in the two sub-groups, indicating that these two subgroups in wheat might be B genome specific and originate from the Ae.searsii snd Ae. speltoides, respectively.
1．山羊草属Sitopsis组LMW-GS基因鉴定及系统进化分析
从山羊草属Sitopsis组的5个物种中共得到69个LMW-GS基因。根据N-末端推导氨基酸序列，可将其主要划分成5种类群，其中MENSHIPGLERP-与小麦中B基因组特异的LMW-GS基因类群属于一类。而METSCIPSLER-、METSHILSLEK-和METSHIPSLEKSL-是A、B、D、G染色体组中没有的新型LMW-GS基因类型，该结果进一步丰富了对LMW-GS多基因家族的认识。系统进化表明，N-末端为METSCIPSLER-的基因与来自A和D染色体组的基因聚为一类，而N-末端为METSHILSLEK-和METSHIPSLEKSL-则与来自小麦B染色体组的LMW=GS基因类群紧密相关。这些结果支持Sitopsis组做为小麦B染色体组供体的观点。然而，这3类基因涵盖了Sitopsis组中除Ae.speltoides的其余4个物种，无法提供究竟哪一个物种更可能是B染色体组的供体种的有力证据。本研究结果显示，Ae．speltoides与其余4个物种的遗传关系可能较远。
2．山羊草属Sitopsis组α-醇溶蛋白基因鉴定及系统进化分析
从山羊草属Sitopsis组3个物种中分离了40个α-醇溶蛋白基因，并与来自Ae．speltoide和Ae．longissima的基因进行比较，发现山羊草属Sitopsis组的α-醇溶蛋白基因存在丰富的多样性，可以划分成12类，其中，Ae．speltoides中的类型最多，有6种，且4种为Ae．speltoides特有。这表明Ae．speltoides与其他4个物种的遗传差异较大。某些(或可能)位于B染色体的α-醇溶蛋白与类型group S3、group S8和group S6聚为1类，表明这些B染色体特异的基因可能来自Sitopsis组物种。此外，某些基因类型与来自A染色体组和D染色体组的基因聚为1类，类似现象也同样在LMW-GS中发现，但出现频率更高，这表明α-醇溶蛋白基因家族更加复杂。
3．山羊草属Sitopsis组γ-醇溶蛋白基因鉴定及系统进化分析
从山羊草属Sitopsis组的5个物种中分离得到55个γ-醇溶蛋白基因。序列比对显示，这些基因与已知γ-醇溶蛋白基因具有很高的序列相似性，但又确实存在差异，具有典型的γ-醇溶蛋白一级结构。根据半胱氨酸残基数目将这些基因分为两类：1类具有8个保守的半胱氨酸残基，另一类在重复区存在1个自由的半胱氨酸残基，可能参与到谷蛋白的多聚物的聚合当中。将本研究所得γ-醇溶蛋白基因与已知来自Sitopsis和A、B、D染色体组的基因进行分类比较和系统进化分析。结果显示，第1类即重复区无半胱氨酸的基因可进一步聚为3类：1类为D基因组特异类型；而定位于1B的基因与部分来自5个Sitopsis物种的基因聚为1类，表明这类基因可能为B基因组特异，且Sitopsis物种可能是其初始来源；而第3类来自AABB和AABBDD基因组物种的基因与3个来自Ae．longissima和Ae.sharonesis的基因与聚为1类，不能断定该类型所属基因组。
来自Ae．bicornis、Ae．longissima和Ae．sharonesis的第2类重复区内有半胱氨酸的基因聚为1类，而来自Ae．searsii和Ae．speltoides的基因分别单独聚为1类。同时，分别与部分来自4倍体(AABB)和(或)6倍体(AABBDD)小麦的基因聚为1类，无1A或1D染色体的基因聚于此类。这些结果可能暗示其为B染色体组特异类型，Ae．searsii或Ae．Speltoides可能是其类基因的原始供体。
The low-molecular-weight glutenin subunits and gliadins are major seed storage proteins of wheat, and closely related to the wheat quality and processing properties. Their coding genes at molecular level had been extensively conducted in wheat and its wild diploid progenitors of A and D genomes. In the present study, the characterization and phylogenetic analysis of LMW-GS (low-molecular-weight glutenin subunits, LMW-GS) and gliadin genes were conducted in the Aegilops sect. sitopsis species, Aegilops longissima (2n=2x=14, S~lS~l), Aegilops. sharonensis (2n=2x=14, S~(sh)S~(sh)), Aegilops searsii (2n=2x=14, S~sS~s), Aegilops bicornis (2n=2x=14, S~bS~b) and Aegilops speltoides (2n=2x=14, SS). The main results obtained were the following:
1. Characterization and phylogenetic analysis of the LMW-GS genes in Aegilops sect. sitopsis species
A total of 69 LMW-GS genes were obtained and could be divided into 5 groups based on the deduced amino acids of the N-terminal domain, in which the MENSHIPGLERP- could belong to a B genome specific gene group in wheat, while the others METSCIPSLER, METSHILSLEK-, and METSHIPSLEKSL were proved to be novel gene groups which had not been observed in the A, B, D, and G genomes, by PCR assaying. Phylogenetic analysis indicated the genes with N-terminal domain of METSCIPSLER were grouped with those from A and D genomes, and METSHIPSLEKSL- and METSHIPSLEKSL were highly homologous to the genes in the B genome. These results supported the Sitopsis, but none of the five species, should be assigned as the donors of B genome of wheat. It was also be speculated that the Ae. Speltoides was more divergent with the other 4 species.
2. Characterization and Phylogenetic analysis of a-gliadin genes in Aegilops sect, sitopsis species
Forty a-gliadin genes were obtained from 3 species in the sect, sitopsis. Sequence comparison with known genes from Ae. speltoide and Ae. longissima indicated that the a-gliadin genes in Aegilops sect, sitopsis could be divided into 12 groups. The genes from Ae. speltoide were involved in 6 groups, among which 4 groups were Ae. Speltoide specific, indicating that Ae. speltoide might be divergent from other 4 species. Some a-gliadin genes in B genome from wheat were clustered with groups of group S3, groupS8, and group S6, indicating that these genes might originate from the species in the sitopsis section. Further more, group S7-1 and group S7-2 were clustered with that from A and D genomes, which was similar in LMW-GS genes.
3. Characterization and Phylogenetic analysis ofγ-gliadin genes in Aegilops sect, sitopsis species
Fifty fiveγ-gliadin genes were obtained from 5 species of Aegilops sect. sitopsis. Sequence analysis showed that these genes were high similar, but not identical to the known genes. Two groups of y-gliadin genes were apparently classified based on the number of the cysteines: group 1 with 8 conserved cysteines and group 2 with an additional free cysteine in the repetitive domain which might be involved in the formation of the glutenin polymer. The phylogenetic analysis showed that the genes with 8 cysteines could be clustered into 3 sub-groups: a distinct sub-group specific to the D genome, a sub-group of B genome specific genes and that from 5 Aegilops sect, sitopsis species, indicating that sitopsis should be considered as the origin of this B-genome specific sub-group, and the 3nd sub-group of genes from species with ABD and AB genomes and sitopsis species Ae. longissima and Aesharonesis, however, it could not be concluded that which genome could be assigned to the this sub-group.
The genes of group 2 from Ae.bicornis, Ae. longissima and Ae. Sharonesis were distinctly clustered from the species, indicating it was a specific sub-group. The genes from Ae.searsii and Ae. Speltoides were clustered with some genes from species with AABBDD and AABB genomes, respectively, and no A and D genome specific genes were present in the two sub-groups, indicating that these two subgroups in wheat might be B genome specific and originate from the Ae.searsii snd Ae. speltoides, respectively.
引文
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