苜蓿中华根瘤菌CCNWSX0020抗铜基因的克隆与功能验证
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摘要
根瘤菌是土壤中广泛分布的一类能与豆科植物共生结瘤固氮的重要农业微生物资源,根瘤菌—豆科植物的共生固氮体系是生物固氮体系的重要组成。苜蓿中华根瘤菌(S. meliloti)CCNWSX0020分离自陕西凤县重金属尾矿的天蓝苜蓿(Medicago lupulin),能抗高达1.8 mM CuSO_4。本文利用Tn5-1063a转座子对S. meliloti CCNWSX0020进行随机突变,建立了一个库容达14000余个突变株库,从中筛选得到在0.8 mM CuSO_4的YMA培养基上不能生长的6株铜敏感性突变株SXa-1、SXa-2、SXc-1、SXc-2、SXn和SXy。重金属抗性表明SXc-1、SXc-2、SXn和SXy对ZnSO_4表现出一定程度的敏感性,而SXa-1和SXa-2对ZnSO_4不多敏感,暗示了SXc-1、SXc-2、SXn和SXy中的突变基因可能与阳离子转运有关或者Zn~(2+)参与这些突变株的某些代谢途径;对Pb(NO_3)_2抗性测定表明只有SXc-1、SXc-2和SXy对Pb(NO_3)_2较为敏感,其余突变株则对Pb(NO_3)_2敏感性较弱,进一步表明这三株突变株突变基因可能与二价阳离子运输有关。
利用Tn5-1063a可以自连成环并具有自我复制的功能,对突变株的全基因组DNA进行酶切并自连,再根据Tn5-1063a两端的已知序列设计引物,对其PCR扩增,PCR产物电泳谱图显示6株突变株都只有单拷贝插入,表明突变株的铜敏感表型是由于转座子插入造成的。进而对其两翼序列进行测序,结果表明,6株突变株是由4个基因突变引起的,其中SXa-1和SXa-2是由lpxXL基因突变所致,该基因编码产物为LpxXL C28-酰基转移酶;SXc-1和SXc-2是由cueR基因突变所致,该基因是MerR家族中一种,是铜抗性基因copA和cueO的调控基;SXn是转座子插入到S. meliloti 1021的SMc02281内部,该基因编码未知膜蛋白,与Agrobacterium sp. H13-3的铜抗性蛋白CopB同源性为74.63%,以此本文把该基因定位copB;SXy由fixI1基因突变造成的,该基因是E1-E2 ATPase家族中一个基因,是一种阳离子泵,且参与细胞有氧呼吸的电子传递过程。
根据突变株的测序结果,PCR扩增得到含启动子的全长片段,将其克隆到穿梭载体pBBR1MCS-5上,三亲杂交将各个基因转入到相应的突变株中,结果发现SXy-1可以在1.2 mM CuSO_4固体平板上恢复生长,其它突变株都可以在1.5 mM CuSO_4平板上恢复生长,进一步证明这些基因与S. meliloti CCNWSX0020的抗铜性有关。
结瘤实验表明,lpxXL和fixI1基因的显著降低了与宿主植物的结瘤数目;盆栽实验表明,当培养基质中CuSO_4含量高于47.36 mg/kg时,与接种野生型菌株相比,所有突变株接种的宿主植物的根茎干重都显著降低,且根茎中的铜含量也显著降低,说明突变株已经影响到宿主植物的生长和对的铜吸收,也可以间接说明突变基因与铜离子的运输有关。
It is well known that symbiotic nitrogen-fixing bacteria are commonly associated with the roots of leguminous plants. Rhizobia colonize the roots of legumes where they fix atmospheric N2, some of which can be utilized for plant growth. A root nodule bacterium, S.meliloti CCNWSX0020, resistant to 1.8 mM CuSO_4 was isolated from Medicago lupulina growing in mine tailings of Fengxian county, China. Six mutants sensitve to copper-resistance were obtained after screening about 14000 mutants by mutagenesis with Tn5-1064a, they were SXa-1, SXa-2, SXc-1, SXc-2, SXn and SXy. Zinc-resistant experiment showed that SXc-1, SXc-2, SXn and SXy were sensitive to zinc, while SXa-1 and SXa-2 were not, which suggested that mutant genes of SXc-1、SXc-2、SXn and SXy were probably involved in cation transport or Zn~(2+) involved in some metabolic pathways in these mutants. Meanwhile, SXc-1、SXc-2 and SXy showed sensitivity to Pb(NO_3)_2 while other mutants were slightly sensitive, which further indicated that the mutant genes of SXc-1、SXc-2 and SXy were probably related to divalent cations transport.
Copper-sensitive phenotype of mutants were identified by the construction and screening of a transposon Tn5-1063a library and PCR. The results showed that single copy inserted into mutants, indicating that coppe-sensitive phenotype was the result of transposon insertion. Six copper-sensitive mutants were obtained and five different genes were indentified by sequencing. SXa-1 and SXa-2 were found with disruptions in the lpxXL gene, encoding LpxXL C28-acyltransferase; SXc-1 and SXc-2 were disrupted with cueR gene, which was found as regulator of copper- tolerance genes copA and cueO; SMc02281 of S. meliloti 1021 was inserted with SXn which encoded unknown membrane protein. It showed 74.63% similarity with the copper-tolerance protein in Agrobacterium sp. H13-3. Therefore, in this paper, SXn gene was lacated in copB and SXy was disrupted with fixI1 gene, belonging to E1-E2 ATPase family, involving in cellular aerobic respiration the electron transfer as a ion pump.
The mutant genes were cloned into the vector, and through the tri-parental conjugation, each mutant gene was transferred into the corresponding mutants. The results showed that SXy-1 completely restored the growth at the concentration of 1.2 mM CuSO_4 and others could grow even at the concentration of 1.5 mM CuSO_4, furtherly suggesting that these mutant genes were related to copper-tolerance of S.meliloti CCNWSX0020.
Nodule tests showed that lpxX and fixI1 mutant genes could remarably reduce the number of nodules of host plants. Comparing with the host plants inoculated wild S.meliloti CCNWSX0020, dry weight and copper accumulation in root as well as shoot of plants inoculated mutants obviously reduced. The findings suggested that mutants in this study affected the growth of host plants as well as absorption to copper, which indirectly indicating mutant genes were related to copper ion transport.
利用Tn5-1063a可以自连成环并具有自我复制的功能,对突变株的全基因组DNA进行酶切并自连,再根据Tn5-1063a两端的已知序列设计引物,对其PCR扩增,PCR产物电泳谱图显示6株突变株都只有单拷贝插入,表明突变株的铜敏感表型是由于转座子插入造成的。进而对其两翼序列进行测序,结果表明,6株突变株是由4个基因突变引起的,其中SXa-1和SXa-2是由lpxXL基因突变所致,该基因编码产物为LpxXL C28-酰基转移酶;SXc-1和SXc-2是由cueR基因突变所致,该基因是MerR家族中一种,是铜抗性基因copA和cueO的调控基;SXn是转座子插入到S. meliloti 1021的SMc02281内部,该基因编码未知膜蛋白,与Agrobacterium sp. H13-3的铜抗性蛋白CopB同源性为74.63%,以此本文把该基因定位copB;SXy由fixI1基因突变造成的,该基因是E1-E2 ATPase家族中一个基因,是一种阳离子泵,且参与细胞有氧呼吸的电子传递过程。
根据突变株的测序结果,PCR扩增得到含启动子的全长片段,将其克隆到穿梭载体pBBR1MCS-5上,三亲杂交将各个基因转入到相应的突变株中,结果发现SXy-1可以在1.2 mM CuSO_4固体平板上恢复生长,其它突变株都可以在1.5 mM CuSO_4平板上恢复生长,进一步证明这些基因与S. meliloti CCNWSX0020的抗铜性有关。
结瘤实验表明,lpxXL和fixI1基因的显著降低了与宿主植物的结瘤数目;盆栽实验表明,当培养基质中CuSO_4含量高于47.36 mg/kg时,与接种野生型菌株相比,所有突变株接种的宿主植物的根茎干重都显著降低,且根茎中的铜含量也显著降低,说明突变株已经影响到宿主植物的生长和对的铜吸收,也可以间接说明突变基因与铜离子的运输有关。
It is well known that symbiotic nitrogen-fixing bacteria are commonly associated with the roots of leguminous plants. Rhizobia colonize the roots of legumes where they fix atmospheric N2, some of which can be utilized for plant growth. A root nodule bacterium, S.meliloti CCNWSX0020, resistant to 1.8 mM CuSO_4 was isolated from Medicago lupulina growing in mine tailings of Fengxian county, China. Six mutants sensitve to copper-resistance were obtained after screening about 14000 mutants by mutagenesis with Tn5-1064a, they were SXa-1, SXa-2, SXc-1, SXc-2, SXn and SXy. Zinc-resistant experiment showed that SXc-1, SXc-2, SXn and SXy were sensitive to zinc, while SXa-1 and SXa-2 were not, which suggested that mutant genes of SXc-1、SXc-2、SXn and SXy were probably involved in cation transport or Zn~(2+) involved in some metabolic pathways in these mutants. Meanwhile, SXc-1、SXc-2 and SXy showed sensitivity to Pb(NO_3)_2 while other mutants were slightly sensitive, which further indicated that the mutant genes of SXc-1、SXc-2 and SXy were probably related to divalent cations transport.
Copper-sensitive phenotype of mutants were identified by the construction and screening of a transposon Tn5-1063a library and PCR. The results showed that single copy inserted into mutants, indicating that coppe-sensitive phenotype was the result of transposon insertion. Six copper-sensitive mutants were obtained and five different genes were indentified by sequencing. SXa-1 and SXa-2 were found with disruptions in the lpxXL gene, encoding LpxXL C28-acyltransferase; SXc-1 and SXc-2 were disrupted with cueR gene, which was found as regulator of copper- tolerance genes copA and cueO; SMc02281 of S. meliloti 1021 was inserted with SXn which encoded unknown membrane protein. It showed 74.63% similarity with the copper-tolerance protein in Agrobacterium sp. H13-3. Therefore, in this paper, SXn gene was lacated in copB and SXy was disrupted with fixI1 gene, belonging to E1-E2 ATPase family, involving in cellular aerobic respiration the electron transfer as a ion pump.
The mutant genes were cloned into the vector, and through the tri-parental conjugation, each mutant gene was transferred into the corresponding mutants. The results showed that SXy-1 completely restored the growth at the concentration of 1.2 mM CuSO_4 and others could grow even at the concentration of 1.5 mM CuSO_4, furtherly suggesting that these mutant genes were related to copper-tolerance of S.meliloti CCNWSX0020.
Nodule tests showed that lpxX and fixI1 mutant genes could remarably reduce the number of nodules of host plants. Comparing with the host plants inoculated wild S.meliloti CCNWSX0020, dry weight and copper accumulation in root as well as shoot of plants inoculated mutants obviously reduced. The findings suggested that mutants in this study affected the growth of host plants as well as absorption to copper, which indirectly indicating mutant genes were related to copper ion transport.
引文
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