Plant species richness,identity and productivity differentially influence key groups of microbes in grassland soils of contrasting fertility

The abundance of microbes in soil is thought to be strongly influenced by plant productivity rather than by plant species richness per se. However, whether this holds true for different microbial groups and under different soil conditions is unresolved. We tested how plant species richness, identity and biomass influence the abundances of arbuscular mycorrhizal fungi (AMF), saprophytic bacteria and fungi, and actinomycetes, in model plant communities in soil of low and high fertility using phospholipid fatty acid analysis. Abundances of saprophytic fungi and bacteria were driven by larger plant biomass in high diversity treatments. In contrast, increased AMF abundance with larger plant species richness was not explained by plant biomass, but responded to plant species identity and was stimulated by Anthoxantum odoratum. Our results indicate that the abundance of saprophytic soil microbes is influenced more by resource quantity, as driven by plant production, while AMF respond more strongly to resource composition, driven by variation in plant species richness and identity. This suggests that AMF abundance in soil is more sensitive to changes in plant species diversity per se and plant species composition than are abundances of saprophytic microbes.     

Exotic cheatgrass and loss of soil biota decrease the performance of a native grass

Soil disturbances can alter microbial communities including arbuscular mycorrhizal (AM) fungi, which may in turn, affect plant community structure and the abundance of exotic species. We hypothesized that altered soil microbial populations owing to disturbance would contribute to invasion by cheatgrass (Bromus tectorum), an exotic annual grass, at the expense of the native perennial grass, squirreltail (Elymus elymoides). Using a greenhouse experiment, we compared the responses of conspecific and heterospecific pairs of cheatgrass and squirreltail inoculated with soil (including live AM spores and other organisms) collected from fuel treatments with high, intermediate and no disturbance (pile burns, mastication, and intact woodlands) and a sterile control. Cheatgrass growth was unaffected by type of soil inoculum, whereas squirreltail growth, reproduction and nutrient uptake were higher in plants inoculated with soil from mastication and undisturbed treatments compared to pile burns and sterile controls. Squirreltail shoot biomass was positively correlated with AM colonization when inoculated with mastication and undisturbed soils, but not when inoculated with pile burn soils. In contrast, cheatgrass shoot biomass was negatively correlated with AM colonization, but this effect was less pronounced with pile burn inoculum. Cheatgrass had higher foliar N and P when grown with squirreltail compared to a conspecific, while squirreltail had lower foliar P, AM colonization and flower production when grown with cheatgrass. These results indicate that changes in AM communities resulting from high disturbance may favor exotic plant species that do not depend on mycorrhizal fungi, over native species that depend on particular taxa of AM fungi for growth and reproduction.     

Establishment and effectiveness of inoculated arbuscular mycorrhizal fungi in agricultural soils

Arbuscular mycorrhizal fungi (AMF) are promoted as biofertilizers for sustainable agriculture. So far, most researchers have investigated the effects of AMF on plant growth under highly controlled conditions with sterilized soil, soil substrates or soils with low available P or low inoculum potential. However, it is still poorly documented whether inoculated AMF can successfully establish in field soils with native AMF communities and enhance plant growth. We inoculated grassland microcosms planted with a grass–clover mixture (Lolium multiflorum and Trifolium pratense) with the arbuscular mycorrhizal fungus Rhizoglomus irregulare. The microcosms were filled with eight different unsterilized field soils that varied greatly in soil type and chemical characteristics and indigenous AMF communities. We tested whether inoculation with AMF enhanced plant biomass and R. irregulare abundance using a species specific qPCR. Inoculation increased the abundance of R. irregulare in all soils, irrespective of soil P availability, the initial abundance of R. irregulare or the abundance of native AM fungal communities. AMF inoculation had no effect on the grass but significantly enhanced clover yield in five out of eight field soils. The results demonstrate that AMF inoculation can be successful, even when soil P availability is high and native AMF communities are abundant.     

Differential responses of soil bacteria,fungi, archaea and protists to plant species richness and plant functional group identity

Plants are known to influence belowground microbial community structure along their roots, but the impacts of plant species richness and plant functional group (FG) identity on microbial communities in the bulk soil are still not well understood. Here, we used 454‐pyrosequencing to analyse the soil microbial community composition in a long‐term biodiversity experiment at Jena, Germany. We examined responses of bacteria, fungi, archaea, and protists to plant species richness (communities varying from 1 to 60 sown species) and plant FG identity (grasses, legumes, small herbs, tall herbs) in bulk soil. We hypothesized that plant species richness and FG identity would alter microbial community composition and have a positive impact on microbial species richness. Plant species richness had a marginal positive effect on the richness of fungi, but we observed no such effect on bacteria, archaea and protists. Plant species richness also did not have a large impact on microbial community composition. Rather, abiotic soil properties partially explained the community composition of bacteria, fungi, arbuscular mycorrhizal fungi (AMF), archaea and protists. Plant FG richness did not impact microbial community composition; however, plant FG identity was more effective. Bacterial richness was highest in legume plots and lowest in small herb plots, and AMF and archaeal community composition in legume plant communities was distinct from that in communities composed of other plant FGs. We conclude that soil microbial community composition in bulk soil is influenced more by changes in plant FG composition and abiotic soil properties, than by changes in plant species richness per se.     

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。     

Mycorrhizal fungi reduce the negative effects of nitrogen enrichment on plant community structure in dune grassland

Nitrogen (N) inputs to ecosystems have increased worldwide, often leading to large changes in plant community structure and reducing plant diversity. Yet, the interaction of increased N availability with other factors that determine plant community composition, are still poorly understood. Here, we test whether the impact of N addition on plant communities depends on the presence of arbuscular mycorrhizal fungi (AMF). AMF are widespread plant symbionts that facilitate growth of many plant species. We hypothesize that AM fungi reduce the negative impact of N addition on plant communities by supporting growth of species that are sensitive to N enrichment.We established experimental grassland microcosms consisting of 18 plant species. These microcosms were subjected to high and low N supply and were inoculated with AMF or remained nonmycorrhizal. Both N addition and AMF had a big impact on plant community composition, but with opposite effects. N addition induced a 2.8‐fold increase in grass biomass and reduced legume biomass. Grasses dominated the microcosms at high N supply, especially when AMF were absent. In contrast, AMF enhanced biomass of all legumes species (on average 6.8‐fold) and reduced the relative abundance of grasses. The proportion of legume biomass out of total shoot biomass at high N supply was 19% with AMF and only 3% without AMF. Our results show that responses of plant communities to N enrichment depend on AMF and that AMF can reduce the negative impact of increased N availability on plant community structure by reducing grass dominance.     

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.     

Species-specific trade-offs between regrowth and mycorrhizas in the face of defoliation and phosphorus addition

Arbuscular mycorrhizal fungi (AMF) enhance nutrient provision in exchange for carbon. However, a shift from nutrient to carbon limitation, induced by repeated or intense defoliation, can represent a trade-off between plant regrowth and the maintenance of mycorrhiza. The combined effects of AMF, phosphorus and defoliation on growth of Agropyron elongatum (C₃ grass, low mycorrhizal responsiveness) and Brachiaria brizantha (C₄ grass, high mycorrhizal responsiveness) were explored. Each species was subjected to a manipulative experiment with AMF inoculation (non-inoculated, inoculated), soluble P supply (low, high) and defoliation (non-defoliated, 60% defoliated). In A. elongatum, at low P supply, mycorrhizal plants showed increased growth rates following defoliation without substantial changes in AMF colonization. At high P supply instead, we found a clear trade-off between regrowth and the maintenance of mycorrhiza evidenced by growth depression (biomass and tillers) and lower AMF activity (reduction of arbuscules). In contrast, in B. brizantha, defoliation effects on plant regrowth were independent from AMF at any P supply (no trade-off). This indicates that cost-benefit relationship in defoliated plants is highly context-dependent and may vary with mycorrhizal responsiveness of species. This variation of responses can play a decisive role on plant recovery in pastures and natural grasslands subjected to herbivory.     

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.     

Contrasting impacts of defoliation on root colonization by arbuscular mycorrhizal and dark septate endophytic fungi of Medicago sativa

Individual plants typically interact with multiple mutualists and enemies simultaneously. Plant roots encounter both arbuscular mycorrhizal (AM) and dark septate endophytic (DSE) fungi, while the leaves are exposed to herbivores. AMF are usually beneficial symbionts, while the functional role of DSE is largely unknown. Leaf herbivory may have a negative effect on root symbiotic fungi due to decreased carbon availability. However, evidence for this is ambiguous and no inoculation-based experiment on joint effects of herbivory on AM and DSE has been done to date. We investigated how artificial defoliation impacts root colonization by AM (Glomus intraradices) and DSE (Phialocephala fortinii) fungi and growth of Medicago sativa host in a factorial laboratory experiment. Defoliation affected fungi differentially, causing a decrease in arbuscular colonization and a slight increase in DSE-type colonization. However, the presence of one fungal species had no effect on colonization by the other or on plant growth. Defoliation reduced plant biomass, with this effect independent of the fungal treatments. Inoculation by either fungal species reduced root/shoot ratios, with this effect independent of the defoliation treatments. These results suggest AM colonization is limited by host carbon availability, while DSE may benefit from root dieback or exudation associated with defoliation. Reductions in root allocation associated with fungal inoculation combined with a lack of effect of fungi on plant biomass suggest DSE and AMF may be functional equivalent to the plant within this study. Combined, our results indicate different controls of colonization, but no apparent functional consequences between AM and DSE association in plant roots in this experimental setup.     

Arbuscular mycorrhizas in a valley-type savanna in southwest China

The arbuscular mycorrhizal (AM) status of 67 plant species in a savanna community in the hot, dry valley of Jinsha River, southwest China was surveyed. It was found that about 95% of the plant species formed AM and 5% possibly formed AM. The composition of AM fungi (AMF) in the rhizosphere soils was also investigated. The AMF spore density ranged from 5 to 6,400 per 100 g soil, with an average of 1,530, and these spores/sporocarps were identified as belonging to six genera. Fungi belonging to the genera Glomus and Acaulospora were the dominant AMF. High densities of AMF spores in the rhizosphere soils, and the intensive colonization of the plant roots, indicated that plants grown in this valley-type savanna may be highly dependent on AM.     

接种AMF对菌根植物和非菌根植物竞争的影响

为了研究丛枝菌根真菌(arbuscular mycorrhizal fungus, AMF)对菌根植物与非菌根植物种间竞争的影响,以玉米(菌根植物)和油菜(非菌根植物)作为供试植物,分别进行间作、尼龙网分隔和单作,模拟这两种植物之间不同的竞争状态,接种丛枝菌根真菌Glomus intraradices和Glomus mosseae,比较菌根植物和非菌根植物的生长和磷营养状况,分析AMF侵染对植物种间竞争作用的影响。结果显示,与单作相比,间作模式下玉米的生物量及磷营养状况有所降低,但其菌根依赖性却有所提高。与不接种相比,接种处理显著降低了间作体系油菜根系的磷含量和磷吸收量,但趋于改善菌根植物玉米的磷营养状况。因此,接种AMF可以降低非菌根植物的磷营养状况及生物量,使得菌根植物的相对竞争能力明显提高,说明AMF在维持物种多样性方面有着重要的作用。     

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.     

Mycorrhizal Fungi: Siderophore Production

Abstract

Mycorrhizal fungi, which commonly occur in natural as well as agricultural soils, are known to enhance plant uptake of nutrients, including metal ions present as trace concentrations. As mycorrhizal infection is a widespread feature of plant communities, it seems appropriate to review the data on mycorrhizal fungi and their potential to produce siderophores.

Based on a bioassay with Aureobacteriumflavescens JG-9 it was shown that a number of ectomycorrhizal fungi (EM) produce hydroxamate siderophores. Also an arbuscular mycorrhizal (AM) grass species, which showed greater iron uptake than nonmycorrhizal controls, tested positively when bioassayed for hydroxamate siderophores. Encoid mycorrhizal fungi, too, have been demonstrated to be capable of producing hydroxamate-type siderophores. However, only in the case of the eridoid mycorrhizal fungi the main siderophores have been isolated and subsequently identified as ferricrocin and fusigen, respectively. The biotechnological and ecological significance of studies of the siderophore biosynthesis by mycorrhizal fungi is discussed.     

Phylogenetically diverse AM fungi from Ecuador strongly improve seedling growth of native potential crop trees

In many deforested regions of the tropics, afforestation with native tree species could valorize a growing reservoir of degraded, previously overused and abandoned land. The inoculation of tropical tree seedlings with arbuscular mycorrhizal fungi (AM fungi) can improve tree growth and viability, but efficiency may depend on plant and AM fungal genotype. To study such effects, seven phylogenetically diverse AM fungi, native to Ecuador, from seven genera and a non-native AM fungus (Rhizophagus irregularis DAOM197198) were used to inoculate the tropical potential crop tree (PCT) species Handroanthus chrysanthus (synonym Tabebuia chrysantha), Cedrela montana, and Heliocarpus americanus. Twenty-four plant-fungus combinations were studied in five different fertilization and AMF inoculation treatments. Numerous plant growth parameters and mycorrhizal root colonization were assessed. The inoculation with any of the tested AM fungi improved seedling growth significantly and in most cases reduced plant mortality. Plants produced up to threefold higher biomass, when compared to the standard nursery practice. AM fungal inoculation alone or in combination with low fertilization both outperformed full fertilization in terms of plant growth promotion. Interestingly, root colonization levels for individual fungi strongly depended on the host tree species, but surprisingly the colonization strength did not correlate with plant growth promotion. The combination of AM fungal inoculation with a low dosage of slow release fertilizer improved PCT seedling performance strongest, but also AM fungal treatments without any fertilization were highly efficient. The AM fungi tested are promising candidates to improve management practices in tropical tree seedling production.     

Inoculation effect of arbuscular mycorrhizal fungi on the growth of Vatica chinensis L.: a critically endangered species of Western Ghats

Pot experiments were conducted to assess the efficacy of three arbuscular mycorrhizal (AM) fungi viz. Glomus fasciculatum, Funneliformis mosseae and Acaulospora laevis on the growth of Vatica chinensis. Plants grown in the presence of AM fungi generally showed an increase in growth and biomass over those grown in the absence of AM fungal inoculation. It was observed that a consortium of AM fungi without organic manure was found to be the best treatment to enhance plant biomass. The plant height, number of leaves and plant dry and fresh matter, plant P and N contents varied significantly with treatments. The AM colonization was higher in the seedling roots inoculated with two or three AM fungi. This study indicated that the inoculation with AM fungi either alone or in combination improved the growth, biomass yield and phosphorus nutrition of V. chinensis compared to that of the control. This information may be helpful to improve the growth of these species in sustainable forestry program.     

The intercropping partner affects arbuscular mycorrhizal fungi and Fusarium oxysporum f. sp. lycopersici interactions in tomato

Arbuscular mycorrhizal fungi (AMF) and their bioprotective aspects are of great interest in the context of sustainable agriculture. Combining the benefits of AMF with the utilisation of plant species diversity shows great promise for the management of plant diseases in environmentally compatible agriculture. In the present study, AMF were tested against Fusarium oxysporum f. sp. lycopersici with tomato intercropped with either leek, cucumber, basil, fennel or tomato itself. Arbuscular mycorrhizal (AM) root colonisation of tomato was clearly affected by its intercropping partners. Tomato intercropped with leek showed even a 20 % higher AM colonisation rate than tomato intercropped with tomato. Positive effects of AMF expressed as an increase of tomato biomass compared to the untreated control treatment could be observed in root as well as in shoot weights. A compensation of negative effects of F. oxysporum f. sp. lycopersici on tomato biomass by AMF was observed in the tomato/leek combination. The intercropping partners leek, cucumber, basil and tomato had no effect on F. oxysporum f. sp. lycopersici disease incidence or disease severity indicating no allelopathic suppression; however, tomato co-cultivated with tomato clearly showed a negative effect on one plant/pot with regard to biomass and disease severity of F. oxysporum f. sp. lycopersici. Nonetheless, bioprotective effects of AMF resulting in the decrease of F. oxysporum f. sp. lycopersici disease severity were evident in treatments with AMF and F. oxysporum f. sp. lycopersici co-inoculation. However, these bioprotective effects depended on the intercropping partner since these effects were only observed in the tomato/leek and tomato/basil combination and for the better developed plant of tomato/tomato. In conclusion, the effects of the intercropping partner on AMF colonisation of tomato are of great interest for crop plant communities and for the influences on each other. The outcome of the bioprotective effects of AMF resulting in the decrease on F. oxysporum f. sp. lycopersici disease severity and/or compensation of plant biomass does not depend on the degree of AM colonisation but more on the intercropping partner.     

Communities of arbuscular mycorrhizal fungi in Stipa krylovii (Poaceae) in the Mongolian steppe

We examined arbuscular mycorrhizal (AM) fungi colonizing the roots of Stipa krylovii, a grass species dominating the grasslands of the steppe zone in Hustai and Uvurkhangai in Mongolia. The AM fungal communities of the collected S. krylovii roots were examined by molecular analysis based on the partial sequences of a small subunit of ribosomal RNA gene as well as AM fungal colonization rates. Almost all AM fungi detected were in Glomus-group A, and were divided into 10 phylotypes. Among them, one phylotype forming a clade with G. intraradices and G. irregulare was the most dominant. Furthermore, it was also found that most of the phylotypes include AM fungi previously detected in high altitude regions in the Eurasian Continent. Significant correlations were found among soil total N, total plant biomass and AM fungal colonization ratio, which suggested that higher plant biomass may be required for the proliferation of AM fungi in the environment. Meanwhile, redundancy analysis on AM fungal distribution and environmental variables suggested that the effect of plant biomass and most soil chemical properties on the AM fungal communities were not significant.     

Diversity of Arbuscular Mycorrhizal Fungus Populations in Heavy-Metal-Contaminated Soils

High concentrations of heavy metals have been shown to adversely affect the size, diversity, and activity of microbial populations in soil. The aim of this work was to determine how the diversity of arbuscular mycorrhizal (AM) fungi is affected by the addition of sewage-amended sludge containing heavy metals in a long-term experiment. Due to the reduced number of indigenous AM fungal (AMF) propagules in the experimental soils, several host plants with different life cycles were used to multiply indigenous fungi. Six AMF ecotypes were found in the experimental soils, showing consistent differences with regard to their tolerance to the presence of heavy metals. AMF ecotypes ranged from very sensitive to the presence of metals to relatively tolerant to high rates of heavy metals in soil. Total AMF spore numbers decreased with increasing amounts of heavy metals in the soil. However, species richness and diversity as measured by the Shannon-Wiener index increased in soils receiving intermediate rates of sludge contamination but decreased in soils receiving the highest rate of heavy-metal-contaminated sludge. Relative densities of most AMF species were also significantly influenced by soil treatments. Host plant species exerted a selective influence on AMF population size and diversity. We conclude based on the results of this study that size and diversity of AMF populations were modified in metal-polluted soils, even in those with metal concentrations that were below the upper limits accepted by the European Union for agricultural soils.     

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.