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1.
丛枝菌根真菌对红三叶草利用不同有机磷源的研究   总被引:5,自引:3,他引:5  
以红三叶草为材料 ,利用三室隔网培养方法 ,施用不同有机磷源 :植酸钠 (Na -Phytate)、核糖核酸 (RNA)和卵磷脂 (Lecithin) ,研究接种菌根真菌Glmous versiforme对土壤及外加有机磷源的利用效率 ,另设无机磷及不施磷作为对照。结果表明 ,接种菌根真菌能明显增加植株干物重、含磷量和吸磷总量。与各有机磷处理相比 ,无机磷处理前期的生长效应较好 ,施用有机磷各处理在不同生长时期均明显促进了植株生长 ,但不同有机磷源之间没有显著差异。在植株吸磷量上 ,植株生长 7周以前 ,磷酸二氢钾处理高于其它处理 ,而植株生长 10周时 ,植酸钠处理高于磷酸二氢钾处理。接种菌根处理由于丛枝菌根活化了土壤有机磷 ,到植株生长 10周时其吸收有机磷的量已占吸磷总量的 76 .7%。  相似文献   

2.
 Plants colonised with the arbuscular mycorrhizal fungi (AMF) Glomus E3 and Glomus tenue were grown in microcosms that permitted separation into root:hyphae and hyphae compartments. Hydrolysed polysaccharides from the hyphae and water-soluble sugars released into the hyphosphere were assayed using chromatography. Total sugars and most monosaccharides were elevated in the hyphosphere of Glomus E3 but not in the hyphosphere of G. tenue. Differences in the levels of sugars did not depend on hyphal surface area. It is suggested the diversity in sugars produced in the hyphosphere of AMF may drive some of the spatial and temporal variation in microbial diversity and function in soils.  相似文献   

3.
Increased phosphate (P) uptake in plants by arbuscular mycorrhizal (AM) fungi is thought to depend mainly on the extension of external hyphae into soil. On the other hand, it is known that the hyphae of some kinds of ectomycorrhizal fungi release organic acids into soil and that they dissolve the insoluble inorganic P. This study collected hyphal exudates of AM fungi within compartmentalized pot culture and clarified their ability to solubilize insoluble inorganic P. Sterilized Andisol was packed in pots that were separated into root and hyphal compartments with a nylon net of 30 μm pore size. Seedlings of Allium cepa inoculated with AM fungi, Gigaspora margarita, or Glomus etunicatum were grown. Control pots were not inoculated. Mullite ceramic tubes were buried in the soil of each compartment and soil solution was collected. The anionic fraction of the soil solution was incubated with iron phosphate (4 mg FePO4 in 1 mL of 0.4 acetate buffer). Solubilized P was measured. The AM colonization of plants inoculated with G. margarita and G. etunicatum was 86% and 54%, respectively. Adhesion of external hyphae was observed on the surface of the mullite ceramic tubes buried in soil of the hyphal compartment. Colonization of both fungi increased shoot P uptake and growth. Soil solution collected from the hyphal compartments of both fungi solubilized more P than did that from uninoculated plants. It is suggested that hyphal exudates can contribute to increased P uptake of colonized plants.  相似文献   

4.
不同AM菌根菌分泌的磷酸酶对根际土壤有机磷的影响   总被引:18,自引:1,他引:18  
苏友波  林春  张福锁  李晓林 《土壤》2003,35(4):334-338,343
以三叶草为材料,利用3室隔网培养方法,研究了4种AM菌根菌侵染三叶草后对根际土壤酸性和碱性磷酸酶活性以及菌丝酶活性对土壤有机P的影响。结果表明,接种AM菌根菌 (9周) 对根际土壤酸性和碱性磷酸酶活性均有增强作用,但作用强度主要取决于菌丝在土壤中的生长状况,Glomus属菌根菌在整个菌丝室 (0~6cm) 都影响土壤磷酸酶的活性,其活性在整个菌丝室中都比Gigaspora的高。同一属不同种的根际土壤磷酸酶活性差异不大。AM菌根根际土壤磷酸酶对土壤有机P的降解有很强的促进作用。  相似文献   

5.
Arbuscular mycorrhizal fungi (AMF) are obligate symbionts of most higher plants. In addition to being a major component of soil microbial biomass, AMF hyphae produce glomalin, a recalcitrant glycoproteinaceous substance highly correlated with soil aggregate water stability. This study addresses the lack of knowledge concerning the decomposition of hyphae and glomalin. We used an experimental design that exploited the lack of saprobic capabilities of AMF hyphae by incubating field soil samples in the dark, and hence in the absence of plant or AMF hyphal growth. In 150 days, hyphal length decreased 60%, while glomalin, quantified by the Bradford protein assay, declined only 25%. Immuno-reactive glomalin decreased 46%. This study serves as a proof-of-concept for further examination of factors that influence decomposition of AMF hyphae using similar experimental designs.  相似文献   

6.
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.  相似文献   

7.
Abstract

In general, according to previous studies, pioneer species do not require arbuscular mycorrhizal fungi (AMF) to increase their growth and survival in tropical systems. The aim of this study was to determine the dependence response to AMF of Heliocarpus appendiculatus, a pioneer species, at different phosphorus (P) levels. In a greenhouse experiment, H. appendiculatus seedlings were grown in pots with a sterile vermiculite-sand mixture (1:1). Two sets of pots were set up: One set was inoculated (150 spores per pot) with indigenous AMF from a tropical rain forest at “Los Tuxtlas” (Veracruz, Mexico); the other set was not inoculated. To each set, 0, 0.02, 0.2, and 2 g L?1P was added. All pots were watered with 250 mL of nutrient solution. Mycorrhizal plants showed a higher total dry weight and relative growth rate in 0.02 g L?1P concentration, while nonmycorrhizal plants responded positively at 0.2 g L?1P; a decrease in plant responses at higher P levels was observed in both treatments. H. appendiculatus showed to have higher relative dependence at lower P concentration (≈50%). As levels of P increased, mycorrhizal colonization decreased. Successful growth of pioneer species during succession process may be improved if there is AMF content in soils, prior to disturbance.  相似文献   

8.
ABSTRACT

This work was conducted to study phosphorus (P) efficiency of two maize genotypes (Zea mays, L.) in calcareous soil grown in potted soil with two levels of P in soil by adding 40 and 270 mg P/kg soil. Half of the pots were inoculated with arbuscular mycorrhizal fungi (AMF) (Rhizoglomus irregulare). The maize genotypes were harvested two times at 35 and 50 days after transplanting. The plant dry matter, root length and Plant P uptake of maize genotype Hagen 1 without mycorrhizal fungi (AMF) increased significantly compared with Hagen 9 at a low P level. In contrast, there was no significant difference between two maize genotypes inoculated with AMF under the same P level. The predicted value increased rapidly with increasing P levels from about 70% up to 97% in both maize genotypes with and without mycorrhizal fungi. At a low P level, the mycorrhizae hyphae contributed by about 31.6% and 30.2% of the predicted total P uptake in maize genotype Hagen 1 and Hagen 9, respectively. The results of this study suggested that the P-inefficient genotype Hagen 9 improved with inoculation with mycorrhizal fungi under a low P level at the same conditions of this experiment. Also, root growth system and mycorrhizal hyphae length would be a suitable plant parameter for studying P efficient maize genotypes, especially under limited P supply. The current study clearly pointed out that the mechanistic simulation model (NST 3.0) provides useful tools for studying the role of AMF in P uptake of plant.  相似文献   

9.
Arbuscular mycorrhizal fungi (AMF) differ in their rate and extent of colonization of both plant roots and soil but the mechanism responsible for these differences is unclear. We compared the external mycelium of three AMF isolates (Glomus intraradices, Glomus etunicatum and Gigaspora gigantea) during early colonization of plant roots. We investigated whether an AMF with the most rapid colonization would have higher numbers of infective structures (i.e., infection hyphae and contact points), an AMF with extensive root colonization would have more infection units, and (3) AMF with extensive soil colonization would have large numbers of all external features (including absorptive hyphae, runner hyphae and hyphal bridges). Using specially designed soil and root observation chambers, we followed the development of the external mycelium for 7 weeks. We found that rapid colonization rate was due, in part, to the presence of more infective structures, in particular more infection hyphae and root contact points. Second, the extensive root colonizer had more, larger infection units. Third, data did not support the hypothesis that the extensive soil colonizer had more external structures. These results show that differences in the architecture of the external mycelium are responsible, in part, for variation in the colonization strategy of AMF.  相似文献   

10.
《Applied soil ecology》2003,22(2):139-148
A glasshouse pot experiment investigated the uptake by arbuscular mycorrhizal (AM) fungi associated with red clover of three organic sources of P added to a sterilized calcareous soil of low P availability. Each pot was separated into a central compartment for plant growth and two outer compartments for external mycelium using 30-μm nylon mesh to restrict the roots but allow hyphal penetration. Plants in the central compartments were inoculated with the AM fungus Glomus versiforme and uninoculated controls were included. Plants were harvested on three occasions: 5, 7 and 10 weeks after sowing. Application of each of the three organic P sources (lecithin, RNA and sodium phytate) or inorganic P (KH2PO4) at 50 mg P kg−1 to the outer compartments of mycorrhizal and uninoculated pots increased the yield, P concentration and total P uptake of red clover compared with pots to which no P was applied, with no differences among P sources in non-mycorrhizal plants but differences observed in mycorrhizal plants both 7 and 10 weeks after sowing suggesting differences in availability of the four P sources to AM mycelium. The contribution of external mycelium to plant uptake of applied P increased with time. The three organic P sources made smaller contributions to plant P nutrition than KH2PO4 at the first and second harvests. At the third harvest, the contribution from KH2PO4 was 23%, while those from lecithin, RNA and sodium phytate were 23, 17 and 31%, respectively. This suggests that with the mediation of AM fungi, soil organic P sources can make a contribution to host plant P nutrition comparable to that of soluble orthophosphate.  相似文献   

11.
丛枝菌根对三叶草根际磷酸酶活性的影响   总被引:8,自引:0,他引:8  
以三叶草为材料,利用三室隔网培养方法,探讨了取自肥料长期定位试验中多年施用与不施用有机肥的田间小区土壤上,接种菌根菌(G.mosseae)对根际土壤酸性和碱性磷酸酶活性的影响。植物生长9周后,收获测定菌丝生长室土壤酸性磷酸酶和碱性磷酸酶活性,并对磷酸酶产生位点进行细胞化学定位。结果表明,接种丛技菌根菌对根际土壤酸性和碱性磷酸酶活性均有增强作用,但作用程度在有机服小区土壤上要大于无机肥小区土壤。根际土壤酸性磷酸酶原位化学定位结果表明,菌丝周围有明显的酸性磷酸酶的反应产物,说明报外丛枝菌根菌丝能直接向外分泌磷酸酶。  相似文献   

12.
【目的】蚯蚓和丛枝菌根真菌处于不同的营养级,但在促进植物生长和提高土壤肥力等方面却都发挥着积极作用。研究蚯蚓菌根互作及其对玉米吸收土壤中的氮、磷养分的影响,可为提升土壤生物肥力及促进农业的可持续发展提供理论依据。【方法】本研究采用田间盆栽方式,以玉米为供试作物,研究蚯蚓(Eisenia fetida)与丛枝菌根真菌(Glomus intraradices)互作及其对玉米养分吸收的影响。试验设置P 25和175 mg/kg两个水平。每个磷水平进行接种与不接种菌根真菌以及添加与不添加蚯蚓,共8个处理。调查了玉米生长、养分吸收以及真菌浸染和土壤养分的有效性。【结果】两个磷水平下,蚯蚓和菌根在增加玉米地上部和根系生物量方面有显著正交互作用(P0.05)。接种菌根真菌的各处理显著增加了玉米的侵染率及泡囊丰度、根内菌丝丰度等菌根指标。同时添加蚯蚓和接种菌根真菌的处理(AM+E)显著提高了菌根的侵染率、菌丝密度、丛枝丰度和根内菌丝丰度但是泡囊丰度有所下降。两种磷水平下,AM+E处理玉米地上部和地下部含氮量和含磷量均显著高于其他三个处理。在低磷条件下,地上部氮磷总量的增加分别是添加蚯蚓和接菌的作用;而地下部磷总量的增加主要是菌根真菌的作用。在高磷条件下,单加蚯蚓显著增加玉米氮磷的总量,而接种菌根真菌对玉米氮磷吸收的影响未达显著性水平。在高磷条件下,单加蚯蚓的处理显著提高玉米地上地下部生物量(P0.05),而单接菌的处理效应不显著,蚯蚓菌根互作通过提高土壤微生物量碳、氮实现对玉米生长和养分吸收的调控。在低磷条件下,单接菌显著提高了玉米的生物量(P0.05),单加蚯蚓的处理具有增加玉米生物量的趋势。菌根真菌主要促进玉米对磷的吸收,蚯蚓主要矿化秸秆和土壤中的氮磷养分增加土壤养分的有效性,蚯蚓菌根互作促进了玉米根系对土壤养分的吸收并形成氮磷互补效应。【结论】无论在高磷还是低磷水平下,蚯蚓菌根相互作用都提高了玉米地上地下部生物量、氮磷吸收量同时提高了土壤微生物量碳、氮。蚯蚓菌根相互作用对植物生长的影响取决于土壤养分条件。在高磷条件下(氮相对不足),蚯蚓菌根互作通过调控土壤微生物量碳、氮调控玉米生长和养分吸收。低磷条件下,菌根主要发挥解磷作用,蚯蚓主要矿化秸秆和土壤中的氮素,蚯蚓和菌根互补调控土壤中氮、磷,从而促进植物的生长和养分吸收。  相似文献   

13.
In this work, a greenhouse experiment, arranged in a randomized block design 2 × 6 factorial with six replicates, was conducted to estimate the effect of inoculation with arbuscular mycorrhizal fungi (AMF) on lead uptake, location and stress attenuation in mycorrhizal Zea mays L. seedlings. Treatments were the mycorrhizal inoculation (+M) or non-mycorrhizal inoculation (?M) and six lead concentrations (0, 50, 100, 200, 500 and 1000 μg/g) to soil. The results showed that AM fungal inoculation could attenuate the oxidative stress of lead to Z. mays seedlings. The higher height, basal diameter and biomass of seedlings were found in mycorrhizal Z. mays seedlings growing in the soil with increasing lead concentrations. Moreover, superoxide dismutase (SOD) activity was higher than that of non-inoculated seedlings. AMF increased accumulation of lead in the root system. In the presence of 200, 500 and 1000 μg/g lead, there were higher lead concentrations in roots of mycorrhizal seedlings than in non-mycorrhizal seedlings. Lead was identified to dominantly deposit in the hyphal wall, the hyphal inner chambers, the hyphal inner-chamber membranes and the vacuole inner-chambers membrane. It is, therefore, hypothesized that lead stress can be decreased through the AM fungal cell. The ability of arbuscular mycorrhiza immobilizing lead can alleviate the phytotoxicity of lead to Z. mays seedlings.  相似文献   

14.
Species composition of arbuscular mycorrhizal fungi (AMF) was analysed in two differently managed mountain grasslands in Thuringia (Germany). Arbuscular mycorrhizal fungi were studied in the roots of 18 dominant plant species from a total of 56 (32%). Additionally, spores of AMF were isolated from soil samples. Arbuscular mycorrhizal fungi species composition was analysed based on 96 sequences of the internal transcribed spacer of the nuclear ribosomal DNA, 72 originated from mycorrhizal roots, and 24 originated from AMF spores. Phylogenetic analyses revealed a total of 19 AMF species representing all genera of the Glomeromycota except Scutellospora and Pacispora. Despite a different farming intensity, resulting in remarkable differences concerning their plant species diversity (27 against 43 plant species), the diversity of AMF was found to be similar with 11 species on the intensively farmed meadow and ten species on the extensively farmed one. Nevertheless, species composition between both sites was clearly different. It thus seems likely that the AMF species composition, but not necessarily the species number, is related to above ground plant biodiversity in the system under study.  相似文献   

15.
Soil aggregation is a principal ecosystem process mediated by soil biota. Collembola and arbuscular mycorrhizal (AM) fungi are important groups in the soil, and can interact in various ways. Few studies have examined collembola effects on soil aggregation, while many have quantified AM effects. Here, we asked if collembola have any effect on soil aggregation, and if they alter AM fungi-mediated effects on soil aggregation.We carried out a factorial greenhouse study, manipulating the presence of both collembola and AM fungi, using two different plant species, Sorghum vulgare and Daucus carota. We measured root length and biomass, AMF (and non-AMF) soil hyphal length, root colonization, and collembolan populations, and quantified water stable soil aggregates (WSA) in four size classes.Soil exposed to growth of AMF hyphae and collembola individually had higher WSA than control treatments. Moreover, the interaction effects between AMF and collembola were significant, with non-additive increases in the combined application compared to the single treatments.Our findings show that collembola can play a crucial role in maintaining ecological sustainability through promoting soil aggregation, and point to the importance of considering organism interactions in understanding formation of soil structure.  相似文献   

16.
《Applied soil ecology》2007,35(2-3):200-208
The temporal and spatial dynamics of arbuscular mycorrhizal fungi (AMF) were investigated in Indian Thar Desert. Soil samples under Mitragyna parvifolia were collected from July 2003 to June 2004. AMF colonization and spore density were used to compare the responses of AMF to different abiotic parameters. The mean percent colonization and spore density of AMF reached maximal values in rainy and summer seasons, respectively. Vesicular and hyphal colonizations were positively correlated with soil organic carbon content. AMF spore density was positively correlated with soil pH and negatively correlated with Olsen P content. A high Shannon–Weiner diversity index of AMF was observed in Thar Desert. A total of fifteen AMF species were associated with M. parvifolia. Percent spore density and species richness suggest that the genus Glomus was the predominant AMF under Thar Desert environment. The reasons for the observed variations are discussed.  相似文献   

17.
Hydrothermal carbonization (HTC) is a method to produce carbonized material at relatively low temperatures (180–250 °C) under pressure and aqueous conditions. The product is called hydrochar and can be used as a soil amendment. However, applied in high dosages it may have detrimental effects on plants or soil biota. The potential impact of hydrochar amendment on beneficial soil organisms such as arbuscular mycorrhizal fungi (AMF) and earthworms and their interactions are not well understood. The goal of the present study was to determine effects of hydrochar on plant growth and soil biota and to evaluate interactions of earthworms and hydrochar on plant and AMF performance and to identify underlying mechanisms. In a greenhouse experiment, we investigated the effect of hydrochar at different addition rates (control, 1% and 10%, v/v) with or without the earthworm Aporrectodea caliginosa on the growth of Plantago lanceolata L. and the performance of its AMF. We observed a positive interaction between earthworms and 10% hydrochar on shoot and root biomass: added as a single treatment hydrochar had a negative effect on plant growth at this dosage, but plant biomass increased significantly when hydrochar was added together with earthworms. Root colonization by AMF increased significantly with increasing concentration of hydrochar, but was not affected by earthworms. Contrastingly, extraradical hyphal length of AMF was reduced by earthworms, but not affected by hydrochar. Thus, hydrochar and earthworms affected the performance of AMF, albeit of different AMF structures and in different directions. Our results indicate that earthworms may play an important role in alleviating the negative impacts of high dosages of hydrochar on plant growth; such interactions should move into focus of future research on potential effects of HTC materials.  相似文献   

18.
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.  相似文献   

19.
Although soil-dwelling Collembola can influence plant growth and nutrient cycling, their specific role in soil food webs is poorly understood. Soil-free microcosm studies suggest that Collembola are primarily fungivores where they feed preferentially on saprophytic fungi (SF) over other fungal types. We directly assessed collembolan consumption of arbuscular mycorrhizal fungi (AMF) and SF using plant-soil mesocosms and natural abundance stable carbon isotope techniques. Mycorrhizal Andropogon gerardii (C4 grass) seedlings were placed in pots containing Collembola and soil from a C3 plant dominated site, while mycorrhizal Pascopyrum smithii (C3 grass) seedlings were placed in pots with Collembola and soil collected at a C4 plant dominated site. After 6 weeks, collembolans assimilated carbon derived from C3 and C4 sources in both A. gerardii and P. smithii treatments. Comparing Collembola isotope values in AMF vs. AMF-suppressed treatments, our data show that both AMF and SF were consumed in these experimental soil environments.  相似文献   

20.
The development of communities of arbuscular mycorrhizal fungi (AMF) was investigated in the subalpine foreland of the glacier Morteratsch located at approx. 1900–2100 m a.s.l. near Pontresina (Engiadin’ Ota, Switzerland). In particular, we asked if the succession of AMF communities follows or precedes the primary plant succession, and we checked the mycorrhizal status of the pioneer plant Epilobium fleischeri. Soil samples were taken at pioneer and dense grassland sites established during the last hundred years representing different periods of glacier retreat: 1875–1900, 1940–1950, 1970–1980 and 1990–2000. Extraradical hyphal length densities and AMF spore populations were analyzed in soil samples. Spore formation and mycorrhizal root colonization were monitored in trap cultures grown on Trifolium pratense, Lolium perenne, Plantago lanceolata and Hieracium pilosella or on E. fleischeri over 14 months. We found that E. fleischeri is strongly arbuscular mycorrhizal, but plants in closest distance to the glacier (glacier retreat in the last 4–6 years before sampling) were non-mycorrhizal. Spore densities and root colonization in trap cultures were generally low in samples from glacier stage 1990–2000. Highest spore density and colonization were found for the sites ice-free since 1970–1980, whilst highest AMF species richness and hyphal length densities were found at the sites ice-free since 1875–1900. Our findings show an establishment of a few AMF pioneer species (e.g. Diversispora versiformis and Acaulospora punctata) within 5–10 years and species rich AMF communities at sites ice-free for 100 years (28 species). Their succession generally follows the succession of the plant communities. We conclude that AMF pioneer species might be mainly distributed by wind transport while other AMF fungi (e.g. Glomus rubiforme and Glomus aureum) rather need a below-ground hyphal network to invade new areas.  相似文献   

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