大豆鲜籽粒蔗糖含量的研究及糖代谢相关基因的克隆与功能分析
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摘要
菜用大豆是指大豆鼓粒末期(R6-R7)籽粒饱满,英色翠绿时采青食用的大豆专用型品种,是一种重要的豆类蔬菜。外观品质和食味品质是菜用大豆两个基础的品质要素,与菜用大豆这两个性状紧密相关的百荚鲜重、百粒鲜重及鲜籽粒蔗糖含量是菜用大豆品质遗传研究的重要性状。依据菜用大豆这些特殊的目标性状,本研究在对我国323份栽培大豆种质资源进行准确评价、鉴定的基础上,通过关联作图和连锁作图方法发掘与大豆百荚鲜重、百粒鲜重及鲜籽粒蔗糖含量相关的QTL或关联位点。同时,通过同源克隆的方法获得大豆蔗糖代谢关键酶转化酶及蔗糖磷酸合成酶的基因,对这些基因进行大豆发育中籽粒及大豆根、茎、叶、花中的表达特性进行分析,并检测了这些组织中相关酶的活性及糖代谢物的含量。进一步将大豆转化酶基因、蔗糖磷酸合成酶基因转化野生型拟南芥,并将大豆转化酶基因转化拟南芥细胞质转化酶基因功能缺失突变体,观测拟南芥转基因株系相应酶的活性,蔗糖、葡萄糖的含量,以及生长发育中幼苗根长、莲座叶直径等性状。以了解这些酶对组织中糖分累积及植物生长发育的影响。本研究主要结果如下：
1.按照菜用大豆品质要求直接通过表型筛选得到一批优异鲜籽粒性状的种质,并通过关联分析方法鉴定得到与百荚鲜重、百粒鲜重、鲜籽粒蔗糖含量相关的58个(次)标记-性状关联位点,涉及36个SSR标记。在这58个关联位点中,有9个蔗糖-SSR关联点,28个百荚鲜重-SSR关联点,21个百粒鲜重-SSR关联点,这49个荚粒鲜重相关位点共涉及29个SSR标记。进一步对这些显著关联位点进行特异等位变异及载体材料发掘,鉴定出一批优异等位变异位点及其典型载体材料。同时用连锁作图的方法共定位到11个(次)QTL与百荚鲜重(5个)、百粒鲜重(4个)、鲜籽粒蔗糖含量(2个)相关。两种不同作图方法检测QTL的结果中均涉及到的SSR标记有4个,分别为Satt136.Satt208.Satt251及Satt445.
2.以拟南芥细胞质转化酶基因及蔗糖磷酸合成酶基因序列为探针,分别同源克隆得到两个大豆细胞质转化酶基因GmCInv1和GmCInv2,两个大豆蔗糖磷酸合成酶基因GmSPS1和GmSPS2。 GmCInv1和GmCInv2的开放阅读框分别为2040bp和1665bp,分别编码680个和555个氨基酸残基。推测两蛋白分子量分别为76.96kD和63.30kD,等电点分别为5.94和6.22。GmSPS1和GmSPS2的开放阅读均为3177bp,编码1059个氨基酸残基。推测两蛋白分子量分别为117.97和117.99kD,等电点分别为5.99和6.09。同源性分析发现GmCInv1和GmCInv2与其他植物细胞质转化酶氨基酸序列相似性较高,并与酸性转化酶基因完全分开。GmSPS1与GmSPS2在同源性分析中与拟南芥ATSPS1F及ATSPS2F分到一组,但亲缘关系最近的却是葡萄和茄。
3.随着大豆籽粒的发育进程,籽粒中蔗糖含量呈先增长后降低的趋势,葡萄糖含量前期降低较快,后期降低较慢。淀粉含量在不同品种中随发育时期变化的趋势不同。在大豆根、茎、叶、花中,蔗糖含量大都低于葡萄糖含量,这与籽粒中情况恰好相反。随着大豆籽粒发育进程,籽粒中细胞质转化酶、液泡转化酶、细胞壁转化酶活性均是逐步下降,且液泡转化酶活性一直高于细胞质转化酶活性高于细胞壁转化酶活性；蔗糖磷酸合成酶活性在不同品种中变化规律不同。非籽粒组织中,一般叶片和花中转化酶活性较高；蔗糖磷酸合成酶活性有的品种是花中最高,有的是根中最高。随着籽粒的发育进程,两个细胞质转化酶基因GmCInv1与GmCInv2的表达模式不同：GmCInv1呈上升趋势,而GmCInv2有下降趋势,对籽粒中转化酶活性具有明显效应的可能是GmCInv2。两细胞质转化酶基因均是在根中表达量高于其他组织。大豆蔗糖磷酸合成酶基因随籽粒发育进程表达量逐步升高,在根中的表达量高于其他非籽粒组织。糖代谢物含量与酶活性、酶活性与酶基因表达量之间均存在一定的影响关系,但没有完全的控制效果。
4.将GmCInv1及GmCInv2分别转化拟南芥野生型(WT)及拟南芥细胞质转化酶基因功能缺失突变体cinv1,将获得纯合阳性转基因株系分别称作W组转基因株系和M组转基因株系。除GmCInv2的M11株系外,GmCInv1、GmCInv2的其他转基因株系均有目的基因的表达及拟南芥自身转化酶AtCinv1基因的表达,WT没有大豆基因表达,突变体cinv1没有大豆基因的表达且自身AtCinv1基因的表达极低。GmCInv1及GmCInv2的表达,专一性的提高了转基因株系的细胞质转化酶活性,对液泡转化酶、细胞壁转化酶活性没有明显效应。转基因株系的蔗糖含量及其蔗糖含量／葡萄糖含量比被降低,表明转化酶活性的提高促进了蔗糖的分解。与对照相比,转基因株系的生长发育受到影响,基因效应包括：促进幼苗根伸长,促进莲座叶生长,促进莲座叶数目增加,增加抽薹数,增加株高,延长开花期。两转化酶基因在某些功能上具有差异。大豆GmCInv1及GmCInv2的表达对拟南芥突变体cinvl的一些表型具有补偿作用,可以恢复其表型,但不一定能完全恢复至WT水平。
5.转GmSPS1、GmSPS2的转基因株系均有目的基因的表达,但两基因的6个转基因株系中仅有1个株系Line5的SPS活性与WT有显著性差异。Line5的蔗糖含量明显提高,与WT相比,Line5株高度及幼苗根长得到显著增加。其他5个株系虽然SPS酶活性没有显著提高,但大部分株系仍在所考察的6个生长发育表型中有某一个或两个与WT产生显著性差异。这些结果表明拟南芥中SPS活性的提高能够促进蔗糖的累积,并引起植株生长表型的改变；大豆SPS基因在拟南芥中表达后可能受到其他因素的抑制,以至不能提高拟南芥SPS的活性。
Vegetable soybean is a specialized soybean (Glycine max (L.) Merrill), harvested in the R6-R7growth stage with full seeds and still green pods. Traits of100-pod weight,100-seed weight and sucrose content are the most important factors of appearance and eating quality for vegetable soybean. Based on the goals of vegetable soybean breeding and genetic improvement, we evaluated323cultivated soybean germplasms in China, and identified quantitative trait loci (QTL) related to100-pod weight,100-seed weight and sucrose content in fresh soybean seeds. The key enzyme genes involved sugar accumulation of vegetable soybean were cloned, then the expression pattern of these genes in soybean tissues were analyzed, together with the determination of sugars contents and enzymes activity. These genes were transformed into wild type and cinvl mutant of Arabidopsis thaliana for functional analysis. Main results are as follows:
1. A number of elite germplasms were identified base on the vegetable breeding objectives. A total of58marker-trait associations related to100-pod weight (28),100-seed weight (21) and sucrose content in fresh seeds (9) were identified by association mapping, involving36different SSR markers, the loci alleles which were significantly associated with the traits were analyzed further. Five, four, and two QTLs were detected relating to100-pod weight,100-seed weight and sucrose content in fresh seeds by linkage mapping, respectively. Four SSRs (Sattl36, Satt208, Satt251and Satt445) were found to be related to the pod and/or seed weights traits using the both mapping methods.
2. Two soybean cytoplasmic invertase (CInv) genes were homologous cloned and designated GmCInvl and GmCInv2, respectively. GmCInvl contains a2040bp open reading frame (ORF), encoding a680amino acid polypeptide (76.96kD). GmCInv2contains a1665bp ORF, encoding a555amino acid polypeptide (63.30kD). The phylogenetic analysis indicated that GmCInvl and GmCInv2were separated with the acid invertase genes, and all the cytoplasmic invertase genes were clustered together. Two sucrose phosphate synthase genes in soybean were identified, and designated GmSPSl and GmSPS2, respectively. GmSPS1and GmSPS2both contain a3177bp ORF, encoding a680amino acid polypeptide. The phylogenetic analysis indicated that GmSPSl and GmSPS2protein had more closed relationship to Vitis than others.
3. The sucrose content in the fresh soybean seeds keeps increasing in the early development stage, and decreased in the later stage, while the glucose content gradually decreased during the whole development stage. The starch content change trends of the developing seeds was different in the three evaluated soybean cultivars. The sucrose content was higher than glucose content in the root, stem, leaf and flower. The activity of cytoplasmic invertase (CInv), vacuolar invertase (VInv) and-cell wall invertase (CWInv) all had a decreased trend during the seed growth. The Inv activity in leaves and flowers was higher than that in stem and root. The activity of sucrose phosphate synthase (SPS) have not a stable change mode in developing seed, and the flower or root usually shows a higher SPS activity. GmCInv1and GmCInv2gene were detected in all different tissues, and expressed strongly in roots. The GmCInv1transcription levels were gradually up-regulated with seed development, which were different from GmCInv2, and GmCInv2is more effective for CInv activity in soybean seeds. GmSPS1and GmSPS2have a similar expression pattern to GmCInv1. The expression level of these genes affected the activity of the corresponding enzymes, and changed the sugar contents, but none of the genes or enzymes could regulate the sucrose or glucose level individually.
4. GmCInv1and GmCInv2were transformed into wild type (WT) and cinvl mutant of Arabidopsis, cinvl, CINV1-deficient plants, had reduced activities of neutral invertase (cytoplasmic invertase). Transcripts of the target genes can be detected in transgenic lines except Line11. The GmCInv1and GmCInv2expression in Arabidopsis have higher CInv activity. The sucrose contents and value of sucrose content/glucose content of GmCInvl and GmCInv2transgenic lines were lower than that of controls. The growth and development of these transgenic lines was effected by GmCInv1and GmCInv2, the root length, plant height, rosettes diameter, rosette leaf number, stem number and flowering time were increased in different transgenic lines. The expression of soybean CInv genes complemented the deficient phenotype of the cinvl mutant in some degree.
5. Five of the six GmSPSl and GmSPS2transgenic lines have not significant differences in SPS activity compared with WT, though the transcript of GmSPSl and GmSPS2can be detected in corresponding lines. The significant line Line5has a higher sucrose level than WT, the length of seedlings roots and plant height was increased too. There were no significant SPS activity in other five lines may due to the effect of some certain repressors in the Arabidopsis plant.
1.按照菜用大豆品质要求直接通过表型筛选得到一批优异鲜籽粒性状的种质,并通过关联分析方法鉴定得到与百荚鲜重、百粒鲜重、鲜籽粒蔗糖含量相关的58个(次)标记-性状关联位点,涉及36个SSR标记。在这58个关联位点中,有9个蔗糖-SSR关联点,28个百荚鲜重-SSR关联点,21个百粒鲜重-SSR关联点,这49个荚粒鲜重相关位点共涉及29个SSR标记。进一步对这些显著关联位点进行特异等位变异及载体材料发掘,鉴定出一批优异等位变异位点及其典型载体材料。同时用连锁作图的方法共定位到11个(次)QTL与百荚鲜重(5个)、百粒鲜重(4个)、鲜籽粒蔗糖含量(2个)相关。两种不同作图方法检测QTL的结果中均涉及到的SSR标记有4个,分别为Satt136.Satt208.Satt251及Satt445.
2.以拟南芥细胞质转化酶基因及蔗糖磷酸合成酶基因序列为探针,分别同源克隆得到两个大豆细胞质转化酶基因GmCInv1和GmCInv2,两个大豆蔗糖磷酸合成酶基因GmSPS1和GmSPS2。 GmCInv1和GmCInv2的开放阅读框分别为2040bp和1665bp,分别编码680个和555个氨基酸残基。推测两蛋白分子量分别为76.96kD和63.30kD,等电点分别为5.94和6.22。GmSPS1和GmSPS2的开放阅读均为3177bp,编码1059个氨基酸残基。推测两蛋白分子量分别为117.97和117.99kD,等电点分别为5.99和6.09。同源性分析发现GmCInv1和GmCInv2与其他植物细胞质转化酶氨基酸序列相似性较高,并与酸性转化酶基因完全分开。GmSPS1与GmSPS2在同源性分析中与拟南芥ATSPS1F及ATSPS2F分到一组,但亲缘关系最近的却是葡萄和茄。
3.随着大豆籽粒的发育进程,籽粒中蔗糖含量呈先增长后降低的趋势,葡萄糖含量前期降低较快,后期降低较慢。淀粉含量在不同品种中随发育时期变化的趋势不同。在大豆根、茎、叶、花中,蔗糖含量大都低于葡萄糖含量,这与籽粒中情况恰好相反。随着大豆籽粒发育进程,籽粒中细胞质转化酶、液泡转化酶、细胞壁转化酶活性均是逐步下降,且液泡转化酶活性一直高于细胞质转化酶活性高于细胞壁转化酶活性；蔗糖磷酸合成酶活性在不同品种中变化规律不同。非籽粒组织中,一般叶片和花中转化酶活性较高；蔗糖磷酸合成酶活性有的品种是花中最高,有的是根中最高。随着籽粒的发育进程,两个细胞质转化酶基因GmCInv1与GmCInv2的表达模式不同：GmCInv1呈上升趋势,而GmCInv2有下降趋势,对籽粒中转化酶活性具有明显效应的可能是GmCInv2。两细胞质转化酶基因均是在根中表达量高于其他组织。大豆蔗糖磷酸合成酶基因随籽粒发育进程表达量逐步升高,在根中的表达量高于其他非籽粒组织。糖代谢物含量与酶活性、酶活性与酶基因表达量之间均存在一定的影响关系,但没有完全的控制效果。
4.将GmCInv1及GmCInv2分别转化拟南芥野生型(WT)及拟南芥细胞质转化酶基因功能缺失突变体cinv1,将获得纯合阳性转基因株系分别称作W组转基因株系和M组转基因株系。除GmCInv2的M11株系外,GmCInv1、GmCInv2的其他转基因株系均有目的基因的表达及拟南芥自身转化酶AtCinv1基因的表达,WT没有大豆基因表达,突变体cinv1没有大豆基因的表达且自身AtCinv1基因的表达极低。GmCInv1及GmCInv2的表达,专一性的提高了转基因株系的细胞质转化酶活性,对液泡转化酶、细胞壁转化酶活性没有明显效应。转基因株系的蔗糖含量及其蔗糖含量／葡萄糖含量比被降低,表明转化酶活性的提高促进了蔗糖的分解。与对照相比,转基因株系的生长发育受到影响,基因效应包括：促进幼苗根伸长,促进莲座叶生长,促进莲座叶数目增加,增加抽薹数,增加株高,延长开花期。两转化酶基因在某些功能上具有差异。大豆GmCInv1及GmCInv2的表达对拟南芥突变体cinvl的一些表型具有补偿作用,可以恢复其表型,但不一定能完全恢复至WT水平。
5.转GmSPS1、GmSPS2的转基因株系均有目的基因的表达,但两基因的6个转基因株系中仅有1个株系Line5的SPS活性与WT有显著性差异。Line5的蔗糖含量明显提高,与WT相比,Line5株高度及幼苗根长得到显著增加。其他5个株系虽然SPS酶活性没有显著提高,但大部分株系仍在所考察的6个生长发育表型中有某一个或两个与WT产生显著性差异。这些结果表明拟南芥中SPS活性的提高能够促进蔗糖的累积,并引起植株生长表型的改变；大豆SPS基因在拟南芥中表达后可能受到其他因素的抑制,以至不能提高拟南芥SPS的活性。
Vegetable soybean is a specialized soybean (Glycine max (L.) Merrill), harvested in the R6-R7growth stage with full seeds and still green pods. Traits of100-pod weight,100-seed weight and sucrose content are the most important factors of appearance and eating quality for vegetable soybean. Based on the goals of vegetable soybean breeding and genetic improvement, we evaluated323cultivated soybean germplasms in China, and identified quantitative trait loci (QTL) related to100-pod weight,100-seed weight and sucrose content in fresh soybean seeds. The key enzyme genes involved sugar accumulation of vegetable soybean were cloned, then the expression pattern of these genes in soybean tissues were analyzed, together with the determination of sugars contents and enzymes activity. These genes were transformed into wild type and cinvl mutant of Arabidopsis thaliana for functional analysis. Main results are as follows:
1. A number of elite germplasms were identified base on the vegetable breeding objectives. A total of58marker-trait associations related to100-pod weight (28),100-seed weight (21) and sucrose content in fresh seeds (9) were identified by association mapping, involving36different SSR markers, the loci alleles which were significantly associated with the traits were analyzed further. Five, four, and two QTLs were detected relating to100-pod weight,100-seed weight and sucrose content in fresh seeds by linkage mapping, respectively. Four SSRs (Sattl36, Satt208, Satt251and Satt445) were found to be related to the pod and/or seed weights traits using the both mapping methods.
2. Two soybean cytoplasmic invertase (CInv) genes were homologous cloned and designated GmCInvl and GmCInv2, respectively. GmCInvl contains a2040bp open reading frame (ORF), encoding a680amino acid polypeptide (76.96kD). GmCInv2contains a1665bp ORF, encoding a555amino acid polypeptide (63.30kD). The phylogenetic analysis indicated that GmCInvl and GmCInv2were separated with the acid invertase genes, and all the cytoplasmic invertase genes were clustered together. Two sucrose phosphate synthase genes in soybean were identified, and designated GmSPSl and GmSPS2, respectively. GmSPS1and GmSPS2both contain a3177bp ORF, encoding a680amino acid polypeptide. The phylogenetic analysis indicated that GmSPSl and GmSPS2protein had more closed relationship to Vitis than others.
3. The sucrose content in the fresh soybean seeds keeps increasing in the early development stage, and decreased in the later stage, while the glucose content gradually decreased during the whole development stage. The starch content change trends of the developing seeds was different in the three evaluated soybean cultivars. The sucrose content was higher than glucose content in the root, stem, leaf and flower. The activity of cytoplasmic invertase (CInv), vacuolar invertase (VInv) and-cell wall invertase (CWInv) all had a decreased trend during the seed growth. The Inv activity in leaves and flowers was higher than that in stem and root. The activity of sucrose phosphate synthase (SPS) have not a stable change mode in developing seed, and the flower or root usually shows a higher SPS activity. GmCInv1and GmCInv2gene were detected in all different tissues, and expressed strongly in roots. The GmCInv1transcription levels were gradually up-regulated with seed development, which were different from GmCInv2, and GmCInv2is more effective for CInv activity in soybean seeds. GmSPS1and GmSPS2have a similar expression pattern to GmCInv1. The expression level of these genes affected the activity of the corresponding enzymes, and changed the sugar contents, but none of the genes or enzymes could regulate the sucrose or glucose level individually.
4. GmCInv1and GmCInv2were transformed into wild type (WT) and cinvl mutant of Arabidopsis, cinvl, CINV1-deficient plants, had reduced activities of neutral invertase (cytoplasmic invertase). Transcripts of the target genes can be detected in transgenic lines except Line11. The GmCInv1and GmCInv2expression in Arabidopsis have higher CInv activity. The sucrose contents and value of sucrose content/glucose content of GmCInvl and GmCInv2transgenic lines were lower than that of controls. The growth and development of these transgenic lines was effected by GmCInv1and GmCInv2, the root length, plant height, rosettes diameter, rosette leaf number, stem number and flowering time were increased in different transgenic lines. The expression of soybean CInv genes complemented the deficient phenotype of the cinvl mutant in some degree.
5. Five of the six GmSPSl and GmSPS2transgenic lines have not significant differences in SPS activity compared with WT, though the transcript of GmSPSl and GmSPS2can be detected in corresponding lines. The significant line Line5has a higher sucrose level than WT, the length of seedlings roots and plant height was increased too. There were no significant SPS activity in other five lines may due to the effect of some certain repressors in the Arabidopsis plant.
引文
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