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Plant communities can be affected both by arbuscular mycorrhizal fungi (AMF) and hemiparasitic plants. However, little is known about the interactive effects of these two biotic factors on the productivity and diversity of plant communities. To address this question, we set up a greenhouse study in which different AMF inocula and a hemiparasitic plant ( Rhinanthus minor) were added to experimental grassland communities in a fully factorial design. In addition, single plants of each species in the grassland community were grown with the same treatments to distinguish direct AMF effects from indirect effects via plant competition. We found that AMF changed plant community structure by influencing the plant species differently. At the community level, AMF decreased the productivity by 15–24%, depending on the particular AMF treatment, mainly because two dominant species, Holcus lanatus and Plantago lanceolata, showed a negative mycorrhizal dependency. Concomitantly, plant diversity increased due to AMF inoculation and was highest in the treatment with a combination of two commercial AM strains. AMF had a positive effect on growth of the hemiparasite, and thereby induced a negative impact of the hemiparasite on host plant biomass which was not found in non-inoculated communities. However, the hemiparasite did not increase plant diversity. Our results highlight the importance of interactions with soil microbes for plant community structure and that these indirect effects can vary among AMF treatments. We conclude that mutualistic interactions with AMF, but not antagonistic interactions with a root hemiparasite, promote plant diversity in this grassland community. [ABSTRACT FROM AUTHOR]

Nitrogen isotopes (15N/14N ratios, expressed as δ15N values) are useful markers of the mycorrhizal role in plant nitrogen supply because discrimination against 15N during creation of transfer compounds within mycorrhizal fungi decreases the 15N/14N in plants (low δ15N) and increases the 15N/14N of the fungi (high δ15N). Analytical models of 15N distribution would be helpful in interpreting δ15N patterns in fungi and plants. To compare different analytical models, we measured nitrogen isotope patterns in soils, saprotrophic fungi, ectomycorrhizal fungi, and plants with different mycorrhizal habits on a glacier foreland exposed during the last 100 years of glacial retreat and on adjacent non-glaciated terrain. Since plants during early primary succession may have only limited access to propagules of mycorrhizal fungi, we hypothesized that mycorrhizal plants would initially be similar to nonmycorrhizal plants in δ15N and then decrease, if mycorrhizal colonization were an important factor influencing plant δ15N. As hypothesized, plants with different mycorrhizal habits initially showed similar δ15N values (−4 to −6‰ relative to the standard of atmospheric N2 at 0‰), corresponding to low mycorrhizal colonization in all plant species and an absence of ectomycorrhizal sporocarps. In later successional stages where ectomycorrhizal sporocarps were present, most ectomycorrhizal and ericoid mycorrhizal plants declined by 5–6‰ in δ15N, suggesting transfer of 15N-depleted N from fungi to plants. The values recorded (−8 to −11‰) are among the lowest yet observed in vascular plants. In contrast, the δ15N of nonmycorrhizal plants and arbuscular mycorrhizal plants declined only slightly or not at all. On the forefront, most ectomycorrhizal and saprotrophic fungi were similar in δ15N (−1 to −3‰), but the host-specific ectomycorrhizal fungus Cortinarius tenebricus had values of up to 7‰. Plants, fungi and soil were at least 4‰ higher in δ15N from the mature site than in recently exposed sites. On both the forefront and the mature site, host-specific ectomycorrhizal fungi had higher δ15N values than ectomycorrhizal fungi with a broad host range. From these isotopic patterns, we conclude:(1) large enrichments in 15N of many ectomycorrhizal fungi relative to co-occurring ectomycorrhizal plants are best explained by treating the plant-fungal-soil system as a closed system with a discrimination against 15N of 8–10‰ during transfer from fungi to plants, (2) based on models of 15N mass balance, ericoid and ectomycorrhizal fungi retain up to two-thirds of the N in the plant-mycorrhizal system under the N-limited conditions at forefront sites, (3) sporocarps are probably enriched in 15N by an additional 3‰ relative to available nitrogen, and (4) host-specific ectomycorrhizal fungi may transfer more N to plant hosts than non-host-specific ectomycorrhizal fungi. Our study confirms that nitrogen isotopes are a powerful tool for probing nitrogen dynamics between mycorrhizal fungi and associated plants. [ABSTRACT FROM AUTHOR]

Extreme arctic-alpine vegetation has relatively low affinity to form mycorrhizal symbiosis. We asked whether the mycorrhizal growth benefit for the host plant is lower at low temperatures. We investigated the role of two root-associated fungi and temperature in growth, carbon–nitrogen relations and germination of an arctic-alpine herb. Seeds of Gnaphalium norvegicum were germinated at 8° or 15show $132#°show $132#C with or without arbuscular mycorrhizal (AM, Glomus claroideum) and dark septate endophytic (DSE, Phialocephala fortinii) inocula in a climate chamber. We found that germination percentage, shoot and root biomass, shoot N% and root AM colonization were lower at 8show $132#°show $132#C than at 15show $132#°show $132#C. P. fortinii inoculation had a positive impact on germination at both temperatures, whereas G. claroideum produced no effect. N% was lower in AM plants at both temperatures. Plant biomass and shoot N content were higher in AM plants than in control plants at 15show $132#°show $132#C, but not at 8show $132#°show $132#C. DSE inoculation tended also to have positive effects on plant biomass and N content at 15show $132#°show $132#C. At 15show $132#°show $132#C, rate of photosynthesis, photosynthetic nutrient use efficiency and specific leaf area were positively affected by G. claroideum, which suggests that G. claroideum formed a carbon sink and possibly enhanced the seedling water economy. The positive effects of P. fortinii were probably due to its saprotrophic function in the substrate because it did not colonize the roots. These results suggest that the effects of AM and DSE on plant growth are affected by temperature and that the mycorrhizal benefit for the host plant was lower at the lower temperature. Low saprotrophic activity and decreased mycorrhiza-mediated nutrient acquisition may thus constrain plant nutrient acquisition in cold environments. Decreased mycorrhizal benefit may be related to the comparatively low mycotrophy of cold environment vegetation. [ABSTRACT FROM AUTHOR]

A large number of herbaceous and woody plants from tropical woodland, savanna, and monsoon forest were analysed to determine the impact of environmental factors (nutrient and water availability, fire) and biological factors (microbial associations, systematics) on plant δ15N values. Foliar δ15N values of herbaceous and woody species were not related to growth form or phenology, but a strong relationship existed between mycorrhizal status and plant δ15N. In woodland and savanna, woody species with ectomycorrhizal (ECM) associations and putative N2-fixing species with ECM/arbuscular (AM) associations had lowest foliar δ15N values (1.0–0.6‰), AM species had mostly intermediate δ15N values (average +0.6‰), while non-mycorrhizal Proteaceae had highest δ15N values (+2.9 to +4.1‰). Similar differences in foliar δ15N were observed between AM (average 0.1 and 0.2‰) and non-mycorrhizal (average +0.8 and +0.3‰) herbaceous species in woodland and savanna. Leguminous savanna species had significantly higher leaf N contents (1.8–2.5% N) than non-fixing species (0.9–1.2% N) indicating substantial N acquisition via N2 fixation. Monsoon forest species had similar leaf N contents (average 2.4% N) and positive δ15N values (+0.9 to +2.4‰). Soil nitrification and plant NO3- use was substantially higher in monsoon forest than in woodland or savanna. In the studied communities, higher soil N content and nitrification rates were associated with more positive soil δ15N and plant δ15N. In support of this notion, Ficus, a high NO3- using taxa associated with NO3- rich sites in the savanna, had the highest δ15N values of all AM species in the savanna. δ15N of xylem sap was examined as a tool for studying plant δ15N relations. δ15N of xylem sap varied seasonally and between differently aged Acacia and other savanna species. Plants from annually burnt savanna had significantly higher δ15N values compared to plants from less frequently burnt savanna, suggesting that foliar 15N natural abundance could be used as marker for assessing historic fire regimes. Australian woodland and savanna species had low leaf δ15N and N content compared to species from equivalent African communities indicating that Australian biota are the more N depauperate. The largest differences in leaf δ15N occurred between the dominant ECM Australian and African savanna (miombo) species, which were depleted and enriched in 15N, respectively. While the depleted δ15N of Australian ECM species are similar to those of previous reports on ECM species in natural plant communities, the 15N-enriched δ15N of African ECM species represent an anomaly. [ABSTRACT FROM AUTHOR]

Although variation in the effectiveness of the plant-mycorrhizal association has been demonstrated among plant species and among crop cultivars, the level of specificity of the plant-mycorrhizal association in natural populations and, in particular, its effect on intraspecific plant interactions have not been explored. To examine the influence of the specificity of the plant-mycorrhizal association on intraspecific interactions in Allium vineale, we surrounded target plants with either genetically identical neighbors, neighbors from the same population, or neighbors from a different population, and planted them in pots either with or without a soil fungi community. Overall, the presence of a soil fungal community was beneficial for plant growth, and thus the soil fungal community effects in this system are dominated by the mutualistic effect of mycorrhizae. In addition, both the target plant and neighbor plants had a relatively greater benefit from the mycorrhizal association if the neighbors were genetically identical or from the same population than if the neighbors came from a different population. Thus, there is specificity in the interaction between A. vineale plants and the soil fungal community at the population level, and this specificity favors intraspecific interactions among plants from the same population. In addition, in the presence of the soil fungal community, the variance in plant size within a pot was greater among neighbors from a different population. Therefore the growth advantage for plants growing with neighbors from the same population was not due to pre-emption of resources by individual plants. This finding provides indirect evidence to support the hypothesis that plants from the same population were able to share a more efficient hyphal network. [ABSTRACT FROM AUTHOR]

Mycorrhizal plants are commonly believed to direct much more of their photosynthates into the soil than non-mycorrhizal plants. As the growth of most organisms of the detrital food web is limited by energy, the flow of C through mycorrhizal plants into the below-ground milieu is widely assumed to nourish a variety of decomposer organisms in soils. In the current experiment, I explored whether some representatives of soil mesofauna, either fungivores or microbi-detritivores, derive benefit from the presence of ectomycorrhizal (EM) fungi growing on the roots of Scots pine (Pinus sylvestris). I also investigated whether the role of soil mesofauna in affecting pine growth depends on the presence of EM fungi in the pine rhizosphere. The study was established in microcosms with a mixture of raw humus and sand. The soil was defaunated, reinoculated with 10 species of soil bacteria and 11 species of saprophytic soil fungi, and pine seedlings, either infected or non-infected with four taxa of EM fungi, were planted in the microcosms. Five treatments with different food web configurations were established: (1) saprophytic microbes alone, (2) as (1) but with the omnivorous enchytraeid species Cognettia sphagnetorum present, (3) as (1) but with Collembola (Hypogastrura assimilis), (4) as (1) but with four species of oribatid mites (Acari) involved, and (5) as 1) but with C. sphagnetorum, H. assimilis and the Acari. The microcosms were incubated in a climate chamber with varying temperature and illumination regimes for two growing periods for the pine. After 60 weeks, pine biomass production was significantly greater in the mycorrhizal systems, the total biomass being 1.43 times higher in the presence than absence of EM fungi. Similarly, almost ten times more fungal biomass was detected on pine roots growing in the mycorrhizal than in the non-mycorrhizal systems. The presence of EM fungi was also associated with significantly lowered pH and percent organic matter of the soil. Despite the clearly larger biomass of both the pines and the fungi on the pine roots, neither the numbers nor biomasses of the mesofauna differed significantly between the EM and non-EM systems. The presence of Collembola and C. sphagnetorum had a positive influence on pine growth, particularly in the absence of EM fungi, whereas oribatid mites had no effects on pine growth. The complexity of the mesofaunal community was not related to the biomass production of the pines in a straightforward manner; for example, the complex systems with each faunal group present did not produce more pine biomass than the simple systems where C. sphagnetorum existed alone. The results of this experiment suggest that the short-term role of EM fungi in fuelling the detrital food web is less significant than generally considered, but that their role as active decomposers and/or stimulators of the activity of saprophytic microbes can be more important than is often believed. [ABSTRACT FROM AUTHOR]