Slowed Biogeochemical Cycling in Sub-arctic Birch Forest Linked to Reduced Mycorrhizal Growth and Community Change after a Defoliation Event.
ABSTRACT: Sub-arctic birch forests (Betula pubescens Ehrh. ssp. czerepanovii) periodically suffer large-scale defoliation events caused by the caterpillars of the geometrid moths Epirrita autumnata and Operophtera brumata. Despite their obvious influence on ecosystem primary productivity, little is known about how the associated reduction in belowground C allocation affects soil processes. We quantified the soil response following a natural defoliation event in sub-arctic Sweden by measuring soil respiration, nitrogen availability and ectomycorrhizal fungi (EMF) hyphal production and root tip community composition. There was a reduction in soil respiration and an accumulation of soil inorganic N in defoliated plots, symptomatic of a slowdown of soil processes. This coincided with a reduction of EMF hyphal production and a shift in the EMF community to lower autotrophic C-demanding lineages (for example, /russula-lactarius). We show that microbial and nutrient cycling processes shift to a slower, less C-demanding state in response to canopy defoliation. We speculate that, amongst other factors, a reduction in the potential of EMF biomass to immobilise excess mineral nitrogen resulted in its build-up in the soil. These defoliation events are becoming more geographically widespread with climate warming, and could result in a fundamental shift in sub-arctic ecosystem processes and properties. EMF fungi may be important in mediating the response of soil cycles to defoliation and their role merits further investigation.
Project description:Loss of photosynthetic area can affect soil microbial communities by altering the availability of fixed carbon. We used denaturing gradient gel electrophoresis (DGGE) and Biolog filamentous-fungus plates to determine the effects of artificial defoliation of pines in a mixed pine-spruce forest on the composition of the fungal community in a forest soil. As measured by DGGE, two fungal species were affected significantly by the defoliation of pines (P < 0.001); the frequency of members of the ectomycorrhizal fungus genus Cenococcum decreased significantly, while the frequency of organisms of an unidentified soil fungus increased. The decrease in the amount of Cenococcum organisms may have occurred because of the formation of extensive hyphal networks by species of this genus, which require more of the carbon fixed by their host, or because this fungus is dependent upon quantitative differences in spruce root exudates. The defoliation of pines did not affect the overall composition of the soil fungal community or fungal-species richness (number of species per core). Biolog filamentous-fungus plate assays indicated a significant increase (P < 0.001) in the number of carbon substrates utilized by the soil fungi and the rate at which these substrates were used, which could indicate an increase in fungal-species richness. Thus, either small changes in the soil fungal community give rise to significant increases in physiological capabilities or PCR bias limits the reliability of the DGGE results. These data indicate that combined genetic and physiological assessments of the soil fungal community are needed to accurately assess the effect of disturbance on indigenous microbial systems.
Project description:The mechanisms linking C/N balance to N uptake and assimilation are central to plant responses to changing soil nutrient levels. Defoliation and subsequent regrowth of grasses both impact C partitioning, thereby creating a significant point of interaction with soil N availability. Using defoliation as an experimental treatment, we investigated the dynamic relationships between plant carbohydrate status and NO3--responsive uptake systems, transporter gene expression, and nitrate assimilation in Lolium perenne L. High- and low-affinity NO3- uptake was reduced in an N-dependent manner in response to a rapid and large shift in carbohydrate remobilization triggered by defoliation. This reduction in NO3- uptake was rescued by an exogenous glucose supplement, confirming the carbohydrate dependence of NO3- uptake. The regulation of NO3- uptake in response to the perturbation of the plant C/N ratio was associated with changes in expression of putative high- and low-affinity NO3- transporters. Furthermore, NO3- assimilation appears to be regulated by the C-N status of the plant, implying a mechanism that signals the availability of C metabolites for NO3- uptake and assimilation at the whole-plant level. We also show that cytokinins may be involved in the regulation of N acquisition and assimilation in response to the changing plant C/N ratio.
Project description:Soils of northern temperate and boreal forests represent a large terrestrial carbon (C) sink. The fate of this C under elevated atmospheric CO2 and climate change is still uncertain. A fundamental knowledge gap is the extent to which ectomycorrhizal fungi (EMF) and saprotrophic fungi contribute to C cycling in the systems by soil organic matter (SOM) decomposition. In this study, we used a novel approach to generate and compare enzymatically active EMF hyphae-dominated and saprotrophic hyphae-enriched communities under field conditions. Fermentation-humus (FH)-filled mesh bags, surrounded by a sand barrier, effectively trapped EMF hyphae with a community structure comparable to that found in the surrounding FH layer, at both trophic and taxonomic levels. In contrast, over half the sequences from mesh bags with no sand barrier were identified as belonging to saprotrophic fungi. The EMF hyphae-dominated systems exhibited levels of hydrolytic and oxidative enzyme activities that were comparable to or higher than saprotroph-enriched systems. The enzymes assayed included those associated with both labile and recalcitrant SOM degradation. Our study shows that EMF hyphae are likely important contributors to current SOM turnover in sub-boreal systems. Our results also suggest that any increased EMF biomass that might result from higher below-ground C allocation by trees would not suppress C fluxes from sub-boreal soils.
Project description:Climate warming at high northern latitudes has caused substantial increases in plant productivity of tundra vegetation and an expansion of the range of deciduous shrub species. However significant the increase in carbon (C) contained within above-ground shrub biomass, it is modest in comparison with the amount of C stored in the soil in tundra ecosystems. Here, we use a 'space-for-time' approach to test the hypothesis that a shift from lower-productivity tundra heath to higher-productivity deciduous shrub vegetation in the sub-Arctic may lead to a loss of soil C that out-weighs the increase in above-ground shrub biomass. We further hypothesize that a shift from ericoid to ectomycorrhizal systems coincident with this vegetation change provides a mechanism for the loss of soil C. We sampled soil C stocks, soil surface CO2 flux rates and fungal growth rates along replicated natural transitions from birch forest (Betula pubescens), through deciduous shrub tundra (Betula nana) to tundra heaths (Empetrum nigrum) near Abisko, Swedish Lapland. We demonstrate that organic horizon soil organic C (SOCorg ) is significantly lower at shrub (2.98 ± 0.48 kg m(-2) ) and forest (2.04 ± 0.25 kg m(-2) ) plots than at heath plots (7.03 ± 0.79 kg m(-2) ). Shrub vegetation had the highest respiration rates, suggesting that despite higher rates of C assimilation, C turnover was also very high and less C is sequestered in the ecosystem. Growth rates of fungal hyphae increased across the transition from heath to shrub, suggesting that the action of ectomycorrhizal symbionts in the scavenging of organically bound nutrients is an important pathway by which soil C is made available to microbial degradation. The expansion of deciduous shrubs onto potentially vulnerable arctic soils with large stores of C could therefore represent a significant positive feedback to the climate system.
Project description:To understand the responses to external disturbance such as defoliation and possible feedback mechanisms at global change in terrestrial ecosystems, it is necessary to examine the extent and nature of effects on aboveground-belowground interactions. We studied a temperate heathland system subjected to experimental climate and atmospheric factors based on prognoses for year 2075 and further exposed to defoliation. By defoliating plants, we were able to study how global change modifies the interactions of the plant-soil system. Shoot production, root biomass, microbial biomass, and nematode abundance were assessed in the rhizosphere of manually defoliated patches of Deschampsia flexuosa in June in a full-factorial FACE experiment with the treatments: increased atmospheric CO 2, increased nighttime temperatures, summer droughts, and all of their combinations. We found a negative effect of defoliation on microbial biomass that was not apparently affected by global change. The negative effect of defoliation cascades through to soil nematodes as dependent on CO 2 and drought. At ambient CO 2, drought and defoliation each reduced nematodes. In contrast, at elevated CO 2, a combination of drought and defoliation was needed to reduce nematodes. We found positive effects of CO 2 on root density and microbial biomass. Defoliation affected soil biota negatively, whereas elevated CO 2 stimulated the plant-soil system. This effect seen in June is contrasted by the effects seen in September at the same site. Late season defoliation increased activity and biomass of soil biota and more so at elevated CO 2. Based on soil biota responses, plants defoliated in active growth therefore conserve resources, whereas defoliation after termination of growth results in release of resources. This result challenges the idea that plants via exudation of organic carbon stimulate their rhizosphere biota when in apparent need of nutrients for growth.
Project description:Fire is a major natural disturbance factor in boreal forests, and the frequency of forest fires is predicted to increase due to climate change. Nitrogen (N) is a key determinant of carbon sequestration in boreal forests because the shortage of N limits tree growth. We studied changes in N pools and fluxes, and the overall N balance across a 155-year non stand-replacing fire chronosequence in sub-arctic Pinus sylvestris forests in Finland. Two years after the fire, total ecosystem N pool was 622 kg ha-1 of which 16% was in the vegetation, 8% in the dead biomass and 76% in the soil. 155 years after the fire, total N pool was 960 kg ha-1, with 27% in the vegetation, 3% in the dead biomass and 69% in the soil. This implies an annual accumulation rate of 2.28 kg ha-1 which was distributed equally between soil and biomass. The observed changes in N pools were consistent with the computed N balance +2.11 kg ha-1 yr-1 over the 155-year post-fire period. Nitrogen deposition was an important component of the N balance. The biological N fixation increased with succession and constituted 9% of the total N input during the 155 post-fire years. N2O fluxes were negligible (? 0.01 kg ha-1 yr-1) and did not differ among post-fire age classes. The number and intensity of microbial genes involved in N cycling were lower at the site 60 years after fire compared to the youngest and the oldest sites indicating potential differences in soil N cycling processes. The results suggest that in sub-arctic pine forests, the non-stand-replacing, intermediate-severity fires decrease considerably N pools in biomass but changes in soil and total ecosystem N pools are slight. Current fire-return interval does not seem to pose a great threat to ecosystem productivity and N status in these sub-arctic forests.
Project description:Background:The post-harvest recovery and sustained productivity of Nothofagus pumilio forests in Tierra del Fuego may be affected by the abundance and composition of ectomycorrhizal fungi (EMF). Timber harvesting alters EMF community structure in many managed forests, but the impacts of harvesting can vary with the management strategy. The implementation of variable retention (VR) management can maintain, increase, or decrease the diversity of many species, but the effects of VR on EMF in the forests of southern Patagonia have not been studied, nor has the role of EMF in the regeneration process of these forests. Methods:We evaluated the effects of VR management on the EMF community associated with N. pumilio seedlings. We quantified the abundance, composition, and diversity of EMF across aggregate (AR) and dispersed (DR) retention sites within VR managed areas, and compared them to primary forest (PF) unmanaged stands. EMF assemblage and taxonomic identities were determined by ITS-rDNA sequencing of individual root tips sampled from 280 seedlings across three landscape replicates. To better understand seedling performance, we tested the relationships between EMF colonization, EMF taxonomic composition, seedling biomass, and VR treatment. Results:The majority of EMF taxa were Basidiomycota belonging to the families Cortinariaceae (n = 29), Inocybaceae (n = 16), and Thelephoraceae (n = 8), which was in agreement with other studies of EMF diversity in Nothofagus forests. EMF richness and colonization was reduced in DR compared to AR and PF. Furthermore, EMF community composition was similar between AR and PF, but differed from the composition in DR. EMF community composition was correlated with seedling biomass and soil moisture. The presence of Peziza depressa was associated with higher seedling biomass and greater soil moisture, while Inocybe fibrillosibrunnea and Cortinarius amoenus were associated with reduced seedling biomass and lower soil moisture. Seedling biomass was more strongly related to retention type than EMF colonization, richness, or composition. Discussion:Our results demonstrate reduced EMF attributes and altered composition in VR treatments relative to PF stands, with stronger impacts in DR compared to AR. This suggests that VR has the potential to improve the conservation status of managed stands by supporting native EMF in AR. Our results also demonstrate the complex linkages between retention treatments, fungal community composition, and tree growth at individual and stand scales.
Project description:Warmer and drier conditions associated with ongoing climate change will increase abiotic stress for plants and mycorrhizal fungi in drylands worldwide, thereby potentially reducing vegetation cover and productivity and increasing the risk of land degradation and desertification. Rhizosphere microbial interactions and feedbacks are critical processes that could either mitigate or aggravate the vulnerability of dryland vegetation to forecasted climate change.We conducted a four-year manipulative study in a semiarid shrubland in the Iberian Peninsula to assess the effects of warming (~2.5ºC; W), rainfall reduction (~30%; RR) and their combination (W+RR) on the performance of native shrubs (Helianthemum squamatum) and their associated mycorrhizal fungi.Warming (W and W+RR) decreased the net photosynthetic rates of H. squamatum shrubs by ~31% despite concurrent increases in stomatal conductance (~33%), leading to sharp decreases (~50%) in water use efficiency. Warming also advanced growth phenology, decreased leaf nitrogen and phosphorus contents per unit area, reduced shoot biomass production by ~36% and decreased survival during a dry year in both W and W+RR plants. Plants under RR showed more moderate decreases (~10-20%) in photosynthesis, stomatal conductance and shoot growth.Warming, RR and W+RR altered ectomycorrhizal fungal (EMF) community structure and drastically reduced the relative abundance of EMF sequences obtained by high-throughput sequencing, a response associated with decreases in the leaf nitrogen, phosphorus and dry matter contents of their host plants. In contrast to EMF, the community structure and relative sequence abundances of other non-mycorrhizal fungal guilds were not significantly affected by the climate manipulation treatments.Synthesis: Our findings highlight the vulnerability of both native plants and their symbiotic mycorrhizal fungi to climate warming and drying in semiarid shrublands, and point to the importance of a deeper understanding of plant-soil feedbacks to predict dryland vegetation responses to forecasted aridification. The interdependent responses of plants and ectomycorrhizal fungi to warming and rainfall reduction may lead to a detrimental feedback loop on vegetation productivity and nutrient pool size, which could amplify the adverse impacts of forecasted climate change on ecosystem functioning in EMF-dominated drylands.
Project description:It is essential to understand how the loss of biodiversity impacts both ecosystem function (EF) and multifunctionality (EMF). Previous studies have mostly focused on predicting how species richness (SR) impacts EMF, while the effect of functional diversity (FD) on EMF remains unclear. Specifically, we know little about the primary functional drivers impacting EMF compared with SR. Therefore, we analysed 8 ecosystem functions within 58 natural secondary forest plots to investigate the effect of FD on both individual EF and EMF. Our results suggest that SR and FD had very significant positive effects on plant phosphorus, soil available phosphorus, and soil total nitrogen. FD explained significantly more variations in these functional responses than SR for individual ecosystem functioning. We also used a multiple threshold approach to test the effect of SR and FD on EMF. We found that FD and SR were positively related to EMF regardless of whether low-level function or high-level function was desired, but FD had a larger effect than SR. Based on the averaging approach, OLS regression, multivariate linear regression model and random forest analysis, we found that SR and FD were both drivers of EMF but that FD had a stronger effect and could explain more variation. As such, we conclude that FD drives ecosystem multifunctionality more than SR.
Project description:The increased demand for harvesting energy wood raises questions about its effects on the functioning of the forest ecosystems, soil processes and biodiversity. Impacts of tree stump removal on ectomycorrhizal fungal (EMF) communities of Norway spruce saplings were studied with 454-pyrosequencing in a 3-year field experiment replicated in 3 geographical areas. This is possibly the most thorough investigation of EMF communities associated with saplings grown on sites subjected to energy wood harvesting. To separate impacts of tree stump and logging residue removal on EMF and plant variables, we used three harvesting treatments with increasing complexity from patch mounding alone (P) to patch mounding combined with logging residue removal (RP), and patch mounding combined with both logging residue and stump removal (SRP). Saplings grown in uncut forests (F) served as references for harvesting treatments. A majority of sequences (>92%) and operational taxonomic units (OTUs, 55%) were assigned as EMF. EMF OTU richness, fungal community composition or sapling growth did not differ between harvesting treatments (P, RP and SRP), while EMF OTU richness, diversity and evenness were highest and sapling growth lowest in the undisturbed reference forests (F). The short study period may partially explain the similarities in fungal and sapling variables in different harvesting treatments. In conclusion, our results indicate that neither stump removal nor logging residue removal have significant additional negative impacts on EMF communities or growth of Norway spruce saplings in the short-term compared with the impacts of more conventional harvesting methods, including clear cutting and patch mounding.