Arbuscular Mycorrhizal Fungi in Plant Production of Temperate Agroecosystems

Abstract

The importance of arbuscular fungi for plant development and health is now widely demonstrated. However, although it is more and more evident that they are not only an integral part of many cultivated plants but also an essential component of soil fertility, their rational use in plant production is still in its infancy. Because of their role as bioregulators, biofertilizers, and biocontrol agents, they represent potentially important tools for new orientations in agriculture, particularly in Europe, where there is increasing demand for development of new plant management techniques that are less dependent on chemical inputs. The discovery of mycmutants turned out to be an excellent tool for better understanding the ecophysiology of arbuscular mycorrhizas under field conditions and for allowing considerable progress in our knowledge on the genes controlling this symbiosis. Progress in this area, together with ongoing generation of specific nucleic acid probes for arbuscular mycorrhizal fungi, appear to be essential for promoting mycorrhizal biotechnology. Despite this, arbuscular mycorrhizal fungi can already be exploited successfully in certain areas of plant production, such as orchards and ornamental nurseries.     

生物因子对AM真菌多样性的影响

丛枝菌根真菌(Arbuscular Mycorrhizas,AM）真菌分布于自然界各生态系统中,生态因子对AM真菌多样性个有举足轻重的影响,其中动物,植物,微生物,人为因素等生物因子的作用,亦日益受到人们的关注,通过介绍该领域最近10年来的研究成果,探讨和分析当前研究中所存在的问题和动向。     

Phytoprotective Effect of Arbuscular Mycorrhizal Fungi Species Against Arsenic Toxicity in Tropical Leguminous Species

Arbuscular mycorrhizal fungi (AMF) improve the tolerance of hosting plants to arsenic (As) in contaminated soils. This work assessed the phytoprotective effect of Glomus etunicatum, Acaulospora morrowiae, Gigaspora gigantea, and Acaulospora sp. on four leguminous species (Acacia mangium, Crotalaria juncea, Enterolobium contortisiliquum, and Stizolobium aterrimum) in an As-contaminated soil from a gold mining area. AMF root colonization, biomass production, As and P accumulation, as well as arsenic translocation index (TI) from roots to shoots were measured. The AMF phytoprotective effect was assessed by the P/As ratio and the activity of plant antioxidant enzymes. The AMF colonization ranged from 24 to 28%. In general, all leguminous species had low As TI when inoculated with AMF species. Inoculation of C. juncea with Acaulospora sp. improved significantly As accumulation in roots, and decreased the activity of ascorbate peroxidase (APX) and superoxide dismutase (SOD), highlighting its phytoprotective effect and the potential use of this symbiosis for phytoremediation of As-contaminated soils. However, S. aterrimum has also shown a potential for phytoremediation irrespectively of AMF inoculation. APX was a good indicator of the phytoprotective effect against As contamination in C. juncea and A. mangium. In general P/As ratio in shoots was the best indicator of the phytoprotective effect of all AMF species in all plant species.     

Global Distributions of Arbuscular Mycorrhizal Fungi

We examined potential large-scale controls over the distribution of arbuscular mycorrhizal (AM) fungi and their host plants. Specifically, we tested the hypothesis that AM fungi should be more prevalent in biomes where nutrients are primarily present in mineral, and not organic, forms. Values of percentage root length colonized (%RLC) by AM fungi, AM abundance, and host plant availability were compiled or calculated from published studies to determine biome-level means. Altogether, 151 geographic locations and nine biomes were represented. Percent RLC differed marginally significantly among biomes and was greatest in savannas. AM abundance (defined as total standing root length colonized by AM fungi) varied 63-fold, with lowest values in boreal forests and highest values in temperate grasslands. Biomes did not differ significantly in the percentage of plant species that host AM fungi, averaging 75%. Contrary to the hypothesis, %RLC, AM abundance, and host plant availability were not related to the size, influx, or turnover rate of soil organic matter pools. Instead, AM abundance was positively correlated with standing stocks of fine roots. The global pool of AM biomass within roots might approach 1.4 Pg dry weight. We note that regions harboring the largest stocks of AM fungi are also particularly vulnerable to anthropogenic nitrogen deposition, which could potentially alter global distributions of AM fungi in the near future.     

The Ecology of Arbuscular Mycorrhizal Fungi

Arbuscular mycorrhiza is a mutually beneficial biological association between species in the fungal phylum Glomeromycota and higher plants roots. The symbiosis is thought to have afforded green plants the opportunity to invade dry land ca 450 Ma ago and the vast majority of extant terrestrial plants retain this association. Arbuscular mycorrhizal (AM) fungi perform various ecological functions in exchange for host photosynthetic carbon that almost always contribute to the fitness of hosts from an individual to community level. Recent AM fungal research, increasingly delving into the ‘Black Box’, suggests that species in this phylum may play a key facilitative role in below-ground micro- and meso-organism community dynamics, even more perhaps, that of a bioengineer. The ubiquitous nature of the symbiosis in extant flora and the fact that variations from the AM symbiosis are recent events suggest that Glomeromycota and plant roots coevolved. This review considers aspects of AM fungal ecology emphasizing past and present importance of the phylum in niche to global ecosystem function. Nutrient exchange, evolution, taxonomy, phenology, below-ground microbial interaction, propagule dissemination, invasive plants interactions, the potential role in phytoremediation and some of the factors affecting AM fungal biology are discussed. We conclude that it is essential to include AM association in any study of higher plants in natural environments in order to provide an holistic understanding of ecosystems.     

丛枝菌根真菌对植物生长影响的研究

菌根是自然界中一种极为普遍和重要的共生现象,其中分布最为广泛的菌根类型就是丛枝菌根,可以增强植物从土壤中获取水分的能力,改善植物根系对磷、镉等矿质元素及养分的吸收,从而促进植物的生长。本文综述了丛枝菌根真菌对植物生长影响的概况。有关丛枝菌根真菌对植物水分和矿质营养的利用,尤其是磷素营养的研究较为深入,而对植物光合特性的研究较少,这些研究工作为深入理解菌根真菌与植物的相互关系提供基础资料。     

丛枝菌根真菌分类最新进展

近10a来,随着分子生物学技术在丛枝菌根(arbuscularmycorrhiza,AM)研究中的应用,AM真菌分类学得到迅速发展。重点介绍AM真菌的最新分类系统,并对其发展简史作一简单回顾。     

不同生境下川麦冬根围土壤丛枝菌根真菌多样性北大核心CSCD

采用高通量(Illumina Miseq)测序技术对栽培和野生2种生境下川麦冬根围的丛枝菌根(AM)真菌多样性和群落结构进行测定,并结合土壤理化因子进行相关性分析,以明确两种生境下川麦冬根围土壤AM真菌多样性和优势群落的分布特点,探讨AM真菌群落分布差异的驱动因子,为AM真菌应用于麦冬生产提供理论依据和技术支持。结果表明:(1)不同生境下川麦冬根围土壤中共鉴定出AM真菌3属10种,其中野生川麦冬根围土壤鉴定出的AM真菌3属7种,分别隶属于无梗囊霉属(Acaulospora)、多孢囊霉属(Diversispora)和球囊霉属(Glomus),而栽培环境下鉴定出AM真菌1属6种,隶属于球囊霉属。2个生境优势属均为球囊霉属。(2)不同生境下川麦冬根围AM真菌之间存在显著差异,野生生境下川麦冬根围土壤AM真菌多样性指数ACE和Shannon均显著高于人工栽培生境,而Simpson指数则相反。(3)相关性分析表明,AM真菌多样性指数及群落组成结构均与土壤理化因子存在相关性,其中全钾(TK)、全磷(TP)、全氮(TN)对AM真菌多样性指数和群落结构组成均存在显著影响。研究认为,不同生境下川麦冬根围AM真菌群落存在显著差异,球囊霉属为川麦冬互利共生的关键属,TK、TP、TN是不同生境川麦冬根围AM真菌群落差异的主要驱动因子。     

石油污染土壤中不同人工林木根际丛枝菌根真菌与球囊霉素的研究

以陕西延长县石油污染区常见的13种人工种植林木为材料,测定了各人工种植林木根际丛枝菌根(AM)真菌发育状况、污染土壤的理化性质、土壤酶活性和球囊霉素含量,探讨AM真菌在石油污染土壤生态修复中的作用。结果表明,13种林木均能形成AM,其定殖率平均为63.2%,孢子密度平均为1.93个.g-1干土,其中受污染程度最低的柠条AM真菌定殖率和孢子密度最高,分别为91.6%和4.73个.g-1干土;毛白杨、狼牙刺和刺槐的根际土壤养分(有机碳、碱解氮、速效磷)含量相对较高;各种人工种植林木的根际土壤球囊霉素含量、多酚氧化酶和过氧化氢酶活性随根际土壤石油烃污染浓度的增加而明显升高,其中刺槐、狼牙刺和酸枣根际土壤的过氧化氢酶和多酚氧化酶活性都较高,同时这3种林木的球囊霉素含量也较高。因此,林木根际土壤球囊霉素含量、多酚氧化酶和过氧化氢酶活性可以作为石油污染的敏感指标。     

长期定位施肥土壤中的丛枝菌根真菌

从连续26年长期定位施肥土壤中的玉米和小麦根围采集土样,从中鉴定出丛枝茵根(Arbuscular Myc- orrguza,AM)真菌5属19种,包括Glomus属11种:G．caledonium,G．constrictum,G．eburneum,G．ebunicatum, G．hyderabadensis,G．mossese,G．sinuosum,G．tortuosum,G．versiforme,G．sp1,G．sp2;Acaulospora属2种: A．excacota,A．sp;Gigaspora属2种:Gi．margarita,Gi．sp;Scutellospora属3种:S．cerradensis,S．coralloidea, S．pellucida;Archaespora属1种:Ar．leptoticha。     

云南松幼苗上红菇类菌根真菌的物种多样性及其菌根形态

以滇中1～2年生云南松幼苗为研究对象,观察鉴定与其共生的红菇属真菌外生菌根。形态观察发现了6种形态型（morphotypes）。本文对这6种形态型的外观和显微特征进行了详细描述,尤其强调了菌套形态特征。对rDNAITS片段比对分析表明,6种形态型对应6种红菇属真菌,它们分别是与Russula li-vescens、R.violeipes、R.densifolia、R.nigricans、R.sanguinea及R.nauseosa相近的红菇种类。本研究表明,形成的菌根及其菌套和囊状体的特征在红菇属真菌的系统分支间表现出较为稳定的差异。这一研究与前人对该属真菌的菌根形态及分类学研究基本吻合。红菇属真菌是云南松1～2年幼苗期的主要共生真菌类群。     

Species Diversity of Arbuscular Mycorrhizal Fungi in the Rhizosphere of <i>Hevea brasiliensis</i> in Hainan Island,China

Hevea brasiliensis is one of the important economic trees with a great economic value for natural rubber production. Symbiosis between roots of H. brasiliensis and arbuscular mycorrhizal fungi (AMF) is widely recognized, and can provide a range of benefits for both of them. Hainan Island harbors is one of the largest plantations of H. brasiliensis in China, whereas the information regarding the diversity of AMF in the rhizosphere of H. brasiliensis on this island is scarce. The diversity of AMF species in the rhizosphere of rubber tree plantations in Hainan was investigated in this study. A total of 72 soil samples from the rhizosphere of H. brasiliensis RY7-33-97 were collected. These included 48 samples from plantations in 11 cities or counties that had been planted for 15–25 years, and 24 samples from a demonstrating plantation site of the China National Rubber Tree Germplasm Repository representing plantations with tree plantation ages from one to 40 year-old. Collectively, a total of 68 morphotypes of AMF, belonging to the genera of Archaeospora (1), Glomus (43), Acaulospora (18), Entrophospora (3), Scutellospora (2), and Gigaspora (1) were isolated and identified, as per morphological characteristics of spores presented in the collected soil samples. Glomus (Frequency, F = 100%) and Acaulospora (F = 100%) were the predominant genera, and A. mellea (F = 63.9%) and A. scrobiculata (F = 63.9%) were the predominant species. AMF species differed significantly among collected sites in spore density (SD, 290.7–2,186.7 spores per 100 g dry soil), species richness (SR, 4.3–12.3), and Shannon-Weiner index of diversity (H, 1.24–2.24). SD was negatively correlated with available phosphorus level in the soil; SR was positively correlated with soil total phosphorus content; and H was positively correlated with levels of soil organic matter and total phosphorus. Similarly, SD, SR, and H were also correlated with H. brasiliensis plantation age, and an increasing trend was observed up to 40 years. These results suggest that the AMF community was complex and ubiquitous in the island plantation ecosystems of H. brasiliensis, with high species abundance and diversity. Soil factors and plantation age dramatically affected AMF diversity at species level.     

3种杓兰属植物菌根真菌系统发育和多样性分析

兰科植物菌根真菌(Orchid mycorrhizal fungi, OrMF)在兰科植物种子萌发和后续生长发育过程中具有重要作用。该研究采用培养(菌丝团分离)和非培养(克隆文库)2种方法获得同一栖息地3种不同杓兰属植物根中菌根真菌ITS序列并划分可操作分类单元(Operational taxonomic units, OTUs),分析其系统发育关系和多样性。结果表明:(1)所有根段中都有菌丝团定植,共分离出菌根真菌64株,其中63株为胶膜菌科(Tulasnellaceae)真菌,1株为角担菌科(Ceratobasidiaceae)真菌;可划分为7个OUT,每个OTU代表菌株的菌丝都能形成OrMF典型的近球形或椭球形链状排列的念珠状细胞;分离出来的菌根真菌均为无性型菌丝且不产生无性孢子。(2) 非培养法得到的3种杓兰属植物的根中OrMF分别隶属于胶膜菌科(Tulasnellaceae),腊壳菌科(Sebacinaceae)、角担菌科(Ceratobasidiaceae)和革菌科(Thelephoraceae),其中胶膜菌科OTU在种类和数量上占有绝对优势,培养和非培养2种方法得到的OrMF OTU类型和数量均为西藏杓兰(Cypripedium tibeticum)>无苞杓兰(C. flavum)>黄花杓兰(C. bardolphianum),但培养法少于非培养法。(3)对胶膜菌进行系统发育分析显示,优势和非优势OTU均分布在系统发育树的3个不同分支上,这种与多种亲缘关系较远的OrMF共生的现象可能与杓兰属植物对环境的适应性有关,且不同杓兰的OrMF物种丰富度没有显著差异,但群落结构存在差异。     

Species Diversity of Russuloid Mycorrhizae forming Fungi on Pinus yunnanensis Seedlings and the Mycorrhizal Morphology

Russuloid ectomycorrhizae on 1-2 years old seedlings of Pinus yunnanensis collected from the central Yunnan, China were investigated. Six morphotypes were recognized by macro and anatomical morphological approaches as well as molecular analyses. The six morphotypes were confirmed to represent six phylotypes by matching rDNA ITS sequences. The fungal partners in the six morpho phylotypes are those closely related with Russula livescens, Rvioleipes ,Rdensifolia ,Rnigricans ,Rsanguinea and Rnauseosa respectively. The correlation between the morphology of mantles and cystidia and the phylogenetic clades is further supported in our work. The framework, in which this report is included, shows that russuloid mycorrhizae are one of the most dominant representative ectomycorrhizae formed on 1-2 years seedlings of Pinus yunnanensis.     

丛枝菌根真菌离体培养研究概况(综述)

本文综述丛枝菌根真菌在离体条件下的生长发育、生长促进物质及生理生化代谢等方面的研究概况。     

Diversity of Rhizosphere Soil Arbuscular Mycorrhizal Fungi in Various Soybean Cultivars under Different Continuous Cropping Regimes

Recent studies have shown that continuous cropping in soybean causes substantial changes to the microbial community in rhizosphere soil. In this study, we investigated the effects of continuous cropping for various time periods on the diversity of rhizosphere soil arbuscular mycorrhizal (AM) fungi in various soybean cultivars at the branching stage. The soybean cultivars Heinong 37 (an intermediate cultivar), Heinong 44 (a high-fat cultivar) and Heinong 48 (a high-protein cultivar) were seeded in a field and continuously cropped for two or three years. We analyzed the diversity of rhizosphere soil AM fungi of these soybean plants at the branching stage using morphological and denaturing gradient gel electrophoresis (DGGE) techniques. The clustering analysis of unweighted pair-group method with arithmetic averages (UPGMA) was then used to investigate the AM fungal community shifts. The results showed that increasing the number of years of continuous cropping can improve the colonization rate of AM fungi in different soybean cultivars at the branching stage. The dominant AM fungi in the experimental fields were Funneliformismosseae and Glomus spp. The number of years of continuous cropping and the soybean cultivar both had obvious effects on the diversity of AM fungi, which was consistent with the results of colonization rate analysis. This study establishes a basis for screening dominant AM fungi of soybean. In addition, the results of this study may be useful for the development of AM fungal inoculants.