Gadgil Effect Research Articles

Field Reduction of Ectomycorrhizal Fungi Has Cascading Effects on Soil Microbial Communities and Reduces the Abundance of Ectomycorrhizal Symbiotic Bacteria.

Specific interactions between bacteria and ectomycorrhizal fungi (EcMF) can benefit plant health, and saprotrophic soil fungi represent a potentially antagonistic guild to these mutualisms. Yet there is little field-derived experimental evidence showing how the relationship among these three organismal groups manifests across time. To bridge this knowledge gap, we experimentally reduced EcMF in forest soils and monitored both bacterial and fungal soil communities over the course of a year. Our analyses demonstrate that soil trenching shifts the community composition of fungal communities towards a greater abundance of taxa with saprotrophic traits, and this shift is linked to a decrease in both EcMF and a common ectomycorrhizal helper bacterial genus, Burkholderia, in a time-dependent manner. These results not only reveal the temporal nature of a widespread tripartite symbiosis between bacteria, EcMF and a shared host tree, but they also refine our understanding of the commonly referenced 'Gadgil effect' by illustrating the cascading effects of EcMF suppression and implicating soil saprotrophic fungi as potential antagonists on bacterial-EcMF interactions.

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.

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.

Carbon cycle in tropical upland ecosystems: a global review

Abstract. Along with habitat transformation, climate change has profound impacts on biodiversity and may alter ecosystem services on which human welfare depends. Many studies of the carbon cycle have focused on lowland tropical forests; however, upland forests have been less explored despite their pivotal role in carbon sequestration. Here, I synthesized the state of knowledge on the allocation of carbon in its different stocks (aboveground, belowground, and soil) as well as in its main fluxes (plant decomposition, respiration, and litterfall) in tropical upland ecosystems of the planet. In November 2020, a systematic review was carried out to identify references published from 2000 to 2020 through a combination of key terms in Google Scholar and Scopus databases, thus analysing bibliographic, geographical, methodological, and carbon cycling information of the global upland tropics (between 23.5∘ N–23.5∘ S). After analysing a total of 1967 references according to inclusion–exclusion criteria, 135 references published in the last 20 years were selected. Most of the studies were conducted in the tropical and subtropical moist broadleaf forest of South America. The main factors studied were elevation and forest type. Forest structure and soil variables were largely associated when studying carbon cycling in these ecosystems. Estimations of carbon stocks comprised three-fourths of the total studies, while the remaining fraction focused on carbon fluxes. Aboveground biomass and carbon in soils were highly investigated, while plant decomposition and respiration were the components that received the least attention. Even though in the last 20 years there was a slight increase in the number of studies on carbon cycle in tropical upland forests, I found bias associated with the biomes and ecoregions studied (especially in the Andes). Elevation was the main factor examined but other essential aspects such as the successional gradient, landscape management, diversity–productivity relationship, faunal and microbial effect, trophic cascades, and Gadgil effect require more attention. The inclusion of different litter species and origins (i.e. roots and stems) and theoretical frameworks including home-field advantage, substrate–matrix interaction, and phenology–substrate match may provide explanatory mechanisms to better understand litter decomposition over these forests. Despite respiration being a paramount link that is closely tied to above- and belowground compartment, this flux constitutes one of the important gaps to fulfil in future research. For a comprehensive understanding of the carbon cycle in upland forests, it is necessary to obtain information on its main fluxes and integrate them into climate change mitigation plans.

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.

Partitioning of forest floor CO2 emissions reveals the belowground interactions between different plant groups in a Scots pine stand in southern Finland

Changes in the climate may have unpredictable effects on belowground carbon processes and thus, the carbon balance of boreal forests. To understand the interactions of these processes in soil and to quantify the potential changes in the carbon cycle, partitioning of forest floor respiration is crucial. For this purpose, we used nine different treatments to separate the sources of forest floor carbon dioxide (CO2) emissions in a mature Scots pine (Pinus sylvestris L.) stand in southern Finland. To partition the belowground CO2 emissions, we used two different trenching methods: 1) to exclude roots and mycorrhizal fungal mycelia (mesh with 1-µm pores) and 2) to exclude roots, but not mycorrhizal hyphae (mesh with 50-µm pores). Additionally, we used 3) a control treatment that included roots and fungal hyphae. To partition the CO2 emissions from the forest floor vegetation, we 1) removed it, 2) left only the dwarf shrubs, or 3) left the vegetation intact. The forest floor CO2 emissions were regularly measured with a flux chamber throughout the growing seasons in 2013–2015. The total forest floor respiration was partitioned into respiration of tree roots (contributing 48%), heterotrophic soil respiration (30%) and respiration of ground vegetation other than shrubs (10%), dwarf shrubs (8%), and hyphae of mycorrhizal fungi (4%). Heterotrophic respiration increased in the trenched treatments without ground vegetation over time, due to the so-called ‘Gadgil effect’. In the absence of tree roots, but when hyphal access was allowed, respiration in the dwarf shrub treatment increased throughout the experiment. This indicated that dwarf shrubs had fungal connections to outside the experimental plots via their ericoid mycorrhiza. At the same time, other ground vegetation, such as mosses, suppressed the dwarf shrub respiration in trenched treatments. Our results show that competition on the forest floor is intense between plant roots and soil microbes.

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.

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.

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.

No “Gadgil effect”: Temperate tree roots and soil lithology are effective predictors of wood decomposition

AbstractThe “Gadgil effect” hypothesizes that root associations may slow down decomposition through pre‐emptive competition. In the context of recalcitrant litter decomposition, specifically coarse wood debris, it is uncertain as to what is the relative importance of soil communities associated with living roots when compared to those without roots. Here, it is hypothesized that the presence of live roots and active photosynthates will enhance wood decomposition. To test this hypothesis, the presence or absence of temperate tree roots was used in this study. Sugar maple (Acer saccharum) and white oak (Quercus alba) roots were manipulated at three sites of either limestone or shale parent rock residuum. At each site, wood substrate was placed in soils beneath the canopy of either A. saccharum or Q. alba, while in the presence of roots (root+). At the same time, wood substrate was placed in the same soil community, but live root exposure was eliminated by trenching (root−). This eliminated active photosynthate supply to the soil microbial community. Results determined that live root exposure promoted faster decomposition and greater mycelial colonization of wood substrate. Also, sites of shale parent rock residuum had higher rates of decomposition in comparison with limestone parent rock residuum. Although additional work is needed to determine the extent in which roots and lithology can facilitate wood decomposition, these findings suggest that living roots impact decomposers and provide a pathway towards humus and soil organic matter formation.

Contrasting effects of ectomycorrhizal fungi on early and late stage decomposition in a boreal forest.

Symbiotic ectomycorrhizal fungi have received increasing attention as regulators of below-ground organic matter storage. They are proposed to promote organic matter accumulation by suppressing saprotrophs, but have also been suggested to play an active role in decomposition themselves. Here we show that exclusion of tree roots and associated ectomycorrhizal fungi in a boreal forest increased decomposition of surface litter by 11% by alleviating nitrogen limitation of saprotrophs-a "Gadgil effect". At the same time, root exclusion decreased Mn-peroxidase activity in the deeper mor layer by 91%. Our results show that ectomycorrhizal fungi may hamper short-term litter decomposition, but also support a crucial role of ectomycorrhizal fungi in driving long-term organic matter oxidation. These observations stress the importance of ectomycorrhizal fungi in regulation of below-ground organic matter accumulation. By different mechanisms they may either hamper or stimulate decomposition, depending upon stage of decomposition and location in the soil profile.

Fungal guilds are evenly distributed along a vertical spruce forest soil profile while individual fungi show pronounced niche partitioning

Saprotrophic and ectomycorrhizal (EcM) forest fungi decompose organic matter and mobilize nutrients for host plants, respectively. Competition between the two guilds may cause the so-called Gadgil effect, i.e., decreased litter decomposition rates resulting in increased carbon storage in soil. The Gadgil effect was supposed to even affect global climate, highlighting the necessity to understand fungal distribution and interactions in soil. Searching for evidence of competition between saprotrophic and mycorrhizal fungi, we analyzed the distribution of fungi along a well-stratified vertical spruce forest soil profile in two seasons, i.e., autumn and the following spring. The different soil strata (i.e., two mineral horizons and two organic layers) underneath the litter layer were colonized by distinct fungal communities, which included roughly consistent proportions of all fungal guilds and phyla at each time. However, the community composition changed quantitatively between the sampling dates. Along the vertical soil profile, it differed mostly between the organic layers and the mineral soil, which is supposed to be due to differences in the predominant energy sources (i.e., aboveground litter and rhizodeposition, respectively). Network analyses revealed co-occurrences (i.e., positive correlations of individual abundances) to outweigh mutual exclusions (i.e., negative correlations) between individual fungi in each soil stratum and season. This also applied for interactions between saprotrophic and EcM fungi. Network analyses therefore provided no indications for a possible Gadgil effect. However, considering individual nutrient use efficiencies might refine insights from network analyses in future studies and facilitate linking community dynamics to ecosystem processes.

Slow decomposition of leaf litter from mature Fagus sylvatica trees promotes offspring nitrogen acquisition by interacting with ectomycorrhizal fungi

Summary Leaf litter chemistry and ectomycorrhizal (ECM) fungi are key drivers of the below‐ground nitrogen (N) cycling within forest ecosystems. Their combined effects on litter decomposition and N competition between microbial decomposers and plants are still uncertain. We conducted a glasshouse microcosm experiment with low or high ECM‐colonized beech (Fagus sylvatica) saplings, growing with litter collected from old or young beech trees growing on the same loamy soil. After 6 months of growth, we investigated litter decomposition rates, microbial respiration and the N pools within leaf litter, soil (different pools), microbial and plant shoot biomass. We found that the mass loss of the litter from young trees was always higher than the litter from the mature trees. The microbial biomass N per unit soil carbon was low when the litter, especially from mature trees, was added, suggesting that the litter provided recalcitrant compounds and limited soil microbial activity. In contrast to the ‘Gadgil effect’ hypothesis, the high ECM‐colonized roots increased the litter decomposition rate and N immobilization in poorly decomposable litter in comparison with the litter incubated with the low ECM‐colonized roots. Finally, the high ECM‐colonized plants that received the poorly decomposable leaf litter exhibited the highest shoot N amount and biomass and were associated with the lowest microbial biomass N. Two‐way anovas revealed that litter and ECM fungi occurrence together impacted final particulate organic N, microbial biomass N, the amount of shoot N and shoot biomass. Four N pools are key drivers of microbial biomass N and shoot N: particulate organic N, total N, soil ammonium concentration and litter N concentration. Synthesis. Our results support the hypothesis that poorly decomposable leaf litter produced by mature beech trees and ECM fungi together decrease microbial N immobilization but increase tree N acquisition. Increasing N retention within the recalcitrant N forms in soil was identified as a key mechanism by which beech alters soil N cycling with potential positive feedbacks on its acquisition by the plant. Our result emphasizes the importance of considering within‐species litter‐trait variability in litter decomposition.

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.

Litter decomposition, ectomycorrhizal roots and the ‘Gadgil’ effect

○Ectomycorrhizas may interact with saprotrophic microorganisms to influence decomposition. These interactions are poorly understood. ○We buried mesh envelopes filled with either litter or F-layer material in the forest floor of a Pinus resinosa plantation. After 16 months we characterized decomposition, C: N ratio, water content, and the density of four ectomycorrhiza morphotypes. ○For both substrates there were negative correlations between ectomycorrhiza density and decomposition, and between ectomycorrhiza density and water content. The relative proportion of morphotypes was significantly different in litter and F-layer material buried at the same depth in the forest floor. ○We conclude that the negative effect of ectomycorrhizas on decomposition could be mediated by extraction of water, particularly in relatively dry years. The 'Gadgil effect' may be explained by this phenomenon. In wetter years, water extraction may be less consequential. Also, variation in substrate quality may influence the structure of ectomycorrhizal fungal communities.

Ectomycorrhizas and retarded decomposition in a Pinus resinosa plantation

Summary Ectomycorrhizas may interact with saprotrophic microorganisms to influence decomposition. These interactions are poorly understood. We buried mesh envelopes filled with either litter or F‐layer material in the forest floor of a Pinus resinosa plantation. After 16 months we characterized decomposition, C : N ratio, water content, and the density of four ectomycorrhiza morphotypes. For both substrates there were negative correlations between ectomycorrhiza density and decomposition, and between ectomycorrhiza density and water content. The relative proportion of morphotypes was significantly different in litter and F‐layer material buried at the same depth in the forest floor. We conclude that the negative effect of ectomycorrhizas on decomposition could be mediated by extraction of water, particularly in relatively dry years. The ‘Gadgil effect’ may be explained by this phenomenon. In wetter years, water extraction may be less consequential. Also, variation in substrate quality may influence the structure of ectomycorrhizal fungal communities.