Arbuscular mycorrhizal (AM) fungi are integral components of grasslands because most plants are associated with interconnected networks of AM hyphae. Mycorrhizae generally facilitate plant uptake of nutrients from the soil. However, mycorrhizal associations are known to vary in their mutualistic function, and there is currently no metric that links AM functioning with fungal colonization of roots. Mycorrhizal structures differ in their physiological and ecological functioning, so changes in AM allocation to intraradical (inside roots) and extraradical (in soil) structures may signal shifts in mycorrhizal function. We hypothesize that the functional equilibrium model applies to AM fungi and that fertilization should reduce allocation to arbuscules, coils, and extraradical hyphae, the fungal structures that are directly involved in nutrient acquisition and transfer to plants. This study compared AM responses to experimental N enrichment at five grasslands distributed across North America. Samples were collected from replicated N‐enriched (and some P‐enriched) and control plots throughout the growing season for three years. Intraradical AM structures were measured in over 1400 root samples, extraradical hyphal density was measured in over 590 soil samples, and spore biovolume was analyzed in over 400 soil samples. There were significant site × N interactions for spore biovolume, extraradical hyphae, intraradical hyphae, and vesicles. Nitrogen enrichment strongly decreased AM structures at Cedar Creek, the site with the lowest soil N:P, and it increased AM structures at Konza Prairie, the site with the highest soil N:P. As predicted by the functional equilibrium model, in soils with sufficient P, relative allocation to arbuscules, coils, and extraradical hyphae was generally reduced by N enrichment. Allocation to spores and hyphae was most sensitive to fertilization. At the mesic sites, this response was associated with a shift in the relative abundance of Gigasporaceae within AM fungal communities. This study demonstrates that N enrichment impacts mycorrhizal allocation across a wide range of grassland ecosystems. Such changes are important because they suggest an alteration in mycorrhizal functioning that, in turn, may impact plant community composition and ecosystem function.

Human activities release tremendous amounts of nitrogenous compounds into the atmosphere. Wet and dry deposition distributes this airborne nitrogen (N) on otherwise pristine ecosystems. This eutrophication process significantly alters the species composition of native grasslands; generally a few nitrophilic plant species become dominant while many other species disappear. The functional equilibrium model predicts that, compared to species that decline in response to N enrichment, nitrophilic grass species should respond to N enrichment with greater biomass allocation aboveground and reduced allocation to roots and mycorrhizas. The mycorrhizal feedback hypothesis states that the composition of mycorrhizal fungal communities may influence the composition of plant communities, and it predicts that N enrichment may generate reciprocal shifts in the species composition of mycorrhizal fungi and plants. We tested these hypotheses with experiments that compared biomass allocation and mycorrhizal function of four grass ecotypes (three species), two that gained and two that lost biomass and cover in response to long-term N enrichment experiments at Cedar Creek and Konza Long-Term Ecological Research grasslands. Local grass ecotypes were grown in soil from their respective sites and inoculated with whole-soil inoculum collected from either fertilized (FERT) or unfertilized (UNFERT) plots. Our results strongly support the functional equilibrium model. In both grassland systems the nitrophilic grass species grew taller, allocated more biomass to shoots than to roots, and formed fewer mycorrhizas compared to the grass species that it replaced. Our results did not fully support the hypothesis that N-induced changes in the mycorrhizal fungal community were drivers of the plant community shifts that accompany N eutrophication. The FERT and UNFERT soil inoculum influenced the growth of the grasses differently, but this varied with site and grass ecotype in both expected and unexpected ways suggesting that ambient soil fertility or other factors may be interacting with mycorrhizal feedbacks.

Disturbance of vegetation and soil may change the species composition of arbuscular mycorrhizal fungi (AMF), which may in turn affect plant species responses to AMF. Seasonal tropical forest in Mexico is undergoing rapid conversion to early-successional forest because of increased wildfire and may require restoration. The responses of six early-and late-successional tree species were tested using early-and late-successional AMF inoculum. The plants were germinated in the shadehouse and received three inoculum treatments: (1) soil from a two-year-old burned site, (2) soil from a mature forest site, or (3) uninoculated controls. They were transplanted as seedlings to a site prepared by burning, and their growth was measured from September 1997 to November 2000. All six species had the greatest growth response to early-seral inoculum, but the response to late-seral inoculum varied. Two tree species, Ceiba pentandra and Guazuma ulmifolia, were smallest with late-seral inoculum, even smaller than the uninoculated plants, and the other species, Brosimum alicastrum, Havardia albicans, Acacia pennatula, and Leucaena leucocephala, had intermediate growth with late-seral inoculum. Of these, Brosimum, Havardia, and Ceiba occur in late-successional forest, and the others are early seral. Of the several growth measurements (height, cover, biomass, stem diameter), stem-diameter responses to inoculum were still significantly different into the third year for four of the species. The uninoculated plants became infected by residual inoculum in the burned experimental site within three months of transplanting, yet mycorrhizal responses persisted. The treatment size differences may be due to different species composition of the inocula. The early-seral inoculum was dominated by small-spored Glomus spp., while the late-seral inoculum had a higher density of large-spored Gigasporaceae. The latter are known from greenhouse experiments to promote a smaller plant-growth response than Glomus. Mature forest trees may withstand the carbon drain from Gigasporaceae better than establishing seedlings, so the growth patterns we observed with inoculum source are consistent with a rapidly growing successional forest, followed by slower-growing mature forest. The results suggest that early-seral AMF should be used when seedlings are inoculated for restoration, even for late-seral tree species.

No abstract

No abstract