丛枝菌根真菌在植物修复重金属污染土壤中的作用

菌根是真菌与植物根系所建立的互惠共生体,其中以丛枝菌根(AM)真菌在自然界中分布最广。在重金属污染条件下,AM真菌可以减轻重金属对植物的毒害,影响植物对重金属的吸收和转运,在重金属污染土壤的植物修复中显示出极大的应用潜力。文章通过讨论菌根植物对重金属修复的作用机制,提出菌根技术在重金属植物修复中应在通过广泛调查、筛选超积累植物的基础上,不断探索植物-菌根体系修复问题,以促进重金属污染土壤的生物修复。     

丛枝菌根对芘污染土壤修复及植物吸收的影响

采用温室盆栽试验方法,研究了两种丛枝菌根真菌Glomus mosseae和 Glomus etunicatum对三叶草(Trifolium subterraneum L.)和辣椒（Capsicum annuum L.）修复芘污染土壤的影响。供试土样中芘初始浓度为0 ~ 75.18 mg/kg。结果表明,接种AMF可促进供试植物对土壤中芘的吸收,并且显著提高三叶草根的芘含量、根系富集系数、根和茎叶的芘积累量,但对辣椒根和茎叶芘含量、根系富集系数的影响不显著,这主要与植物的菌根侵染率和“菌根依赖度”不同有关。接种AMF土壤中芘的削减率高于普通植物修复,但植物吸收积累对修复的贡献率小于0.2%;因此推测,AM作用下良好的根际环境对土壤微生物数量和活性的提高、进而对土壤中芘降解的促进可能是菌根修复的主要机理。     

丛枝菌根与土壤修复

菌根是真菌与植物根系所建立的互惠共生体, 其中以丛枝菌根在自然界中分布最广。近年来,随着菌根研究的发展,丛枝菌根在土壤修复中的应用日益受到人们的关注。本文综述了丛枝菌根在土壤重金属污染、有机污染、放射污染以及土壤退化修复中的作用,并对当前研究中存在的问题和未来发展前景作了探讨。     

砷污染土壤中接种丛枝菌根真菌和蚯蚓对土壤线虫群落和植物砷吸收的影响

The influences of arbuscular mycorrhizal fungi (MF, Acaulospora spp. and Glomus spp.), rice straw and earthworms (RE, Eisenia foetida) on nematode communities and arsenic (As) uptake by maize (Zea mays L.) in As-contaminated soils were examined in a field experiment conducted in Wujiang, Jiangsu Province, China. The experiment was designed as a 2 × 2 factorial with the factors of MF (inoculated or uninoculated) and RE (added or not added). The results demonstrated that MF inoculation led to significantly higher root colonization of MF and root dry weight. Plants inoculated with both MF and RE had the highest As concentrations in root. The number of total nematodes increased with MF inoculation when RE was absent, and decreased with RE addition when MF was inoculated. The improved abundance of nematodes with the MF treatment implied that the tested MF acted as food sources for fungivores. The abundances of omnivores-predators and plant parasites were reduced by earthworm activity. Twenty-seven genera of nematodes were identified, with Filenchus dominant in all treatments. Trophic diversity (TD), Shannon-Weaver diversity (H′), Simpson dominance index (λ), and species richness (SR) indicated higher species diversity, more proportionate species composition, evenly distributed species, and more food sources in the MF, RE, and their interaction treatments. Maturity index (MI) showed a moderately disturbed environment due to As pollution. Besides enhancing plant uptake of contaminants, MF and RE amendments could also improve soil health by restoring the structure of soil communities, as reflected by more stable nematode community structure.     

丛枝菌根真菌对西藏高原草地植物和土壤环境的影响

采用三室隔网装置，就Glomus etunicatum、Glomus intraradices、Glomus mossecte对2种高山草地植物和土壤环境的影响进行了研究。结果表明：（1）接种AM真菌对草地植物的侵染和生长均具显著效应，植株地上部、根系干物重以及含磷量、吸磷量均显著高于不接种处理，菌根菌丝对植株吸磷的贡献率达47.8％-69．5％。其中，Glomus intraradices、Glomus mosseae分别对穗序剪股颖、紫羊毛吸收土壤磷索更具促进作用。（2）各接种处理中室土壤中各类微生物数量均显著高于边室土壤，但边室土壤中放线菌，特别是细菌、真菌的平均增幅均远高于中室土壤，表明菌根际、菌丝际土壤中各类微生物的数量差异趋于明显缩小，微生物区系构成得以平衡与改善；解磷细菌（芽孢杆菌）亦呈同一趋势。（3）2—1 mm粒径团聚体在土壤水稳性团聚体构成中占有绝对比重，菌根菌丝对距根表不同距离处2—1 mm团聚体形成的贡献率均在70％以上，但距根表2—4cm处菌根菌丝贡献率明显低于0-2、4-6cm处，并未表现出随菌丝密度增加而提高的趋势；5—2mm水稳性团聚体仅距根表较远处有少量形成（菌丝贡献率达100％），0-2、2—4cm处则未见分布。（4）同一、不同AM真菌对不同或同一草地植物的侵染及所产生的菌根效应具有不同程度的差异，穗序剪股颖各接种处理普遍优于紫羊毛，紫羊毛＋Glomus mosseae、穗序剪股颖＋Glomus intraradices优于同组其他接种处理的趋势较为明显。     

丛枝菌根真菌促进植物摄取土壤磷的作用机制

磷在土壤中易被固定沉淀,在植物磷利用率低的情况下,过度施肥会造成磷肥浪费,可能通过地表径流、地下水溶解等方式,造成水体富营养化产生面源污染,对人类生产生活造成较大影响。丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)和植物结合所形成的共生菌根可以显著增强植株对磷的吸收利用。通过AMF可以提高宿主植株对磷的吸收转运的特性,从AMF促进植株对磷元素的摄取机制、AMF促进植物磷摄取分子机理、AMF作用下根系分泌物对植株磷利用的影响与根际微生物对AMF磷元素利用的影响4个方面的研究进展进行分析总结。AMF可以通过改变宿主植株的根系形态和菌丝网络的形成,扩大植株对养分吸收范围;释放有机酸、磷酸酶和质子等根系分泌物改变土壤结构和理化性质,与根际微生物共同作用降解土壤中难溶性磷酸盐;诱导相关磷转运蛋白基因的特异性表达,提高植株对磷的转运能力而促进其吸收。     

丛枝菌根真菌对紫花苜蓿吸收土壤中镉和锌的影响

采用温室盆栽试验方法,研究了镉（Cd）、锌（Zn）污染土壤中,8种不同丛枝菌根真菌（AMF）Glomus lamellosum（G.la）、Acaulospora mellea（A.m）、Glomus mosseae（G.m）、Glomus intraradices（G.i）、Glomus etunicatum（G.e）、Glomus constrictum（G.c）、Diversispora spurcum（D.s）、Glomus aggregatum（G.a）对紫花苜蓿（Medicagosativa L.）吸收Cd、Zn的影响。结果表明,Cd、Zn污染下AMF仍然明显侵染紫花苜蓿,并促进紫花苜蓿对Cd、Zn的吸收积累,但不同AMF影响的效应和植株不同部位对重金属的吸收积累规律存在差异。AMF处理下紫花苜蓿根部Cd、Zn含量和积累量明显增加,但地上部Cd、Zn的含量则降低,地上部Zn的积累量也减小,这表明AMF处理减弱了Cd、Zn由根部向地上部的运移,减轻了植物地上部毒害。接种AMF条件下,植株尤其是根部生物量增加是Cd、Zn在其体内含量和积累量增加的重要因素,不同种类AMF促进植株生物量增加的幅度不同,导致植株对Cd、Zn的积累和抗性存在差异。     

丛枝菌根真菌(Glomus caledonium)对铜污染土壤生物修复机理初探

利用盆栽试验,研究了丛枝菌根真菌(Glomus.caledonium)在不同程度铜污染土壤上对玉米苗期生长的影响。结果表明,即使在土壤施铜量达150mg/kg时,菌根真菌对玉米仍有近55%的侵染率;接种菌根真菌,能显著促进玉米根系的生长。菌根玉米的根系生物量和根系长度,平均较未接种处理分别提高108.4%和58.8%;接种处理的植株地上部生物量达到每盆(3株)10.58g,显著高于不施铜的非菌根玉米。这些结果表明,丛枝菌根真菌对铜污染具有较好的抗性;并且由于菌根的形成,使宿主植物明显地改善了对磷的吸收和运输,并能通过抑制土壤酸化、降低土壤可溶态铜的浓度等机制,增强宿主植物对铜污染的抗(耐)性。在150mg/kg施铜水平时,与非菌根玉米相比,菌根玉米地上部和根系铜浓度分别降低24.3%和24.1%,吸铜量分别提高了28.2%和60.0%,表明菌根植物对铜污染土壤具有一定的生物修复作用。     

贵州务川汞矿区土壤丛枝菌根真菌多样性研究

对贵州务川汞矿区不同水平汞污染植物根际土壤丛枝菌根(Arbuscular Mycorrhizal,AM)真菌资源分布及多样性进行分析，结果发现：本研究选取的务川汞矿区3个样地土壤汞含量在94.1～268.0 mg/kg，远高于国家规定的土壤汞含量，汞污染极其严重。随着汞含量增加，AM真菌侵染率、孢子密度显著降低，AM真菌群落丰富度、多样性及均匀度均有所降低，但差异未达显著水平，而仍有部分对汞污染耐受性较强的AM真菌有较高的分布。另外，管柄囊霉属AM真菌在5个样品中均有较高丰度，球囊霉属AM真菌在火棘根际的相对丰度随着汞含量升高而升高，因此，火棘–管柄囊霉属AM真菌或火棘–球囊霉属AM真菌联合进行汞污染修复，有一定的应用价值。     

Molecular diversity of arbuscular mycorrhizal fungi associated with an Mn hyperaccumulator—Phytolacca americana,in Mn mining area

Arbuscular mycorrhizal fungi (AMF) have great potential for assisting metal-hyperaccumulating plants in the remediation of contaminated soils. However, little information is available about the symbiosis and community composition of AMF associated with manganese (Mn) hyperaccumulator, such as Phytolacca americana, growing on Mn-contaminated soils under natural conditions. Therefore, the objective of this study was to analyze AMF diversity and community composition in P. americana roots growing at an Mn mining site. Molecular techniques were used to analyze AMF community composition and phylogenetic relationship in P. americana roots sampled from three Mn mine spoils and one adjacent reference areas. Results obtained showed that mycorrhizal symbionts successfully established even in the most heavily Mn-polluted sites. Root colonization and AMF diversity were significantly negatively correlated with total and extractable Mn concentrations. Principal component analysis (PCA) revealed that Mn contamination impacted AMF diversity, and shaped AMF community structure. Phylogenetic analyses demonstrated that all species were affiliated with Glomus, suggesting that Glomus was the dominant genus in this AMF community. Some unique sequences that occurred exclusively in heavily polluted sites associated with P. americana may belong to symbiotic fungi with great potential for improving the phytoremediation efficiency of Mn-contaminated soils.     

N,P Contribution and soil adaptability of four arbuscular mycorrhizal fungi

Abstract

A glasshouse study was conducted to investigate the symbiotic efficiency and soil adaptability of four AMF using glass-bead cultivation systems. The results showed that efficiency and adaptability of four fungi varied among three soils. Particularly, efficiency of BEG167 shifted from positive in Beijing soil to negative in Guangdong soil. Furthermore, BEG167 had high adaptability in all three soils. Intraspecific differences of BRG168 and BEG221 were found in efficiency and adaptability in three soils. Taking efficiency and adaptabilty into consideration, it was concluded that BEG167, BEG168 and BEG221 in Beijing soil, BEG151 in Hubei soil, and BEG151 and BEG168 in Guangdong soil were effective AMF for maize.     

AM真菌对桑树根围土壤团聚体的影响机制

为揭示丛枝菌根 (Arbuscular mycorrhizal,AM)真菌对植桑土壤的影响及机制,采用盆栽试验研究接种摩西管柄囊霉 (Funneliformis mosseae,Fm)和根内根生囊霉 (Rhizophagus intraradices,Ri)对土壤有机碳(Soil organic carbon, SOC)、球囊霉素相关土壤蛋白 (Glomalin related soil protein, GRSP)及团聚体组成与稳定性的影响。结果表明：⑴ 接种Ri显著增加土壤大团聚体百分比,并提高平均质量直径 (Mean weight diameter, MWD)和几何平均直径 (Geometric mean diameter, GMD)、显著降低团聚体破坏率 (Percentage of aggregate destruction, PAD)。⑵ 接种Fm和Ri均显著增加微团聚体SOC含量,接种Fm显著降低大团聚体总GRSP含量,而接种Ri却显著增加大团聚体和微团聚体总GRSP含量及易提取GRSP含量。⑶ 接种AM真菌对整体SOC的效应为负,土壤总GRSP对SOC占比在25.5%~76.5%之间,土壤易提取GRSP对SOC占比在4.87%~5.93%之间,且Ri的接种效应高于Fm。⑷ 总GRSP、易提取GRSP和SOC对团聚体组成表现均为正向显著影响,其中易提取GRSP是主要驱动因子,而总GRSP是土壤团聚体稳定性的主要影响因子。综上,AM真菌作用下桑树根围土壤团聚体得以改善并趋于稳定,Ri的接种效应明显大于Fm;土壤团聚体的形成主要依赖易提取GRSP,而其稳定性主要受总GRSP影响。     

客土改良技术及其在砷污染土壤修复中的应用展望

砷是一种毒性很强且对人体健康威胁很大的金属元素,土壤砷污染与修复长期以来受到各国政府和科学家的广泛关注。客土改良技术是污染土壤修复中较常用和有效的方法之一,近年来逐渐受到关注。本文在简述土壤中砷来源及其危害的基础上,重点对近年来该技术在土壤改良及污染土壤修复中的应用进行了系统整理,并以此为基础,比较了砷污染土壤的物理、化学及生物修复的效果。结合我国部分地区耕地砷污染较严重的现状,认为客土改良技术见效快、改良较彻底、且具有较高的实用性。同时,论文还对客土技术在砷污染土壤修复中的研究重点及应用前景等进行了展望。     

Arbuscular mycorrhizal fungi reduce decomposition of woody plant litter while increasing soil aggregation

The decomposition of plant organic matter and the stability of soil aggregates are important components of soil carbon cycling, and the relationship between decomposition rate and arbuscular mycorrhizal fungi (AMF) has recently received considerable attention. The interaction of AMF with their associated microorganisms and the consequences for litter decomposition and soil aggregation still remain fairly unclear. In a laboratory pot experiment we simultaneously tested the single and combined effects of one AMF species (Rhizophagus irregularis) and a natural non-AMF microbial community on the decomposition of small wooden sticks and on soil aggregation. To disentangle effects of hyphae and roots we placed mesh bags as root exclusion compartments in the soil. The decomposition of the wooden sticks in this compartment was significantly reduced in the presence of AMF, but not with the non-AMF microbial community only, compared to the control, while aggregation was increased in all treatments compared to the control. We suggest that AMF directly (via localized nutrient removal or altered moisture conditions) or indirectly (by providing an alternative carbon source) inhibited the activity of decomposers, leading to different levels of plant litter degradation under our experimental settings. Reduced decomposition of woody litter in presence of AMF can be important for nutrient cycling in AMF-dominated forests and in the case of woody plants and perennials that develop lignified roots in grasslands.     

Dual plant host effects on two arbuscular mycorrhizal fungi

Mycorrhizal fungi may simultaneously associate with multiple plant hosts, and the implications of this for the fungi involved are not well understood. To address this question, two arbuscular mycorrhizal fungi (AMF), Glomus clairoideum (a treatment referred to as “Glo”) and Scutellospora fulgida (a treatment referred to as “Scut”), were grown separately in pots that each consisted of two plant compartments separated by a root-free-compartment (RFC). Fungi within each two-plant-compartment pot were exposed to either two individuals of indiangrass (Sorghastrum nutans), two individuals of big bluestem (Andropogon gerardii), or one of each. A non-inoculated treatment (“Non”) was included to help gauge the potential influence of greenhouse contaminant fungi, cross-contamination, or any misidentification of non-AMF hyphae. The two host species had additive effects on the growth of AM hyphae in plant compartments of Scut, Glo, and Non pots, and in the RFCs of Scut pots. In Glo RFCs, however, they were antagonistic in their effects. Synergism between hosts in Non RFCs suggested that any potential contaminants or misidentification could not explain this result. Underyielding was not seen in shoot weight, root weight, or root length in dual host pots, and also therefore could not explain the result. Hyphal growth in the Scut treatment was evenly distributed between the RFC and plant compartments (or marginally skewed toward the RFC), while hyphal growth in the Glo treatment was skewed toward plant compartments (nearer roots). However, hyphal lengths were more highly correlated across plant compartments within a common pot in the Glo treatment, suggesting that this AMF bridged the RFC to experience the entire two-host pot as a single environment to a greater extent than Scut did. These AMF differed in how they responded to both the species composition of the two-host environment and its spatial structure; potential implications for mycorrhizal community dynamics are discussed.     

Community structure of arbuscular mycorrhizal fungi associated with Robinia pseudoacacia in uncontaminated and heavy metal contaminated soils

The significance of arbuscular mycorrhizal fungi (AMF) in soil remediation has been widely recognized because of their ability to promote plant growth and increase phytoremediation efficiency in heavy metal (HM) polluted soils by improving plant nutrient absorption and by influencing the fate of the metals in the plant and soil. However, the symbiotic functions of AMF in remediation of polluted soils depend on plant–fungus–soil combinations and are greatly influenced by environmental conditions. To better understand the adaptation of plants and the related mycorrhizae to extreme environmental conditions, AMF colonization, spore density and community structure were analyzed in roots or rhizosphere soils of Robinia pseudoacacia. Mycorrhization was compared between uncontaminated soil and heavy metal contaminated soil from a lead–zinc mining region of northwest China. Samples were analyzed by restriction fragment length polymorphism (RFLP) screening with AMF-specific primers (NS31 and AM1), and sequencing of rRNA small subunit (SSU). The phylogenetic analysis revealed 28 AMF group types, including six AMF families: Glomeraceae, Claroideoglomeraceae, Diversisporaceae, Acaulosporaceae, Pacisporaceae, and Gigasporaceae. Of all AMF group types, six (21%) were detected based on spore samples alone, four (14%) based on root samples alone, and five (18%) based on samples from root, soil and spore. Glo9 (Rhizophagus intraradices), Glo17 (Funneliformis mosseae) and Acau3 (Acaulospora sp.) were the three most abundant AMF group types in the current study. Soil Pb and Zn concentrations, pH, organic matter content, and phosphorus levels all showed significant correlations with the AMF species compositions in root and soil samples. Overall, the uncontaminated sites had higher species diversity than sites with heavy metal contamination. The study highlights the effects of different soil chemical parameters on AMF colonization, spore density and community structure in contaminated and uncontaminated sites. The tolerant AMF species isolated and identified from this study have potential for application in phytoremediation of heavy metal contaminated areas.     

Microbiota accompanying different arbuscular mycorrhizal fungal isolates influence soil aggregation

Arbuscular mycorrhizal fungi (AMF) are key components of ecosystems through their influence on plant communities and ecosystem processes. A major source of information regarding the importance of AMF species richness on process rates are mesocosm experiments using different levels of diversity of AMF as provided by single-species cultures of AMF. Since AMF inocula are generally made available in the form of non-sterile pot culture material, it is possible that AMF symbiosis-associated microbiota are at least partially responsible for some effects hitherto directly attributed to the AMF mycelium. Here, we provide evidence that microbiota associated with single-species cultures of AMF (after long-term pot culture enrichment of 7–8 years) can strongly affect the ecosystem process of soil aggregation. This effect occurred in an AMF isolate specific manner, but in the absence of live and active AMF mycelium. We additionally documented large differences in microbiota communities associated with the different AMF inocula (using PLFA analyses), suggesting that these differences were at least partly responsible for the observed changes in soil aggregation. This result points to AMF–microbiota interactions as a largely unexplored mechanism underlying soil aggregation (and potentially other ecosystem processes). We suggest that a reinterpretation of previous experiments using greenhouse-derived AMF cultures may be necessary, and the need to consider AMF symbiosis-associated microbiota in mechanistic studies of AMF and mycorrhizae in general is emphasized.     

丛枝菌根与植物寄生性线虫相互作用及抗性机制

丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)能够与大多数陆地植物互惠共生,促进植物对养分的吸收,提高植物对各种生物和非生物胁迫的抗逆性,对植物健康生长有重要的作用。在土壤中丛枝菌根真菌与植物寄生性线虫共同依靠寄主植物根系完成生命循环,但二者对寄主植物作用完全相反,引起研究者广泛兴趣,成为菌根研究的热点和焦点之一。本文分析了丛植菌根真菌与植物寄生线虫的相互作用,并探讨了菌根提高植物对线虫抗性的可能机制:菌根真菌改善植物的生长和营养状况、改变植物根系形态结构、影响根系分泌物和根际微生物区系、诱导寄主植物产生防御反应等,旨在深入挖掘丛枝菌根真菌的生物学功能,进一步发挥其在农业生产中的应用潜力。     

AM菌对三叶草吸收、累积重金属的影响

采用4室根箱培养系统,探讨了Cu、Zn、Pb、Cd 4种重金属复合污染土壤中,丛枝菌根菌对三叶草生长及吸收、累积重金属的作用,结果表明:重金属Cu 100mg/kg、Zn 600mg/kg、Pb 300mg/kg、Cd 10mg/kg的复合污染对三叶草生物量影响较小,但土壤重金属处理使丛枝菌根菌Glomus intraradices和Glomus caledonium对三叶草的侵染率分别降低53%和56%,菌种G.intraradice的菌丝密度降低73%;接种菌根真菌能明显减少重金属复合污染土壤中三叶草对Cu、Cd和Pb的吸收,并强化根系在限制重金属Pb和Cd向地上部运输中的作用,地上部Pb和Cd含量分别下降24.2%~55.3%和65%~97.9%,使三叶草地上部Cd和Pb含量均低于我国牧草重金属安全含量,提高了三叶草可食部分的质量;不同菌根真菌对三叶草吸收、累积及分配重金属的影响有明显差异,Glomus intraradices对减少三叶草对重金属的吸收及其在地上部可食部分的累积的作用大于Glomus caledonium。丛枝菌根菌对于强化三叶草根系对重金属的固持作用,调节生态系统中重金属的生物循环,减轻重金属对食物链的污染风险方面起着重要作用。