禾草内生真菌作为生防因子的潜力分析

早熟禾亚科多种禾草可与Neotyphodium内生真菌形成禾草-内生真菌共生体, 这种植物-微生物共生体性状较为稳定, 且在自然界中广泛存在。禾草-内生真菌共生体稳定的互利共生关系不但保证了内生真菌所需的全部营养物质, 而且共生体产生的次生代谢物又可显著提高宿主禾草对生物胁迫的抗逆性。众多研究表明, 内生真菌的侵染可显著提高宿主禾草对虫害、病害及伴生植物等多种生物胁迫的抗性。据不完全统计, 禾草内生真菌对蛛形纲、线虫纲、昆虫纲3个纲至少79个种的害虫表现出较明显的抗性, 对至少22个种的病原真菌表现出明显的抗性。尽管利用内生真菌进行禾草品种选育及其品质改良的技术日趋成熟, 但是内生真菌在不同宿主禾草之间高效的替代转化技术, 及其在宿主体内遗传的稳定性仍有待于进一步深入探索。研究者把禾草内生真菌作为生防手段, 在未来的应用过程中不应只考虑其与宿主禾草之间的共生特异性, 而应更全面地分析禾草-内生真菌-生态环境之间的相互关系, 让内生真菌更好地为人类服务。     

内生真菌对氮添加羽茅根际土壤特性和微生物群落的影响

内生真菌不仅能改变与其共生植物的生理和生长指标,还可通过宿主植物间接对土壤的理化性质和微生物群落结构产生影响。以天然禾草——羽茅(Achnatherum sibiricum)为研究材料,探究在不同施氮水平下,内生真菌的种类对不同基因型的宿主植物根际土壤理化性质和微生物群落产生何种影响。结果表明,内生真菌侵染显著提高了羽茅根际土壤的pH值和微生物总量,但降低了土壤中真菌与细菌的比值。同时,土壤的pH值还受到了内生真菌种类的影响,其中感染Epichlo3 sibirica的羽茅根际土壤pH显著高于感染Epichlo3 gansuense-1的羽茅,而感染Epichlo3 gansuense-2的羽茅根际土壤pH与感染E.gansuensis-1、E.sibirica菌的羽茅相比没有显著差异。另外,内生真菌感染与否、内生真菌种类、施氮量以及宿主植物基因型对土壤总碳、总氮、微生物及碳矿化能力均无显著影响。     

植物根际促生菌及丛枝菌根真菌协助植物修复重金属污染土壤的机制

植物修复是一种前景广阔的重金属污染土壤的主要修复技术,在微生物的协助下效果更为显著。植物根际促生菌可通过分泌吲哚-3-乙酸(IAA)、产铁载体、固氮溶磷等方式促进植物生长、改善植物重金属耐受性,从而有效提高重金属污染土壤的植物修复效率。菌根真菌是土壤-植物系统中重要的功能菌群之一,可侵染植物根系改变根系形态和矿质营养状况,通过菌丝体吸附重金属,也可产生球囊霉素、有机酸、植物生长素等次生代谢产物改变重金属生物有效性。植物根际促生菌与丛枝菌根真菌可对植物产生协同促生作用,在重金属污染土壤修复中具有一定应用潜力。目前,国内外关于植物根际促生菌和丛枝菌根真菌互作已有大量研究,而二者的相互作用机理仍处于探索阶段。本文综述了近年来国内外植物根际促生菌和丛枝菌根真菌在重金属污染土壤植物修复中的作用机制,并对其研究前景进行展望。     

丛枝菌根真菌与植物硫素营养研究进展

丛枝菌根真菌是土壤微生物群落的重要组成部分,是最常见的地下共生菌,对植物和土壤具有多种有益作用。本文阐述了近年来丛枝菌根真菌对植物吸收土壤硫素的最新进展,在目前耕地缺硫状况下,着重分析了丛枝菌根真菌改善植物硫素营养以及丛枝菌根真菌利用硫素的分子调控机制,总结了影响菌根硫代谢的因素,并指出该研究方向仍存在的一些问题以及未来的研究侧重点。     

丛枝菌根真菌在寄主植物抵御生物和非生物胁迫中的作用

丛枝菌根真菌(arbuscular mycorrhiza fungi,AMF)是生态系统中普遍存在的土壤微生物,能与绝大多数植物形成共生关系,它在寄主植物抵御生物和非生物胁迫中所起的作用逐渐引起国内外学者的关注.论文综述了丛枝菌根真菌在植物抵御非生物胁迫(重金属污染、有机污染、盐胁迫和干旱胁迫)以及生物胁迫(致病菌和线虫侵染)中的作用,并在此基础上提出了未来该领域值得进一步研究的方向.     

盐碱胁迫对染内生菌和不染菌苇状羊茅根系丛枝菌根真菌群落多样性和组成的影响

香柱菌属Epichloë内生真菌存在于宿主植物地上部组织,不仅能提高宿主植物对生物与非生物逆境的抗性,而且能对周围环境中的微生物产生影响。该研究以染内生菌(endophyte-infected,EI)和不染菌(endophyte-free,EF)苇状羊茅Festuca arundinacea为实验材料,探究内生真菌和不同水平盐碱胁迫处理对宿主根系丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)群落多样性和组成的影响。结果表明,内生真菌和盐碱胁迫处理对苇状羊茅根系AMF多样性影响存在交互作用。EF苇状羊茅根系AMF多样性随盐碱胁迫处理水平的增加而降低,内生真菌的存在缓解了这一效应,在200和400 mmol/L盐碱胁迫处理下,内生真菌感染增加了苇状羊茅根系AMF多样性;此外,内生真菌感染改变了苇状羊茅根系AMF群落组成,降低了优势属Funneliformis相对多度,增加了Claroideoglomus、Glomus和unclassified AMF相对多度。结构方程模型结果表明,内生真菌通过间接增加土壤总磷浓度对苇状羊茅根系AMF多样性产生影响。本研究为筛选盐碱污染区生态修复的植物-微生物共生体提供基础。     

内生真菌和丛枝菌根真菌对羊草生长的影响

该研究比较了摩西球囊霉(Glmous mosseae)和幼套球囊霉(Glmous etunicatum)两种丛枝菌根真菌和内生真菌单独及混合接种对羊草(Leymus chinensis)生长的影响。结果表明, 内生真菌对2种菌根真菌的侵染均无显著影响, 内生真菌可极显著增加羊草的分蘖数、地上生物量、总生物量。内生真菌与菌根真菌之间的相互作用因菌根真菌种类而不同, 幼套球囊霉对宿主植物生长无明显影响且和内生真菌之间也无明显的相互作用; 单独接种摩西球囊霉显著增加羊草的地上、地下和总生物量, 当其与内生真菌共同存在时, 二者之间存在一定的拮抗作用。冗余分析结果表明, 在内生真菌-AM真菌-羊草共生体中, 内生真菌对宿主植物生长的影响最大, 摩西球囊霉对宿主植物生长也有一定的贡献, 幼套球囊霉对宿主植物生长无明显影响。     

丛枝菌根真菌对植物根腐病的抑制效应及其机制

根腐病是一类危害严重的土传病害,常常导致作物产量和品质降低。丛枝菌根(AM)真菌是一类重要的土壤微生物,通过与植物根系建立共生体而发挥重要的生理生态功能。研究表明,AM真菌通过调节宿主植物一系列生理生化响应,诱导植物增强根腐病抗性。当前,利用AM真菌开展根腐病等土传病害的生物防治是植物与微生物互作领域的研究热点。本文全面梳理了AM真菌对宿主植物根腐病病原物的抑制效应,系统总结了AM真菌改变宿主植物根系形态结构、改善植物营养水平、与病原物竞争生态位点、激活植物防御体系、调节根系分泌物等方面的研究结果,分析了AM真菌抑制根腐病危害的作用机制,展望了AM真菌抑制根腐病危害的潜在机制和AM真菌高效利用面临的现实问题,旨在为利用AM真菌开展植物根腐病的生物防治提供理论依据。     

丛枝菌根真菌形态结构、物种多样性和群落组成对氮沉降响应研究进展

工业革命以来,人类活动输入到生态系统中的氮迅速增加,已突破地球所能承受的氮循环阈值。过量氮沉降会造成生物多样性丧失等一系列危害,严重影响生态系统结构和功能。丛枝菌根(AM)真菌能够与大约70%-80%的陆地植物种类形成共生关系,在宿主植物养分吸收、抵抗外界不良环境压力、群落动态和物种共存、生物地球化学循环等方面具有重要的作用。探究AM真菌对氮沉降的响应对认识和把握菌根真菌缓解氮沉降的负面后果,维持生态系统的结构和功能具有重要意义。本文综述了AM真菌的形态结构、物种多样性和群落组成等对氮沉降的响应机制。前人研究表明氮沉降通常降低AM真菌的根系定殖率,减少根外菌丝密度和土壤孢子密度,改变菌丝生长的时间动态;降低AM真菌多样性,改变AM真菌群落组成。氮沉降主要通过缓解植物氮限制、降低植物对菌根的依赖性、减少植物对菌根的碳分配、改变根系和土壤中菌根生物量比率、在植物根内维持稳定的菌根真菌组成作为应对未来扰动的“保险”、改变土壤资源有效性及土壤酸度等直接和间接途径影响AM真菌结构和功能。我们建议在未来研究中整合多组学手段、开展学科交叉,聚焦复杂的生物互作体系对氮沉降的响应机制,以及AM真菌对氮沉降响应的生态后果。     

菌根真菌在生态系统中的作用

 菌根是一种植物营养根与土壤真菌形成的共生体,在自然界中分布广泛。本文着重从以下几个方面介绍相关的研究进展：1) 菌根真菌作为生态系统的重要组成部分,具有不可忽视的生物量,并成为连接绿色植物和食真菌者食物链的重要一环;2) 菌根真菌通过参与凋落物的酶降解过程影响有机物的循环,通过促进生物固氮、加速土壤磷的风化、提高土壤溶液离子的有效性以及直接吸收等过程影响氮、磷、钾、钙、镁等元素的无机循环;3) 菌根真菌与土壤微生物间存在有益的或拮抗的相互作用,并可以直接或间接地影响根际生物区系的组成和数量;菌根真菌通过对宿主植物的有益作用而影响植物的种间竞争,通过菌根网络而形成的种团可以在同种或不同种植物间实现资源的重新分配和共享;由于对种间关系的作用和对食物链的影响,菌根真菌对群落的物种构成和多样性的维持具有重要的作用;菌根真菌是群落演替过程的指示者,也是这一过程的参与者和推动者,并且菌根真菌的存在也有利于提高土壤团聚体的稳定性及促进灰壤的形成;4) 菌根真菌的种类和数量可以指示生态系统中自然的或人类活动引起的变化,并可以在生态系统的保护、恢复或重建过程中发挥重要作用。文章的最后还介绍了最新的研究热点和发展趋势。     

Consequences of simultaneous interactions of fungal endophytes and arbuscular mycorrhizal fungi with a shared host grass

Plants simultaneously associate with multiple microbial symbionts throughout their lifetimes. To address the question of whether the effects of simultaneous symbionts are contingent on the specific identities, we conducted a greenhouse experiment manipulating the presence and identities of arbuscular mycorrhizal fungi (AMF) and fungal endophytes on the shared host grass Elymus hystrix. Each plant host was inoculated with one of two AMF species having varying effects on host growth, or a sterile soil control. Further, we used naturally occurring endophyte‐infected (E+) and uninfected (E–) individuals from two populations of the endophyte Epichloë elymi that varied in their interaction with E. hystrix. We then measured responses of plants, AMF, and fungal endophytes. Overall, we found that the combined effects of AMF and fungal endophytes on plant growth were additive, reflecting the mutualistic quality of each symbiont independently interacting with host plants. However, fungal endophyte infection differentially altered hyphal colonization of the two AMF species and the identity of the coinfecting AMF species affected fungal endophyte fitness traits. The results of this study demonstrate that the outcome of interspecific symbiotic interactions varies with partner identity such that the effects of simultaneous symbioses can not be generalized.     

Evidence for leaf endophyte regulation of root symbionts: effect of <Emphasis Type="Italic">Neotyphodium</Emphasis> endophytes on the pre-infective state of mycorrhizal fungi

Neotyphodium endophytes and arbuscular mycorrhizal (AM) fungi are common constituents of natural grasslands. The plant–endophyte symbiosis can introduce changes in soil conditions that affect the density and activity of different functional groups of soil organisms. In the present work we performed in vitro assays to evaluate the effect of root and endophyte exudates on the pre-infective state of mycorrhizal fungi (Gigaspora margarita and G. rosea). Plant roots of Bromus setifolius from populations of Patagonia, and four strains of Neotyphodium were used to obtain the exudates. Root exudates of infected plants, at a high concentration, significantly increased AMF hyphal branches and length relative to exudates from naturally endophyte free plants. The effect of Neotyphodium endophyte exudates on AMF mycelial length varied depending on strain and the concentration used, suggesting a differential interaction between endophyte and AMF species. AMF hyphal branches were increased by Neotyphodium fungal exudates in both mycorrhizal species. A few previous studies have suggested that Neotyphodium endophytes can reduce mycorrhizal sporulation and colonization of host roots in commonly-cultivated agronomic hosts. In this study we report the opposite effect in B. setifolius. This study reports the direct and positive effect of root exudates from plants in symbiosis with Neotyphodium, on AMF pre-infective state. Further, identical effects were detected using exudates from Neotyphodium endophytes.     

<Emphasis Type="Italic">Epichloë</Emphasis> exudates promote in vitro and in vivo arbuscular mycorrhizal fungi development and plant growth

Background and aims

We studied, through exudates employment, the effect of Epichloë (endophytic fungi), both independently and in association with Bromus auleticus (grass), on arbuscular mycorrhizal fungi (AMF) colonization, host and neighbouring plants biomass production and soil changes.

Methods

Through in vitro and greenhouse experiments, Epichloë endophytes effect on AMF development was evaluated. In vitro studies of exudates effect on Gigaspora rosea and Rhizophagus intraradices were performed using root or endophyte exudates. A 6-month greenhouse experiment was conducted to determine Bromus auleticus endophytic status effect and endophyte exudates role in biomass production, neighbouring plants mycorrhizal colonization and soil properties.

Results

Endophyte exudates and E+ plant root exudates promoted in vitro AMF development in the pre-infective stage of G. rosea and in carrot root culture mycelium of R. intraradices in a dose-response relationship, while control media and E- plants exudates had no effect. R. intraradices colonization and plant growth was clearly increased by endophytes and their exudates.

Conclusions

This is the first work evidencing the direct effect of Epichloë endophytes and infected plants root exudates on AMF extramatrical development. While higher levels of AMF colonization were observed in E+ plants, no clear effect was detected in neighbouring plants colonization, plant biomass or soil properties.

     

Can seed-borne endophytes promote grass invasion by reducing host dependence on mycorrhizas?

Symbiotic interactions between plants and microorganisms have recently become the focus of research on biological invasions. However, the interaction between different symbionts and their consequences in host-plant invasion have been seldom explored. Here, we propose that vertically transmitted fungal endophytes could reduce the dependency of invasive grasses on mycorrhizal fungi allowing host establishment in those environments where the specific mutualist may be not present. Through analyzing published studies on nine grass species, we evaluated the effect of seed-borne Epichloë endophytes on the relationship of invasive and non-invasive grasses with arbuscular mycorrhizal fungi (AMF), a symbiosis known to be fundamental for plant fitness and invasion success. The endophyte effect on AMF colonization differed between invasive and non-invasive grasses, reducing mycorrhization only on invasive species but with no impact on their biomass. These results allowed us to propose that Epichloë endophytes could reduce the dependency of host plants on the mutualism with AMF, promoting host grass establishment and subsequent invasion. Simultaneous interactions with different types of mutualists may have profound effects on the host-plant fitness facilitating its range expansion. Our findings suggest that some specific mutualistic fungi such as epichloid endophytes facilitate host invasion by reducing the requirements of the benefits derived from other mutualisms.     

Epichloë endophyte affects the root colonization pattern of belowground symbionts in a wild grass

Plants host multiple symbionts that interact with each other affecting plant performance and regulating their establishment. Here, we analyzed how the association with Epichloë endophytes affects belowground colonization by Dark Septate Endophytes (DSE) and arbuscular mycorrhizal fungi (AMF) in the grass Bromus auleticus. Epichloë-symbiotic (E+) and Epichloë-non symbiotic (E−) plants were sampled from a long-term experimental plot and colonization structures were analyzed in the roots. We also examined the influence of Epichloë exudates on the in vitro growth of DSE Microdochium bolleyi isolated from roots. Epichloë symbiosis increased AMF colonization, although differences were not significant. Despite the lack of differences in total DSE colonization, in concordance with in vitro findings, a higher significant abundance of microsclerotia was observed in E+ plants. A negative correlation between total mycorrhizal and DSE was found. Our findings show a more uniform root colonization pattern in E+ plants, suggesting a root symbiosis modulating role.     

Contrasting diurnal variations in soil organic carbon decomposition and root respiration due to a hysteresis effect with soil temperature in a Gossypium s. (cotton) plantation

Cool-season grasses commonly harbor fungal endophytes in their aerial tissues. However the effects of these symbionts on soil microbial communities have rarely been investigated. Our objective was to explore microbial community responses in soils conditioned by plants of the annual grass Lolium multiflorum with contrasting levels of infection with the endophyte Neotyphodium occultans. At the end of the host growing season, we estimated the functional capacity of soil microbial communities (via catabolic response profiles), the contribution of fungi and bacteria to soil activity (via selective inhibition with antibiotics), and the structure of both microbial communities by molecular analyses. Soil conditioning by highly infected plants affected soil catabolic profiles and tended to increase soil fungal activity. We detected a shift in bacterial community structure while no changes were observed for fungi. Soil responses became evident even without changes in host plant biomass or soil organic carbon or total nitrogen content, suggesting that the endophyte modified host rhizodepositions during the conditioning phase. Our results have implications for the understanding of the reciprocal interactions between above and belowground communities, suggesting that plant-soil feedbacks can be mediated by this symbiosis.     

Balancing multiple mutualists: asymmetric interactions among plants, arbuscular mycorrhizal fungi, and fungal endophytes

Most organisms engage in beneficial interactions with other species; however, little is known regarding how individuals balance the competing demands of multiple mutualisms. Here we examine three-way interactions among a widespread grass, Schedonorus phoenix , a protective fungal endophyte aboveground, Neotyphodium coenophialum , and nutritional symbionts (arbuscular mycorrhizal fungi) belowground. In a greenhouse experiment, we manipulated the presence/absence of both fungi and applied a fertilizer treatment to individual plants. Endophyte presence in host plants strongly reduced mycorrhizal colonization of roots. Additionally, for plants with the endophyte, the density of endophyte hyphae was negatively correlated with mycorrhizal colonization, suggesting a novel role for endophyte abundance in the interaction between the symbionts. Endophyte presence increased plant biomass, and there was a positive correlation between endophyte hyphal density and plant biomass. The effects of mutualists were asymmetric: mycorrhizal fungi treatments had no significant impact on the endophyte and negligible effects on plant biomass. Fertilization affected all three species – increasing plant biomass and endophyte density, but diminishing mycorrhizal colonization. Mechanisms driving negative effects of endophytes on mycorrhizae may include inhibition via endophyte alkaloids, altered nutritional requirements of the host plant, and/or temporal and spatial priority effects in the interactions among plants and multiple symbionts.     

Effects of Resident Soil Fungi and Land Use History Outweigh Those of Commercial Mycorrhizal Inocula: Testing a Restoration Strategy in Unsterilized Soil

Arbuscular mycorrhizal fungi (AMF) have numerous effects on temperate grassland ecosystems, but prairie restorations are frequently located in sites with depauperate AMF communities. In this greenhouse study, four native species (Schizachyrium scoparium, Elymus canadensis, Monarda punctata, and Aster ericoides) and an invasive grass (Bromus inermis) were grown in unsterilized field soils and treated with two types of commercial AMF inoculum. Inocula were applied at one and two times the manufacturers' suggested rate. Soil was collected from a meadow enrolled in the Conservation Reserve Program (CRP), and from an active agricultural field. Inoculum addition had no effect on biomass or percent colonization by AMF for any grass species, regardless of soil type. Inoculum type significantly affected Aster biomass and percent colonization, although pairwise comparisons of treated individuals and controls were not significant. The overall lack of effectiveness of the commercial inocula may reflect the small number of propagules added, even when used at twice the recommended rate. Higher rates of fungal colonization in all three grasses and increased biomass in the native grasses were observed in individuals grown in the CRP soil. Plants were also colonized by dark septate endophytic fungi; for Schizachyrium, endophyte colonization was significantly greater in tilled than CRP soil. Our results indicate that an existing soil fungal community promotes colonization by AMF more than the addition of commercial inocula, and that soil characteristics associated with land use history significantly affect the growth of native species in a restoration setting.     

Effects of the endophyte Epichloë coenophiala on the root microbial community and growth performance of tall fescue in different saline-alkali soils

Soil salinization is detrimental to plant growth and yield in agroecosystems worldwide. Epichloë endophytes, a class of clavicipitaceous fungi, enhance the resistance of host plants to saline-alkali stress. This study explored the effects of the systemic fungal endophyte Epichloë coenophiala on the root microbial community and growth performance of tall fescue (Lolium arundinaceum) growing under different saline-alkali stress conditions. Structural equation modeling (SEM) was conducted to analyze the direct and indirect effects (mediated by root microbial community diversity and soil properties) of the endophyte on the growth of tall fescue under saline-alkali stress. The endophyte-infected plants produced higher shoot and root biomass compared to endophyte-free plants under saline-alkali stress (200 and 400 mM). Endophyte infection increased the fungal community diversity and altered its composition in the roots, decreasing the relative abundance of Ascomycota and increasing that of Glomeromycota. Furthermore, endophyte infection decreased the bacterial community diversity and the relative abundance of dominant Proteobacteria. SEM showed that endophyte infection increased the shoot and root biomass under saline-alkali stress (200 and 400 mM) by increasing the arbuscular mycorrhizal fungal diversity in the roots, and soil total nitrogen and phosphorus concentrations. Therefore, it is important to examine aboveground microbes as factors influencing plant growth in saline-alkali stress by affecting belowground microbes and soil chemical properties.