The extent of mycorrhizal colonization of roots and its influence on plant growth and phosphorus content

Aims: The most common metric of arbuscular mycorrhizal fungal (AMF) abundance is percent root length colonized (PRLC) by mycorrhizal structures. Frequently, plants with greater PRLC are assumed to receive more nutrients (such as phosphorus, P) from their mycorrhizal symbionts, leading to greater plant growth. Nevertheless, the functional significance of this metric remains controversial. In this review, I discuss whether manipulations of PRLC generally led to changes in plant biomass and P content, and whether AMF taxa and plant functional groups influence these relationships. Methods: I conducted a meta-analysis of laboratory- and field-based trials in which mycorrhizal colonization was directly altered compared to unmanipulated controls. For each trial, I calculated (1) the difference in PRLC (ΔPRLC) between the treatments, and (2) the response ratio of plant biomass. In a subset of these studies, the response ratio of P content of host plants could also be calculated. Results: The response ratio of plant biomass and P content rose significantly and exponentially as ΔPRLC increased. Nevertheless, ΔPRLC explained only a fraction of the variation in response ratios in each case. Moreover, AMF taxa varied in their effects on biomass per unit ΔPRLC. In addition, plant functional groups differed in effects on plant P content per unit ΔPRLC, with C4 grasses responding most strongly. Conclusions: It appears that as the extent to which plant roots are colonized by AMF increases, plant growth and P content often increase, although substantial variability exists among trials. As others have found, a likely mechanism for this relationship is increased transfer of P (and perhaps other nutrients) through the more-prevalent mycorrhizal structures. © 2013 Springer Science+Business Media Dordrecht

Climate change feedbacks to microbial decomposition in boreal soils

Boreal ecosystems store 10-20% of global soil carbon and may warm by 4-7°C over the next century. Higher temperatures could increase the activity of boreal decomposers and indirectly affect decomposition through other ecosystem feedbacks. For example, permafrost melting will likely alleviate constraints on microbial decomposition and lead to greater soil CO2 emissions. However, wet boreal ecosystems underlain by permafrost are often CH4 sources, and permafrost thaw could ultimately result in drier soils that consume CH4, thereby offsetting some of the greenhouse warming potential of soil CO2 emissions. Climate change is also likely to increase winter precipitation and snow depth in boreal regions, which may stimulate decomposition by moderating soil temperatures under the snowpack. As temperatures and evapotranspiration increase in the boreal zone, fires may become more frequent, leading to additional permafrost loss from burned ecosystems. Although post-fire decomposition could also increase due to higher soil temperatures, reductions in microbial biomass and activity may attenuate this response. Other feedbacks such as soil drying, increased nutrient mineralization, and plant species shifts are either weak or uncertain. We conclude that strong positive feedbacks to decomposition will likely depend on permafrost thaw, and that climate feedbacks will probably be weak or negative in boreal ecosystems without permafrost. However, warming manipulations should be conducted in a broader range of boreal systems to validate these predictions. © 2011 Elsevier Ltd and The British Mycological Society

Mycorrhizal fungi have a potential role in soil carbon storage under elevated CO2 and nitrogen deposition.

In this review, we discuss the potential for mycorrhizal fungi to act as a source or sink for carbon (C) under elevated CO2 and nitrogen deposition. Mycorrhizal tissue has been estimated to comprise a significant fraction of soil organic matter and below-ground biomass in a range of systems. The current body of literature indicates that in many systems exposed to elevated CO2, mycorrhizal fungi might sequester increased amounts of C in living, dead and residual hyphal biomass in the soil. Through this process, the fungi might serve as a negative feedback on the rise in atmospheric CO2 levels caused by fossil fuel burning and deforestation. By contrast, a few preliminary studies suggest that N deposition might increase turnover rates of fungal tissue and negate CO2 effects on hyphal biomass. If these latter responses are consistent among ecosystems, C storage in hyphae might decline in habitats surrounding agricultural and urban areas. When N additions occur without CO2 enrichment, effects on mycorrhizal growth are inconsistent. We note that analyses of hyphal decomposition under elevated CO2 and N additions are extremely sparse but are critical in our understanding of the impact of global change on the cycling of mycorrhizal C. Finally, shifts in the community composition of arbuscular and ectomycorrhizal fungi with increasing CO2 or N availability are frequently documented. Since mycorrhizal groups vary in growth rate and tissue quality, these changes in species assemblages could produce unforeseeable impacts on the productivity, survivorship, or decomposition of mycorrhizal biomass

Direct nitrogen and phosphorus limitation of arbuscular mycorrhizal fungi: a model and field test.

Since mycorrhizal fungi constitute an important component of the soil–plant interface, their responses to changes in nutrient availability may mediate shifts in ecosystem function. We tested the hypothesis that initial soil nutrient availability may determine effects of nitrogen (N) and phosphorus (P) additions on the growth and community of arbuscular mycorrhizal (AM) fungi. Extraradical hyphal lengths and degree of root colonization of AM fungi were measured in control and fertilized plots along a soil fertility gradient in Hawaii. Responses of individual AM genera were assessed through immunofluorescent labeling. The AM biomass was increased by N and P additions in the N- and P-limited sites, respectively, and reduced by P fertilization in the fertile site only. The abundance of Scutellospora was lower under N than under P fertilization, whereas the incidence of Glomus was higher in the fertile site than the N-limited site. Gigaspora and Acaulospora did not vary among sites or treatments. Our results indicate that a decrease in AM abundance following nutrient additions cannot be assumed to occur and the effects may differ among AM genera and ecosystems with varying soil nutrients. Limitation of N and P may be one possible explanation

Using lipid analysis and hyphal length to quantify AM and saprotrophic fungal abundance along a soil chronosequence.

We evaluate the use of signature fatty acids and direct hyphal counts as tools to detect and quantify arbuscular mycorrhizal (AM) and saprotrophic fungal (SF) biomass in three Hawaiian soils along a natural soil fertility gradient. Phospholipids16:1omega5c and 18:2omega6,9c were used as an index of AM and saprotrophic fungal biomass, respectively. Both phospholipid analysis and hyphal length indicated that the biomass of AMF was greatest at the highest fertility site, and lowest where phosphorus limits plant growth. Saprotrophic fungal biomass did not vary. Hyphal length counts appeared to under-estimate SF abundance, while the phospholipid AMF:SF ratio was in line with expectations. This study indicates that phospholipids may be a valuable and reliable tool for studying the abundance, distribution, and interactions between AM and saprotrophic fungi in soil

Absorption of ant-provided carbon dioxide and nitrogen by a tropical epiphyte

ALTHOUGH ant-plant mutualisms have been described in many ecosystems, the magnitude of the direct benefits from such relationships are hard to quantify. In Bako National Park, Sarawak, Malaysia, stunted ‘kerangas’ forests occur on nutrient-poor sandstone hills1-3. As trees are widely spaced and have a sparse leaf area, a significant amount of light reaches the tree trunks and enables a diverse community of epiphytes to thrive there4. One of these epiphytes, Dischidia major (Vahl) Merr. (Asclepiadaceae), has evolved unusual methods for enhancing carbon and nitrogen acquisition. We show here that a mutualistic relationship exists between ants of the genus Philidris and their host, D. major. Using stable isotope analysis, we calculate that 39% of the carbon in occupied host plant leaves is derived from ant-related respiration, and that 29% of the host nitrogen is derived from debris deposited into the leaf cavities by ants. © 1995 Nature Publishing Groups. All Rights Reserved

Mycorrhizal dynamics under elevated CO2 and nitrogen fertilization in a warm temperate forest

We examined the response of mycorrhizal fungi to free-air CO2 enrichment (FACE) and nitrogen (N) fertilization in a warm temperate forest to better understand potential influences over plant nutrient uptake and soil carbon (C) storage. In particular, we hypothesized that mycorrhizal fungi and glomalin would become more prevalent under elevated CO2 but decrease under N fertilization. In addition, we predicted that N fertilization would mitigate any positive effects of elevated CO2 on mycorrhizal abundance. Overall, we observed a 14% increase in ectomycorrhizal (ECM) root colonization under CO2 enrichment, which implies that elevated CO2 results in greater C investments in these fungi. Arbuscular mycorrhizal (AM) hyphal length and glomalin stocks did not respond substantially to CO2 enrichment, and effects of CO2 on AM root colonization varied by date. Nitrogen effects on AM fungi were not consistent with our hypothesis, as we found an increase in AM colonization under N fertilization. Lastly, neither glomalin concentrations nor ECM colonization responded significantly to N fertilization or to an N-by-CO2 interaction. A longer duration of N fertilization may be required to detect effects on these parameters. © 2007 Springer Science+Business Media B.V

Extracellular enzyme activity in the mycorrhizospheres of a boreal fire chronosequence

Saprotrophic microbes are typically credited with producing extracellular enzymes that recycle organic matter, though roots and mycorrhizal fungi also can contribute and may compete with the saprotrophs. We examined extracellular enzyme activity associated with the mycorrhizospheres of arbuscular mycorrhizal, ectomycorrhizal, dual-colonized (arbuscular and ectomycorrhizal), and ericoid mycorrhizal plants in a fire chronosequence in Alaska. Bulk soil and soil from beneath host plants were gathered in July 2004 and assayed for five enzymes that target organic C, P, and N substrates. Compared to bulk soil, activities of the C-targeting enzymes β-1,4-glucosidase and peroxidase were lower in arbuscular mycorrhizospheres and ericoid mycorrhizospheres, respectively. Moreover, extracellular enzyme activity varied among mycorrhizosphere types. Specifically, N-targeting leucine aminopeptidase was highest in arbuscular mycorrhizospheres, followed by ericoid and ectomycorrhizal/dual-colonized mycorrhizospheres; β-1,4-glucosidase had the reverse pattern. In addition, enzymatic stoichiometry suggested that extracellular enzyme producers invested more in C-acquisition than in N-acquisition in recent fire scars compared to mature forests. These data extend previous findings that roots and mycorrhizal fungi compete with saprotrophs by showing that the strength of this effect varies by mycorrhizal host. As a result the community composition of mycorrhizal host plants might mediate enzymatic activity in boreal soils. © 2011 Elsevier GmbH

Arbuscular mycorrhizal fungi as mediators of ecosystem responses to nitrogen deposition: A trait-based predictive framework

Anthropogenic nitrogen (N) deposition is exposing plants and their arbuscular mycorrhizal fungi (AMFs) to elevated N availability, often leading to shifts in communities of AMF. However, physiological trade-offs among AMF taxa in their response to N enrichment vs the ability to acquire other soil nutrients could have negative effects on plant and ecosystem productivity. It follows that information on the functional traits of AMF taxa can be used to generate predictions of their potential role in mediating ecosystem responses to N enrichment. Arbuscular mycorrhizal fungi taxa that produce extensive networks of external hyphae should forage for N and phosphorus (P) more effectively, but these services incur greater carbon (C) costs to the plant. If N enrichment ameliorates plant nutrient limitation, then plants may reduce C available for AMF, which in turn could eliminate AMF taxa with large extensive external hyphae from the soil community. As a result, the remaining AMF taxa may confer less P benefit to their host plants. Using a synthesis of data from the literature, we found that the ability of a taxon to persist in the face of increasing soil N availability was particularly high in isolates from the genus Glomus, but especially low among the Gigasporaceae. Across AMF genera, our data support the prediction that AMF with a tolerance for high soil N may confer a lower P benefit to their host plant. Relationships between high N tolerance and production of external hyphae were mixed. Synthesis. If the relationship between N tolerance and plant P benefit is widespread, then shifts in arbuscular mycorrhizal fungi communities associated with N deposition could have negative consequences for the ability of plants to acquire P and possibly other nutrients via a mycorrhizal pathway. Based on this relationship, we predict that arbuscular mycorrhizal fungi responses could constrain net primary productivity in P-limited ecosystems exposed to N enrichment. This prediction could be tested in future empirical and modelling studies

The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls. © 2008 Springer Science+Business Media B.V