对苜蓿中华根瘤菌c-di-GMP合成及分解酶的功能和LuxR家族转录调控蛋白ExpR的研究
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摘要
空气中含有约80%的氮气,然而它们并不能为高等动物、植物所直接利用。生物固氮指固氮微生物具有在化合态氮素缺乏时将氮气转化为能被植物吸收利用的氨类化合物的能力,在农业生产中发挥着举足轻重的作用。根瘤菌与豆科植物之间形成的共生体系由于其固氮作用的天然、无污染和高效率,始终是生物固氮研究的焦点之一。同时根瘤菌与豆科植物之间的信号识别、传递及基因的顺序性表达和调控对根瘤的形成、发育和固氮作用的效率等有着复杂的联系。因此以根瘤菌为主要研究对象,以根瘤菌和豆科植物的相互关系作为研究微生物——植物相互作用的模式将是我们研究的主要目标,对于降低农业生产成本、保持农业和环境的可持续发展有着重要的科学价值。
鸟苷酸环化酶和磷酸二酯酶是分别含有GGDEF或EAL结构域的蛋白质,可合成或分解细菌二级信号分子——环二鸟苷酸(c-di-GMP)。基因组DNA序列分析表明这些蛋白质分布于几乎所有的细菌当中。但是,对于可以进行共生固氮作用的土壤细菌——根瘤菌而言,其GGDEF/EAL蛋白的研究仍是空白。本研究通过生物信息学方法在苜蓿中华根瘤菌(Sinorhizbium meliloti)中鉴定到19个GGDEF和EAL蛋白基因,其中编码GGDEF蛋白的基因5个,编码EAL蛋白的基因4个,同时编码GGDEF和EAL双结构域蛋白的基因10个。通过质粒插入失活的方法成功构建到14个基因突变菌株。所有14个基因的突变株在基本培养基中均表现出生长缺陷和运动性缺陷的表型,其中11个基因的突变株可以合成较多的胞外多糖,但其在紫花苜蓿上的竞争结瘤能力有所降低。进一步的研究发现,这些基因突变影响了根瘤菌对碳源和氮源的利用。我们的研究表明,GGDEF和EAL蛋白参与调节根瘤菌的多种生理过程,其催化产物——c-di-GMP在根瘤菌与宿主豆科植物相互作用过程中发挥了重要作用。
此外,我们发现具有完整expR基因的苜蓿中华根瘤菌在四倍体紫花苜蓿(Medicago sativa)和二倍体蒺藜苜蓿(M. truncatula)上具有不同的结瘤固氮表型,即该菌在紫花苜蓿结瘤固氮与野生型没有差别,但是在蒺藜苜蓿上却只能诱导低效固氮根瘤。我们对引起这种现象的原因进行了初步的探讨,发现该菌携带一个新的突变位点,其机制有待进一步研究。
About80%of the atmosphere is the nitrogen, but it can not be used by higher animals and plants directly. Biological nitrogen fixation affords the most part of nitrogen input.It is a hotspot in current study. Establishing this symbiosis requires a response of molecular signals, gene expression and nutrient exchange between the legumes and the rhizobia. Therefore, analyse the rhizobia and the relationship between rhizobia and legumes will be the main goal of our study, which has important scientific value in reducing agricultural production costs, maintaining the sustainable development of agriculture and environment.
A new bacterial secondary messenger, bis (3'-5')-cyclic dimeric guanosine monophosphate (c-di-GMP), is usually synthesized or decomposed by proteins containing a GGDEF or EAL domain. These proteins often act as a cyclase or phosphodiesterase of c-di-GMP, and their genes are distributed among almost all bacteria according to known genomic DNA sequences. However, the systematic identification of GGDEF and EAL protein genes remains unclear in rhizobia, soil bacteria that interact with compatible legumes to form nitrogen-fixing nodules. In this study,19putative GGDEF and EAL protein genes were identified in a model rhizobium, Sinorhizobium meliloti, by bioinformatic analysis (encoding5GGDEF proteins,4EAL proteins, and10GGDEF and EAL double-domain proteins). Null mutants of14genes were constructed through systematic plasmid insertion. All14gene mutants showed deficient growth in minimal medium and defective motility, and11gene mutants produced a lot more exopolysaccharide and displayed less competitive nodulation on the host plant, alfalfa. We also find that these mutants had different capacity to utilize sugars and amino acids as carbon and nitrogen sources respectively. Our results suggest that GGDEF and EAL proteins may play different roles in the control of S. meliloti physiology, although they contain conserved catalytic (GGDEF or EAL) domains. Our finding also implies that c-di-GMP may play an important role in the interactions between this rhizobium and its host plants to establish efficient symbiosis.
Additionally, S. meliloti strain carrying an intact expR gene induced the same efficiently nitrogen-fixing nodules on Medicago sativa as the wild type strain; however, it only induced lower efficient nodules on M. truncatula. And a now mutation locus was proposed, and the detail will be elucidated in the future.
鸟苷酸环化酶和磷酸二酯酶是分别含有GGDEF或EAL结构域的蛋白质,可合成或分解细菌二级信号分子——环二鸟苷酸(c-di-GMP)。基因组DNA序列分析表明这些蛋白质分布于几乎所有的细菌当中。但是,对于可以进行共生固氮作用的土壤细菌——根瘤菌而言,其GGDEF/EAL蛋白的研究仍是空白。本研究通过生物信息学方法在苜蓿中华根瘤菌(Sinorhizbium meliloti)中鉴定到19个GGDEF和EAL蛋白基因,其中编码GGDEF蛋白的基因5个,编码EAL蛋白的基因4个,同时编码GGDEF和EAL双结构域蛋白的基因10个。通过质粒插入失活的方法成功构建到14个基因突变菌株。所有14个基因的突变株在基本培养基中均表现出生长缺陷和运动性缺陷的表型,其中11个基因的突变株可以合成较多的胞外多糖,但其在紫花苜蓿上的竞争结瘤能力有所降低。进一步的研究发现,这些基因突变影响了根瘤菌对碳源和氮源的利用。我们的研究表明,GGDEF和EAL蛋白参与调节根瘤菌的多种生理过程,其催化产物——c-di-GMP在根瘤菌与宿主豆科植物相互作用过程中发挥了重要作用。
此外,我们发现具有完整expR基因的苜蓿中华根瘤菌在四倍体紫花苜蓿(Medicago sativa)和二倍体蒺藜苜蓿(M. truncatula)上具有不同的结瘤固氮表型,即该菌在紫花苜蓿结瘤固氮与野生型没有差别,但是在蒺藜苜蓿上却只能诱导低效固氮根瘤。我们对引起这种现象的原因进行了初步的探讨,发现该菌携带一个新的突变位点,其机制有待进一步研究。
About80%of the atmosphere is the nitrogen, but it can not be used by higher animals and plants directly. Biological nitrogen fixation affords the most part of nitrogen input.It is a hotspot in current study. Establishing this symbiosis requires a response of molecular signals, gene expression and nutrient exchange between the legumes and the rhizobia. Therefore, analyse the rhizobia and the relationship between rhizobia and legumes will be the main goal of our study, which has important scientific value in reducing agricultural production costs, maintaining the sustainable development of agriculture and environment.
A new bacterial secondary messenger, bis (3'-5')-cyclic dimeric guanosine monophosphate (c-di-GMP), is usually synthesized or decomposed by proteins containing a GGDEF or EAL domain. These proteins often act as a cyclase or phosphodiesterase of c-di-GMP, and their genes are distributed among almost all bacteria according to known genomic DNA sequences. However, the systematic identification of GGDEF and EAL protein genes remains unclear in rhizobia, soil bacteria that interact with compatible legumes to form nitrogen-fixing nodules. In this study,19putative GGDEF and EAL protein genes were identified in a model rhizobium, Sinorhizobium meliloti, by bioinformatic analysis (encoding5GGDEF proteins,4EAL proteins, and10GGDEF and EAL double-domain proteins). Null mutants of14genes were constructed through systematic plasmid insertion. All14gene mutants showed deficient growth in minimal medium and defective motility, and11gene mutants produced a lot more exopolysaccharide and displayed less competitive nodulation on the host plant, alfalfa. We also find that these mutants had different capacity to utilize sugars and amino acids as carbon and nitrogen sources respectively. Our results suggest that GGDEF and EAL proteins may play different roles in the control of S. meliloti physiology, although they contain conserved catalytic (GGDEF or EAL) domains. Our finding also implies that c-di-GMP may play an important role in the interactions between this rhizobium and its host plants to establish efficient symbiosis.
Additionally, S. meliloti strain carrying an intact expR gene induced the same efficiently nitrogen-fixing nodules on Medicago sativa as the wild type strain; however, it only induced lower efficient nodules on M. truncatula. And a now mutation locus was proposed, and the detail will be elucidated in the future.
引文
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