Mycorrhizal fungal growth responds to soil characteristics,but not host plant identity,during a primary lacustrine dune succession

Soil factors and host plant identity can both affect the growth and functioning of mycorrhizal fungi. Both components change during primary succession, but it is unknown if their relative importance to mycorrhizas also changes. This research tested how soil type and host plant differences among primary successional stages determine the growth and plant effects of arbuscular mycorrhizal (AM) fungal communities. Mycorrhizal fungal community, plant identity, and soil conditions were manipulated among three stages of a lacustrine sand dune successional series in a fully factorial greenhouse experiment. Late succession AM fungi produced more arbuscules and soil hyphae when grown in late succession soils, although the community was from the same narrow phylogenetic group as those in intermediate succession. AM fungal growth did not differ between host species, and plant growth was similarly unaffected by different AM fungal communities. These results indicate that though ecological filtering and/or adaptation of AM fungi occurs during this primary dune succession, it more strongly reflects matching between fungi and soils, rather than interactions between fungi and host plants. Thus, AM fungal performance during this succession may not depend directly on the sequence of plant community succession.     

Interactions between biochar and mycorrhizal fungi in a water-stressed agricultural soil

Biochar may alleviate plant water stress in association with arbuscular mycorrhizal (AM) fungi but research has not been conclusive. Therefore, a glasshouse experiment was conducted to understand how interactions between AM fungi and plants respond to biochar application under water-stressed conditions. A twin chamber pot system was used to determine whether a woody biochar increased root colonisation by a natural AM fungal population in a pasture soil (‘field’ chamber) and whether this was associated with increased growth of extraradical AM fungal hyphae detected by plants growing in an adjacent (‘bait’) chamber containing irradiated soil. The two chambers were separated by a mesh that excluded roots. Subterranean clover was grown with and without water stress and harvested after 35, 49 and 63 days from each chamber. When biochar was applied to the field chamber under water-stressed conditions, shoot mass increased in parallel with mycorrhizal colonisation, extraradical hyphal length and shoot phosphorus concentration. AM fungal colonisation of roots in the bait chamber indicated an increase in extraradical mycorrhizal hyphae in the field chamber. Biochar had little effect on AM fungi or plant growth under well-watered conditions. The biochar-induced increase in mycorrhizal colonisation was associated with increased growth of extraradical AM fungal hyphae in the pasture soil under water-stressed conditions.     

Indigenous arbuscular mycorrhizal fungal assemblages protect grassland host plants from pathogens

Plant roots can establish associations with neutral, beneficial and pathogenic groups of soil organisms. Although it has been recognized from the study of individual isolates that these associations are individually important for plant growth, little is known about interactions of whole assemblages of beneficial and pathogenic microorganisms associating with plants.We investigated the influence of an interaction between local arbuscular mycorrhizal (AM) fungal and pathogenic/saprobic microbial assemblages on the growth of two different plant species from semi-arid grasslands in NE Germany (Mallnow near Berlin). In a greenhouse experiment each plant species was grown for six months in either sterile soil or in sterile soil with one of three different treatments: 1) an AM fungal spore fraction isolated from field soil from Mallnow; 2) a soil pathogen/saprobe fraction consisting of a microbial community prepared with field soil from Mallnow and; 3) the combined AM fungal and pathogen/saprobe fractions. While both plant species grew significantly larger in the presence of AM fungi, they responded negatively to the pathogen/saprobe treatment. For both plant species, we found evidence of pathogen protection effects provided by the AM fungal assemblages. These results indicate that interactions between assemblages of beneficial and pathogenic microorganisms can influence the growth of host plants, but that the magnitude of these effects is plant species-specific.     

Arbuscular mycorrhizal fungal communities change among three stages of primary sand dune succession but do not alter plant growth

Plant interactions with soil biota could have a significant impact on plant successional trajectory by benefiting plants in a particular successional stage over others. The influence of soil mutualists such as mycorrhizal fungi is thought to be an important feedback component, yet they have shown benefits to both early and late successional plants that could either retard or accelerate succession. Here we first determine if arbuscular mycorrhizal (AM) fungi differ among three stages of primary sand dune succession and then if they alter growth of plants from particular successional stages. We isolated AM fungal inoculum from early, intermediate or late stages of a primary dune succession and compared them using cloning and sequencing. We then grew eight plant species that dominate within each of these successional stages with each AM fungal inoculum. We measured fungal growth to assess potential AM functional differences and plant growth to determine if AM fungi positively or negatively affect plants. AM fungi isolated from early succession were more phylogenetically diverse relative to intermediate and late succession while late successional fungi consistently produced more soil hyphae and arbuscules. Despite these differences, inocula from different successional stages had similar effects on the growth of all plant species. Host plant biomass was not affected by mycorrhizal inoculation relative to un‐inoculated controls. Although mycorrhizal communities differ among primary dune successional stages and formed different fungal structures, these differences did not directly affect the growth of plants from different dune successional stages in our experiment and therefore may be less likely to directly contribute to plant succession in sand dunes.     

Host plant species effects on arbuscular mycorrhizal fungal communities in tallgrass prairie

Symbiotic associations between plants and arbuscular mycorrhizal (AM) fungi are ubiquitous in many herbaceous plant communities and can have large effects on these communities and ecosystem processes. The extent of species-specificity between these plant and fungal symbionts in nature is poorly known, yet reciprocal effects of the composition of plant and soil microbe communities is an important assumption of recent theoretical models of plant community structure. In grassland ecosystems, host plant species may have an important role in determining development and sporulation of AM fungi and patterns of fungal species composition and diversity. In this study, the effects of five different host plant species [Poa pratensis L., Sporobolus heterolepis (A. Gray) A. Gray, Panicum virgatum L., Baptisia bracteata Muhl. ex Ell., Solidago missouriensis Nutt.] on spore communities of AM fungi in tallgrass prairie were examined. Spore abundances and species composition of fungal communities of soil samples collected from patches within tallgrass prairie were significantly influenced by the host plant species that dominated the patch. The AM fungal spore community associated with B. bracteata showed the highest species diversity and the fungi associated with Pa. virgatum showed the lowest diversity. Results from sorghum trap cultures using soil collected from under different host plant species showed differential sporulations of AM fungal species. In addition, a greenhouse study was conducted in which different host plant species were grown in similar tallgrass prairie soil. After 4 months of growth, AM fungal species composition was significantly different beneath each host species. These results strongly suggest that AM fungi show some degree of host-specificity and are not randomly distributed in tallgrass prairie. The demonstration that host plant species composition influences AM fungal species composition provides support for current feedback models predicting strong regulatory effects of soil communities on plant community structure. Differential responses of AM fungi to host plant species may also play an important role in the regulation of species composition and diversity in AM fungal communities. Received: 29 January 1999 / Accepted: 20 October 1999     

Plant interspecific differences in arbuscular mycorrhizal colonization as a result of soil carbon addition

Soil nutrient availability and colonization by arbuscular mycorrhizal fungi are important and potentially interacting factors shaping vegetation composition and succession. We investigated the effect of carbon (C) addition, aimed at reducing soil nutrient availability, on arbuscular mycorrhizal colonization. Seedlings of 27 plant species with different sets of life-history traits (functional group affiliation, life history strategy and nitrophilic status) were grown in pots filled with soil from a nutrient-rich set-aside field and amended with different amounts of C. Mycorrhizal colonization was progressively reduced along the gradient of increasing C addition in 17 out of 27 species, but not in the remaining species. Grasses had lower colonization levels than forbs and legumes and the decline in AM fungal colonization was more pronounced in legumes than in other forbs and grasses. Mycorrhizal colonization did not differ between annual and perennial species, but decreased more rapidly along the gradient of increasing C addition in plants with high Ellenberg N values than in plants with low Ellenberg N values. Soil C addition not only limits plant growth through a reduction in available nutrients, but also reduces mycorrhizal colonization of plant roots. The effect of C addition on mycorrhizal colonization varies among plant functional groups, with legumes experiencing an overproportional reduction in AM fungal colonization along the gradient of increasing C addition. We therefore propose that for a better understanding of vegetation succession on set-aside fields one may consider the interrelationship between plant growth, soil nutrient availability and mycorrhizal colonization of plant roots.     

Plant–fungus competition for nitrogen erases mycorrhizal growth benefits of Andropogon gerardii under limited nitrogen supply

Considered to play an important role in plant mineral nutrition, arbuscular mycorrhizal (AM) symbiosis is a common relationship between the roots of a great majority of plant species and glomeromycotan fungi. Its effects on the plant host are highly context dependent, with the greatest benefits often observed in phosphorus (P)‐limited environments. Mycorrhizal contribution to plant nitrogen (N) nutrition is probably less important under most conditions. Moreover, inasmuch as both plant and fungi require substantial quantities of N for their growth, competition for N could potentially reduce net mycorrhizal benefits to the plant under conditions of limited N supply. Further compounded by increased belowground carbon (C) drain, the mycorrhizal costs could outweigh the benefits under severe N limitation. Using a field AM fungal community or a laboratory culture of Rhizophagus irregularis as mycorrhizal inoculants, we tested the contribution of mycorrhizal symbiosis to the growth, C allocation, and mineral nutrition of Andropogon gerardii growing in a nutrient‐poor substrate under variable N and P supplies. The plants unambiguously competed with the fungi for N when its supply was low, resulting in no or negative mycorrhizal growth and N‐uptake responses under such conditions. The field AM fungal communities manifested their potential to improve plant P nutrition only upon N fertilization, whereas the R. irregularis slightly yet significantly increased P uptake of its plant host (but not the host's growth) even without N supply. Coincident with increasing levels of root colonization by the AM fungal structures, both inoculants invariably increased nutritional and growth benefits to the host with increasing N supply. This, in turn, resulted in relieving plant P deficiency, which was persistent in non‐mycorrhizal plants across the entire range of nutrient supplies.     

Arbuscular Mycorrhizal Fungi Promote the Growth of Ceratocarpus arenarius (Chenopodiaceae) with No Enhancement of Phosphorus Nutrition

The mycorrhizal status of plants in the Chenopodiaceae is not well studied with a few controversial reports. This study examined arbuscular mycorrhizal (AM) colonization and growth response of Ceratocarpus arenarius in the field and a greenhouse inoculation trial. The colonization rate of AM fungi in C. arenarius in in-growth field cores was low (around 15%). Vesicles and intraradical hyphae were present during all growth stages, but no arbuscules were observed. Sequencing analysis of the large ribosomal rDNA subunit detected four culturable Glomus species, G. intraradices, G. mosseae, G. etunicatum and G. microaggregatum together with eight unculturable species belong to the Glomeromycota in the root system of C. arenarius collected from the field. These results establish the mycotrophic status of C. arenarius. Both in the field and in the greenhouse inoculation trial, the growth of C. arenarius was stimulated by the indigenous AM fungal community and the inoculated AM fungal isolates, respectively, but the P uptake and concentration of the mycorrhizal plants did not increase significantly over the controls in both experiments. Furthermore, the AM fungi significantly increased seed production. Our results suggest that an alternative reciprocal benefit to carbon-phosphorus trade-off between AM fungi and the chenopod plant might exist in the extremely arid environment.     

Adaptation of plant‐mycorrhizal interactions to moisture availability in prairie restoration

The strength and direction of plant response to inoculation with arbuscular mycorrhizal fungi (AM fungi) is dependent on both abiotic and biotic contexts, often generating patterns of AM fungal mediation of plant adaptation. However, knowledge of plant‐community level effects of these interactions in grassland restoration is limited. We conducted a field inoculation experiment by inoculating five plant species native to a drier prairie and five plant species native to a moister prairie with mycorrhizal fungal communities from each prairie type. Species were paired by genus or family to account for phylogenetic effects. The inoculated plants were transplanted to study plots seeded with a restoration seed mix. Plots were manipulated to create either moister or drier conditions similar to environments of the plant species and mycorrhizal communities. In both transplanted and seeded plant species, we found that only drier prairie‐range species benefited from moisture‐regime matched AM fungal inoculum. Other seeded prairie plant species demonstrated a negative response to inoculation, likely due to the earlier successional stage of these species. Additionally, nonseeded plants benefited from inoculation in different ways: native nonseeded plants had highest cover with drier prairie inoculum in drier conditions, while nonnative plants had highest cover with moister prairie‐origin inoculum. These results suggest that use of local AM fungi may be particularly important in restorations at drier sites, even at relatively small differences in moisture availability. Further, specific knowledge of relative responsiveness of seeded plant species and nonseeded plant species to AM fungal inoculation will be useful in planning restorations.     

Arbuscular mycorrhizal fungal species suppress inducible plant responses and alter defensive strategies following herbivory

In a greenhouse experiment using Plantago lanceolata, plants grown with different arbuscular mycorrhizal (AM) fungal species differed in constitutive levels of chemical defense depending on the species of AM fungi with which they were associated. AM fungal inoculation also modified the induced chemical response following herbivory by the specialist lepidopoteran herbivore Junonia coenia, and fungal species varied in how they affected induced responses. On average, inoculation with AM fungi substantially reduced the induced chemical response as compared with sterile controls, and inoculation with a mixture of AM fungi suppressed the induced response of P. lanceolata to herbivory. These results suggest that AM fungi can exert controlling influence over plant defensive phenotypes, and a portion of the substantial variation among experimental tests of induced chemical responses may be attributable to AM fungi.     

Patterns and regulation of mycorrhizal plant and fungal diversity

The diversity of mycorrhizal fungi does not follow patterns of plant diversity, and the type of mycorrhiza may regulate plant species diversity. For instance, coniferous forests of northern latitudes may have more than 1000 species of ectomycorrhizal (EM) fungi where only a few ectomycorrhizal plant species dominate, but there are fewer than 25 species of arbuscular mycorrhizal (AM) fungi in tropical deciduous forest in Mexico with 1000 plant species. AM and EM fungi are distributed according to biome, with AM fungi predominant in arid and semiarid biomes, and EM fungi predominant in mesic biomes. In addition, AM fungi tend to be more abundant in soils of low organic matter, perhaps explaining their predominance in moist tropical forest, and EM fungi generally occur in soils with higher surface organic matter.EM fungi are relatively selective of host plant species, while AM tend to be generalists. Similar morphotypes of AM fungi collected from different sites confer different physiological benefits to the same plant species. While the EM fungi have taxonomic diversity, the AM fungi must have physiological diversity for individual species to be so widespread, as supported by existing studies. The environmental adaptations of mycorrhizal fungi are often thought to be determined by their host plant, but we suggest that the physiology and genetics of the fungi themselves, along with their responses to the plant and the environment, regulates their diversity. We observed that one AM plant species,Artemisia tridentata, was associated with different fungal species across its range, indicating that the fungi can respond to the environment directly and must not do so indirectly via the host. Different species of fungi were also active during different times of the growing season on the same host, again suggesting a direct response to the environment.These patterns suggest that even within a single functional group of microorganisms, mycorrhizal fungi, considerable diversity exists. A number of researchers have expressed the concept of functional redundancy within functional groups of microorganisms, implying that the loss of a few species would not be detectable in ecosystem functioning. However, there may be high functional diversity of AM fungi within and across habitats, and high species diversity as well for EM fungi. If one species of mycorrhizal fungus becomes extinct in a habitat, field experimental data on AM fungi suggest there may be significant shifts in how plants acquire resources and grown in that habitat.     

Interactions between functionally diverse fungal mutualists inconsistently affect plant performance and competition

Plants form mutualistic relationship with a variety of belowground fungal species. Such a mutualistic relationship can enhance plant growth and resistance to pathogens. Yet, we know little about how interactions between functionally diverse groups of fungal mutualists affect plant performance and competition. We experimentally determined the effects of interaction between two functional groups of belowground fungi that form mutualistic relationship with plants, arbuscular mycorrhizal (AM) fungi and Trichoderma, on interspecific competition between pairs of closely related plant species from four different genera. We hypothesized that the combination of two functionally diverse belowground fungal species would allow plants and fungi to partition their symbiotic relationships and relax plant–plant competition. Our results show that: 1) the AM fungal species consistently outcompeted the Trichoderma species independent of plant combinations; 2) the fungal species generally had limited effects on competitive interactions between plants; 3) however, the combination of fungal species relaxed interspecific competition in one of the four instances of plant–plant competition, despite the general competitive superiority of AM fungi over Trichoderma. We highlight that the competitive outcome between functionally diverse fungal species may show high consistency across a broad range of host plants and their combinations. However, despite this consistent competitive hierarchy, the consequences of their interaction for plant performance and competition can strongly vary among plant communities.     

The differential behavior of arbuscular mycorrhizal fungi in interaction with Astragalus sinicus L. under salt stress

Three arbuscular mycorrhizal (AM) fungi (Glomus mosseae, Glomus claroideum, and Glomus intraradices) were compared for their root colonizing ability and activity in the root of Astragalus sinicus L. under salt-stressed soil conditions. Mycorrhizal formation, activity of fungal succinate dehydrogenase, and alkaline phosphatase, as well as plant biomass, were evaluated after 7 weeks of plant growth. Increasing the concentration of NaCl in soil generally decreased the dry weight of shoots and roots. Inoculation with AM fungi significantly alleviated inhibitory effect of salt stress. G. intraradices was the most efficient AM fungus compared with the other two fungi in terms of root colonization and enzyme activity. Nested PCR revealed that in root system of plants inoculated with a mix of the three AM fungi and grown under salt stress, the majority of mycorrhizal root fragments were colonized by one or two AM fungi, and some roots were colonized by all the three. Compared to inoculation alone, the frequency of G. mosseae in roots increased in the presence of the other two fungal species and highest level of NaCl, suggesting a synergistic interaction between these fungi under salt stress.     

亚热带草地中植物优势种与从属种对AM真菌的差异性生长反应

AM真菌能够影响植物生态系统的群落结构.以亚热带草地生态系统为研究对象,调查了两块草地中优势种和从属种的菌根,并在盆栽试验中比较了优势种和从属种对AM真菌的土著菌种和外源菌种Glomus mosseae的生长反应、养分吸收.结果表明,两块草地各自的优势种藿香蓟和两耳草对土著菌种的菌根依赖性分别是41.5%和77.4%,远远高于从属种莎草和毛蓼（16.0%和7.9%）;但是它们对Glomus mosseae的菌根依赖性有所变化,分别是79.6%、44.2%、74.1%和24.9%.这表明,土著菌种是优势种和从属种的形成机制之一,而外源菌种可能改变基于土著菌种而形成的植物群落结构.植株磷营养的分析结果表明,AM真菌对优势种和从属种生长的促进与对磷吸收的促进高度相关,表明AM真菌促进养分吸收是其影响植物群落结构的机制之一.     

The interactive effects of plant microbial symbionts: a review and meta-analysis

In nature, plants often associate with multiple symbionts concurrently, yet the effects of tripartite symbioses are not well understood. We expected synergistic growth responses from plants associating with functionally distinct symbionts. In contrast, symbionts providing similar benefits to a host may reduce host plant growth. We reviewed studies investigating the effect of multiple interactions on host plant performance. Additionally, we conducted a meta-analysis on the studies that performed controlled manipulations of the presence of two microbial symbionts. Using response ratios, we investigated the effects on plants of pairs of symbionts (mycorrhizal fungi, fungal endophytes, and nitrogen-fixers). The results did not support the view that arbuscular mycorrhizal (AM) fungi and rhizobia should interact synergistically. In contrast, we found the joint effects of fungal endophytes and arbuscular mycorrhizal fungi to be greater than expected given their independent effects. This increase in plant performance only held for antagonistic endophytes, whose negative effects were alleviated when in association with AM fungi, while the impact of beneficial endophytes was not altered by infection with AM fungi. Generalizations from the meta-analysis were limited by the substantial variation within types of interactions and the data available, highlighting the need for more research on a range of plant systems.     

Mycorrhizal colonization and drought stress affect Δ13C in CO2‐labeled lettuce plants

We studied the role of different arbuscular‐mycorrhizal (AM) fungi on lettuce (Lactuca sativa L.) plant carbon metabolism under drought stress. Plants were grown in pots maintained at two levels of soil moisture and labeled during photosynthesis with CO₂. P‐fertilized plants were used as a non‐mycorrhizal control. Well‐watered mycorrhizal plants showed similar growth to that of P‐fertilized plants. The level of mycorrhizal root infection was not significantly affected by fungal species or by water treatment. In contrast, important differences in Δ¹³C between P‐fertilized and AM plants were found in shoot and root tissues as a consequence of both water limitation and fungal presence. Δ¹³C in shoots and roots increased in non‐mycorrhizal treatment as compared with the well‐watered plants, whereas this parameter decreased significantly in mycorrhizal plants. Photosynthetic activity was increased in AM plants in well‐watered and droughted plants. G. deserticola was the most beneficial endophyte for water use efficiency in both water treatments. Transpiration rate was not affected by any of the treatments. On the basis of total C in plant tissues, in AM plants the newly fixed C seemed to be preferentially utilized for fungal activity rather than being stored in roots.     

Availability and function of arbuscular mycorrhizal and ectomycorrhizal fungi during revegetation of dewatered reservoirs left after dam removal

Revegetation following dam removal projects may depend on recovery of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal communities, which perform valuable ecosystem functions. This study assessed the availability and function of AM and EM fungi for plants colonizing dewatered reservoirs following a dam removal project on the Elwha River, Olympic Peninsula, Washington, United States. Availability was assessed via AM fungal spore density in soils and EM root tip colonization of Salix sitchensis (Sitka willow) in an observational field study. The effect of mycorrhizal fungi from 4 sources (reservoir soils, commercial inoculum, and 2 mature plant community soils) on growth and nutrient status of S. sitchensis was quantified in a greenhouse study. AM fungal spores and EM root tips were present in all field samples. In the greenhouse, plants receiving reservoir soil inoculum had only incipient mantle formation, while plants receiving inoculum from mature plant communities had fully formed EM root tips. EM formation corresponded with alleviation of phosphorus stress in plants (lower shoot nitrogen:phosphorus). Thus, revegetating plants have access to AM and EM fungi following dam removal, and EM formation may be especially important for plant P uptake in reservoir soils. However, availability of mycorrhizal fungi declines with distance from established plant communities. Furthermore, EM fungal communities in recently dewatered reservoirs may not be as effective at forming beneficial mycorrhizae as those from mature plant communities. Whole soil inoculum from mature plant communities may be important for the success of revegetating plants and recovery of mycorrhizal fungal communities.