Unraveling root and rhizosphere traits in temperate maize landraces and modern cultivars: Implications for soil resource acquisition and drought adaptation.

A holistic understanding of plant strategies to acquire soil resources is pivotal in achieving sustainable food security. However, we lack knowledge about variety-specific root and rhizosphere traits for resource acquisition, their plasticity and adaptation to drought. We conducted a greenhouse experiment to phenotype root and rhizosphere traits (mean root diameter [Root D], specific root length [SRL], root tissue density, root nitrogen content, specific rhizosheath mass [SRM], arbuscular mycorrhizal fungi [AMF] colonization) of 16 landraces and 22 modern cultivars of temperate maize (Zea mays L.). Our results demonstrate that landraces and modern cultivars diverge in their root and rhizosphere traits. Although landraces follow a 'do-it-yourself' strategy with high SRLs, modern cultivars exhibit an 'outsourcing' strategy with increased mean Root Ds and a tendency towards increased root colonization by AMF. We further identified that SRM indicates an 'outsourcing' strategy. Additionally, landraces were more drought-responsive compared to modern cultivars based on multitrait response indices. We suggest that breeding leads to distinct resource acquisition strategies between temperate maize varieties. Future breeding efforts should increasingly target root and rhizosphere economics, with SRM serving as a valuable proxy for identifying varieties employing an outsourcing resource acquisition strategy.

The plant root economics space in relation to nutrient limitation in Eurasian herbaceous plant communities.

Plant species occupy distinct niches along a nitrogen-to-phosphorus (N:P) gradient, yet there is no general framework for belowground nutrient acquisition traits in relation to N or P limitation. We retrieved several belowground traits from databases, placed them in the "root economics space" framework, and linked these to a dataset of 991 plots in Eurasian herbaceous plant communities, containing plant species composition, aboveground community biomass and tissue N and P concentrations. Our results support that under increasing N:P ratio, belowground nutrient acquisition strategies shift from "fast" to "slow" and from "do-it-yourself" to "outsourcing", with alternative "do-it-yourself" to "outsourcing" strategies at both ends of the spectrum. Species' mycorrhizal capacity patterns conflicted with root economics space predictions based on root diameter, suggesting evolutionary development of alternative strategies under P limitation. Further insight into belowground strategies along nutrient stoichiometry is crucial for understanding the high abundance of threatened plant species under P limitation.

Responses of rhizosphere fungi to the root economics space in grassland monocultures of different age.

Recent studies on root traits have shown that there are two axes explaining trait variation belowground: the collaboration axis with mycorrhizal partners and the conservation ('fast - slow') axis. However, it is yet unknown whether these trait axes affect the assembly of soilborne fungi. We expect saprotrophic fungi to link to the conservation axis of root traits, whereas pathogenic and arbuscular mycorrhizal fungi link to the collaboration axis, but in opposite directions, as arbuscular mycorrhizal fungi might provide pathogen protection. To test these hypotheses, we sequenced rhizosphere fungal communities and measured root traits in monocultures of 25 grassland plant species, differing in age. Within the fungal guilds, we evaluated fungal species richness, relative abundance and community composition. Contrary to our hypotheses, fungal diversity and relative abundance were not strongly related to the root trait axes. However, saprotrophic fungal community composition was affected by the conservation gradient and pathogenic community composition by the collaboration gradient. The rhizosphere AMF community composition did not change along the collaboration gradient, even though the root trait axis was in line with the root mycorrhizal colonization rate. Overall, our results indicate that in the long term, the root trait axes are linked with fungal community composition.

Shift of root nitrogen-acquisition strategy with tree age is mediated by root functional traits along the collaboration gradient of the root economics space.

Root nitrogen (N)-uptake rate and uptake preference, and their association with root morphological and chemical traits are important to characterize root N-acquisition strategies of trees. However, how the root N-acquisition strategy varies with tree age, especially for those species that coexist at a common site, remains unknown. In this study, a field isotopic hydroponic method was used to determine the uptake rate and contribution of NH4+, NO3- and glycine, for three coexisting ectomycorrhizal coniferous species [Pinus koraiensis (Korean pine), Picea koraiensis (Korean spruce) and Abies nephrolepis (smelly fir)] at three age classes (young, middle-aged and mature) in a temperate forest. Concurrently, root morphological and chemical traits, as well as mycorrhizal colonization rate were determined. Our results show that the root uptake rate of total N and NH4+ gradually decreased across all three species with increasing tree age. The three species at all age classes preferred NH4+, except for middle-aged Korean spruce and mature smelly fir, which preferred glycine. In contrast, all three species showed the lowest acquisition of NO3-. According to the conceptual framework of 'root economics space', only a 'collaboration' gradient (i.e. dimension of root diameter vs specific root length or area) was identified for each species, in which root N-uptake rate loaded heavily on the side of 'do-it-yourself' (i.e. foraging N more by roots). Young trees of all species tended to exhibit the 'do-it-yourself' strategy for N uptake, and mature trees had an 'outsourcing' strategy (i.e. foraging N by a mycorrhizal partner), whereas middle-aged trees showed a balanced strategy. These findings suggest that shifts of root N-acquisition strategy with tree age in these species are mainly mediated by root traits along the 'collaboration' gradient, which advances our understanding of belowground competition, species coexistence and N cycling in temperate forests.

An alternative risk assessment framework for tropical soil multi-metal contamination using arbuscular mycorrhizal fungi.

Arbuscular mycorrhizal fungi (AMF) are a key group of soil organisms involved in several ecosystem services, but they had not been explored in retrospective ecological risk assessment (ERA) schemes, partially due to a lack of experimental protocols. We aimed to perform a site-specific test to assess the effects of multiple metal contaminated soil (Zn, Cu, Cd, and Pb) on several ecotypes of AMF and evaluate the conceptual fitness of the performed AMF test to include in the TRIAD approach of ERA schemes. The results demonstrated that increasing metal contaminated soil proportions that inhibited 50 % (IC50) of spore germination varied from 28 to >80. Ecotypes such as Claroideoglomus etunicatum (IC50 > 80) and Racocetra gregaria (IC50 > 80) experienced 50 % reduction in spore germination at metals concentrations of 10,776.3 for Zn, 1015.2 Cu, 65.5 Cd, 140.2 mg dm-3 Pb, that are 3 times higher than those for Acaulospora mellea CMM101 (IC50 28 [16.2-39.8]) (3441.7 Zn, 333.9 Cu, 17.8 Cd, 56.5 mg dm-3 Pb). In the evaluation of the suitability of the AMF ecotoxicological test to ERA, both spore germination and germinative tube growth were best evaluated and thus suitable in the following descending order: Tier III, Tier II, and Tier I. Variable effects of multiple-metal contamination on the ecotypes indicates how AMF community is affected in its pre-symbiotic structures. The ecotoxicological test allowed the selection of two species with the greatest sensitivity (Ambispora appendicula and Rhizophagus clarus CMM103) to the metal matrix, with the potential to best fit ERA objectives. The site-specific ecotoxicological test with AMF ex-situ proved adequate as an alternative test for Tiers II and III of TRIAD ERA schemes for metal contaminated areas. Data generated through test results, such as the inhibition concentrations (ICs), could be incorporated into ERAs risk indexes, increasing its ecological relevance, and reducing overall uncertainties.

Assessment of beneficial fungal microorganism's bio-efficacy in stimulating morphological and physiological parameters of Allium cepa plants grown in soil amended with fish wastes.

BACKGROUND: The increase in the human consumption of fish results in the production of organic fish wastes (FW). For enhanced soil fertility and plant growth at a lower cost and without the negative impacts of chemical fertilizers, these wastes could be employed as a valuable organic fertilizer. To determine the synergistic bio-efficacy of Trichoderma sp. and arbuscular mycorrhizal (AM) fungi in stimulating the morphological and physiological characteristics of FW-fertilized Alium cepa, as well as to investigate their involvement in boosting soil fertility, the current study was carried out. Overall, eight treatments were applied as follows: AM, Trichoderma sp., AM + Trichoderma sp., FW, AM + FW, Trichoderma sp. + FW, AM + Trichoderma sp. + FW, and control. Growth and physiological assessments of onion plants were taken after 8 weeks from FW application. RESULTS: Our results showed that FW application combined with AM fungi and Trichoderma sp. inoculations increased aggregate stability of the soil (glomalin content) and soil chitinase activity. Moreover, using the bio-inoculations along with FW amendments significantly (p < 0.05) improved the photosynthetic pigments, protein, carbohydrates, and nutrients content of onion plants. It's interesting to note that the triple interaction of AM + Trichoderma sp. + FW led to the greatest increase in plant height, root length, number of leaves, and leaf area as well as total fresh and dry weights of shoots and roots. Besides, AM fungal colonization was at its highest percentage with Trichoderma sp. inoculation, although this percentage decreased with FW addition. CONCLUSION: We concluded that the combined treatments of AM fungi and Trichoderma sp. along with FW application to the soil can be proposed as a successful strategy for plant performance in nutrient-deficient soils as both fungal inoculants are capable of degrading these wastes and converting them into manure suitable for farming so plants can uptake the minerals effortlessly.

[Comparison and health risk assessment of mineral elements in stems and leaves of Dendrobium officinale cultivated with conventional method and mycorrhizal fungi].

This study aims to analyze the variation of the content of mineral elements in stems and leaves of Dendrobium officinale cultivated with conventional method and mycorrhizal fungi, which is expected to lay a basis for safety of stems and leaves of D. officinale. A total of 7 samples from Jiangsu, Fujian, Shanghai, and Zhejiang were collected, which were then cultivated with conventional method and mycorrhizal fungi, separately. The content of 17 mineral elements in stems and leaves was measured by inductively coupled plasma-mass spectrometry(ICP-MS), and the content changes of the mineral elements were analyzed. The health risks of Pb, Cd, Hg, and As in stems were assessed by target hazard quotient(THQ). The results showed that the content of polluting elements in stems and leaves of D. officinale was low, and the content in the plants cultivated with mycorrhizal fungi was reduced. The content of K, Ca, Mg, and P was high in stems and leaves of the species, suggesting that cultivation with mycorrhizal fungi improved the content of other elements irregularly. According to the THQ, the safety risk of stems of D. officinale cultivated with either conventional method or mycorrhizal fungi was low, particularly the D. officinale cultivated mycorrhizal fungi. The results indicated that cultivation with mycorrhizal fungi influenced the element content in stems and leaves of D. officinale. It is necessary to study the culture substrate, processing technology, and the mechanism of the increase or decrease in mineral elements of D. officinale in the future.

Assessment of mycorrhizal association of a threatened medicinal plant Clerodendrum indicum (L.) O. Kuntze (Verbenaceae) in different ecological variations.

Mycorrhizae association is reported to enhance the survivability of the host plant under adverse environmental conditions. The present study aims to explore the mycorrhizal association in the roots of different ecotypes of a threatened medicinal plant, Clerodendrum indicum (L.) O. Kuntze (Verbenaceae), collected from W.B., India, which correlates the degree of root colonization to the nutritional status of the native soil. Ten ecotypes of C. indicum having diverse morphological variations were collected. The mycorrhizae were characterized by both morphological and molecular methods. The nutritional status of the native soils was estimated. The study revealed that all the ecotypes have an association with mycorrhizal forms like hyphae, arbuscules, and vesicles. The molecular analysis showed Glomus intraradices and Rhizophagus irregularis as the associated arbuscular mycorrhizal fungi (AMF). A significant variation in arbuscule and vesicle formation was found growing in the varied nutritional statuses concerning soil parameters. The arbuscule was found negatively correlated with pH, conductivity, and potassium and positively correlated with organic carbon, nitrogen, and phosphorus. The vesicle was found positively correlated with pH, organic carbon, and potassium and negatively correlated with conductivity, nitrogen, and phosphorus. The interaction between conductivity: nitrogen, conductivity: phosphorus, organic-carbon: nitrogen, and pH: conductivity was significant in influencing vesicle formation. However, none of the interactions between parameters was found significant in influencing arbuscule formation. Thus, the study concludes that G. intraradices and R. irregularis are the principle mycorrhizae forming the symbiotic association with the threatened medicinal plant, C. indicum. They form vesicles and arbuscules based on their soil nutritive factors. Therefore, a large-scale propagation through a selective AMF association would help in the conservation of this threatened species from extinction.

Mycorrhizal symbiosis pathway and edaphic fertility frame root economics space among tree species.

The root economics space (RES) is multidimensional and largely shaped by belowground biotic and abiotic influences. However, how root-fungal symbioses and edaphic fertility drive this complexity remains unclear. Here, we measured absorptive root traits of 112 tree species in temperate and subtropical forests of China, including traits linked to functional differences between arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) hosts. Our data, from known mycorrhizal tree species, revealed a 'fungal-symbiosis' dimension distinguishing AM from ECM species. This divergence likely resulted from the contrasting mycorrhizal evolutionary development of AM vs ECM associations. Increased root tissue cortical space facilitates AM symbiosis, whereas increased root branching favours ECM symbiosis. Irrespective of mycorrhizal type, a 'root-lifespan' dimension reflecting aspects of root construction cost and defence was controlled by variation in specific root length and root tissue density, which was fully independent of root nitrogen content. Within this function-based RES, we observed a substantial covariation of axes with soil phosphorus and nitrate levels, highlighting the role played by these two axes in nutrient acquisition and conservation. Overall, our findings demonstrate the importance of evolved mycorrhizal symbiosis pathway and edaphic fertility in framing the RES, and provide theoretical and mechanistic insights into the complexity of root economics.

Combined application of marble waste and beneficial microorganisms: toward a cost-effective approach for restoration of heavy metals contaminated sites.

Heavy metal (HM) pollution and the need to preserve the environment have gathered increasing scientific attention. The immobilization of HMs into less-soluble, less mobile, and less toxic forms in addition to the improvement of Medicago sativa L. growth and HMs accumulation were evaluated after the application of marble waste (MW) and/or beneficial PGP rhizobacteria and mycorrhizae to the mining soil compost. A greenhouse assay was conducted to elucidate the influence of both amendment and beneficial microorganisms. The application of marble waste to the soil-compost resulted in decreasing the bioavailability of metals (Cu, Zn, Pb, and Cd), thus ameliorating the installation of the vegetal cover for 6 months of culture. Cultivation of M. sativa under 5% MW-amended soil for 6 months increased the shoot dry weight by almost twofold, while the inoculation with rhizobacteria-mycorrhizae combined with the application of 15% MW resulted in an improvement of 3.5-fold in case of shoot dry weight. In addition, the application of marble waste amendment or their combination with metallo-resistant bacteria resulted in decreasing HM accumulation leading to HM content below the threshold recommended for animal grazing. Thus, the application of amendments and beneficial microorganisms appeared to guarantee the safe cultivation of alfalfa for 6 months of culture. The dual combination amendments and beneficial microorganisms showed the good potential to restore HM polluted soils and could stand as a novel approach for restoration of HM-contaminated soils.

The Costs and Benefits of Plant-Arbuscular Mycorrhizal Fungal Interactions.

The symbiotic interaction between plants and arbuscular mycorrhizal (AM) fungi is often perceived as beneficial for both partners, though a large ecological literature highlights the context dependency of this interaction. Changes in abiotic variables, such as nutrient availability, can drive the interaction along the mutualism-parasitism continuum with variable outcomes for plant growth and fitness. However, AM fungi can benefit plants in more ways than improved phosphorus nutrition and plant growth. For example, AM fungi can promote abiotic and biotic stress tolerance even when considered parasitic from a nutrient provision perspective. Other than being obligate biotrophs, very little is known about the benefits AM fungi gain from plants. In this review, we utilize both molecular biology and ecological approaches to expand our understanding of the plant-AM fungal interaction across disciplines.

Root phosphatase activity aligns with the collaboration gradient of the root economics space.

The adoption of diverse resource acquisition strategies is critical for plant growth and species coexistence. Root phosphatase is of particular importance in the acquisition of soil phosphorus (P), yet it is often overlooked in studies of root trait syndromes. Here, we evaluated the role of root phosphatase activity (RPA) within the root economics space and the order-based variation of RPA, as well as the correlations between RPA and a suite of leaf traits and soil properties over a range of evergreen tree species in a subtropical forest. Root phosphatase activity exhibited a high degree of inter-specific variation. We found that there were two leading dimensions of the multidimensional root economics space, the root diameter-specific root length axis (collaboration trait gradient) and the root tissue density-root nitrogen concentration axis (classical trait gradient), and RPA aligned with the former. Root phosphatase activity is used as a 'do it yourself' strategy of soil P acquisition, and was found to be inversely correlated with mycorrhizal colonization, which suggests a trade-off in plant P acquisition strategies. Compared with soil and foliar nutrient status, root traits mattered most for the large inter-specific changes in RPA. Furthermore, RPA generally decreased from first- to third-order roots. Taken together, such diverse P-acquisition strategies are conducive to plant coexistence within local forest communities. The use of easily measurable root traits and their tight correlations with RPA could be a feasible and promising approach to estimating species-specific RPA values, which would be helpful for better understanding plant P acquisition and soil P cycling.

Linking root exudation to belowground economic traits for resource acquisition.

The concept of a root economics space (RES) is increasingly adopted to explore root trait variation and belowground resource-acquisition strategies. Much progress has been made on interactions of root morphology and mycorrhizal symbioses. However, root exudation, with a significant carbon (C) cost (c. 5-21% of total photosynthetically fixed C) to enhance resource acquisition, remains a missing link in this RES. Here, we argue that incorporating root exudation into the structure of RES is key to a holistic understanding of soil nutrient acquisition. We highlight the different functional roles of root exudates in soil phosphorus (P) and nitrogen (N) acquisition. Thereafter, we synthesize emerging evidence that illustrates how root exudation interacts with root morphology and mycorrhizal symbioses at the level of species and individual plant and argue contrasting patterns in species evolved in P-impoverished vs N-limited environments. Finally, we propose a new conceptual framework, integrating three groups of root functional traits to better capture the complexity of belowground resource-acquisition strategies. Such a deeper understanding of the integrated and dynamic interactions of root morphology, root exudation, and mycorrhizal symbioses will provide valuable insights into the mechanisms underlying species coexistence and how to explore belowground interactions for sustainable managed systems.

Above- and belowground biodiversity jointly tighten the P cycle in agricultural grasslands.

Experiments showed that biodiversity increases grassland productivity and nutrient exploitation, potentially reducing fertiliser needs. Enhancing biodiversity could improve P-use efficiency of grasslands, which is beneficial given that rock-derived P fertilisers are expected to become scarce in the future. Here, we show in a biodiversity experiment that more diverse plant communities were able to exploit P resources more completely than less diverse ones. In the agricultural grasslands that we studied, management effects either overruled or modified the driving role of plant diversity observed in the biodiversity experiment. Nevertheless, we show that greater above- (plants) and belowground (mycorrhizal fungi) biodiversity contributed to tightening the P cycle in agricultural grasslands, as reduced management intensity and the associated increased biodiversity fostered the exploitation of P resources. Our results demonstrate that promoting a high above- and belowground biodiversity has ecological (biodiversity protection) and economical (fertiliser savings) benefits. Such win-win situations for farmers and biodiversity are crucial to convince farmers of the benefits of biodiversity and thus counteract global biodiversity loss.

Mallocybe heimii ectomycorrhizae with Cistus creticus and Pinus halepensis in Mediterranean littoral sand dunes - assessment of phylogenetic relationships to M. arenaria and M. agardhii.

Ectomycorrhizal symbiosis appears extensively in the Northern Hemisphere, where Mediterranean ecosystems constitute an important ecological area of considerable biodiversity value. Littoral sand dunes are among high-risk habitats, and ectomycorrhizal lifestyle contributes significantly to supporting life in such regions. Mallocybe heimii (Bon) Matheny & Esteve-Rav. (Inocybaceae, Basidiomycota) and the very similar M. arenaria (Bon) Matheny & Esteve-Rav. grow in poor, usually sandy soils, in association with angiosperms or gymnosperms. Basidiomata originally identified under these names were collected from littoral sand dunes of Greece, and their morpho-anatomical characteristics were examined in conjunction with material derived from other European regions. Sequences from basidiomata and root tips corresponding to the nuclear rDNA internal transcribed spacer region (ITS) and large subunit (LSU) were obtained and analyzed. Phylogenetic results demonstrated that material identified as M. heimii or M. arenaria form a single well-supported group, while M. agardhii (N. Lund) Matheny & Esteve-Rav. is confirmed to be distinct from M. arenaria (the latter was initially described as a variety of the former, i.e., I. agardhii var. arenaria Bon). A detailed tree of the genus Mallocybe was generated on the basis of concatenated ITS and LSU sequences, and relationships of selected taxa are discussed in the light of morphological and sequence data. In addition, the first morphotype descriptions of M. heimii ectomycorrhizae with Cistus creticus L. and Pinus halepensis Miller are hereby provided. Both morphotypes exhibited the typical characteristics of Inocybe/Mallocybe ectomycorrhizae; however, differences were noted, the most significant being the presence of clamps on mantle hyphae and the type of anastomoses.

Genetic variability assessment of 127 Triticum turgidum L. accessions for mycorrhizal susceptibility-related traits detection.

Positive effects of arbuscular mycorrhizal fungi (AMF)-wheat plant symbiosis have been well discussed by research, while the actual role of the single wheat genotype in establishing this type of association is still poorly investigated. In this work, the genetic diversity of Triticum turgidum wheats was exploited to detect roots susceptibility to AMF and to identify genetic markers in linkage with chromosome regions involved in this symbiosis. A tetraploid wheat collection of 127 accessions was genotyped using 35K single-nucleotide polymorphism (SNP) array and inoculated with the AMF species Funneliformis mosseae (F. mosseae) and Rhizoglomus irregulare (R. irregulare), and a genome-wide association study (GWAS) was conducted. Six clusters of genetically related accessions were identified, showing a different mycorrhizal colonization among them. GWAS revealed four significant quantitative trait nucleotides (QTNs) involved in mycorrhizal symbiosis, located on chromosomes 1A, 2A, 2B and 6A. The results of this work enrich future breeding activities aimed at developing new grains on the basis of genetic diversity on low or high susceptibility to mycorrhization, and, possibly, maximizing the symbiotic effects.

Comprehensive Phytotoxicity Assessment Protocol for Engineered Nanomaterials.

In order for nanotechnology to be sustainably applied in agriculture, emphasis should be on comprehensive assessment of multiple endpoints, including biouptake and localization of engineered nanomaterials (ENMs), potential effects on food nutrient quality, oxidative stress responses, and crop yield, before ENMs are routinely applied in consumer and agronomic products. This chapter succinctly outlines a protocol for conducting nanophytotoxicity studies focusing on nanoparticle purification and characterization, arbuscular mycorrhizal fungi (AMF)/symbiont inoculation, biouptake and translocation/localization, varied endpoints of oxidative stress responses, and crop yield.

Demonstrational gardens with EDTA-washed soil. Part II: Soil quality assessment using biological indicators.

In this study, we evaluated the impact of washing of Pb, Zn and Cd contaminated soil using EDTA-based technology (ReSoil®) on soil biological properties by measuring some of the most commonly used/sensitive biological indicators of soil perturbation. We estimated the temporal dynamics of the soil respiration, the activities of soil enzymes (dehydrogenase, ß-glucosidase, urease, acid and alkaline phosphatase), and the effect of the remediation process on arbuscular mycorrhizal (AM) fungi in original (Orig), remediated (Rem) and remediated vitalized (Rem+V) soils during a more than one-year garden experiment. ReSoil® technology initially affected the activity level of soil microbial respiration and all enzyme activities except urease and reduced AM fungal potential in the soil. However, after one year of vegetable cultivation and standard gardening practices, soil microbial respiration, acid and alkaline phosphatase in the Rem and Rem+V reached similar activities as in the Orig. Only the activities of dehydrogenase and ß-glucosidase remained lower in the remediated soil compared to the Orig. The frequency of arbuscular mycorrhiza in the root system, arbuscular density in the colonized root fragment, and the intensity of mycorrhizal colonization in the colonized root fragments in the remediated treatments increased with time; at the end of the experiment, no consistent differences in these parameters of mycorrhizal colonization were found among the treatments. Our results suggest a restored biological functioning of the remediated soil after one year of vegetable cultivation. In general, no differences were found between the Rem and Rem+V treatments, indicating that simple common garden practices are sufficient to restore soil functioning after remediation.

Walrasian equilibrium behavior in nature.

The interaction between land plants and mycorrhizal fungi (MF) forms perhaps the world's most prevalent biological market. Most plants participate in such markets, in which MF collect nutrients from the soil and trade them with host plants in exchange for carbon. In a recent study, M. D. Whiteside et al. [Curr. Biol. 29, 2043-2050.e8 (2019)] conducted experiments that allowed them to quantify the behavior of arbuscular MF when trading phosphorus with their host roots. Their experimental techniques enabled the researchers to infer the quantities traded under multiple scenarios involving different amounts of phosphorus resources initially held by different MF patches. We use these observations to confirm a revealed preference hypothesis, which characterizes behavior in Walrasian equilibrium, a centerpiece of general economic equilibrium theory.

Phosphorus deficiencies invoke optimal allocation of exoenzymes by ectomycorrhizas.

Ectomycorrhizal (EM) fungi can acquire phosphorus (P) through the production of extracellular hydrolytic enzymes (exoenzymes), but it is unclear as to the manner and extent native EM fungal communities respond to declining soil P availability. We examined the activity of six exoenzymes (xylosidase, N-acetyl glucosaminidase, ß-glucosidase, acid phosphomonoesterase, acid phosphodiesterase [APD], laccase) from EM roots of Pseudotsuga menzesii across a soil podzolization gradient of coastal British Columbia. We found that APD activity increased fourfold in a curvilinear association with declining inorganic P. Exoenzyme activity was not related to organic P content, but at a finer resolution using 31P-NMR, there was a strong positive relationship between APD activity and the ratio of phosphodiesters to orthophosphate of surface organic horizons (forest floors). Substantial increases (two- to fivefold) in most exoenzymes were aligned with declining foliar P concentrations of P. menzesii, but responses were statistically better in relation to foliar nitrogen (N):P ratios. EM fungal species with consistently high production of key exoenzymes were exclusive to Podzol plots. Phosphorus deficiencies in relation to N limitations may provide the best predictor of exoenzyme investment, reflecting an optimal allocation strategy for EM fungi. Resource constraints contribute to species turnover and the assembly of distinct, well-adapted EM fungal communities.