Comparison of boreal ecosystem model sensitivity to variability in climate and forest site parameters

Ecosystem models are useful tools for evaluating environmental controls on carbon and water cycles under past or future conditions. In this paper we compare annual carbon and water fluxes from nine boreal spruce forest ecosystem models in a series of sensitivity simulations. For each comparison, a single climate driver or forest site parameter was altered in a separate sensitivity run. Driver and parameter changes were prescribed principally to be large enough to identify and isolate any major differences in model responses, while also remaining within the range of variability that the boreal forest biome may be exposed to over a time period of several decades. The models simulated plant production, autotrophic and heterotrophic respiration, and evapotranspiration (ET) for a black spruce site in the boreal forest of central Canada (56°N). Results revealed that there were common model responses in gross primary production, plant respiration, and ET fluxes to prescribed changes in air temperature or surface irradiance and to decreased precipitation amounts. The models were also similar in their responses to variations in canopy leaf area, leaf nitrogen content, and surface organic layer thickness. The models had different sensitivities to certain parameters, namely the net primary production response to increased CO2 levels, and the response of soil microbial respiration to precipitation inputs and soil wetness. These differences can be explained by the type (or absence) of photosynthesis-CO2 response curves in the models and by response algorithms of litter and humus decomposition to drying effects in organic soils of the boreal spruce ecosystem. Differences in the couplings of photosynthesis and soil respiration to nitrogen availability may also explain divergent model responses. Sensitivity comparisons imply that past conditions of the ecosystem represented in the models\u27 initial standing wood and soil carbon pools, including historical climate patterns and the time since the last major disturbance, can be as important as potential climatic changes to prediction of the annual ecosystem carbon balance in this boreal spruce forest

Functional indicators of response mechanisms to nitrogen deposition, ozone, and their interaction in two Mediterranean tree species

The effects of nitrogen (N) deposition, tropospheric ozone (O3) and their interaction were investigated in two Mediterranean tree species, Fraxinus ornus L. (deciduous) and Quercus ilex L. (evergreen), having different leaf habits and resource use strategies. An experiment was conducted under controlled condition to analyse how nitrogen deposition affects the ecophysiological and biochemical traits, and to explore how the nitrogen-induced changes influence the response to O3. For both factors we selected realistic exposures (20 kg N ha-1 yr-1 and 80 ppb h for nitrogen and O3, respectively), in order to elucidate the mechanisms implemented by the plants. Nitrogen addition resulted in higher nitrogen concentration at the leaf level in F. ornus, whereas a slight increase was detected in Q. ilex. Nitrogen enhanced the maximum rate of assimilation and ribulose 1,5-bisphosphate regeneration in both species, whereas it influenced the light harvesting complex only in the deciduous F. ornus that was also affected by O3 (reduced assimilation rate and accelerated senescence-related processes). Conversely, Q. ilex developed an avoidance mechanism to cope with O3, confirming a substantial O3 tolerance of this species. Nitrogen seemed to ameliorate the harmful effects of O3 in F. ornus: the hypothesized mechanism of action involved the production of nitrogen oxide as the first antioxidant barrier, followed by enzymatic antioxidant response. In Q. ilex, the interaction was not detected on gas exchange and photosystem functionality; however, in this species, nitrogen might stimulate an alternative antioxidant response such as the emission of volatile organic compounds. Antioxidant enzyme activity was lower in plants treated with both O3 and nitrogen even though reactive oxygen species production did not differ between the treatments

Mechanistic Modeling of Microtopographic Impacts on CO2 and CH4 Fluxes in an Alaskan Tundra Ecosystem Using the CLM-Microbe Model

Spatial heterogeneities in soil hydrology have been confirmed as a key control on CO2 and CH4 fluxes in the Arctic tundra ecosystem. In this study, we applied a mechanistic ecosystem model, CLM-Microbe, to examine the microtopographic impacts on CO2 and CH4 fluxes across seven landscape types in Utqiaġvik, Alaska: trough, low-centered polygon (LCP) center, LCP transition, LCP rim, high-centered polygon (HCP) center, HCP transition, and HCP rim. We first validated the CLM-Microbe model against static-chamber measured CO2 and CH4 fluxes in 2013 for three landscape types: trough, LCP center, and LCP rim. Model application showed that low-elevation and thus wetter landscape types (i.e., trough, transitions, and LCP center) had larger CH4 emissions rates with greater seasonal variations than high-elevation and drier landscape types (rims and HCP center). Sensitivity analysis indicated that substrate availability for methanogenesis (acetate, CO2 + H2) is the most important factor determining CH4 emission, and vegetation physiological properties largely affect the net ecosystem carbon exchange and ecosystem respiration in Arctic tundra ecosystems. Modeled CH4 emissions for different microtopographic features were upscaled to the eddy covariance (EC) domain with an area-weighted approach before validation against EC-measured CH4 fluxes. The model underestimated the EC-measured CH4 flux by 20% and 25% at daily and hourly time steps, suggesting the importance of the time step in reporting CH4 flux. The strong microtopographic impacts on CO2 and CH4 fluxes call for a model-data integration framework for better understanding and predicting carbon flux in the highly heterogeneous Arctic landscape

Climatic, edaphic, and biotic controls over storage and turnover of carbon in soils

Soil carbon, a major component of the global carbon inventory, has significant potential for change with changing climate and human land use. We applied the Century ecosystem model to a series of forest and grassland sites distributed globally to examine large-scale controls over soil carbon. Key site-specific parameters influencing soil carbon dynamics are soil texture and foliar lignin content; accordingly, we perturbed these variables at each site to establish a range of carbon concentrations and turnover times. We examined the simulated soil carbon stores, turnover times, and C:N ratios for correlations with patterns of independent variables. Results showed that soil carbon is related linearly to soil texture, increasing as clay content increases, that soil carbon stores and turnover time are related to mean annual temperature by negative exponential functions, and that heterotrophic respiration originates from recent detritus (∼50%), microbial turnover (∼30%), and soil organic matter (∼20%) with modest variations between forest and grassland ecosystems. The effect of changing temperature on soil organic carbon (SOC) estimated by Century is dSOC/dT= 183e−0.034T. Global extrapolation of this relationship leads to an estimated sensitivity of soil C storage to a temperature of −11.1 Pg° C−1, excluding extreme arid and organic soils. In Century, net primary production (NPP) and soil carbon are closely coupled through the N cycle, so that as temperatures increase, accelerated N release first results in fertilization responses, increasing C inputs. The Century-predicted effect of temperature on carbon storage is modified by as much as 100% by the N cycle feedback. Century-estimated soil C sensitivity (−11.1 Pg° C−1) is similar to losses predicted with a simple data-based calculation (−14.1 Pg° C−1). Inclusion of the N cycle is important for even first-order predictions of terrestrial carbon balance. If the NPP-SOC feedback is disrupted by land use or other disturbances, then SOC sensitivity can greatly exceed that estimated in our simulations. Century results further suggest that if climate change results in drying of organic soils (peats), soil carbon loss rates can be high

Forest Net Primary Production Resistance Across a Gradient of Moderate Disturbance

The global carbon (C) balance is vulnerable to disturbances that alter terrestrial C uptake and loss. Moderate disturbances that kill or defoliate only a subset of canopy trees such as insect defoliation, drought, and age-related senescence are increasing in extent and frequency; yet, little is known about the effect of moderate disturbance on forest production and the mechanisms sustaining or supporting the recovery of the C cycle across a range of moderate disturbance severities. We used a broad plot-scale gradient of upper canopy tree mortality within a large manipulation of forest disturbance to: 1) quantify how aboveground wood net primary production (ANPPw) responds to a range of moderate disturbance severities and; 2) identify the primary mechanisms supporting ANPPw resistance or resilience following moderate disturbance. We found that ANPPw was highly resistant to moderate disturbance, with production levels sustained following the senescence of 9 to \u3e 60 % of the upper canopy tree basal area. As upper canopy gap fraction increased with rising disturbance severity, greater light availability to the subcanopy enhanced leaf-level C uptake and the growth of this formerly light-limited canopy stratum, compensating for upper canopy production losses. As a result, whole-ecosystem production efficiency (ANPPw/LAI) increased at high levels of disturbance severity and leaf area loss. These findings provide a mechanistic explanation for sustained ANPPw across the disturbance gradient, in which the physiological and growth enhancement of undisturbed vegetation was proportional to the level of disturbance severity. Our results have important ecological and management implications, showing that moderate disturbances may minimally alter ecosystem functions such as C storage

Carbon fluxes in a mature deciduous forest under elevated CO₂

This PhD thesis addressed several major aspects of the carbon (C) cycle in a c. 100-year-old, mixed deciduous forest under elevated CO₂ with an emphasis on below-ground processes. The aim was to assess the responses of tree fine roots and soil respiration to canopy CO₂ enrichment (? 550 ppm) in this tallest forest studied to date. Furthermore, leaf gas-exchange of the five study species was examined to ascertain the long-term response of photosynthetic carbon uptake to elevated atmospheric CO₂. Investigations at the Swiss Canopy Crane (SCC) experimental site were guided by the following key questions: (1) Does below-ground C allocation to fine root production increase in response to CO₂ enrichment in order to acquire more nutrients to match the enhanced C supply in the forest canopy? (2) Is below-ground metabolism enhanced and therefore forest soil respiration stimulated by canopy CO₂ enrichment? (3) Is leaf-level photosynthesis persistently stimulated by elevated CO₂ in this stand or had these mature broad-leaved trees reduced their carbon up- take by photosynthetic down-regulation under long-term CO₂ enrichment? Findings from earlier studies at the SCC site, including 13C isotope tracing, all point towards an in- creased flux of C through CO₂-enriched trees to the soil but neither fine root biomass nor soil respiration were stimulated by elevated CO₂. Surprisingly, fine root biomass in bulk soil and ingrowth cores showed strong reductions by ? 30% in year five and six but were unaffected in the following seventh year of CO₂ enrichment. Given the absence of a positive biomass response of fine roots, we assumed that the extra C assimilated in the CO₂-enriched forest canopy was largely respired back to the atmosphere via increases in fine root and rhizosphere respiration and the metabolization of increased root derived exudates by soil microbes. Indeed, 52% higher soil air CO₂ concentration during the growing season and 14% greater soil microbial biomass both in- dicated enhanced below-ground metabolism in soil under CO₂-enriched trees. However, this did not translate into a persistent stimulation of soil respiration. At times of high or continuous precipitation soil water savings under CO₂-exposed trees (resulting from reduced sapflow) led to excessive soil moisture (> 45 vol.-%) impeding soil gas-exchange and thus soil respiration. Depending on the interplay between soil temperature and the consistently high soil water content in this stand, instantaneous rates of soil respiration were periodically reduced or increased under elevated CO₂ but on a diel scale and integrated over the growing season soil CO₂ emissions were similar under CO₂-enriched and control trees. Soil respiration could therefore not explain the fate of the extra C. The lacking sink capacity for additional assimilates led us to assume downward adjustment of photosynthetic capacity in CO₂-enriched trees thereby reducing carbon uptake in the forest canopy. Photosynthetic acclimation cannot completely eliminate the CO₂-driven stimulation in carbon uptake, but a reduction could hamper the detection of a CO₂ effect considering the low statistical power inevitably involved with such large-scale experiments. However, after eight years of CO₂ enrichment we found sustained stimulation in leaf photosynthesis (42-49%) indicating a lack of closure in the carbon budget for this stand under elevated atmospheric CO₂

Optogenetic manipulation of stomatal kinetics improves carbon assimilation, water use, and growth

Stomata serve dual and often conflicting roles, facilitating carbon dioxide influx into the plant leaf for photosynthesis and restricting water efflux via transpiration. Strategies for reducing transpiration without incurring a cost for photosynthesis must circumvent this inherent coupling of carbon dioxide and water vapor diffusion. We expressed the synthetic, light-gated K+ channel BLINK1 in guard cells surrounding stomatal pores in Arabidopsis to enhance the solute fluxes that drive stomatal aperture. BLINK1 introduced a K+ conductance and accelerated both stomatal opening under light exposure and closing after irradiation. Integrated over the growth period, BLINK1 drove a 2.2-fold increase in biomass in fluctuating light without cost in water use by the plant. Thus, we demonstrate the potential of enhancing stomatal kinetics to improve water use efficiency without penalty in carbon fixation

Carbon allocation and carbon isotope fluxes in the plant-soil-atmosphere continuum: a review

The terrestrial carbon (C) cycle has received increasing interest over the past few decades, however, there is still a lack of understanding of the fate of newly assimilated C allocated within plants and to the soil, stored within ecosystems and lost to the atmosphere. Stable carbon isotope studies can give novel insights into these issues. In this review we provide an overview of an emerging picture of plant-soil-atmosphere C fluxes, as based on C isotope studies, and identify processes determining related C isotope signatures. The first part of the review focuses on isotopic fractionation processes within plants during and after photosynthesis. The second major part elaborates on plant-internal and plant-rhizosphere C allocation patterns at different time scales (diel, seasonal, interannual), including the speed of C transfer and time lags in the coupling of assimilation and respiration, as well as the magnitude and controls of plant-soil C allocation and respiratory fluxes. Plant responses to changing environmental conditions, the functional relationship between the physiological and phenological status of plants and C transfer, and interactions between C, water and nutrient dynamics are discussed. The role of the C counterflow from the rhizosphere to the aboveground parts of the plants, e.g. via CO<sub>2</sub> dissolved in the xylem water or as xylem-transported sugars, is highlighted. The third part is centered around belowground C turnover, focusing especially on above- and belowground litter inputs, soil organic matter formation and turnover, production and loss of dissolved organic C, soil respiration and CO<sub>2</sub> fixation by soil microbes. Furthermore, plant controls on microbial communities and activity via exudates and litter production as well as microbial community effects on C mineralization are reviewed. A further part of the paper is dedicated to physical interactions between soil CO<sub>2</sub> and the soil matrix, such as CO<sub>2</sub> diffusion and dissolution processes within the soil profile. Finally, we highlight state-of-the-art stable isotope methodologies and their latest developments. From the presented evidence we conclude that there exists a tight coupling of physical, chemical and biological processes involved in C cycling and C isotope fluxes in the plant-soil-atmosphere system. Generally, research using information from C isotopes allows an integrated view of the different processes involved. However, complex interactions among the range of processes complicate or currently impede the interpretation of isotopic signals in CO<sub>2</sub> or organic compounds at the plant and ecosystem level. This review tries to identify present knowledge gaps in correctly interpreting carbon stable isotope signals in the plant-soil-atmosphere system and how future research approaches could contribute to closing these gaps