Belowground Components Of Ecosystems Research Articles

Opposing responses of temporal stability of aboveground and belowground net primary productivity to water and nitrogen enrichment in a temperate grassland

AbstractChanges in water and nitrogen availability, as important elements of global environmental change, are known to affect the temporal stability of aboveground net primary productivity (ANPP). However, evidences for their effects on the temporal stability of belowground net primary productivity (BNPP), and whether such effects are consistent between belowground and aboveground, are rather scarce. Here, we investigated the responses of temporal stability of both ANPP and BNPP to water and nitrogen addition based on a 9‐year manipulative experiment in a temperate grassland in northern China. The results showed that the temporal stability of ANPP increased with water addition but decreased with nitrogen addition. By contrast, the temporal stability of BNPP decreased with water addition but increased with nitrogen enrichment. The temporal stability of ANPP was mainly determined by the soil moisture and inorganic nitrogen, which modulated species asynchrony, as well as by the stability of dominant species. On the other hand, the temporal stability of BNPP was mainly driven by the soil moisture and inorganic nitrogen that modulated ANPP of grasses, and by the direct effect of soil water availability. Our study provides the first evidence on the opposite responses of aboveground and belowground grassland temporal stability to increased water and nitrogen availability, highlighting the importance of considering both aboveground and belowground components of ecosystems for a more comprehensive understanding of their dynamics.

Contrasting fungal responses to wildfire across different ecosystem types.

Wildfire affects our planet's biogeochemistry both by burning biomass and by driving changes in ecological communities and landcover. Some plants and ecosystem types are threatened by increasing fire pressure while others respond positively to fire, growing in local and regional abundance when it occurs regularly. However, quantifying total ecosystem response to fire demands consideration of impacts not only on aboveground vegetation, but also on soil microbes like fungi, which influence decomposition and nutrient mineralization. If fire-resistant soil fungal communities co-occur with similarly adapted plants, these above- and belowground ecosystem components should shift and recover in relative synchrony after burning. If not, fire might decouple ecosystem processes governed by these different communities, affecting total functioning. Here, we use a natural experiment to test whether fire-dependent ecosystems host unique, fire-resistant fungal communities. We surveyed burned and unburned areas across two California ecosystem types with differing fire ecologies in the immediate aftermath of a wildfire, finding that the soil fungal communities of fire-dependent oak woodlands differ from those of neighbouring mixed evergreen forests. We discovered furthermore that the latter are more strongly altered compositionally by fire than the former, suggesting that differences in fungal community structure support divergent community responses to fire across ecosystems. Our results thus indicate that fire-dependent ecosystems may host fire-resistant fungal communities.

Enlightenments from potted landscapes (Bonsai and Penjing) for urban revegetation in the built environment

Impervious surfaces in modern cities can be used for revegetation. However, the ecological quality of belowground ecosystem components and constrained soil space are crucial factors that limit habitat restoration. The long-term development of Bonsai (Penjing) trees has resulted in mature theories and techniques that suggest optimal root-canopy ratios. Considering the similarities with constrained pot landscapes, some “enlightenments”, or key findings, derived from Bonsai theory can also be applied to urban special revegetation (USR). In this study, we analyzed the root-shoot correlations of the different styles of Bonsai trees and provide and assessment on the relationships between Bonsai theory and USR. Using an on-site survey and biomass experiment, nine “enlightenments” from Bonsai theory are proposed: positive root-canopy correlation, environmental stress increasing the root-shoot ratio, connectivity of the tree bark “water line”, rebalancing after disruption of the root-canopy balance, precise root and canopy design, artificial auxiliary fixation of the root, control of growth, horticultural reshaping skills, and plant diversity. In particular, for the successful establishment of urban rebuilt environments, it is essential to understand the mechanisms of root-canopy correlations. Therefore, we suggest that the four main types of USR (green roofs, vertical greening, container gardening, and street trees) should follow the core principle based on the root-shoot balance to effectively design both the root and canopy.

Above and belowground community strategies respond to different global change drivers

Environmental changes alter the diversity and structure of communities. By shifting the range of species traits that will be successful under new conditions, environmental drivers can also dramatically impact ecosystem functioning and resilience. Above and belowground communities jointly regulate whole-ecosystem processes and responses to change, yet they are frequently studied separately. To determine whether these communities respond similarly to environmental changes, we measured taxonomic and trait-based responses of plant and soil microbial communities to four years of experimental warming and nitrogen deposition in a temperate grassland. Plant diversity responded strongly to N addition, whereas soil microbial communities responded primarily to warming, likely via an associated decrease in soil moisture. These above and belowground changes were associated with selection for more resource-conservative plant and microbe growth strategies, which reduced community functional diversity. Functional characteristics of plant and soil microbial communities were weakly correlated (P = 0.07) under control conditions, but not when above or belowground communities were altered by either global change driver. These results highlight the potential for global change drivers operating simultaneously to have asynchronous impacts on above and belowground components of ecosystems. Assessment of a single ecosystem component may therefore greatly underestimate the whole-system impact of global environmental changes.

Tree ring-based reconstruction of the long-term influence of wildfires on permafrost active layer dynamics in Central Siberia

Although it has been recognized that rising temperatures and shifts in the hydrological cycle affect the depth of the seasonally thawing upper permafrost stratum, it remains unclear to what extent the frequency and intensity of wildfires, and subsequent changes in vegetation cover, influence the soil active layer on different spatiotemporal scales. Here, we use ring width measurements of the subterranean stem part of 15 larch trees from a Sphagnum bog site in Central Siberia to reconstruct long-term changes in the thickness of the active layer since the last wildfire occurred in 1899. Our approach reveals a three-step feedback loop between above- and belowground ecosystem components. After all vegetation is burned, direct atmospheric heat penetration over the first ~20 years caused thawing of the upper permafrost stratum. The slow recovery of the insulating ground vegetation reverses the process and initiates a gradual decrease of the active layer thickness. Due to the continuous spreading and thickening of the peat layer during the last decades, the upper permafrost horizon has increased by 0.52 cm/year. This study demonstrates the strength of annually resolved and absolutely dated tree-ring series to reconstruct the effects of historical wildfires on the functioning and productivity of boreal forest ecosystems at multi-decadal to centennial time-scale. In so doing, we show how complex interactions of above- and belowground components translate into successive changes in the active permafrost stratum. Our results are particularly relevant for improving long-term estimates of the global carbon cycle that strongly depends on the source and sink behavior of the boreal forest zone.

Dynamics of soil CO2 efflux under varying atmospheric CO2 concentrations reveal dominance of slow processes.

We evaluated the effect on soil CO2 efflux (FCO2 ) of sudden changes in photosynthetic rates by altering CO2 concentration in plots subjected to +200ppmv for 15years. Five-day intervals of exposure to elevated CO2 (eCO2 ) ranging 1.0-1.8 times ambient did not affect FCO2 . FCO2 did not decrease until 4months after termination of the long-term eCO2 treatment, longer than the 10days observed for decrease of FCO2 after experimental blocking of C flow to belowground, but shorter than the ~13months it took for increase of FCO2 following the initiation of eCO2 . The reduction of FCO2 upon termination of enrichment (~35%) cannot be explained by the reduction of leaf area (~15%) and associated carbohydrate production and allocation, suggesting a disproportionate contraction of the belowground ecosystem components; this was consistent with the reductions in base respiration and FCO2 -temperature sensitivity. These asymmetric responses pose a tractable challenge to process-based models attempting to isolate the effect of individual processes on FCO2 .

Leaf endophytic fungus interacts with precipitation to alter belowground microbial communities in primary successional dunes.

Understanding interactions between above- and belowground components of ecosystems is an important next step in community ecology. These interactions may be fundamental to predicting ecological responses to global change because indirect effects occurring through altered species interactions can outweigh or interact with the direct effects of environmental drivers. In a multiyear field experiment (2010-2015), we tested how experimental addition of a mutualistic leaf endophyte (Epichloë amarillans) associated with American beachgrass (Ammophila breviligulata) interacted with an altered precipitation regime (±30%) to affect the belowground microbial community. Epichloë addition increased host root biomass at the plot scale, but reduced the length of extraradical arbuscular mycorrhizal (AM) fungal hyphae in the soil. Under ambient precipitation alone, the addition of Epichloë increased root biomass per aboveground tiller and reduced the diversity of AM fungi in A. breviligulata roots. Furthermore, with Epichloë added, the diversity of root-associated bacteria declined with higher soil moisture, whereas in its absence, bacterial diversity increased with higher soil moisture. Thus, the aboveground fungal mutualist not only altered the abundance and composition of belowground microbial communities but also affected how belowground communities responded to climate, suggesting that aboveground microbes have potential for cascading influences on community dynamics and ecosystem processes that occur belowground.

Above- and belowground linkages of a nitrogen and phosphorus co-limited tropical mountain pasture system – responses to nutrient enrichment

Little is known about how N and P co-limited ecosystems respond to single nutrient enrichment. This work assesses the susceptibility of above- and belowground ecosystem components and of their linkages in an N and P co-limited pasture to N- and P-enrichment. We tested if the plants’ responses can be explained by the concept of serially linked nutrients introduced by Agren (Ecol Lett 7:185–191, 2004). In this concept, the control of the growth rate by one nutrient is assumed to depend on the control of a different cellular process by another nutrient. We investigated the responses of shoot and root biomass and C:N:P stoichiometry of the grass Setaria sphacelata (Schumach.) to moderate N, P, and N + P application over 5 years. In addition, the effects of nutrient enrichment on soil nutrient pools, on arbuscular mycorrhizal fungi (AMF) as well as on microbial biomass, activity, and community structure (phospholipid fatty acids: PLFA) were tested. In order to evaluate the importance of different factors explaining microbial responses, we applied a likelihood-based information-theoretic approach. The application of N + P increased aboveground grass biomass (+61 %). Root biomass was stimulated by P-treatment (+45 %). Grass C:N:P stoichiometry responded by altering the P-uptake (P-treatment) or by translocating P from shoot to root (N-treatment). In particular, root C:N and C:P stoichiometry decreased in P- and in N-treatment. Extractable fractions of soil C, N, and P were significantly affected by nutrient enrichment. P application increased the biomass of Gram-positive bacteria (+22 %) and the abundance of AMF (+46 %), however, results of the IT-approach suggested indirect effects of nutrient enrichment on microbes. The responses of the N and P co-limited pasture to particular nutrient enrichment support the concept of serially linked nutrients. The present study provides evidence for the fundamental importance of P for controlling resource allocation of plants in responses to nutrient enrichment. Resource allocation of the grass rather than direct effects of nutrient additions drives changes in AMF, microbial biomass, community structure, and activity.

Multi-set indicators to assess environmental quality using soil microarthropods, plants and humus

The present study used a multi-set indicator approach to evaluate environmental quality in complex ecosystem. The simultaneous analysis of information deriving from different ecosystem scales and/or taxonomic groups was used to achieve a better understanding of the interconnected above- and below-ground ecosystem components. Three indicator sets were utilized in eight sampling sites representative of the main habitats of a Mediterranean area in Central Italy, the Castelporziano Presidential Estate: (1) plant indicators derived from Ellenberg and Hemeroby models; (2) fauna indicators from soil microarthropods; and (3) humus forms and soil parameters from direct measurements. Principal component analysis and Spearman correlation test were performed on the data indicators/parameters matrix to evaluate the agreement of the multi-set in describing environmental quality. Results highlighted significant correlations among the different indicators/parameters. In particular, vascular plants and soil microarthropods represent two independent gradients in assessing ecosystem functioning, whilst humus forms provided similar information on habitat characteristics.

Long-term resilience of above- and below ground ecosystem components among contrasting ecosystems.

While several studies have explored how short-term ecological responses to disturbance vary among ecosystems, experimental studies of how contrasting ecosystems recover from disturbance in the longer term are few. We performed a simple long-term experiment on each of 30 contrasting forested islands in northern Sweden that vary in size; as size decreases, time since fire increases, soil fertility and ecosystem productivity declines, and plant species diversity increases. We predicted that resilience of understory plant community properties would be greatest on the larger, more productive islands, and that this would be paralleled by greater resilience of soil biotic and abiotic properties. For each island, we applied three disturbance treatments of increasing intensity to the forest understory once in 1998, i.e., light trimming, heavy trimming, and burning; a fourth treatment was an undisturbed control. We measured recovery of the understory vascular plant community annually over the following 14 years, and at that time also assessed recovery of mosses and several belowground variables. Consistent with our predictions, vascular plant whole-community variables (total cover, species richness, diversity [Shannon's H'], and community composition) recovered significantly more slowly on the smaller (least fertile) than the larger islands, but this difference was not substantial, and only noticeable in the most severely disturbed treatment. When an index of resilience was used, we were unable to detect effects of island size on the recovery of any property. We found that mosses and one shrub species (Empetrum hermaphroditum) recovered particularly slowly, and the higher abundance of this shrub on small islands was sufficient to explain any slower recovery of whole-ecosystem variables on those islands. Further, several belowground variables had not fully recovered from the most intense disturbance after 14 yr, and counter to our predictions, the degree of their recovery was never influenced by island size. While several studies have shown large variation among plant communities in their short-term response (notably resistance) to environmental perturbations, our results reveal that when perturbations are applied equally to highly contrasting ecosystems, differences in resilience among them in the longer term can be relatively minor, regardless of the severity of disturbance.

On ecology of post-fire soil fungi: assessing impact of disturbance using species-abundance models as measure of community organization

Mediterranean ecosystems are among those most significantly modified by fires. Such fires lead to evident disturbance of the above- and below-ground ecosystem components, at various spatial and temporal scales, including soil microfungi. The ecological parameters used to measure the effects of disturbance on soil fungal communities include species-abundance distribution patterns, which reflect changes in the relationships between species numbers and their relative abundance, and serve as a critical measure of community organization. Species-abundance distribution patterns were used to assess the disturbance impact of experimental fires on soil fungal communities in Mediterranean maquis (southern Italy) in the short- to mid-term. The trend in the distribution patterns of heat-stimulated and xerotolerant soil fungi over time, their varying responses to low- and high-intensity fire, the efficiency of the soil germplasm bank, and the pivotal role of Neosartorya spp. in post-fire community structure in Mediterranean burned soils may all be used as tools to accurately assess the effects of fire on soil mycobiota.

Seasonal exposure to drought and air warming affects soil Collembola and mites.

Global environmental changes affect not only the aboveground but also the belowground components of ecosystems. The effects of seasonal drought and air warming on the genus level richness of Collembola, and on the abundance and biomass of the community of Collembola and mites were studied in an acidic and a calcareous forest soil in a model oak-ecosystem experiment (the Querco experiment) at the Swiss Federal Research Institute WSL in Birmensdorf. The experiment included four climate treatments: control, drought with a 60% reduction in rainfall, air warming with a seasonal temperature increase of 1.4°C, and air warming + drought. Soil water content was greatly reduced by drought. Soil surface temperature was slightly increased by both the air warming and the drought treatment. Soil mesofauna samples were taken at the end of the first experimental year. Drought was found to increase the abundance of the microarthropod fauna, but reduce the biomass of the community. The percentage of small mites (body length 0.20 mm) increased, but the percentage of large mites (body length >0.40 mm) decreased under drought. Air warming had only minor effects on the fauna. All climate treatments significantly reduced the richness of Collembola and the biomass of Collembola and mites in acidic soil, but not in calcareous soil. Drought appeared to have a negative impact on soil microarthropod fauna, but the effects of climate change on soil fauna may vary with the soil type.

Global diversity and distribution of arbuscular mycorrhizal fungi

Arbuscular mycorrhizal (AM) fungi form associations with most land plants and can control carbon, nitrogen, and phosphorus cycling between above- and belowground components of ecosystems. Current estimates of AM fungal distributions are mainly inferred from the individual distributions of plant biomes, and climatic factors. However, dispersal limitation, local environmental conditions,and interactions among AM fungal taxa may also determine local diversity and global distributions. We assessed the relative importance of these potential controls by collecting 14,961 DNA sequences from 111 published studies and testing for relationships between AM fungal community composition and geography, environment, and plant biomes. Our results indicated that the global species richness of AM fungi was up to six times higher than previously estimated, largely owing to high beta diversity among sampling sites. Geographic distance, soil temperature and moisture, and plant community type were each significantly related to AM fungal community structure, but explained only a small amount of the observed variance. AM fungal species also tended to be phylogenetically clustered within sites, further suggesting that habitat filtering or dispersal limitation is a driver of AM fungal community assembly. Therefore, predicted shifts in climate and plant species distributions under global change may alter AM fungal communities.

Invasion by an exotic tree alters above and belowground ecosystem components

With the widespread introduction and invasion of exotic plants there is a need for studies that quantify alterations of basic ecosystem structure and function. Ecosystem invasion by Melaleuca quinquenervia significantly altered both above- and belowground ecosystem components in this study. We measured the quantity and nutrient concentration of the litterfall, litter layer, and soil; microbial biomass pools; and rates of potentially mineralizable nitrogen and soil oxygen demand. Annual litterfall was 4.9 times higher in the non-invaded sites and contained 1.9 times more phosphorus than invaded sites. Non-invaded plots contained a larger litter layer compared to invaded plots: 2.4 ± 1.2 kg m−2 and 0.62 ± 0.3 kg m−2, respectively. Lower nutrient concentration and quantity of the litter layer in the invaded plots led to changes in the aboveground storage of nutrients. In the invaded plots there was four times less carbon, seven times less nitrogen, and ten times less phosphorus stored in the organic litter layer compared to the non-invaded plots. Microbial biomass nutrient pools were consistently lower at both the 0–5 cm and 5–15 cm depth in the invaded soils compared to non-invaded soils, indicating a plant mediated change. Although M. quinquenervia altered microbial community structure, microbial activities were not different between invaded and non-invaded plots at either depth as measured by rates of soil oxygen demand and potentially mineralizable nitrogen. These changes may affect both native plant growth and water quality, and may act to promote and maintain site dominance by M. quinquenervia.

Linking the Aboveground and Belowground Components of Ecosystems

Linking the Aboveground and Belowground Components of Ecosystems

Carbon budgets of temperate forest ecosystems.

A summary of carbon-budget data from 30 forest stands is presented together with information on both above- and belowground ecosystem components. Only 10 of the stands had complete biomass data, 21 had complete productivity data and six had heterotrophic, or autotrophic respiration data, or both. The most comprehensive stand-level data have been collected for Pinus spp., Pseudotsuga menziesii, Abies amabilis and Liriodendron tulipifera. Only incomplete carbon budgets are available for most ecosystems studied, because measurements have been limited to one or several ecosystem processes (e.g., decay, productivity, storage). Based on the few complete C-budget studies that have been published, it is concluded that tree species differ in annual turnover of tissues.