豆科植物共生结瘤的分子基础和调控研究进展

豆科植物与根瘤菌共生互作的结果导致了一个新的植物器官――根瘤的形成, 根瘤菌生活在根瘤中, 它们具有将氮气转化为能被植物同化的氨的能力。该文阐述了根瘤的形成过程和类型, 并主要以模式豆科植物蒺藜苜蓿(Medicago truncatula)和日本百脉根(Lotus japonicus)为例, 对近年来共生结瘤过程中宿主植物对根瘤菌结瘤因子的识别和信号传递、侵入线形成和固氮的分子基础, 以及宿主植物对根瘤形成的自主调控机制、环境中氮素营养对结瘤的影响研究进行了综述, 指出当前豆科植物与根瘤菌共生互作研究存在的问题, 并对今后的研究方向作了分析与展望。     

根瘤菌NOD因子的感知与信号传导

根瘤菌是一类引起豆科植物结瘤固氮的土壤细菌。根瘤中的类菌体固定空气中的氮气为宿主植物提供充足的氮源。共生体系的建立始于细菌与宿主植物间复杂的信号交换过程。植物产生类黄酮诱导相应的根瘤菌合成分泌结瘤因子 ,后者进而诱导宿主植物根系形态变化以及早期根瘤素基因表达。以下将就宿主植物结瘤因子的特异识别和早期信号传导进行讨论。     

豆科植物早期共生信号转导的研究进展

豆科植物与根瘤菌建立特异的共生关系,在寄主根部产生固氮根瘤。此过程包含了共生信号识别与传递、根瘤菌侵染、根瘤形成以及固氮功能实现等生物学事件。研究人员已经从2种豆科模式植物蒺藜苜蓿（Medicago truncatula）和百脉根（Lotus japonicus）的共生固氮体系中,筛选到许多与根瘤菌共生相关的突变体及其相对应的功能基因,建立起包含结瘤因子识别、共生信号传递和转录响应在内的早期共生信号途径。该文对豆科植物早期共生信号途径的新进展进行了综述。     

豆科植物早期共生信号转导的研究进展北大核心CSCD

豆科植物与根瘤菌建立特异的共生关系,在寄主根部产生固氮根瘤。此过程包含了共生信号识别与传递、根瘤菌侵染、根瘤形成以及固氮功能实现等生物学事件。研究人员已经从2种豆科模式植物蒺藜苜蓿(Medicago truncatula)和百脉根(Lotus japonicus)的共生固氮体系中,筛选到许多与根瘤菌共生相关的突变体及其相对应的功能基因,建立起包含结瘤因子识别、共生信号传递和转录响应在内的早期共生信号途径。该文对豆科植物早期共生信号途径的新进展进行了综述。     

硝酸盐调控豆科植物与根瘤菌共生固氮的机制研究

氮是植物生长发育所需的大量营养元素之一。硝态氮不仅可以被植物直接吸收利用,还可以作为重要的信号分子调控植物对氮素的响应、吸收、代谢相关基因的表达,从而影响植物的生长和发育。豆科植物可以通过与根瘤菌共生互作来获得生长所需的氮,但共生固氮是一个耗费植物能量的过程。当土壤中存在高浓度的氮素时,氮作为信号分子会影响共生固氮基因的功能从而抑制共生固氮过程。目前的研究表明,硝酸盐通过局部和系统的调控方式抑制共生固氮过程;结瘤自主调控(Autoregulation of nodulation,AON)和NLPs(NIN-like proteins)转录因子在硝酸盐抑制豆科植物根瘤形成中有着重要的作用。本文结合最近的研究进展,重点讨论NLPs转录因子和AON途径在硝酸盐抑制共生固氮过程的作用。     

根瘤菌脂壳寡糖结瘤因子研究概况

共生固氮是根瘤菌与豆科植物相互作用的结果,它在农业上有重要意义。结瘤与固氮包括一系列复杂的生物学过程,它涉及微生物与植物间专一性识别、信息交换和基因协同表达等方面。近些年研究已经揭示出根瘤菌与豆科植物相互作用分子基础的基本框架。在根瘤的形成过程中,植物与根瘤菌之间首先进行信息交换,促使根瘤菌产生脂壳寡糖类物质。这类脂壳寡糖类物质能引起植物形成根瘤,因此被称为脂壳寡糖结瘤因子（Ｌｉｐｏｃｈｉｔｉｎｏｌｉｇｏｓａｃｃｈａｒｉｄｅｓ）或结瘤因子（ｎｏｄｆａｃｔｏｒｓ）［１］。脂壳寡糖结瘤因子的发现、结…     

高氮抑制豆科植物结瘤固氮机制研究进展

豆科植物通过与根瘤菌共生,形成根瘤并进行生物固氮。豆科植物的结瘤固氮作用在农业上具有减肥增效、改良土壤等重大意义。然而,高氮会抑制豆科植物结瘤固氮,形成"氮阻遏"效应。着重论述了高氮抑制豆科植物结瘤的分子机制,包括氮素通过结瘤自调控AON(Autoregulation of nodulation)信号、NLP(NIN-like protein)转录因子、植物激素信号等途径抑制根瘤数目和发育的最新进展;并探讨了高氮抑制根瘤固氮活性的假说及争议,包括亚硝酸盐毒性和碳饥饿等,以期为提高豆科植物应对"氮阻遏"效应提供理论基础。     

结瘤素及其基因表达

一、前言: 根瘤菌与豆科植物相互作用产生固氮根瘤要经过一系列复杂的过程,这些过程中涉及到许多植物及根瘤菌基因的表达。在根瘤菌方面,除有编码固氮酶的基因以及参与其表达调节的nif、fix基因外,还需要有参与结瘤过程的nod基因及参与细菌胞外。多糖合成的exo基因,这些基因的任何一个发生变异都会影响共生固氮过程。     

豆科植物SHR-SCR模块——根瘤“奠基细胞”的命运推手

豆科植物-根瘤菌共生固氮是可持续性农业氮肥的最重要来源。根瘤作为豆科植物共生固氮的一种特化植物侧生器官, 提供了根瘤菌生物固氮必需的微环境, 是根瘤菌的安身之本, 因此, 根瘤的正常发育是实现豆科植物-根瘤菌共生固氮的结构基础。根瘤器官的从头发生主要起始于根瘤菌诱导的根皮层细胞分裂。通常认为豆科植物的根皮层具备有别于非豆科植物根皮层的某种特异属性, 从而响应根瘤菌并与之建立固氮共生, 但长期以来该属性决定的分子机制一直不明确。近日, 中国科学院分子植物科学卓越创新中心王二涛团队以蒺藜苜蓿(Medicago truncatula)等豆科植物和拟南芥(Arabidopsis thaliana)等非豆科植物为研究对象, 发现豆科植物中保守的SHR-SCR干细胞模块决定了其皮层细胞分裂潜能从而赋予根瘤器官发生的命运。该研究揭示了豆科植物根瘤发育的全新机制, 提供了研究和理解植物-根瘤菌固氮共生进化的重要线索, 对提高豆科作物固氮效率和非豆科作物固氮工程具有重要意义。     

The Legume SHR-SCR Module Predetermines Nodule Founder Cell Identity

豆科植物-根瘤菌共生固氮是可持续性农业氮肥的最重要来源。根瘤作为豆科植物共生固氮的一种特化植物侧生器官, 提供了根瘤菌生物固氮必需的微环境, 是根瘤菌的安身之本, 因此, 根瘤的正常发育是实现豆科植物-根瘤菌共生固氮的结构基础。根瘤器官的从头发生主要起始于根瘤菌诱导的根皮层细胞分裂。通常认为豆科植物的根皮层具备有别于非豆科植物根皮层的某种特异属性, 从而响应根瘤菌并与之建立固氮共生, 但长期以来该属性决定的分子机制一直不明确。近日, 中国科学院分子植物科学卓越创新中心王二涛团队以蒺藜苜蓿(Medicago truncatula)等豆科植物和拟南芥(Arabidopsis thaliana)等非豆科植物为研究对象, 发现豆科植物中保守的SHR-SCR干细胞模块决定了其皮层细胞分裂潜能从而赋予根瘤器官发生的命运。该研究揭示了豆科植物根瘤发育的全新机制, 提供了研究和理解植物-根瘤菌固氮共生进化的重要线索, 对提高豆科作物固氮效率和非豆科作物固氮工程具有重要意义。     

Nitric oxide in legume-rhizobium symbiosis

Nitric oxide (NO) is a gaseous signaling molecule with a broad spectrum of regulatory functions in plant growth and development. NO has been found to be involved in various pathogenic or symbiotic plant-microbe interactions. During the last decade, increasing evidence of the occurrence of NO during legume-rhizobium symbioses has been reported, from early steps of plant-bacteria interaction, to the nitrogen-fixing step in mature nodules. This review focuses on recent advances on NO production and function in nitrogen-fixing symbiosis. First, the potential plant and bacterial sources of NO, including NO synthase-like, nitrate reductase or electron transfer chains of both partners, are presented. Then responses of plant and bacterial cells to the presence of NO are presented in the context of the N₂-fixing symbiosis. Finally, the roles of NO as either a regulatory signal of development, or a toxic compound with inhibitory effects on nitrogen fixation, or an intermediate involved in energy metabolism, during symbiosis establishment and nodule functioning are discussed.     

根瘤菌共生固氮能力的进化模式

根瘤菌-豆科植物共生固氮体系对农业的可持续性发展至关重要,也是研究原核与真核生物互利共生的模式体系之一。长期以来,根瘤菌共生固氮相关研究主要集中在结瘤因子与固氮酶合成及调控等少数关键基因,但仅获得这些关键基因却不能保证细菌获得结瘤固氮能力。随着比较和功能基因组学的快速发展和应用,越来越多的研究发现根瘤菌使用了很多系统发育分支特异的遗传机制与豆科植物建立有效的共生关系,进一步揭示了双方互利共生的复杂性。本综述总结了近年来比较基因组学、遗传学以及实验进化等方面的相关研究进展,在此基础上讨论根瘤菌共生固氮能力的进化模式。     

Symbiosis specificity in the legume: rhizobial mutualism

Legume plants are able to engage in root nodule symbiosis with nitrogen-fixing soil bacteria, collectively called rhizobia. This mutualistic association is highly specific, such that each rhizobial species/strain interacts with only a specific group of legumes, and vice versa. Symbiosis specificity can occur at multiple phases of the interaction, ranging from initial bacterial attachment and infection to late nodule development associated with nitrogen fixation. Genetic control of symbiosis specificity is complex, involving fine-tuned signal communication between the symbiotic partners. Here we review our current understanding of the mechanisms used by the host and bacteria to choose their symbiotic partners, with a special focus on the role that the host immunity plays in controlling the specificity of the legume - rhizobial symbiosis.     

Control of NO level in rhizobium-legume root nodules: Not only a plant globin story

Nitric oxide (NO) is a gaseous signaling molecule which plays both regulatory and defense roles in animals and plants. In the symbiosis between legumes and rhizobia, NO has been shown to be involved in bacterial infection and nodule development steps as well as in mature nodule functioning. We recently showed that an increase in NO level inside Medicago truncatula root nodules also could trigger premature nodule senescence. Here we discuss the importance of the bacterial Sinorhizobium meliloti flavohemoglobin to finely tune the NO level inside nodules and further, we demonstrate that S. meliloti possesses at least two non redundant ways to control NO and that both systems are necessary to maintain efficient nitrogen fixing activity.     

Recent development and new insight of diversification and symbiosis specificity of legume rhizobia: mechanism and application

Currently, symbiotic rhizobia (sl., rhizobium) refer to the soil bacteria in α- and β-Proteobacteria that can induce root and/or stem nodules on some legumes and a few of nonlegumes. In the nodules, rhizobia convert the inert dinitrogen gas (N₂) into ammonia (NH₃) and supply them as nitrogen nutrient to the host plant. In general, this symbiotic association presents specificity between rhizobial and leguminous species, and most of the rhizobia use lipochitooligosaccharides, so called Nod factor (NF), for cooperating with their host plant to initiate the formation of nodule primordium and to inhibit the plant immunity. Besides NF, effectors secreted by type III secretion system (T3SS), exopolysaccharides and many microbe-associated molecular patterns in the rhizobia also play important roles in nodulation and immunity response between rhizobia and legumes. However, the promiscuous hosts like Glycine max and Sophora flavescens can nodulate with various rhizobial species harbouring diverse symbiosis genes in different soils, meaning that the nodulation specificity/efficiency might be mainly determined by the host plants and regulated by the soil conditions in a certain cases. Based on previous studies on rhizobial application, we propose a ‘1+n−N’ model to promote the function of symbiotic nitrogen fixation (SNF) in agricultural practice, where ‘1’ refers to appreciate rhizobium; ‘+n’ means the addition of multiple trace elements and PGPR bacteria; and ‘−N’ implies the reduction of chemical nitrogen fertilizer. Finally, open questions in the SNF field are raised to future think deeply and researches.