Differential modulation of host plant delta13C and delta18O by native and nonnative arbuscular mycorrhizal fungi in a semiarid environment.

Native, drought-adapted arbuscular mycorrhizal fungi (AMF) often improve host-plant performance to a greater extent than nonnative AMF in dry environments. However, little is known about the physiological basis for this differential plant response. Seedlings of Olea europaea and Rhamnus lycioides were inoculated with either a mixture of eight native Glomus species or with the nonnative Glomus claroideum before field transplanting in a semiarid area. Inoculation with native AMF produced the greatest improvement in nutrient and water status as well as in long-term growth for both Olea and Rhamnus. Foliar delta18O measurements indicated that native AMF enhanced stomatal conductance to a greater extent than nonnative AMF in Olea and Rhamnus.delta13C data showed that intrinsic water-use efficiency in Olea was differentially stimulated by native AMF compared with nonnative AMF. Our results suggest that modulation of leaf gas exchange by native, drought-adapted AMF is critical to the long-term performance of host plants in semiarid environments. delta18O can provide a time-integrated measure of the effect of mycorrhizal infection on host-plant water relations.

Species-specific measurements of ectomycorrhizal turnover under N-fertilization: combining isotopic and genetic approaches.

Ectomycorrhizal fungi play a significant role in the transfer of nutrients between plant and soil pools. Here we combine natural abundance (14)C measurements with restriction fragment length polymorphism (RFLP) to study the effects of nitrogen fertilization on the residence time of carbon within ectomycorrhizal species. We show that the carbon in ectomycorrhizal fungi turns over every 4-5 years, indicating that these fungi are relatively long-lived. Moreover, ectomycorrhizal fungi responded in a species-specific way to fertilization. Cenococcum geophilum contained younger carbon on average in nitrogen-fertilized plots than in control plots, even though turnover rates of the community as a whole did not shift significantly. Our results suggest that the response of ectomycorrhizal fungi to N availability is complex, and alterations in tissue turnover within this microbial pool may vary depending on community structure.

Ecology of mycorrhizae: a conceptual framework for complex interactions among plants and fungi.

Mycorrhizae regulate elemental and energy flows in terrestrial ecosystems. We understand much of how mycorrhizae work, but integrating all possible mechanisms into a whole has proven elusive. Multiple evolutionary events and the long evolutionary history mean that different plants and fungi bring independent characteristics to the symbiosis. This variety results in extensive physiological variation. How do we integrate functional responses with diversity to understand the role of mycorrhizae in ecosystems? We review ecophysiological mechanisms of mycorrhizae and organize these into functional groups. Species-area relationships are not curvilinear, but resemble the "broken stick" model. We coupled functional groups with a metacommunity analysis to show how complex behavior can be generated using a simple matrix model of resource exchange. This approach provides insights into how we might integrate diversity and function across landscapes.

Reconstruction of the historical changes in mycorrhizal fungal communities under anthropogenic nitrogen deposition.

Archived soil samples (1937-1999) and historic air quality data from the Los Angeles Basin were used for reconstructing the record of change between atmospheric NO(x) loads, soil delta(15)N values and the diversity of arbuscular mycorrhizae (AM), which are ubiquitous plant-fungus mutualists that control plant community productivity. A tripling of atmospheric NO(x) loads between 1937 and the 1970s was paralleled by soil nitrogen enrichment (delta(15)N = 3.18). From 1975 onwards, atmospheric NO(x) declined, but soils became nitrogen saturated (delta(15) N = -4 and NO(3)-nitrogen = 171mgkg(-1)). The shifts in the AM community followed 28 years of atmospheric nitrogen enrichment and coincided with the onset of soil nitrogen saturation. Such changes were manifest in the loss of AM productivity, species richness (one species per year), three genera (Acaulospora, Scutellospora and Gigaspora) in the spore community and Gigaspora within the roots. Nitrogen enrichment also enhanced the proliferation of potentially less mutualistic species of Glomus. Autoregressive models implied that such patterns will persist and be driven by soil nitrogen cycling patterns. Chronic nitrogen enrichment from air pollution thus alters the diversity and mutualistic functioning of AM communities, which, in turn, may influence the plant community.

Endo- and ectomycorrhizas in Quercus agrifolia Nee. (Fagaceae): patterns of root colonization and effects on seedling growth.

We documented the patterns of root occupancy by Glomalean and ectomycorrhizal (EM) fungi in Quercus agrifolia, and host plant responses to inoculation with each mycorrhizal type alone or in combination. Glomalean hyphae, coils and vesicles, and EM root tips were recorded. Colonization patterns conformed to a succession from Glomalean and EM fungi in 1-year-old seedlings to predominantly EM in saplings (>11 years old); both mycorrhizal types were rarely detected within the same root segment. Inoculation of Q. agrifolia seedlings with EM or Glomalean fungi (AM) alone or in combination (EM+AM) altered the cost:benefit relationship of mycorrhizas to the host plant. Seedling survival, plant biomass, foliar nitrogen (N), and phosphorus (P) status were greatest in EM- or AM-only inoculated seedlings. Seedlings inoculated with both mycorrhizal types (AM+EM) exhibited the lowest survival rates, biomass, foliar N, and P levels. Roots of these plants were highly colonized by both EM (38% root length colonized) and Glomalean fungi (34%). Because these levels of colonization were similar to those detected in 1-year-old field seedlings, the presence of both mycorrhizal types may be a carbon cost and, in turn, less beneficial to oaks during establishment in the field. However, the shift to EM colonization in older plants suggests that mycorrhizal effects may become positive with time.

The effects of oxalates produced by Salsola tragus on the phosphorus nutrition of Stipa pulchra.

Oxalic acid is produced by some species of plants and mycorrhizal fungi and it may solubilize unavailable soil phosphorus (P) bound by cations (Ca++, Al++, Fe+++). Field and greenhouse experiments were conducted to show whether oxalate produced by the annual Salsola tragus or added oxalic acid would solubilize P from the inorganic-bound soil P pool, making it available for uptake by Stipa pulchra. Oxalate could be leached in the laboratory from the senescent canopy of Salsola, and leaching by rainfall was hypothesized to be a source of potential increased soil P under the Salsola canopy. Both oxalate leached from the canopy of Salsola and added oxalic acid increased the availability of soil P in greenhouse experiments. The source of the increase in available soil P in the greenhouse experiment was shown to be the inorganic-bound P pool, as the total P concentration of the soil decreased with increasing oxalate. There were significant increases in Stipa shoot P in response to Salsola leachates and in response to added oxalate in the greenhouse studies. These results suggest an important role for oxalate in P cycling. On disturbed sites where Salsola invades it may act to facilitate the establishment of later seral species like Stipa by creating a nutrient island of available P.

Temperature and structural effects on transfer of double-stranded RNA among isolates of the chestnut blight fungus (Cryphonectria parasitica).

Cryphonectria parasitica is a unique fungus which can serve as a model for understanding transfer of genes between eukaryotic microorganisms. We studied transfer of double-stranded RNA (dsRNA) between compatible and incompatible strains of C. parasitica to determine whether hyphal types or temperature could restrict that exchange. Hyphal connections between incompatible strains occurred at about 30% of the frequency of connections between compatible strains and differed morphologically. Gel electrophoresis and in situ hybridization confirmed that dsRNA was transferred through substrate hyphae but not through aerial hyphae. Freezing temperatures resulted in the loss of dsRNA from the new mycelium of the donor colony and stimulated the production of virulent pycnidiospores. These temperature and structural restrictions may help to explain the lack of spread of the dsRNA despite its presence in the field.

WATER RELATIONS OF XERIC GRASSES IN THE FIELD: INTERACTIONS OF MYCORRHIZAS AND COMPETITION.

The effects of vesicular-arbuscular mycorrhizal inoculum and competition from non-mycorrhizal annual plants were tested on Agropyron smithii Rydb. and Agropyron dasystachyum (Hook.) Scribn. growing in an inoculum-poor soil. Mycorrhizal A. smithii decreased stomatal resistance (rs ) and increased leaf water potential compared to the non-inoculated plants, but only during the driest portion of the growing season. The presence of annuals caused increased rs of A. smithii, but only of non-inoculated plants. Mycorrhizal plants had the same rs with or without annuals, indicating that mycorrhizas may alleviate the detrimental effects of competition on rs of A. smithii. By contrast, neither inoculation nor annuals significantly affected water relations of A. dasystachyum. Mycorrhizas also did not increase N and P concentrations or percentage cover of either grass, but both had delayed phenology when mycorrhizal. This research was done during a wet year when the effects of mycorrhizas may be subtle. Long-term observations are necessary to determine the importance of mycorrhizas under field conditions.

Population Dynamics of Sugar Beets, Rhizoctonia solani, and Laetisaria arvalis: Responses of a Host, Plant Pathogen, and Hyperparasite to Perturbation in the Field.

Rhizoctonia solani causes crown rot of sugar beets, a severe disease that has destroyed up to 60% of the plants in a test field in western Nebraska. Laetisaria arvalis, a natural hyperparasite of Rhizoctonia spp., was isolated from fields in western Nebraska. To test for the potential for biological control of R. solani, in November 1980 (following harvest) we applied various combinations of a nematicide (Telone II; Dow Chemical Co.), a nutrition source (sugar beet pulp), and an inoculum of L. arvalis in a randomized block design. Populations of R. solani, L. arvalis, and sugar beets were monitored monthly through October 1981 (just after harvest). In control and nematicide plots, the R. solani population did not change significantly through time. In plots inoculated with L. arvalis, the R. solani populations declined through March, concomitant with an increase in L. arvalis. L. arvalis then declined with a corresponding increase in the R. solani populations. Beet plant numbers declined significantly in all treatments. We suggest that reduction of the R. solani populations with the hyperparasite L. arvalis is possible but that a stable equilibrium naturally exists.

Competition for phosphorus: differential uptake from dual-isotope--labeled soil interspaces between shrub and grass.

Two species of Agropyron grass differed strikingly in their capacity to compete for phosphate in soil interspaces shared with a common competitor, the sagebrush Artemisia tridentata. Of the total phosphorus-32 and -33 absorbed by Artemisia, 86 percent was from the interspace shared with Agropyron spicatum and only 14 percent from that shared with Agropyron desertorum. Actively absorbing mycorrhizal roots of Agropyron and Artemisia were present in both interspaces, where competition for the labeled phosphate occurred. The results have important implications about the way in which plants compete for resources below ground in both natural plant communities and agricultural intercropping systems.