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。     

丛枝真菌对互花米草和芦苇氮磷吸收的影响

互花米草(Spartina alterniflora Loisel.)是我国海滨盐沼的入侵植物,与土著种芦苇(Phragmites australis)形成了广泛的竞争;已知丛枝菌根(AMF)对不同植物的生长存在差异性影响;但其对互花米草与芦苇之间的种间关系,是否对互花米草入侵芦苇群体产生作用值得探讨.研究对两物种进行了丛枝菌根接种处理,种植模式处理和盐度处理的三因素实验.结果表明:盐度增加使得单种时芦苇、混种时互花米草的AMF侵染率显著下降(p<0.05),而混种时芦苇和单种时的互花米草AMF侵染率受盐度影响不显著(p>0.05).混种时,两种植物的丛枝菌根形成均受对方影响,并且盐度升高使两种植物之间对AMF侵染率的影响发生变化,在淡水生境下混种时,芦苇的AMF侵染率比单种时降低40.5%,互花米草的AMF侵染率比单种时提高了86.9%,均差异显著(p<0.05);在低盐度下混种时芦苇的AMF侵染率比单种时降低24.7%,差异显著(p<0.05),而对互花米草的影响不显著;在高盐度下混种对芦苇的AMF侵染率影响不显著,而使互花米草的AMF侵染率显著降低,降低率比例达78.7%.在淡水生境下,丛枝菌根对芦苇和互花米草的N、P吸收均有显著的促进作用;但是在咸水生境下生长时芦苇的N、P含量主要受盐度的显著影响(p<0.05),随盐度增加而增加;虽然在咸水生境下丛枝菌根仍旧促进芦苇的N、P吸收,但其影响远小于盐度的影响,并且促进效果受到盐度的抑制;但互花米草的N、P含量不受盐度影响.由此可见,接种AMF对这两种植物的氮磷吸收有着不同程度的促进,其作用大小与侵染程度有关,且受到盐度和种植模式的影响.     

丛枝菌根对有机污染土壤的修复作用及机理

丛枝菌根(AM)是丛枝菌根真菌(AMF)与植物根系相互作用的互惠共生体,能改良土壤结构,增强植物抗性.自然界中已知的AMF有170多种,分布广泛,且可与大多数植物共生.利用AM修复有机污染土壤正成为一个崭新的研究方向.本文综述了AM对多环芳烃、酞酸脂、石油和农药等一些典型有机污染物污染土壤的修复作用.AM修复有机污染土壤的机理主要包括:AMF代谢有机污染物;AM分泌酶,降解污染物;AM影响根系分泌作用,并促进根际微生物对有机污染物的降解;AMF宿主植物吸收积累污染物.AM修复研究中,高效AMF的筛选、复合菌种效应、土壤老化、AM作用下植物对有机污染物的吸收积累等几方面仍有待于深入研究.     

典型农田土壤中丛枝菌根真菌-根际细菌互作及与氮磷利用的关系

在农业生态系统中,土壤微生物是土壤-作物系统养分循环的重要驱动力,其中丛枝菌根真菌(Arbuscularmycorrhizalfungi,AMF)能够促进作物对养分的吸收,适应逆境胁迫。【目的】进一步揭示AMF和根际细菌群落的跨界网络互作,挖掘与作物氮磷利用显著相关的关键微生物类群,揭示关键类群的生态网络特征。【方法】利用Illumina测序技术对3种典型农田旱地土壤(黑土、潮土和红壤)中AMF和根际细菌群落结构进行分析;构建互作网络并利用偏冗余分析、相关性分析探究了与氮磷利用相关的潜在关键类群。【结果】3种土壤中AMF与根际细菌均以正相互作用为主。不同土壤中AMF与根际细菌互作关系差异明显,在红壤中跨界互作最为密切,其中球囊霉属真菌(Glomus)与根际细菌中的放线菌(Actinobacteria)和变形菌(Proteobacteria)之间的交互作用最多。而在黑土中主要体现为根际细菌的界内互作。与氮磷利用率显著相关的关键微生物类群主要属于球囊霉属真菌、放线菌和α变形菌。【结论】典型旱地土壤中AMF与根际细菌的正相互关系对作物氮磷利用有潜在促进作用,关键类群在有机质和养分贫乏的红壤中可能起到更重要的作用。     

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（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．     

土壤-植物-大气连续体系中氮的研究进展

氮素是植物最需要的重要养分元素之一.近年来,土壤-植物-大气这一连续体系(SPAC)中的氮循环成为研究的热点之一.大气中的氮素可以通过生物固定和N沉降等作用进入土壤和植物内,同时土壤和植物内的氮素又会以氨挥发和氮氧化物等方式排放到大气中.氮素通过生物固持和植物吸收等方式进入植物体内,植物器官脱落使植物损失一部分的氮素,另外雨水的淋洗和植物溢出液也会造成植物的N损失.植物氮素在植物体内的积累和分布随着生长时期和各营养器官而有所不同.另外,植物吸收氮素的过程又受到大气状况和土壤状况的制约.土壤中氮素经过矿化作用、硝化作用和反硝化作用进行转化,一部分把氮素转化成植物能吸收的营养形态,另一部分则从土壤中损失.凋落物的分解和N沉降能补充土壤中的氮素,而植物吸收、微生物固持、水文流失和N溢出等方式使氮素从土壤中损失出去.另外,凋落物的分解和根际土壤、CO2浓度和臭氧对氮素循环有着重要的作用.N污染、N沉降、碳氮循环的耦合作用是今后研究的热点问题.     

紫茎泽兰(Ageratina adenophora)入侵对土壤微生物群落和理化性质的影响

外来入侵植物与入侵地土壤微生物群落的互作关系是影响外来植物入侵力和生态系统可入侵性的一个重要领域。因此,研究外来植物入侵对入侵地土壤微生物群落及其理化性质的影响不仅可以全面地评估入侵植物对生态系统的影响,而且对于探索外来植物入侵的土壤微生物学机制尤为重要。采用磷脂脂肪酸(PLFAs)和传统培养相结合的方法研究了外来入侵植物紫茎泽兰对入侵地土壤微生物群落结构的影响;同时研究了紫茎泽兰入侵对11种土壤理化因子的影响。结果表明紫茎泽兰入侵改变了土壤微生物群落结构,提高了土壤自生固氮菌、氨氧化细菌和真菌的数量;同时,显著地提高了土壤的有效磷、速效钾、硝态氮、氨态氮和土壤有机碳含量,降低了土壤总钾含量和pH值。土壤微生物不同生理类群的变化与土壤中植物可直接吸收利用养分的变化显著相关。紫茎泽兰在入侵地成功定殖后,可能通过改变土壤微生物群落结构,特别是增加了与土壤养分循环相关的微生物功能类群数量,进而提高了土壤可利用的养分水平,创造对自身生长有利的土壤环境。紫茎泽兰改变土壤微生物群落是其入侵的一部分,这种改变进而加速了土壤养分循环,可能增强了紫茎泽兰的养分吸收,进而促进其生长、竞争和扩张。     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

土壤解磷微生物促进植物磷素吸收策略研究进展

随着全球磷肥需求增加和磷矿资源储量短缺矛盾逐渐加重,土壤解磷微生物在促进磷循环方面的重要性日益突显,因此有必要对其种类和促进植物磷素吸收策略进行全面梳理总结。荟萃分析了土壤解磷微生物种类并构建了系统进化树,重点论述了土壤解磷微生物促进植物磷素吸收主要策略。土壤解磷微生物主要通过矿化和溶解作用直接活化难溶性磷,但其也能与植物根系互作间接活化磷素。其间接途径主要包括解磷微生物与根系分泌物和丛枝菌根真菌互作,它们通过碳磷交换间接活化磷素;其次包括一些解磷微生物可以通过固氮作用使植物生长受到磷的限制,从而调节植物磷酸盐转运系统间接活化土壤磷;解磷微生物也能通过分泌植物激素和生物防治剂促进植物根系生长间接促进植物磷素吸收。解磷微生物还能通过磷素固定减少磷流失,也可以通过加速自身磷素周转促进植物磷素吸收。对完善和发展解磷微生物主导的土壤磷循环和植物磷素吸收利用理论体系具有重要意义。     

Assess suitability of hydroaeroponic culture to establish tripartite symbiosis between different AMF species, beans, and rhizobia

Background  

Like other species of the Phaseoleae tribe, common bean (Phaseolus vulgaris L.) has the potential to establish symbiosis with rhizobia and to fix the atmospheric dinitrogen (N₂) for its N nutrition. Common bean has also the potential to establish symbiosis with arbuscular mycorrhizal fungi (AMF) that improves the uptake of low mobile nutrients such as phosphorus, from the soil. Both rhizobial and mycorrhizal symbioses can act synergistically in benefits on plant.     

Effects of co-inoculation with arbuscular mycorrhizal fungi and rhizobia on soybean growth as related to root architecture and availability of N and P

Soybean plants can form tripartite symbiotic associations with rhizobia and arbuscular mycorrhizal (AM) fungi, but little is known about effects of co-inoculation with rhizobia and AM fungi on plant growth, or their relationships to root architecture as well as nitrogen (N) and phosphorus (P) availability. In the present study, two soybean genotypes contrasting in root architecture were grown in a field experiment to evaluate relationships among soybean root architecture, AMF colonization, and nodulation under natural conditions. Additionally, a soil pot experiment in greenhouse was conducted to investigate the effects of co-inoculation with rhizobia and AM fungi on soybean growth, and uptake of N and P. Our results indicated that there was a complementary relationship between root architecture and AMF colonization in the field. The deep root soybean genotype had greater AMF colonization at low P, but better nodulation with high P supply than the shallow root genotype. A synergistic relationship dependent on N and P status exists between rhizobia and AM fungi on soybean growth. Co-inoculation with rhizobia and AM fungi significantly increased soybean growth under low P and/or low N conditions as indicated by increased shoot dry weight, along with plant N and P content. There were no significant effects of inoculation under adequate N and P conditions. Furthermore, the effects of co-inoculation were related to root architecture. The deep root genotype, HN112, benefited more from co-inoculation than the shallow root genotype, HN89. Our results elucidate new insights into the relationship between rhizobia, AM fungi, and plant growth under limitation of multiple nutrients, and thereby provides a theoretical basis for application of co-inoculation in field-grown soybean.     

Arbuscular mycorrhizal fungi enhance P uptake and alter plant morphology in the invasive plant Microstegium vimineum

Invasive plant species can interact with native soil microbes in ways that change how they use nutrients and allocate biomass. To examine whether Microstegium vimineum form symbiotic associations with arbuscular mycorrhizal fungi (AMF) and whether AMF mediate nutrient acquisition and growth of the plant, we conducted a field survey in Raleigh, NC and Hangzhou, China and two experiments in growth chambers. This is the first report that M. vimineum is mycorrhizal, with colonization rates of 47 and 21 % in its native and invaded range, respectively. In the growth chamber, addition of an AMF inoculum mixture significantly promoted M. vimineum biomass accumulation in both field and sterilized soils, particularly after 64 days of growth. Arbuscular mycorrhizal fungi also increased plant phosphorous (P) uptake but did not consistently affect total plant nitrogen (N) acquisition, leading to decreases in plant N:P ratios. More interestingly, AMF significantly altered plant morphology, increasing the number of stolons and aerial roots per individual (59 and 723 %), aerial roots per gram aboveground biomass (374 %) and aerial roots per stolon (404 %). Our results suggest that mycorrhizal enhancement of plant growth by stimulating tillering may serve as another mechanism by which M. vimineum can quickly take over new territory. Future studies on invasive plant-microbial interactions are needed to understand the mechanisms through which microbes contribute to the competitive ability of invasive plants.     

Carbon for nutrient exchange between arbuscular mycorrhizal fungi and wheat varies according to cultivar and changes in atmospheric carbon dioxide concentration

Arbuscular mycorrhizal fungi (AMF) form symbioses with most crops, potentially improving their nutrient assimilation and growth. The effects of cultivar and atmospheric CO₂ concentration ([CO₂]) on wheat–AMF carbon‐for‐nutrient exchange remain critical knowledge gaps in the exploitation of AMF for future sustainable agricultural practices within the context of global climate change. We used stable and radioisotope tracers (¹⁵N, ³³P, ¹⁴C) to quantify AMF‐mediated nutrient uptake and fungal acquisition of plant carbon in three wheat (Triticum aestivum L.) cultivars. We grew plants under current ambient (440 ppm) and projected future atmospheric CO₂ concentrations (800 ppm). We found significant ¹⁵N transfer from fungus to plant in all cultivars, and cultivar‐specific differences in total N content. There was a trend for reduced N uptake under elevated atmospheric [CO₂]. Similarly, ³³P uptake via AMF was affected by cultivar and atmospheric [CO₂]. Total P uptake varied significantly among wheat cultivars and was greater at the future than current atmospheric [CO₂]. We found limited evidence of cultivar or atmospheric [CO₂] effects on plant‐fixed carbon transfer to the mycorrhizal fungi. Our results suggest that AMF will continue to provide a route for nutrient uptake by wheat in the future, despite predicted rises in atmospheric [CO₂]. Consideration should therefore be paid to cultivar‐specific AMF receptivity and function in the development of climate smart germplasm for the future.     

High effectiveness of exotic arbuscular mycorrhizal fungi is reflected in improved rhizobial symbiosis and trehalose turnover in Cajanus cajan genotypes grown under salinity stress

Arbuscular mycorrhizal fungi (AMF) improve functioning of legume-Rhizobium symbiosis under salinity. However, plant responses to mycorrhization vary depending on the plant and fungal species. The current study aimed to compare the effectiveness of a native inoculum from saline soil and two exotic isolates, Funneliformis mosseae and Rhizophagus irregularis on two Cajanus cajan (pigeonpea) genotypes (Paras, Pusa 2002) subjected to NaCl stress. Salinity depleted nodulation and nutrient status in both genotypes with higher negative effects in Paras. Although all AM fungi improved growth, R. irregularis performed better by promoting higher biomass accumulation, nodulation, N₂ fixation and N, P uptake which correlated with higher AM colonization. R. irregularis inoculated plants also accumulated higher trehalose in nodules due to decreased trehalase and increased trehalose-6-P synthase, trehalose-6-phosphatase activities. The results suggest that higher stability of R. irregularis-pigeonpea symbiosis under salt stress makes it an effective ameliorator for overcoming salt stress in pigeonpea.     

Mechanistic understanding of interspecific interaction between a C4 grass and a C3 legume via arbuscular mycorrhizal fungi,as influenced by soil phosphorus availability using a 13C and 15N dual-labelled organic patch

Arbuscular mycorrhizal fungi (AMF) can improve plant nutrient acquisition, either by directly supplying nutrients to plants or by promoting soil organic matter mineralization, thereby affecting interspecific plant relationships in natural communities. We examined the mechanism by which the addition of P affects interspecific interactions between a C4 grass (Bothriochloa ischaemum, a dominant species in natural grasslands) and a C3 legume (Lespedeza davurica, a subordinate species in natural grasslands) via AMF and plant growth, by continuous ¹³C and ¹⁵N labelling, combined with soil enzyme analyses. The results of ¹⁵N labelling revealed that P addition affected the shoot uptake of N via AMF by B. ischaemum and L. davurica differently. Specifically, the addition of P significantly increased the shoot uptake of N via AMF by B. ischaemum but significantly decreased that by L. davurica. Interspecific plant interactions via AMF significantly facilitated the plant N uptake via AMF by B. ischaemum but significantly inhibited that by L. davurica under P-limited soil conditions, whereas the opposite effect was observed in the case of excess P. This was consistent with the impact of interspecific plant interaction via AMF on arbuscular mycorrhizal (AM) benefit for plant growth. Our data indicate that the capability of plant N uptake via AMF is an important mechanism that influences interspecific relationships between C4 grasses and C3 legumes. Moreover, the effect of AMF on the activities of the soil enzymes responsible for N and P mineralization substantially contributed to the consequence of interspecific plant interaction via AMF for plant growth.     

Effects of arbuscular mycorrhizal fungi on photosynthesis,carbon content,and calorific value of black locust seedlings

Arbuscular mycorrhizal fungi (AMF) form symbioses with many plants. Black locust (Robinia pseudoacacia L.) is an important energy tree species that can associate with AMF. We investigated the effects of AMF (Rhizophagus irregularis and Glomus versiforme) on the growth, gas exchange, chlorophyll (Chl) fluorescence, carbon content, and calorific value of black locust seedlings in the greenhouse. The total biomass of the arbuscular mycorrhizal (AM) seedlings was 4 times greater than that of the nonmycorrhizal (NM) seedlings. AMF greatly promoted the photosynthesis of black locust seedlings. AM seedlings had a significantly greater leaf area, higher carboxylation efficiency, Chl content, and net photosynthetic rate (P _N) than NM seedlings. AMF also significantly increased the effective photochemical efficiency of PSII and significantly enhanced the carbon content and calorific value of black locust seedlings. Seedlings inoculated with G. versiforme had the largest leaf area and highest biomass, Chl content, P _N, and calorific value.     

接种AM真菌对玉米和油菜种间竞争及土壤无机磷组分的影响

在温室盆栽条件下,分别模拟单作、间作和尼龙网分隔种植,比较接种丛枝菌根(arbuscular mycorrhizal, AM)真菌Glomus intraradices和Glomus mosseae对菌根植物玉米和非菌根植物油菜生长和磷吸收状况的影响,并分析土壤中各无机磷组分的变化。结果发现,接种AM真菌可以促进土壤中难溶性磷(Ca₁₀-P和O-P)向有效态磷转化,并显著降低总无机磷含量 (P<0.05),显著提高菌根植物玉米的生物量和磷吸收量(P<0.05),特别是在间作体系中使玉米的磷营养竞争比率显著提高了45.0%-104.1% (P<0.05),显著降低了油菜的生物量和磷吸收量(P<0.05),从而增强了了菌根植物的竞争优势,降低了非菌根植物与菌根植物的共存能力。揭示了石灰性土壤中AM真菌对植物物种多样性的影响,有助于更加全面地理解AM真菌在农业生态系统中的作用。     

The role of arbuscular mycorrhizas in improving plant zinc nutrition under low soil zinc concentrations: a review

Many of the world’s soils are zinc (Zn) deficient. Consequently, many crops experience reduced growth, yield and tissue Zn concentrations. Reduced concentrations of Zn in the edible portions of crops have important implications for human Zn nutrition; this is a cause of global concern. Most terrestrial plant species form arbuscular mycorrhizas (AM) with a relatively limited number of specialized soil fungi. Arbuscular mycorrhizal fungi (AMF) can take up nutrients, including Zn, and transfer them to the plant, thereby enhancing plant nutrition. Under high soil Zn concentrations the formation of AM can also ‘protect’ against the accumulation of Zn in plant tissues to high concentrations. Here, a short review focusing on the role of AM in enhancing plant Zn nutrition, principally under low soil Zn concentrations, is presented. Effects of Zn on the colonisation of roots by AMF, direct uptake of Zn by AMF, the role of AM in the Zn nutrition of field grown plants, and emerging aspects of Zn molecular physiology of AM, are explored. Emergent knowledge gaps are identified and discussed in the context of potential future research.