丛枝菌根共生体的氮代谢运输及其生态作用

丛枝菌根真菌能与80%的陆生维管植物形成互惠共生关系,共生体的存在对促进植物营养吸收和提高抗逆性具有重要意义.丛枝菌根真菌从宿主植物获取其光合产物碳水化合物的同时,通过外生菌丝吸收各种氮源,有效增强了宿主植物对氮素的吸收,以及氮在植物居群和群落水平上的交流,改善了植物营养代谢,增强了植物应对外界环境胁迫的能力.而共生体对氮的吸收、转运,以及氮从真菌到宿主植物的传输、代谢机制至今仍有许多问题亟待解决.本文综述了当前丛枝菌根共生体中氮传输代谢的主要机制,以及碳、磷对共生体氮传输代谢的影响;从群落和生态系统水平,简要阐述了丛枝菌根真菌在植物中氮分配的作用和对宿主植物的生态学意义,并提出共生体中氮代谢的一些需要深入研究的问题.
     

丛枝菌根利用氮素研究进展

氮素是植物需求量最大的元素,丛枝菌根真菌与植物形成共生体后能从土壤中获取无机氮、简单的氨基酸,还能利用一些复杂的有机态氮.考虑到NH+4在土壤中的移动性低及丛枝菌根真菌的专性共生菌的特点,丛枝菌根真菌吸收NH+4对植物的贡献较大.近年来的研究发现丛枝菌根真菌内存在与氮素代谢有关的鸟氨酸循环,而精氨酸则是菌丝内氮素转移的主要形式.综述最近的AMF对氮素的吸收、转运、同化、交换等方面的文献,旨在揭示丛枝菌根真菌氮素利用特点,阐明丛枝菌根真菌在氮循环系统中的重要作用.     

丛枝菌根真菌在土壤氮素循环中的作用

作为植物需求量最大的营养元素,氮素是陆地生态系统初级生产力的主要限制因子。丛枝菌根真菌能与地球上80%以上的陆生植物形成菌根共生体,帮助宿主植物吸收土壤中的P、N等矿质养分。目前,丛枝菌根真菌与氮素循环相关研究侧重于真菌对氮素的吸收形态以及共生体中氮的传输代谢机制,却忽略了丛枝菌根真菌在固氮过程、矿化与吸收过程、硝化过程、反硝化过程以及氮素淋洗过程等土壤氮素循环过程中所起到的潜在作用,并且越来越多的证据也表明丛枝菌根真菌是影响土壤氮素循环过程的重要因子。总结了丛枝菌根真菌可利用的氮素形态及真菌的氮代谢转运相关基因的研究现状;重点分析了丛枝菌根真菌在调控土壤氮素循环过程中的潜在作用以及在生态系统中的重要生态学意义,同时提出了丛枝菌根真菌在土壤氮素循环过程中一些需要深入研究的问题。     

丛枝菌根真菌脂类代谢对共生信号调控的响应和反馈

丛枝菌根(AM)真菌是自然生态系统中分布最为广泛的真菌之一,在自然界物质循环和能量流动中发挥着重要作用。经过长期的协同进化,AM真菌和宿主植物之间形成了完美的互惠互利的共生关系,而真菌的脂类代谢可能是揭示共生秘密的关键所在。本文综述了AM真菌脂类代谢在共生关系建立和维持中关键作用的最新研究进展,重点探讨了AM真菌脂类代谢对共生信号调控的响应和反馈机制,主要包括:AM真菌脂类存储和释放对共生和非共生状态的响应,以及脂类代谢产物变化与共生营养传递之间的关系;脂类分解过程在共生建立初期对信号分子调控发生的响应,以及相应的物质转化和能量代谢;菌根共生互惠互利关系维持中,真菌脂类代谢与信号分子交流通道的相互渗透和影响。本文对于理解菌根共生机制,促进菌根在生产中的应用具有促进作用。     

植物与丛枝菌根真菌在共生早期的信号交流

摘要:丛枝菌根真菌(Arbuscular Mycorrhizal Fungi,AMF)与大多数陆生植物互利共生具有广泛的生理生态学意义,而这一生态学功能背后的共生机制我们知之甚少。已探明AM形成前宿主植物根分泌的独脚金内酯促进AMF菌丝分支,分泌的角质单体促进AMF 在宿主根中定植;同时,菌根真菌的分支菌丝释放出脂质几丁糖(lipochitooligosaccharides,LCOs)和短链几丁质寡聚物(short-chain chitin oligomers,COs)信号分子诱导宿主基因表达、侧根发育以及形成Ca2+振荡,它们相互作用共同促进AM形成。在能同时形成菌根和根瘤的蒺藜苜蓿(Medicago truncatula)和日本百脉根(Lotus corniculatus)植物中,根瘤共生体形成过程所需的若干基因与菌根形成所需的基因有关。这些研究成果为全面揭示菌根共生体发生过程的信号转导奠定了基础。本文对目前国内外宿主植物与AM真菌之间的信号物质及其功能、相关基因及其调控功能等进行了综述,旨在为AM真菌共生早期的信号交流研究提供有价值的参考。     

丛枝菌根真菌与植物共生对植物水分关系的影响及机理

自1885年Frank首次提到菌根(mykorhiza)概念以来,大量的试验证实了丛枝菌根真菌(AMF)与植物根系之间形成具有一定结构和功能的共生体,促进植物生长并提高干旱耐受能力,在干旱生态系统中发挥重要的作用。该研究多集中在对宿主植物生理生态的影响及其机制方面,然而菌根共生对宿主植物水分吸收和信号产生、传递的影响研究少而分散,缺少系统总结。综述了最近四十多年丛枝菌根真菌与植物共生体对宿主植物干旱适应性影响研究进展,讨论了菌根共生对植物根冠通讯的影响及机理。干旱胁迫下AMF与植物共生,通过影响宿主植物一系列生理生态过程,提高宿主植物横向根压和纵向蒸腾拉力。经典的Ohm吸水模型是该方向最有代表性的研究成果,该模型揭示了菌根共生的根外菌丝具有不同于根细胞的细胞结构和水分运输性能,这为宿主植物提供一种特殊的快速吸水方式,可提高植物对土壤水分的吸收和运输能力。研究表明,AMF会影响宿主植物根冠通讯过程,如诱发信号级联反应,诱导根系尽早感知水分胁迫并产生非水力根源信号,提高宿主对干旱的耐受性。讨论了AMF在根冠通讯分子机制研究方面存在的问题及可能的解决途径,展望了AMF在干旱农业生产中的应用潜力。     

丛枝菌根形成过程及其信号转导途径

丛枝菌根是由一类土壤中古老的丛枝菌根真菌与植物根系形成的互利互惠共生体。通过共生作用丛枝菌根真菌帮助宿主植物提高水和矿质营养（特别是磷）的吸收效率。作为回报,大约20%的光合作用产物被转移到丛枝菌根真菌中,供其完成自身的生活史。丛枝菌根形成的过程中,需要植物与丛枝菌根真菌之间进行一系列信号分子的识别、交换以及信号转导作用,这一过程由一系列植物和菌根真菌的基因控制。首先,植物会分泌一种植物激素——独角金内酯来诱导菌根真菌加速分支,而菌根真菌也会分泌脂质几丁寡糖促进植物与其形成菌根。加速分支的菌根真菌接触到植物根部以后,会附着在植物根的表皮并形成附着胞,通过附着胞穿透植物根的表皮,最后进入维管组织附近的皮层细胞并在其中不断进行二叉分支,形成特有的丛枝结构。通过对模式植物共生现象的研究,已经发现很多植物基因参与到共生形成的信号转导过程中,包括早期植物反应的基因、菌根与根瘤共生共同需要的转导因子以及菌根特异的信号分子等。本文对菌根的形成过程及信号转导途径进行详细的介绍,为人们深入研究菌根关系提供参考。     

菌根真菌促进植物吸收利用氮素机制的研究进展

作为自然界最为普遍的一种植物共生体,菌根能够极大地促进植物对氮素的吸收和利用,其中菌根真菌在共生结构功能中发挥了重要作用。本文分别从菌根解剖构造、生理生化和分子生物学方面系统总结了菌根真菌促进植物吸收和利用氮素的研究现状。重点介绍了菌根真菌可利用的氮素形态及影响其利用的主要因素、菌根真菌的氮代谢途径GS-GOGAT以及菌根真菌中存在的鸟氨酸循环途径,指出精氨酸是菌丝内氮转运的主要形式,NH3可能为菌根真菌和植物界面质外体的主要转运形式。     

丛枝菌根网络介导的植物间信号交流研究进展及展望

丛枝菌根(arbuscular mycorrhizal, AM)真菌是一类能够与绝大多数陆地植物形成共生关系的土壤真菌, 其根外菌丝可以侵染不同植物根系且可以进行菌丝融合, 从而形成丛枝菌根网络(arbuscular mycorrhizal networks, AMNs)。AMNs可以在植物之间转运水分及营养元素如碳(C)、氮(N)、磷(P)等, 最近研究表明AMNs还可以在植物遭受环境胁迫时向邻近植物传递防御信号, 对周围植物起到“预警”作用。目前, 关于环境胁迫条件下AMNs介导的信号物质传递研究仍处于起步阶段, 许多问题亟待回答。该文首先回顾了目前有关AMNs介导的信号物质传递研究进展, 继而梳理了这一研究领域值得进一步探究的科学问题, 包括AMNs在植物间传递防御信号的可能途径及相关机制, AMNs介导的信号传递对菌根共生体系的可能影响, 以及AMNs研究中常用的技术及其发展, 最后讨论了AMNs介导的信号物质传递在作物保护等方面的可能应用。     

菌根真菌的碳氮循环功能研究进展

菌根(Mycorrhiza)是土壤真菌与植物根系形成的共生体(Symbiont),真菌一方面从植物获取碳水化合物,同时帮助植物吸收氮等矿质养分,因此,菌根真菌在生态系统的碳氮循环过程中发挥重要的作用.研究结果表明,菌根真菌可利用约4％-26％的植物净光合固定的碳水化合物,而其生物量和分泌物(如球囊霉素)具有重要的土壤碳汇功能;同时菌根真菌可参与土壤复杂有机质的降解过程.在菌根共生体系中,氮从根外菌丝到根内菌丝的传输经历了一个“无机-有机-无机”的转变过程.本文重点总结分析了菌根真菌在碳氮代谢功能与机理等方面的国内外最新研究进展,以及目前存在的主要问题与未来的研究重点.     

Arbuscular mycorrhizal fungi can transfer substantial amounts of nitrogen to their host plant from organic material

Nitrogen (N) capture by arbuscular mycorrhizal (AM) fungi from organic material is a recently discovered phenomenon. This study investigated the ability of two Glomus species to transfer N from organic material to host plants and examined whether the ability to capture N is related to fungal hyphal growth. Experimental microcosms had two compartments; these contained either a single plant of Plantago lanceolata inoculated with Glomus hoi or Glomus intraradices, or a patch of dried shoot material labelled with (15)N and (13)carbon (C). In one treatment, hyphae, but not roots, were allowed access to the patch; in the other treatment, access by both hyphae and roots was prevented. When allowed, fungi proliferated in the patch and captured N but not C, although G. intraradices transferred more N than G. hoi to the plant. Plants colonized with G. intraradices had a higher concentration of N than controls. Up to one-third of the patch N was captured by the AM fungi and transferred to the plant, while c. 20% of plant N may have been patch derived. These findings indicate that uptake from organic N could be important in AM symbiosis for both plant and fungal partners and that some AM fungi may acquire inorganic N from organic sources.     

植物菌根共生磷酸盐转运蛋白

大多数植物能和丛枝菌根（arbuscular mycorrhiza, AM）真菌形成菌根共生体。AM能够促进植物对土壤中矿质营养的吸收,尤其是磷的吸收。磷的吸收和转运由磷酸盐转运蛋白介导。总结了植物AM磷酸盐转运蛋白及其结构特征,分析其分类及系统进化,并综述了AM磷酸盐转运蛋白介导的磷的吸收和转运过程及其基因的表达调控。植物AM磷酸盐转运蛋白属于Pht1家族成员,它不仅对磷的吸收和转运是必需的,而且对AM共生也至关重要,为进一步了解菌根形成的分子机理及信号转导途径提供了理论基础。     

Plant–fungus competition for nitrogen erases mycorrhizal growth benefits of Andropogon gerardii under limited nitrogen supply

Considered to play an important role in plant mineral nutrition, arbuscular mycorrhizal (AM) symbiosis is a common relationship between the roots of a great majority of plant species and glomeromycotan fungi. Its effects on the plant host are highly context dependent, with the greatest benefits often observed in phosphorus (P)‐limited environments. Mycorrhizal contribution to plant nitrogen (N) nutrition is probably less important under most conditions. Moreover, inasmuch as both plant and fungi require substantial quantities of N for their growth, competition for N could potentially reduce net mycorrhizal benefits to the plant under conditions of limited N supply. Further compounded by increased belowground carbon (C) drain, the mycorrhizal costs could outweigh the benefits under severe N limitation. Using a field AM fungal community or a laboratory culture of Rhizophagus irregularis as mycorrhizal inoculants, we tested the contribution of mycorrhizal symbiosis to the growth, C allocation, and mineral nutrition of Andropogon gerardii growing in a nutrient‐poor substrate under variable N and P supplies. The plants unambiguously competed with the fungi for N when its supply was low, resulting in no or negative mycorrhizal growth and N‐uptake responses under such conditions. The field AM fungal communities manifested their potential to improve plant P nutrition only upon N fertilization, whereas the R. irregularis slightly yet significantly increased P uptake of its plant host (but not the host's growth) even without N supply. Coincident with increasing levels of root colonization by the AM fungal structures, both inoculants invariably increased nutritional and growth benefits to the host with increasing N supply. This, in turn, resulted in relieving plant P deficiency, which was persistent in non‐mycorrhizal plants across the entire range of nutrient supplies.     

Phosphate in the arbuscular mycorrhizal symbiosis: transport properties and regulatory roles

In response to the colonization by arbuscular mycorrhizal (AM) fungi, plants reprioritize their phosphate (Pi)-uptake strategies to take advantage of nutrient transfer via the fungus. The mechanisms underlying Pi transport are beginning to be understood, and recently, details of the regulation of plant and fungal Pi transporters in the AM symbiosis have been revealed. This review summarizes recent advances in this area and explores current data and hypotheses of how the plant Pi status affects the symbiosis. Finally, suggestions of an interrelationship of Pi and nitrogen (N) in the AM symbiosis are discussed.     

AM fungi root colonization increases the production of essential isoprenoids vs. nonessential isoprenoids especially under drought stress conditions or after jasmonic acid application

Previous studies have shown that root colonization by arbuscular mycorrhiza (AM) fungi enhances plant resistance to abiotic and biotic stressors and finally plant growth. However, little is known about the effect of AM on isoprenoid foliar and root content. In this study we tested whether the AM symbiosis affects carbon resource allocation to different classes of isoprenoids such as the volatile nonessential isoprenoids (monoterpenes and sesquiterpenes) and the non-volatile essential isoprenoids (abscisic acid, chlorophylls and carotenoids). By subjecting the plants to stressors such as drought and to exogenous application of JA, we wanted to test their interaction with AM symbiosis in conditions where isoprenoids usually play a role in resistance to stress and in plant defence. Root colonization by AM fungi favoured the leaf production of essential isoprenoids rather than nonessential ones, especially under drought stress conditions or after JA application. The increased carbon demand brought on by AM fungi might thus influence not only the amount of carbon allocated to isoprenoids, but also the carbon partitioning between the different classes of isoprenoids, thus explaining the not previously shown decrease of root volatile isoprenoids in AM plants. We propose that since AM fungi are a nutrient source for the plant, other carbon sinks normally necessary to increase nutrient uptake can be avoided and therefore the plant can devote more resources to synthesize essential isoprenoids for plant growth.     

Nutrient limitation drives response of <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis> to arbuscular mycorrhiza in primary succession

Little is known about the functioning of arbuscular mycorrhizal (AM) symbiosis over the course of primary succession, where soil, host plants, and AM fungal communities all undergo significant changes. Over the course of succession at the studied post-mining site, plant cover changes from an herbaceous community to the closed canopy of a deciduous forest. Calamagrostis epigejos (Poaceae) is a common denominator at all stages, and it dominates among AM host species. Its growth response to AM fungi was studied at four distinctive stages of natural succession: 12, 20, 30, and 50 years of age, each represented by three spatially separated sites. Soils obtained from all 12 studied sites were γ-sterilized and used in a greenhouse experiment in which C. epigejos plants were (1) inoculated with a respective community of native AM fungi, (2) inoculated with reference AM fungal isolates from laboratory collection, or (3) cultivated without AM fungi. AM fungi strongly boosted plant growth during the first two stages but not during the latter two, where the effect was neutral or even negative. While plant phosphorus (P) uptake was generally increased by AM fungi, no contribution of mycorrhizae to nitrogen (N) uptake was recorded. Based on N:P in plant biomass, we related the turn from a positive to a neutral/negative effect of AM fungi on plant growth, observed along the chronosequence, to a shift in relative P and N availability. No functional differences were found between native and reference inocula, yet root colonization by the native AM fungi decreased relative to the reference inoculum in the later succession stages, thereby indicating shifts in the composition of AM fungal communities reflected in different functional characteristics of their members.     

丛枝菌根（AM）生物技术在现代农业体系中的生态意义

菌根是植物根系与特定的土壤真菌形成的共生体,有利于生态系统中养分循环,协助植物抵御不良环境胁迫．自然条件下,大多数植物表现一定的菌根依赖性,在植株根系发育过程中如能与适宜的菌根真菌形成良好的菌根结构,可提高产量,改善品质,其中丛枝菌根是最普遍的类型．丛枝菌根帮助植物抵御不良环境胁迫及病虫害,促进植物健康生长,可减少化学肥料、杀虫剂施用量,以减少对环境、生态不利的化学物质施用量．丛枝菌根共生体可加速根系生长,提高对移动性低的无机离子吸收,加速养分循环利用,增强植物对不良胁迫(生物与非生物)因素的耐受力,形成良好的土壤结构,提高植物群体的多样性．文章综述了丛枝菌根真菌生态特征,丛枝菌根对寄主植物的影响,丛枝菌根生物技术应用于农业体系的生态意义及其应用潜力．     

Arbuscular mycorrhizal fungi-enhanced resistance against Phytophthora sojae infection on soybean leaves is mediated by a network involving hydrogen peroxide, jasmonic acid, and the metabolism of carbon and nitrogen

The arbuscular mycorrhizal fungi (AMF) enhance the resistance to pathogen infection in host plant. However, it is unclear how the AMF are involved in the systemic acquired resistance of host plant against pathogen. Here, an experiment was carried out to clarify the role of the AMF in soybean’s defense against the infection from pathogen Phytophthora sojae. It was found that the AMF contributed to the resistance of soybean against Phytophthora sojae by the release of hydrogen peroxide and by the accumulation of jasmonic acid in response to pathogenic invasion. Furthermore, the trade of nitrogen (N) from the fungus for carbon from the host was accelerated in the AM symbiosis in the defense reaction, which was indicated by the increased soluble sugar level, NO content and enzyme activities involved in N metabolism in the AM symbiosis.