Facilitation of tree establishment by ectomycorrhizal (EM) networks (MNs) may become increasingly important as drought stress increases with climate change in some forested regions of North America. The objective of this study was to determine (1) whether temperature, CO2 concentration ([CO2]), soil moisture, and MNs interact to affect plant establishment success, such that MNs facilitate establishment when plants are the most water stressed, and (2) whether transfer of C and water between plants through MNs plays a role in this. We established interior Douglas-fir (Pseudotsuga menziesiivar.glauca) seedlings in root boxes with and without the potential to form MNs with nearby conspecific seedlings that had consistent access to water via their taproots. We varied temperature, [CO2], and soil moisture in growth chambers. Douglas-fir seedling survival increased when the potential existed to form an MN. Growth increased with MN potential under the driest soil conditions, but decreased with temperature at 800 ppm [CO2]. Transfer of 13C to receiver seedlings was unaffected by potential to form an MN with donor seedlings, but deuterated water (D2O) transfer increased with MN potential under ambient [CO2]. Chlorophyll fluorescence was reduced when seedlings had the potential to form an MN under high [CO2] and cool temperatures. We conclude that Douglas-fir seedling establishment in laboratory conditions is facilitated by MN potential where Douglas-fir seedlings have consistent access to water. Moreover, this facilitation appears to increase as water stress potential increases and water transfer via networks may play a role in this. These results suggest that conservation of MN potential may be important to forest regeneration where drought stress increases with climate change.
Ectomycorrhizal (EM) networks are hypothesized to facilitate regeneration under abiotic stress. We tested the role of networks in interactions between P. menziesii var. glauca trees and conspecific seedlings along a climatic moisture gradient to: (1) determine the effects of climatic factors on network facilitation of Pseudotsuga menziesii (Mirb.) Franco var. glauca (Mayr) seedling establishment, (2) infer the changing importance of P. menziesii var. glauca parent trees in conspecific regeneration with climate, and (3) parse the competitive from facilitative effects of P. menziesii var. glauca trees on seedlings. When drought conditions were greatest, seedling growth increased when seedlings could form a network with trees in the absence of root competition, but was reduced when unable to form a network. Survival was maximized when seedlings were able to form a network in the absence of root competition. Seedling stem natural abundance d 13 C increased with drought due to increasing water use efficiency, but was unaffected by distance from tree or network potential. We conclude that P. menziesii seedlings may benefit from the presence of established P. menziesii trees when growing under climatic drought, but that this benefit is contingent upon the establishment of an EM network prior to the onset of summer drought. These results suggest that networks are an important mechanism for EM plants establishing in a pattern consistent with the stress-gradient hypothesis, and therefore the importance of EM networks to facilitation in regeneration of EM trees is expected to increase with drought.
Ectomycorrhizal (EM) networks (MN) are thought to be an important mode of EM fungal colonization of coniferous seedlings. How MNs affect EM communities on seedlings, and how this varies with biotic and abiotic factors, is integral to understanding their importance in seedling establishment. We examined EM fungal community similarity between mature trees and conspecific interior Douglas-fir (Pseudotsuga menziesii var. glauca) seedlings in two experiments where seed and nursery-grown seedlings originating from different locations were planted at various distances from trees along a climatic gradient. At harvest, trees shared 60% of their fungal taxa in common with outplanted seedlings and 77% with germinants, indicating potential for seedlings to join the network of residual trees. In both experiments, community similarity between trees and seedlings increased with drought. However, community similarity was lower among nursery seedlings growing at 2.5 m from trees when they were able to form an MN, suggesting MNs reduced seedling EM fungal richness. For field germinants, MNs resulted in lower community similarity in the driest climates. Distance from trees affected community similarity of nursery seedlings to trees, but there was no interaction of provenance with MNs in their effect on similarity in either nursery seedlings or field germinants as hypothesized. We conclude that MNs of trees influence EM colonization patterns of seedlings, and the strength of these effects increases with climatic drought.
We assessed the total length of external arbuscular mycorrhizal hyphae as a function of plant species and functional form richness in restored northern tallgrass prairies. Total hyphal length increased with species and functional form richness. Hyphal length also increased when plant communities were dominated by species with high root density, high root to shoot ratios, and high nitrogen use efficiency. Hyphal length was positively correlated with the biomass of late successional C 4 grasses (Andropogon gerardi Vitman, Panicum virgatum L., Schizachyrium scoparium [Michx.] Nash-Gould, and Sorghastrum nutans L.), which are obligately mycorrhizal and characterized by high root to shoot ratios, and high root surface area per unit of root biomass. We thus conclude that in order to recover extraradical arbuscular mycorrhizal hyphal length in restored northern tallgrass prairies, at least three factors need to be given priority: 1) achieving high levels of species and functional form richness; 2) making sure that late successional C 4 grasses are present; and 3) making sure that the seed mixture includes species that are characterized by high root to shoot ratio, high root density, and high nitrogen-use efficiency. Resumen En este experimento evaluamos la relacion entre la longitud externa del micro micelio arbuscular simbiotico (mycorrhiza) y la diversidad de especies y formas funcionales de plantas en una pradera alta restaurada en el norte de los Estados Unidos. La longitud total del micro micelio subio con el incremento en la diversidad de especies y formas funcionales de plantas. La longitud total del micro micelio tambien subio cuando la comunidad de plantas estaba dominada por especies caracterizadas por una elevada densidades the raices, altas proportion de raices, y alta eficiencia en el uso del nitrogeno. Econtramos tambien que la longitud total del micro micelio estaba correlacionada con la biomasa de herbajes C 4 tipicas del los finales estados de sucesion en estas praderas (Andropogon gerardi Vitman, Panicum virgatum L., Schizachyrium scoparium [Michx.] Nash-Gould, y Sorghastrum nutans L.). Estos herbajes se caracterizan por ser dependientes en micelio arbuscular simbiotico, altas proporciones de raices, y superficie de raices por unidad the masa de raices. Concluimos que para reestablecer adquados niveles de micro micelio arbuscular simbiotico en las praderas altas del norte de los Estados Unidos al menos tres elementos son necesarios: 1) altos niveles de diversidad the especies y formas funcionales de plantas; 2) la presencia de herbajes C 4 tipicas del los finales estados de sucesion; y 3) asegurar que la mesclas de especies plantadas incluye especies caracterizadas por una alta proporcion de raices, alta densidad de raices, y alta efeciencia en la utilizacion del nitrogeno.
Two, two-factor experiments manipulated species and functional form plant richness and the spatial scaling of either nitrogen (N) or phosphorous (P) in restored tallgrass prairie in North Dakota, USA. Nitrate (NO 3 -) leaching was measured in these plots and analyzed for its response to the treatment factors and measured plant community parameters. Nitrate extracted from anion exchange resin was regressed against the first principal component of species and functional form richness, the spatial scaling of N or P, the measured biomass of the functional forms used and the plot values for plant parameters based on weighted averages by species biomass. The treatments applied in the N and P experiments were 1, 2, 5, 10, or 20 plant species taxa, and the application of fertilizer in a random fractal pattern with either finescale or coarse-scale heterogeneity. Nitrate leaching decreased with plant diversity and increased by a factor of two going from fine-scale to coarse-scale N.It was also related to a number of plant functional parameters, and was positively correlated with the biomass of late successional C 3 grasses (Koeleria cristata (Lam.) Beauv., Poa pratensis L., Stipa comata Trin. & Rupr., and Stipa viridula Trin.), which are known from previous studies to have negative mycorrhizal responsiveness and are characterized by high root lateral spread per unit of root biomass. Our results show that while plant diversity has a highly significant influence on plant community uptake of NO 3 -, this effect is mediated by the scaling of soil N and the functional traits of the species comprising the plant assemblage.
Commercial nursery practices usually fail to promote mycorrhization of interior Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.) Franco] seedlings in British Columbia, which may account for their poor performance following planting in the field. We tested the effects of four nursery cultivation factors (nitrogen fertilization, phosphorus fertilization, watering, and soil aeration) and field soil addition on mycorrhization, survival, growth, and biomass allocation of interior Douglas-fir seedlings in a series of greenhouse experiments. Where field soil was added to the growing medium, mycorrhization and root/shoot ratios were maximized at lower levels of mineral nutrient application and aeration. Where field soil was not added, mycorrhization was negligible across all fertilization and aeration treatments, but root/shoot ratio was maximized at lower levels of mineral nutrients and the highest level of aeration. Regardless of whether field soil was added, intermediate levels of soil water resulted in the best mycorrhizal colonization and root/shoot ratios. However, field soil addition reduced seedling mortality at the two lowest water levels. A cluster analysis placed ectomycorrhizal morphotypes into three groups (Mycelium radicis-atrovirens Melin, Wilcoxina, and mixed) based on their treatment response, with all but two morphotypes in the mixed group whose abundance was maximized under conditions common to advanced seedling establishment. For maximal mycorrhization and root development of interior Douglas-fir seedlings, nurseries should minimize addition of nitrogen and phosphorus nutrients, maximize aeration, provide water at moderate rates, and, where possible, add small amounts of field soil to the growing medium.
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