Plants interact simultaneously with each other and with soil biota, yet the relative importance of competition vs. plant-soil feedback (PSF) on plant performance is poorly understood. Using a meta-analysis of 38 published studies and 150 plant species, we show that effects of interspecific competition (either growing plants with a competitor or singly, or comparing inter- vs. intraspecific competition) and PSF (comparing home vs. away soil, live vs. sterile soil, or control vs. fungicide-treated soil) depended on treatments but were predominantly negative, broadly comparable in magnitude, and additive or synergistic. Stronger competitors experienced more negative PSF than weaker competitors when controlling for density (inter- to intraspecific competition), suggesting that PSF could prevent competitive dominance and promote coexistence. When competition was measured against plants growing singly, the strength of competition overwhelmed PSF, indicating that the relative importance of PSF may depend not only on neighbour identity but also density. We evaluate how competition and PSFs might interact across resource gradients; PSF will likely strengthen competitive interactions in high resource environments and enhance facilitative interactions in low-resource environments. Finally, we provide a framework for filling key knowledge gaps and advancing our understanding of how these biotic interactions influence community structure.
Summary1. Divergent hypotheses have been proposed that suggest plant invasions either enhance or degrade the mutualism between plants and arbuscular mycorrhizal (AM) fungi, but their relative support remains unknown. 2. We conducted a meta-analysis using 67 publications, involving 70 native and 55 invasive plant species to assess support for the enhanced mutualism hypothesis, the degraded mutualism hypothesis and an alternative hypothesis that factors other than invasive status (such as plant functional group) better predict AM function following invasion. We used multiple measurements to test these hypotheses: AM fungal colonization, growth responses to AM fungi and AM fungal-mediated shifts in competitive interactions among native and invasive plants. Additionally, we assessed whether invasive plants alter AM associations in native plants and whether native and invasive plants host different AM fungal abundances and communities. 3. Arbuscular mycorrhizal fungal colonization (%) and average growth responses did not differ between native and invasive plants. However, growth responses (AE) were dampened among invasive plants, and the positive correlation between AM fungal colonization and growth response in native plants was absent in invasive plants. Rather than plant invasive status, plant functional group was a significant explanatory factor; forbs were generally more colonized and exhibited positive growth responses (when grown alone and in competition), whereas grass responses were neutral to negative. 4. Arbuscular mycorrhizal fungal abundance (measured by percentage colonization, extraradical hyphal and spore densities, as well as neutral lipid fatty acid and glomalin concentrations) did not differ between native and invasive plants, but invasive plants hosted different AM fungal communities in 78% of studies. AM fungal colonization of native plants was lower when grown with, or after, invasive plants, likely due to the prevalence of non-mycorrhizal plants in studies of neighbour and legacy effects. 5. Synthesis. Neither the degraded nor the enhanced mutualism hypothesis was supported, suggesting that invasions do not select for directional shifts in AM associations. Instead, our results indicate that AM fungi are most likely to influence invasion trajectories when native and invasive plants belong to different functional groups.
Biotic interactions can affect the distribution of species across environmental gradients, and as air and soil temperatures increase, plant community response may depend on interactions with symbionts. We measured the effect of elevated soil temperatures on mycorrhizal function and on the response of both plant and fungal symbionts, using fungal inoculum isolated from either high-temperature thermal or nonthermal grassland soils. Our source for thermal soils was Yellowstone National Park, USA, where plants experience rooting zone temperatures of 45 degrees C or more. In the greenhouse, we grew three plant species (Dichanthelium lanuginosum, Agrostis scabra, and Mimulus guttatus) with three arbuscular mycorrhizal fungal (AMF) treatments (no AMF, nonthermal AMF, thermal AMF) and two soil temperatures (ambient, elevated). Biomass of the facultative thermal plants Agrostis scabra and Mimulus guttatus decreased by 50% in elevated-temperature soils, and AMF had no effect on measured plant traits. In contrast, the biomass and total root length of the obligate thermal plant Dichanthelium lanuginosum were greater at elevated soil temperatures, but only when mycorrhizal. Both mycorrhizal colonization levels and length of extraradical hyphae (ERH) increased with soil temperature across all host species. The source of the AMF inoculum, on the other hand, did not affect colonization level, ERH length, host plant biomass, or flowering for all host species in either temperature treatment, suggesting that AMF from thermal soils are not specifically adapted to higher temperatures. In the field we collected soil cores to measure in situ depth distributions of D. lanuginosum roots and ERH, and to determine which AMF species were active in plants growing in thermal soils. Roots were limited to soils with an average temperature < or =30 degrees C, while ERH existed in the hottest soils we sampled, averaging 35 degrees C. Molecular analyses of roots indicated that thermal AMF communities were composed of both generalist and possibly unique fungal species. The increase in host plant allocation to AMF, apparent lack of temperature adaptation by AMF, and differential host response to AMF suggest that AMF could be significant drivers of plant community response to increased soil temperature associated with global change.
Field and laboratory column experiments were performed to assess the effect of elevated pH and reduced ionic strength on the mobilization of natural colloids in a ferric oxyhydroxide-coated aquifer sediment. The field experiments were conducted as natural gradient injections of groundwater amended by sodium hydroxide additions. The laboratory experiments were conducted in columns of undisturbed, oriented sediments and disturbed, disoriented sediments. In the field, the breakthrough of released colloids coincided with the pH pulse breakthrough and lagged the bromide tracer breakthrough. The breakthrough behavior suggested that the progress of the elevated pH front controlled the transport of the mobilized colloids. In the laboratory, about twice as much colloid release occurred in the disturbed sediments as in the undisturbed sediments. The field and laboratory experiments both showed that the total mass of colloid release increased with increasing pH until the concurrent increase in ionic strength limited release. A decrease in ionic strength did not mobilize significant amounts of colloids in the field. The amount of colloids released normalized to the mass of the sediments was similar for the field and the undisturbed laboratory experiments.
Controlled experiments show that arbuscular mycorrhizal fungi (AMF) can increase competitiveness of exotic plants, potentially increasing invasion success. We surveyed AMF abundance and community composition in Centaurea stoebe and Potentilla recta invasions in the western USA to assess whether patterns were consistent with mycorrhizal-mediated invasions. We asked whether (1) AMF abundance and community composition differ between native and exotic forbs, (2) associations between native plants and AMF shift with invading exotic plants, and (3) AMF abundance and/or community composition differ in areas where exotic plants are highly invasive and in areas where they are not. We collected soil and roots from invaded and native forb communities along invasion gradients and in regions with different invasion densities. We used AMF root colonization as a measure of AMF abundance and characterized AMF communities in roots using 454-sequencing of the LSU-rDNA region. All plants were highly colonized (>60%), but exotic forbs tended to be more colonized than natives (P < 0.001). We identified 30 AMF operational taxonomic units (OTUs) across sites, and community composition was best predicted by abiotic factors (soil texture, pH). Two OTUs in the genera Glomus and Rhizophagus dominated in most communities, and their dominance increased with invasion density (r = 0.57, P = 0.010), while overall OTU richness decreased with invasion density (r = −0.61, P = 0.006). Samples along P. recta invasion gradients revealed small and reciprocal shifts in AMF communities with >45% fungal OTUs shared between neighboring native and P. recta plants. Overall, we observed significant, but modest, differences in AMF colonization and communities between co-occurring exotic and native forbs and among exotic forbs across regions that differ in invasion pressure. While experimental manipulations are required to assess functional consequences, the observed patterns are not consistent with those expected from strong mycorrhizal-mediated invasions.
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