Spatiotemporal variability and origin of CO2 and CH4 tree stem fluxes in an upland forest.

The exchange of multiple greenhouse gases (i.e., CO2 and CH4 ) between tree stems and the atmosphere represents a knowledge gap in the global carbon cycle. Stem CO2 and CH4 fluxes vary across time and space and are unclear, which are their individual or shared drivers. Here we measured CO2 and CH4 fluxes at different stem heights combining manual (biweekly; n = 678) and automated (hourly; n > 38,000) measurements in a temperate upland forest. All trees showed CO2 and CH4 emissions despite 20% of measurements showing net CH4 uptake. Stem CO2 fluxes presented clear seasonal trends from manual and automated measurements. Only automated measurements captured the high temporal variability of stem CH4 fluxes revealing clear seasonal trends. Despite that temporal integration, the limited number of automated chambers made stand-level mean CH4 fluxes sensitive to "hot spots," resulting in mean fluxes with high uncertainty. Manual measurements provided better integration of spatial variability, but their lack of temporal variability integration hindered the detection of temporal trends and stand-level mean fluxes. These results highlight the potential bias of previous studies of stem CH4 fluxes solely based on manual or automated measurements. Stem height, temperature, and soil moisture only explained 7% and 11% of the stem CH4 flux variability compared to 42% and 81% for CO2 (manual and automated measurements, respectively). This large unexplained variability, in combination with high CH4 concentrations in the trees' heartwood, suggests that stem CH4 fluxes might be more influenced by gas transport and diffusivity through the wood than by drivers of respiratory CO2 flux, which has crucial implications for developing process-based ecosystem models. We postulate that CH4 is likely originated within tree stems because of lack of a consistent vertical pattern in CH4 fluxes, evidence of CH4 production in wood incubations, and low CH4 concentration in the soil profile but high concentrations within the trees' heartwood.

Experimental influence of storm-surge salinity on soil greenhouse gas emissions from a tidal salt marsh.

Storm surges can substantially alter the water level and salinity in tidal salt marshes. Little is known about how changes experienced during storm surges affect greenhouse gas emissions (GHG; CO2, CH4, N2O) from tidal salt marsh soils. Understanding how storm surges influence ecosystem processes is critical for evaluating the ecosystem's sensitivity to sea level rise. To explore how hurricane-induced changes in salinity affect GHG emissions, we exposed intact soil mesocosms (0-9 cm depth) from a Mid-Atlantic temperate salt marsh to pulse changes in salinity experienced at the site before, during, and after Hurricane Joaquin in 2015. Soil temperature, oxygen, and water level were kept constant to avoid confounding effects throughout the experiment. Automated measurements (hourly resolution) of soil GHG emissions were recorded in control (i.e., no salinity changes) and treatment mesocosms, and combined with soil pore water chemistry (i.e., SO42-, S2-, Fe2+, TNb, redox potential, pH) to characterize the biogeochemical responses. Using mixed effects models, we found that the role of different biogeochemical processes, such as sulfur cycling, changed throughout the experiment, underscoring the complex nature of GHG emissions in tidal salt marsh soils. Overall, soils subjected to a salinity decrease had greater GHG emissions than control soils, which were maintained at 17 ppt. The treatment soils had a 24% and 23% increase in global warming potential (20- and 100-year scenarios, respectively) indicating that storm surges can produce pulses of GHG emissions. However, both CH4 and N2O emissions returned to baseline values (following hysteresis responses) when initial conditions were reestablished. The results support the fact that tidal salt marshes are resilient ecosystems, as soil GHG emissions recovered relatively quickly from the pulse event.

Greenhouse Gas Fluxes From Tree Stems.

Tree stems exchange carbon dioxide, methane, and nitrous oxide with the atmosphere. The biophysical mechanisms controlling these fluxes are not fully understood, and consequently are not included in process-based models. We highlight advances and opportunities that will allow quantification of the role of these plant structures in the local-to-global balance of greenhouse gases (GHGs).

Automated measurements of greenhouse gases fluxes from tree stems and soils: magnitudes, patterns and drivers.

Tree stems exchange CO2, CH4 and N2O with the atmosphere but the magnitudes, patterns and drivers of these greenhouse gas (GHG) fluxes remain poorly understood. Our understanding mainly comes from static-manual measurements, which provide limited information on the temporal variability and magnitude of these fluxes. We measured hourly CO2, CH4 and N2O fluxes at two stem heights and adjacent soils within an upland temperate forest. We analyzed diurnal and seasonal variability of fluxes and biophysical drivers (i.e., temperature, soil moisture, sap flux). Tree stems were a net source of CO2 (3.80 ± 0.18 µmol m-2 s-1; mean ± 95% CI) and CH4 (0.37 ± 0.18 nmol m-2 s-1), but a sink for N2O (-0.016 ± 0.008 nmol m-2 s-1). Time series analysis showed diurnal temporal correlations between these gases with temperature or sap flux for certain days. CO2 and CH4 showed a clear seasonal pattern explained by temperature, soil water content and sap flux. Relationships between stem, soil fluxes and their drivers suggest that CH4 for stem emissions could be partially produced belowground. High-frequency measurements demonstrate that: a) tree stems exchange GHGs with the atmosphere at multiple time scales; and b) are needed to better estimate fluxes magnitudes and understand underlying mechanisms of GHG stem emissions.

Adjustments and coordination of hydraulic, leaf and stem traits along a water availability gradient.

Trait variability in space and time allows plants to adjust to changing environmental conditions. However, we know little about how this variability is distributed and coordinated at different organizational levels. For six dominant tree species in northeastern Spain (three Fagaceae and three Pinaceae) we quantified the inter- and intraspecific variability of a set of traits along a water availability gradient. We measured leaf mass per area (LMA), leaf nitrogen (N) concentration, carbon isotope composition in leaves (δ13 C), stem wood density, the Huber value (Hv, the ratio of cross-sectional sapwood area to leaf area), sapwood-specific and leaf-specific stem hydraulic conductivity, vulnerability to xylem embolism (P50 ) and the turgor loss point (Ptlp ). Differences between families explained the largest amount of variability for most traits, although intraspecific variability was also relevant. Species occupying wetter sites showed higher N, P50 and Ptlp , and lower LMA, δ13 C and Hv. However, when trait relationships with water availability were assessed within species they held only for Hv and Ptlp . Overall, our results indicate that intraspecific adjustments along the water availability gradient relied primarily on changes in resource allocation between sapwood and leaf area and in leaf water relations.

Methane emissions from tree stems: a new frontier in the global carbon cycle.

Tree stems from wetland, floodplain and upland forests can produce and emit methane (CH4 ). Tree CH4 stem emissions have high spatial and temporal variability, but there is no consensus on the biophysical mechanisms that drive stem CH4 production and emissions. Here, we summarize up to 30 opportunities and challenges for stem CH4 emissions research, which, when addressed, will improve estimates of the magnitudes, patterns and drivers of CH4 emissions and trace their potential origin. We identified the need: (1) for both long-term, high-frequency measurements of stem CH4 emissions to understand the fine-scale processes, alongside rapid large-scale measurements designed to understand the variability across individuals, species and ecosystems; (2) to identify microorganisms and biogeochemical pathways associated with CH4 production; and (3) to develop a mechanistic model including passive and active transport of CH4 from the soil-tree-atmosphere continuum. Addressing these challenges will help to constrain the magnitudes and patterns of CH4 emissions, and allow for the integration of pathways and mechanisms of CH4 production and emissions into process-based models. These advances will facilitate the upscaling of stem CH4 emissions to the ecosystem level and quantify the role of stem CH4 emissions for the local to global CH4 budget.

Microbes as Engines of Ecosystem Function: When Does Community Structure Enhance Predictions of Ecosystem Processes?

Microorganisms are vital in mediating the earth's biogeochemical cycles; yet, despite our rapidly increasing ability to explore complex environmental microbial communities, the relationship between microbial community structure and ecosystem processes remains poorly understood. Here, we address a fundamental and unanswered question in microbial ecology: 'When do we need to understand microbial community structure to accurately predict function?' We present a statistical analysis investigating the value of environmental data and microbial community structure independently and in combination for explaining rates of carbon and nitrogen cycling processes within 82 global datasets. Environmental variables were the strongest predictors of process rates but left 44% of variation unexplained on average, suggesting the potential for microbial data to increase model accuracy. Although only 29% of our datasets were significantly improved by adding information on microbial community structure, we observed improvement in models of processes mediated by narrow phylogenetic guilds via functional gene data, and conversely, improvement in models of facultative microbial processes via community diversity metrics. Our results also suggest that microbial diversity can strengthen predictions of respiration rates beyond microbial biomass parameters, as 53% of models were improved by incorporating both sets of predictors compared to 35% by microbial biomass alone. Our analysis represents the first comprehensive analysis of research examining links between microbial community structure and ecosystem function. Taken together, our results indicate that a greater understanding of microbial communities informed by ecological principles may enhance our ability to predict ecosystem process rates relative to assessments based on environmental variables and microbial physiology.

Strong resilience of soil respiration components to drought-induced die-off resulting in forest secondary succession.

How forests cope with drought-induced perturbations and how the dependence of soil respiration on environmental and biological drivers is affected in a warming and drying context are becoming key questions. The aims of this study were to determine whether drought-induced die-off and forest succession were reflected in soil respiration and its components and to determine the influence of climate on the soil respiration components. We used the mesh exclusion method to study seasonal variations in soil respiration (R S) and its components: heterotrophic (R H) and autotrophic (R A) [further split into fine root (R R) and mycorrhizal respiration (R M)] in a mixed Mediterranean forest where Scots pine (Pinus sylvestris L.) is undergoing a drought-induced die-off and is being replaced by holm oak (Quercus ilex L.). Drought-induced pine die-off was not reflected in R S nor in its components, which denotes a high functional resilience of the plant and soil system to pine die-off. However, the succession from Scots pine to holm oak resulted in a reduction of R H and thus in an important decrease of total respiration (R S was 36 % lower in holm oaks than in non-defoliated pines). Furthermore, R S and all its components were strongly regulated by soil water content-and-temperature interaction. Since Scots pine die-off and Quercus species colonization seems to be widely occurring at the driest limit of the Scots pine distribution, the functional resilience of the soil system over die-off and the decrease of R S from Scots pine to holm oak could have direct consequences for the C balance of these ecosystems.

Changes in soil bacterial community triggered by drought-induced gap succession preceded changes in soil C stocks and quality.

The aim of this study was to understand how drought-induced tree mortality and subsequent secondary succession would affect soil bacterial taxonomic composition as well as soil organic matter (SOM) quantity and quality in a mixed Mediterranean forest where the Scots pine (Pinus sylvestris) population, affected by climatic drought-induced die-off, is being replaced by Holm-oaks (HO; Quercus ilex). We apply a high throughput DNA pyrosequencing technique and (13)C solid-state Nuclear Magnetic Resonance (CP-MAS (13)C NMR) to soils within areas of influence (defined as an surface with 2-m radius around the trunk) of different trees: healthy and affected (defoliated) pines, pines that died a decade ago and healthy HOs. Soil respiration was also measured in the same spots during a spring campaign using a static close-chamber method (soda lime). A decade after death, and before aerial colonization by the more competitive HOs have even taken place, we could not find changes in soil C pools (quantity and/or quality) associated with tree mortality and secondary succession. Unlike C pools, bacterial diversity and community structure were strongly determined by tree mortality. Convergence between the most abundant taxa of soil bacterial communities under dead pines and colonizer trees (HOs) further suggests that physical gap colonization was occurring below-ground before above-ground colonization was taken place. Significantly higher soil respiration rates under dead trees, together with higher bacterial diversity and anomalously high representation of bacteria commonly associated with copiotrophic environments (r-strategic bacteria) further gives indications of how drought-induced tree mortality and secondary succession were influencing the structure of microbial communities and the metabolic activity of soils.