Mycorrhizal fungi modify decomposition: a meta-analysis.

It has been proposed that ectomycorrhizal fungi can reduce decomposition while arbuscular mycorrhizal fungi may enhance it. These phenomena are known as the 'Gadgil effect' and 'priming effect', respectively. However, it is unclear which one predominates globally. We evaluated whether mycorrhizal fungi decrease or increase decomposition, and identified conditions that mediate this effect. We obtained decomposition data from 43 studies (97 trials) conducted in field or laboratory settings that controlled the access of mycorrhizal fungi to substrates colonized by saprotrophs. Across studies, mycorrhizal fungi promoted decomposition of different substrates by 6.7% overall by favoring the priming effect over the Gadgil effect. However, we observed significant variation among studies. The substrate C : N ratio and absolute latitude influenced the effect of mycorrhizal fungi on decomposition and contributed to the variation. Specifically, mycorrhizal fungi increased decomposition at low substrate C : N and absolute latitude, but there was no discernable effect at high values. Unexpectedly, the effect of mycorrhizal fungi was not influenced by the mycorrhizal type. Our findings challenge previous assumptions about the universality of the Gadgil effect but highlight the potential of mycorrhizal fungi to negatively influence soil carbon storage by promoting the priming effect.

Los hongos ectomicorrízicos puden reducir la descomposición mientras que los hongos micorrízico­arbusculares pueden potenciarla. Ambos fenómenos son conocidos como "Gadgil effect" y "priming effect", respectivamente. Sin embargo, no es claro cuál predomina mundialmente. En este trabajo evaluamos si los hongos micorrízicos disminuyen o promueven la descomposición, e identificamos las condiciones que regulan este efecto. Para ello, recopilamos datos de descomposición de 43 estudios (97 observaciones) realizados en condiciones de campo o laboratorio que controlaron el acceso de los hongos micorrízicos a sustratos colonizados por saprótrofos. Los hongos micorrízicos promovieron la descomposición de diferentes sustratos en un 6.7%. Sin embargo, observamos una variación significativa entre estudios. La relación C : N del sustrato y la latitud influyeron en el efecto de los hongos micorrícicos sobre la descomposición y contribuyeron a la variabilidad. Específicamente, los hongos micorrízicos aumentaron la descomposición a valores bajos de C : N del sustrato y latitud, pero no hubo un efecto discernible en valores altos. Inesperadamente, el tipo de micorriza no influyó en el efecto de los hongos micorrízicos. Nuestros hallazgos cuestionan la universalidad del Gadgil effect, y resaltan el potencial de los hongos micorrízicos para influir negativamente en el almacenamiento de carbono del suelo al promover el priming effect.

Soil fertility determines whether ectomycorrhizal fungi accelerate or decelerate decomposition in a temperate forest.

Ectomycorrhizal (ECM) fungi can both accelerate and decelerate decomposition of organic matter in forest soils, but a mechanistic understanding of this differential influence is limited. Here, we tested how ECM fungi affect decomposition along a natural fertility gradient in a temperate forest of European beech. Trees were girdled to reduce belowground carbon supply to the soil. Girdling shifted soil fungal community composition and decreased hyphal biomass production and soil CO2 efflux, indicating a reduced ECM fungal activity. Girdling also affected decomposition processes, but the effects depended on fertility. Our results indicate that ECM fungi decelerate decomposition under conditions of low fertility while under conditions of high fertility ECM fungi and their host roots have an accelerating effect. We conclude that both acceleration and deceleration of decomposition of organic matter by ECM fungi can occur within a forest, with soil fertility determining the direction and magnitude of these effects. We suggest a positive feedback between fertility, stand productivity and soil carbon and nitrogen dynamics that is mediated to a large extent by ECM fungi.

Fine roots and mycorrhizal fungi accelerate leaf litter decomposition in a northern hardwood forest regardless of dominant tree mycorrhizal associations.

●Fine roots and mycorrhizal fungi may either stimulate leaf litter decomposition by providing free-living decomposers with root-derived carbon, or may slow decomposition through nutrient competition between mycorrhizal and saprotrophic fungi. ●We reduced the presence of fine roots and their associated mycorrhizal fungi in a northern hardwood forest in New Hampshire, USA by soil trenching. Plots spanned a mycorrhizal gradient from 96% arbuscular mycorrhizal (AM) associations to 100% ectomycorrhizal (ECM)-associated tree basal area. We incubated four species of leaf litter within these plots in areas with reduced access to roots and mycorrhizal fungi and in adjacent areas with intact roots and mycorrhizal fungi. ●Over a period of 608 d, we found that litter decayed more rapidly in the presence of fine roots and mycorrhizal hyphae regardless of the dominant tree mycorrhizal association. Root and mycorrhizal exclusion reduced the activity of acid phosphatase on decomposing litter. ●Our results indicate that both AM- and ECM-associated fine roots stimulate litter decomposition in this system. These findings suggest that the effect of fine roots and mycorrhizal fungi on litter decay in a particular ecosystem likely depends on whether interactions between mycorrhizal roots and saprotrophic fungi are antagonistic or facilitative.

Soil fertility relates to fungal-mediated decomposition and organic matter turnover in a temperate mountain forest.

Fungi are known to exert a significant influence over soil organic matter (SOM) turnover, however understanding of the effects of fungal community structure on SOM dynamics and its consequences for ecosystem fertility is fragmentary. Here we studied soil fungal guilds and SOM decomposition processes along a fertility gradient in a temperate mountain beech forest. High-throughput sequencing was used to investigate fungal communities. Carbon and nitrogen stocks, enzymatic activity and microbial respiration were measured. While ectomycorrhizal fungal abundance was not related to fertility, saprotrophic ascomycetes showed higher relative abundances under more fertile conditions. The activity of oxidising enzymes and respiration rates in mineral soil were related positively to fertility and saprotrophic fungi. In addition, organic layer carbon and nitrogen stocks were lower on the more fertile plots, although tree biomass and litter input were higher. Together, the results indicated a faster SOM turnover at the fertile end of the gradient. We suggest that there is a positive feedback mechanism between SOM turnover and fertility that is mediated by soil fungi to a significant extent. By underlining the importance of fungi for soil fertility and plant growth, these findings furthermore emphasise the dependency of carbon cycling on fungal communities below ground.

Lower relative abundance of ectomycorrhizal fungi under a warmer and drier climate is linked to enhanced soil organic matter decomposition.

The aboveground impacts of climate change receive extensive research attention, but climate change could also alter belowground processes such as the delicate balance between free-living fungal decomposers and nutrient-scavenging mycorrhizal fungi that can inhibit decomposition through a mechanism called the Gadgil effect. We investigated how climate change-induced reductions in plant survival, photosynthesis and productivity alter soil fungal community composition in a mixed arbuscular/ectomycorrhizal (AM/EM) semiarid shrubland exposed to experimental warming (W) and/or rainfall reduction (RR). We hypothesised that increased EM host plant mortality under a warmer and drier climate might decrease ectomycorrhizal fungal (EMF) abundance, thereby favouring the proliferation and activity of fungal saprotrophs. The relative abundance of EMF sequences decreased by 57.5% under W+RR, which was accompanied by reductions in the activity of hydrolytic enzymes involved in the acquisition of organic-bound nutrients by EMF and their host plants. W+RR thereby created an enhanced potential for soil organic matter (SOM) breakdown and nitrogen mineralisation by decomposers, as revealed by 127-190% increases in dissolved organic carbon and nitrogen, respectively, and decreasing SOM content in soil. Climate aridification impacts on vegetation can cascade belowground through shifts in fungal guild structure that alter ecosystem biogeochemistry and accelerate SOM decomposition by reducing the Gadgil effect.

Degradation activity of fungal communities on avocado peel (Persea americana Mill.) in a solid-state process: mycobiota successions and trophic guild shifts.

To explore the capability of soil mycobiota to degrade avocado peel waste and identify relevant successions and trophic guild shifts, fungal communities from three environments with different land uses were evaluated in a solid-state process. Soil samples used as inoculum were collected from a pristine mature tropical forest, a traditionally managed Mayan land, and an intensively managed monospecific avocado plantation. Soil-substrate mixes were evaluated for 52 weeks to evaluate organic matter decay and the carbon-to-nitrogen ratio. Amplicon-based high-throughput sequencing from internally transcribed spacer (ITS) analysis revealed significant differences in fungal communities widely dominated by Fusarium sp. and Clonostachys sp.; however, less represented taxa showed relevant shifts concomitantly with organic matter content drops. Trophic guild assignment revealed different behaviors in fungal communities between treatments over the 52 weeks, suggesting distinct preconditioning of fungal communities in these environments. Overall, the results lead to the identification of promising degradation moments and inoculum sources for further consortia enrichment or bioprospecting efforts.

Mycorrhizal roots slow the decay of belowground litters in a temperate hardwood forest.

There is increasing evidence that plant roots and mycorrhizal fungi, whether living or dead, play a central role in soil carbon (C) cycling. Root-mycorrhizal-microbial interactions can both suppress and enhance litter decay, with the net result dependent upon belowground nutrient acquisition strategies and soil nutrient availability. We measured the net effect of living roots and mycorrhizal fungi on the decay of dead roots and fungal hyphae in a hardwood forest dominated by either sugar maple (Acer saccharum) or white oak (Quercus alba) trees. Root and fungal litter were allowed to decompose within root-ingrowth bags and root-exclusion cores. In conjunction with root effects on decay, we assessed foraging responses and root induced changes in soil moisture, nitrogen (N) availability and enzyme activity. After 1 year, maple root production increased, and mycorrhizal fungal colonization decreased in the presence of decaying litter. In addition, we found that actively foraging roots suppressed the decay of root litter (- 14%) more than fungal litter (- 3%), and suppression of root decay was stronger for oak (- 20%) than maple roots (- 8%). Suppressive effects of oak roots on decay were greatest when roots also reduced soil N availability, which corresponded with reductions in hydrolytic enzyme activity and enhanced oxidative enzyme activities. These findings further our understanding of context-dependent drivers of root-mycorrhizal-microbial interactions and demonstrate that such interactions can play an underappreciated role in soil organic matter accumulation and turnover in temperate forests.

Decelerated carbon cycling by ectomycorrhizal fungi is controlled by substrate quality and community composition.

Interactions between symbiotic ectomycorrhizal (EM) and free-living saprotrophs can result in significant deceleration of leaf litter decomposition. While this phenomenon is widely cited, its generality remains unclear, as both the direction and magnitude of EM fungal effects on leaf litter decomposition have been shown to vary among studies. Here we explicitly examine how contrasting leaf litter types and EM fungal communities may lead to differential effects on carbon (C) and nitrogen (N) cycling. Specifically, we measured the response of soil nutrient cycling, litter decay rates, litter chemistry and fungal community structure to the reduction of EM fungi (via trenching) with a reciprocal litter transplant experiment in adjacent Pinus- or Quercus-dominated sites. We found clear evidence of EM fungal suppression of C and N cycling in the Pinus-dominated site, but no suppression in the Quercus-dominated site. Additionally, in the Pinus-dominated site, only the Pinus litter decay rates were decelerated by EM fungi and were associated with decoupling of litter C and N cycling. Our results support the hypothesis that EM fungi can decelerate C cycling via N competition, but strongly suggest that the 'Gadgil effect' is dependent on both substrate quality and EM fungal community composition. We argue that understanding tree host traits as well as EM fungal functional diversity is critical to a more mechanistic understanding of how EM fungi mediate forest soil biogeochemical cycling.

Resource-ratio theory predicts mycorrhizal control of litter decomposition.

Ecosystems with ectomycorrhizal plants have high soil carbon : nitrogen ratios, but it is not clear why. The Gadgil effect, where competition between ectomycorrhizal and saprotrophic fungi for nitrogen slows litter decomposition, may increase soil carbon. However, experimental evidence for the Gadgil effect is equivocal. Here, we apply resource-ratio theory to assess whether interguild fungal competition for different forms of organic nitrogen can affect litter decomposition. We focus on variation in resource input ratios and fungal resource use traits, and evaluate our model's predictions by synthesizing prior experimental literature examining ectomycorrhizal effects on litter decomposition. In our model, resource input ratios determined whether ectomycorrhizal fungi suppressed saprotrophic fungi. Recalcitrant litter inputs favored the former over the latter, allowing the Gadgil effect only when such inputs predominated. Although ectomycorrhizal fungi did not always hamper litter decomposition, ectomycorrhizal nitrogen uptake always increased carbon : nitrogen ratios in litter. Our meta-analysis of empirical studies supports our theoretical results: ectomycorrhizal fungi appear to slow decomposition of leaf litter only in forests where litter inputs are highly recalcitrant. We thus find that the specific contribution of the Gadgil effect to high soil carbon : nitrogen ratios in ectomycorrhizal ecosystems may be smaller than predicted previously.

Anthropogenic nitrogen enrichment enhances soil carbon accumulation by impacting saprotrophs rather than ectomycorrhizal fungal activity.

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal forest soils. However, it is unclear how free-living saprotrophs (bacteria and fungi, SAP) and ectomycorrhizal (EM) fungi responses to N addition impact soil C dynamics. Our aim was to investigate how SAP and EM communities are impacted by N enrichment and to estimate whether these changes influence decay of litter and humus. We conducted a long-term experiment in northern Sweden, maintained since 2004, consisting of ambient, low N additions (0, 3, 6, and 12 kg N ha-1  year-1 ) simulating current N deposition rates in the boreal region, as well as a high N addition (50 kg N ha-1  year-1 ). Our data showed that long-term N enrichment impeded mass loss of litter, but not of humus, and only in response to the highest N addition treatment. Furthermore, our data showed that EM fungi reduced the mass of N and P in both substrates during the incubation period compared to when only SAP organisms were present. Low N additions had no effect on microbial community structure, while the high N addition decreased fungal and bacterial biomasses and altered EM fungi and SAP community composition. Actinomycetes were the only bacterial SAP to show increased biomass in response to the highest N addition. These results provide a mechanistic understanding of how anthropogenic N enrichment can influence soil C accumulation rates and suggest that current N deposition rates in the boreal region (≤12 kg N ha-1  year-1 ) are likely to have a minor impact on the soil microbial community and the decomposition of humus and litter.

Below-ground organic matter accumulation along a boreal forest fertility gradient relates to guild interaction within fungal communities.

Plant-soil interactions link ecosystem fertility and organic matter accumulation below ground. Soil microorganisms play a central role as mediators of these interactions, but mechanistic understanding is still largely lacking. Correlative data from a coniferous forest ecosystem support the hypothesis that interactions between fungal guilds play a central role in regulating organic matter accumulation in relation to fertility. With increasing ecosystem fertility, the proportion of saprotrophic basidiomycetes increased in deeper organic layers, at the expense of ectomycorrhizal fungal species. Saprotrophs correlated positively with the activity of oxidative enzymes, which in turn favoured organic matter turnover and nitrogen recycling to plants. Combined, our findings are consistent with a fungus-mediated feedback loop, which results in a negative correlation between ecosystem fertility and below-ground carbon storage. These findings call for a shift in focus from plant litter traits to fungal traits in explaining organic matter dynamics and ecosystem fertility in boreal forests.

Revisiting the 'Gadgil effect': do interguild fungal interactions control carbon cycling in forest soils?

In forest ecosystems, ectomycorrhizal and saprotrophic fungi play a central role in the breakdown of soil organic matter (SOM). Competition between these two fungal guilds has long been hypothesized to lead to suppression of decomposition rates, a phenomenon known as the 'Gadgil effect'. In this review, we examine the documentation, generality, and potential mechanisms involved in the 'Gadgil effect'. We find that the influence of ectomycorrhizal fungi on litter and SOM decomposition is much more variable than previously recognized. To explain the inconsistency in size and direction of the 'Gadgil effect', we argue that a better understanding of underlying mechanisms is required. We discuss the strengths and weaknesses of each of the primary mechanisms proposed to date and how using different experimental methods (trenching, girdling, microcosms), as well as considering different temporal and spatial scales, could influence the conclusions drawn about this phenomenon. Finally, we suggest that combining new research tools such as high-throughput sequencing with experiments utilizing natural environmental gradients will significantly deepen our understanding of the 'Gadgil effect' and its consequences on forest soil carbon and nutrient cycling.

Checklist of Macrofungi Associated with Nine Different Habitats of Taburno-Camposauro Massif in Campania, Southern Italy.

The checklist serves as an informative method for evaluating the diversity, geography, and ecology of established and reproducing macrofungi. Additionally, considering macrofungi as bioindicator species, their census should be incorporated into efforts to monitor the state of health of ecosystems and directly applied to conservation policies. Between 2019 and 2023, a census of macrofungal species was conducted in Taburno-Camposauro Regional Park (Campania, Italy) across nine distinct habitats. A total of 453 fungal taxa were identified, including several new records for the Campania region. The fungal diversity exhibited significant variations based on the dominant plant species in each habitat. Fagacean tree species and Carpinus spp. shared similar fungal communities. Equally, coniferous tree species displayed a comparable fungal composition. In Abies alba and mixed broad-leaved forests, low levels of ectomycorrhizal taxa were observed alongside a concurrent increase in saprotrophs, indicating a disturbed habitat and a reduction in the Gadgil effect. Notably, lower fungal diversity was documented in the grassland habitat, suggesting the potential implications of wildlife imbalance and excessive grazing. The provided checklist constitutes a valuable resource for local management authorities, providing insights to formulate specific management policies.

Interguild fungal competition in litter and soil inversely modulate microbial necromass accumulation during Loess Plateau forest succession.

Microbial interactions determine ecosystem carbon (C) and nutrient cycling, yet it remains unclear how interguild fungal interactions modulate microbial residue contribution to soil C pools (SOC) during forest succession. Here, we present a region-wide investigation of the relative dominance of saprophytic versus symbiotic fungi in litter and soil compartments, exploring their linkages to soil microbial residue pools and potential drivers along a chronosequence of secondary Chinese pine (Pinus tabulaeformis) forests on the Loess Plateau. Despite minor changes in C and nitrogen (N) stocks in the litter or soil layers across successional stages, we found significantly lower soil phosphorus (P) stocks, higher ratios of soil C: N, soil N: P and soil C: P but lower ratios of litter C: N and litter C: P in old (>75 years) than young stands (<30 years). Pine stand development altered the saprotroph: symbiotroph ratios of fungal communities to favor the soil symbiotrophs versus the litter saprotrophs. The dominance of saprotrophs in litter is positively related to microbial necromass contribution to SOC, which is negatively related to the dominance of symbiotrophs in soils. Antagonistic interguild fungal competition in litter and soil layers, in conjunction with increased fungal but decreased bacterial necromass contribution to SOC, jointly contribute to unchanged total necromass contribution to SOC with stand development. The saprotroph: symbiotroph ratios in litter and soil layers are mainly driven by soil P stocks and stand parameters (e.g., stand age and slope), respectively, while substrate stoichiometries primarily regulate microbial necromass accumulation and fungal: bacterial necromass ratios. These results provide novel insights into how microbial interactions at local spatial scales modulate temporal changes in SOC pools, with management implications for mitigating regional land degradation.

Shifts in the Abundances of Saprotrophic and Ectomycorrhizal Fungi With Altered Leaf Litter Inputs.

Ectomycorrhizal (EcM) and saprotrophic fungi interact in the breakdown of organic matter, but the mechanisms underlying the EcM role on organic matter decomposition are not totally clear. We hypothesized that the ecological relations between EcM and saprotroph fungi are modulated by resources availability and accessibility, determining decomposition rates. We manipulated the amount of leaf litter inputs (No-Litter, Control Litter, Doubled Litter) on Trenched (root exclusion) and Non-Trenched plots (with roots) in a temperate deciduous forest of EcM-associated trees. Resultant shifts in soil fungal communities were determined by phospholipid fatty acids and DNA sequencing after 3 years, and CO2 fluxes were measured throughout this period. Different levels of leaf litter inputs generated a gradient of organic substrate availability and accessibility, altering the composition and ecological relations between EcM and saprotroph fungal communities. EcM fungi dominated at low levels of fresh organic substrates and lower organic matter quality, where short-distances exploration types seem to be better competitors, whereas saprotrophs and longer exploration types of EcM fungi tended to dominate at high levels of leaf litter inputs, where labile organic substrates were easily accessible. We were, however, not able to detect unequivocal signs of competition between these fungal groups for common resources. These results point to the relevance of substrate quality and availability as key factors determining the role of EcM and saprotroph fungi on litter and soil organic matter decay and represent a path forward on the capacity of organic matter decomposition of different exploration types of EcM fungi.

Restriction of plant roots in boreal forest organic soils affects the microbial community but does not change the dominance from ectomycorrhizal to saprotrophic fungi.

Boreal forest soils store significant amounts of carbon and are cohabited by saprotrophic and ectomycorrhizal fungi (ECM). The 'Gadgil effect' implies antagonistic interactions between saprotrophic fungi and ECM. Plant photosynthates support the competitive fitness of the ECM, and may also shape the soil bacterial communities. Many 'Gadgil effect' experiments have focused on litter layer (OL) or have litter and root-fragments present, and thus possibly favor the saprotrophs. We compared how the restriction of plant roots and exudates affect soil microbial community structures in organic soil (mixed OF and OH). For this, we established a 3-yr field experiment with 3 different mesh treatments affecting the penetration of plant roots and external fungal hyphae. Exclusion of plant photosynthates induced modest changes in both fungal and bacterial community structures, but not to potential functionality of the microbial community. The microbial community was resilient towards rather short-term disturbances. Contrary to the 'Gadgil effect', mesh treatments restricting the entrance of plant roots and external fungal hyphae did not favor saprotrophs that originally inhabited the soil. Thus, we propose that different substrate preferences (fresh litter vs. fermented or humified soil), rather than antagonism, maintain the spatial separation of saprotrophs and mycorrhizal fungi in boreal forest soils.

Degradation of the polycyclic aromatic hydrocarbon (PAH) fluorene is retarded in a Scots pine ectomycorrhizosphere.

• Polycyclic aromatic hydrocarbons (PAHs) are an important class of persistent organic pollutants (POPs) in the environment and accumulate in forest soils. These soils are often dominated by ectomycorrhizal (EcM) roots, but little is known about how EcM fungi degrade PAHs, or the overall effect of field colonized EcM roots on the fate of PAHs. • The ability of eight EcM fungi to degrade PAHs in liquid culture spiked with 14 C labelled PAHs was investigated. Microcosms were used to determine the impact of naturally colonized mycorrhizal pine seedlings on PAH mineralization and volatilization. • Only two EcM fungi (Thelephora terrestris and Laccaria laccata) degraded at least one PAH and none were able to mineralize the PAHs in pure culture. Where degradation occurred, the compounds were only mono-oxygenated. EcM pine seedlings did not alter naphthalene mineralization or volatilization but retarded fluorene mineralization by 35% compared with unplanted, ectomycorrhizosphere soil inoculated, microcosms. • The EcM fungi possessed limited PAH degrading abilities, which may explain why EcM dominated microcosms retarded fluorene mineralization. This observation is considered in relation to the 'Gadgil-effect', where retarded litter decomposition has been observed in the presence of EcM roots.