RESUMO
Hydraulic stress in plants occurs under conditions of low water availability (soil moisture; θ) and/or high atmospheric demand for water (vapor pressure deficit; D). Different species are adapted to respond to hydraulic stress by functioning along a continuum where, on one hand, they close stomata to maintain a constant leaf water potential (ΨL) (isohydric species), and on the other hand, they allow ΨL to decline (anisohydric species). Differences in water-use along this continuum are most notable during hydrologic stress, often characterized by low θ and high D; however, θ and D are often, but not necessarily, coupled at time scales of weeks or longer, and uncertainty remains about the sensitivity of different water-use strategies to these variables. We quantified the effects of both θ and D on canopy conductance (Gc) among widely distributed canopy-dominant species along the isohydric-anisohydric spectrum growing along a hydroclimatological gradient. Tree-level Gc was estimated using hourly sap flow observations from three sites in the eastern United States: a mesic forest in western North Carolina and two xeric forests in southern Indiana and Missouri. Each site experienced at least 1 year of substantial drought conditions. Our results suggest that sensitivity of Gc to θ varies across sites and species, with Gc sensitivity being greater in dry than in wet sites, and greater for isohydric compared with anisohydric species. However, once θ limitations are accounted for, sensitivity of Gc to D remains relatively constant across sites and species. While D limitations to Gc were similar across sites and species, ranging from 16 to 34% reductions, θ limitations to Gc ranged from 0 to 40%. The similarity in species sensitivity to D is encouraging from a modeling perspective, though it implies that substantial reduction to Gc will be experienced by all species in a future characterized by higher D.
Assuntos
Solo , Árvores , Secas , Florestas , Folhas de Planta , Transpiração Vegetal , ÁguaRESUMO
Land-use/cover change (LUCC) is an important driver of environmental change, occurring at the same time as, and often interacting with, global climate change. Reforestation and deforestation have been critical aspects of LUCC over the past two centuries and are widely studied for their potential to perturb the global carbon cycle. More recently, there has been keen interest in understanding the extent to which reforestation affects terrestrial energy cycling and thus surface temperature directly by altering surface physical properties (e.g., albedo and emissivity) and land-atmosphere energy exchange. The impacts of reforestation on land surface temperature and their mechanisms are relatively well understood in tropical and boreal climates, but the effects of reforestation on warming and/or cooling in temperate zones are less certain. This study is designed to elucidate the biophysical mechanisms that link land cover and surface temperature in temperate ecosystems. To achieve this goal, we used data from six paired eddy-covariance towers over co-located forests and grasslands in the temperate eastern United States, where radiation components, latent and sensible heat fluxes, and meteorological conditions were measured. The results show that, at the annual time scale, the surface of the forests is 1-2°C cooler than grasslands, indicating a substantial cooling effect of reforestation. The enhanced latent and sensible heat fluxes of forests have an average cooling effect of -2.5°C, which offsets the net warming effect (+1.5°C) of albedo warming (+2.3°C) and emissivity cooling effect (-0.8°C) associated with surface properties. Additional daytime cooling over forests is driven by local feedbacks to incoming radiation. We further show that the forest cooling effect is most pronounced when land surface temperature is higher, often exceeding -5°C. Our results contribute important observational evidence that reforestation in the temperate zone offers opportunities for local climate mitigation and adaptation.
Assuntos
Ecossistema , Florestas , Atmosfera , Mudança Climática , TemperaturaRESUMO
We evaluated the effect on soil CO2 efflux (FCO2 ) of sudden changes in photosynthetic rates by altering CO2 concentration in plots subjected to +200 ppmv for 15 years. 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 4 months after termination of the long-term eCO2 treatment, longer than the 10 days observed for decrease of FCO2 after experimental blocking of C flow to belowground, but shorter than the ~13 months 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 .
Assuntos
Dióxido de Carbono , Ecossistema , Solo/química , Fotossíntese , Folhas de PlantaRESUMO
In mountainous areas, cold air drainage from high to low elevations has pronounced effects on local temperature, which is a critical driver of many ecosystem processes, including carbon uptake and storage. Here, we leverage new approaches for interpreting ecosystem carbon flux observations in complex terrain to quantify the links between macro-climate condition, drainage flows, local microclimate, and ecosystem carbon cycling in a southern Appalachian valley. Data from multiple long-running climate stations and multiple eddy covariance flux towers are combined with simple models for ecosystem carbon fluxes. We show that cold air drainage into the valley suppresses local temperature by several degrees at night and for several hours before and after sunset, leading to reductions in growing season respiration on the order of ~8%. As a result, we estimate that drainage flows increase growing season and annual net carbon uptake in the valley by >10% and >15%, respectively, via effects on microclimate that are not be adequately represented in regional- and global-scale terrestrial ecosystem models. Analyses driven by chamber-based estimates of soil and plant respiration reveal cold air drainage effects on ecosystem respiration are dominated by reductions to the respiration of aboveground biomass. We further show that cold air drainage proceeds more readily when cloud cover and humidity are low, resulting in the greatest enhancements to net carbon uptake in the valley under clear, cloud-free (i.e., drought-like) conditions. This is a counterintuitive result that is neither observed nor predicted outside of the valley, where nocturnal temperature and respiration increase during dry periods. This result should motivate efforts to explore how topographic flows may buffer eco-physiological processes from macroscale climate change.
Assuntos
Ciclo do Carbono , Temperatura Baixa , Ecossistema , Microclima , Região dos Apalaches , Dióxido de Carbono , ÁrvoresRESUMO
Uncertainties in ecophysiological responses to environment, such as the impact of atmospheric and soil moisture conditions on plant water regulation, limit our ability to estimate key inputs for ecosystem models. Advanced statistical frameworks provide coherent methodologies for relating observed data, such as stem sap flux density, to unobserved processes, such as canopy conductance and transpiration. To address this need, we developed a hierarchical Bayesian State-Space Canopy Conductance (StaCC) model linking canopy conductance and transpiration to tree sap flux density from a 4-year experiment in the North Carolina Piedmont, USA. Our model builds on existing ecophysiological knowledge, but explicitly incorporates uncertainty in canopy conductance, internal tree hydraulics and observation error to improve estimation of canopy conductance responses to atmospheric drought (i.e., vapor pressure deficit), soil drought (i.e., soil moisture) and above canopy light. Our statistical framework not only predicted sap flux observations well, but it also allowed us to simultaneously gap-fill missing data as we made inference on canopy processes, marking a substantial advance over traditional methods. The predicted and observed sap flux data were highly correlated (mean sensor-level Pearson correlation coefficient = 0.88). Variations in canopy conductance and transpiration associated with environmental variation across days to years were many times greater than the variation associated with model uncertainties. Because some variables, such as vapor pressure deficit and soil moisture, were correlated at the scale of days to weeks, canopy conductance responses to individual environmental variables were difficult to interpret in isolation. Still, our results highlight the importance of accounting for uncertainty in models of ecophysiological and ecosystem function where the process of interest, canopy conductance in this case, is not observed directly. The StaCC modeling framework provides a statistically coherent approach to estimating canopy conductance and transpiration and propagating estimation uncertainty into ecosystem models, paving the way for improved prediction of water and carbon uptake responses to environmental change.
Assuntos
Ecossistema , Modelos Teóricos , Transpiração Vegetal/fisiologia , Carbono , Meio Ambiente , ÁguaRESUMO
The southeastern United States is experiencing a rapid regional increase in the ratio of pine to deciduous forest ecosystems at the same time it is experiencing changes in climate. This study is focused on exploring how these shifts will affect the carbon sink capacity of southeastern US forests, which we show here are among the strongest carbon sinks in the continental United States. Using eight-year-long eddy covariance records collected above a hardwood deciduous forest (HW) and a pine plantation (PP) co-located in North Carolina, USA, we show that the net ecosystem exchange of CO2 (NEE) was more variable in PP, contributing to variability in the difference in NEE between the two sites (ΔNEE) at a range of timescales, including the interannual timescale. Because the variability in evapotranspiration (ET) was nearly identical across the two sites over a range of timescales, the factors that determined the variability in ΔNEE were dominated by those that tend to decouple NEE from ET. One such factor was water use efficiency, which changed dramatically in response to drought and also tended to increase monotonically in nondrought years (P < 0.001 in PP). Factors that vary over seasonal timescales were strong determinants of the NEE in the HW site; however, seasonality was less important in the PP site, where significant amounts of carbon were assimilated outside of the active season, representing an important advantage of evergreen trees in warm, temperate climates. Additional variability in the fluxes at long-time scales may be attributable to slowly evolving factors, including canopy structure and increases in dormant season air temperature. Taken together, study results suggest that the carbon sink in the southeastern United States may become more variable in the future, owing to a predicted increase in drought frequency and an increase in the fractional cover of southern pines.
Assuntos
Dióxido de Carbono/metabolismo , Sequestro de Carbono , Mudança Climática , Florestas , Árvores/fisiologia , Biodiversidade , Secas , North Carolina , Pinus/fisiologia , Transpiração VegetalRESUMO
Soil CO2 efflux (Fsoil ) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO2] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long-term effects of these factors on Fsoil are less clear. Expanding on previous studies at the Duke Free-Air CO2 Enrichment (FACE) site, we quantified the effects of elevated [CO2] and N fertilization on Fsoil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual Fsoil increased under elevated [CO2] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO2] reduced Fsoil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment-induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual Fsoil from potential values - estimated based on temperature alone assuming nonlimiting soil water content - by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO2]. Variability in soil N availability among plots accounted for the spatial variability in Fsoil , showing a decrease of ca. 114 g C m(-2) yr(-1) per 1 g m(-2) increase in soil N availability, with consistently higher Fsoil in elevated [CO2] plots ca. 127 g C per 100 ppm [CO2] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO2] and N fertilization on Fsoil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.
Assuntos
Dióxido de Carbono , Nitrogênio , Solo , Árvores , Dióxido de Carbono/farmacologia , North Carolina , Pinus taeda , Folhas de Planta , Solo/química , Temperatura , ÁguaRESUMO
A number of recent studies have attributed a large proportion of soil respiration (R(soil)) to recently photoassimilated carbon (C). Time lags (tau(PR)) associated with these pulses of photosynthesis and responses of R(soil) have been found on time scales of hours to weeks for different ecosystems, but most studies find evidence for tau(PR) on the order of 1-5 d. We showed that such time scales are commensurate with CO(2) diffusion time scales from the roots to the soil surface, and may thus be independent from photosynthetic pulses. To further quantify the role of physical (i.e. edaphic) and biological (i.e. vegetative) controls on such lags, we investigated tau(PR) at adjacent planted pine (PP) and hardwood (HW) forest ecosystems over six and four measurement years, respectively, using both autocorrelation analysis on automated soil surface flux measurements and their lagged cross-correlations with drivers for and surrogates of photosynthesis. Evidence for tau(PR) on the order of 1-3 d was identified in both ecosystems and using both analyses, but this lag could not be attributed to recently photoassimilated C because the same analysis yielded comparable lags at HW during leaf-off periods. Future efforts to model ecosystem C inputs and outputs in a pulse-response framework must combine measurements of transport in the physical and biological components of terrestrial ecosystems.
Assuntos
Carbono/metabolismo , Ecossistema , Árvores/metabolismo , Fotossíntese , Estações do Ano , Sudeste dos Estados Unidos , Árvores/fisiologiaRESUMO
Orthonormal wavelet transformation (OWT) is a computationally efficient technique for quantifying underlying frequencies in nonstationary and gap-infested time series, such as eddy-covariance-measured net ecosystem exchange of CO2 (NEE). We employed OWT to analyze the frequency characteristics of synchronously measured and modeled NEE at adjacent pine (PP) and hardwood (HW) ecosystems. Wavelet cospectral analysis showed that NEE at PP was more correlated to light and vapor pressure deficit at the daily time scale, and NEE at HW was more correlated to leaf area index (LAI) and temperature, especially soil temperature, at seasonal time scales. Models were required to disentangle the impacts of environmental drivers on the components of NEE, ecosystem carbon assimilation (Ac) and ecosystem respiration (RE). Sensitivity analyses revealed that using air temperature rather than soil temperature in RE models improved the modeled wavelet spectral frequency response on time scales longer than 1 day at both ecosystems. Including LAI improved RE model fit on seasonal time scales at HW, and incorporating parameter variability improved the RE model response at annual time scales at both ecosystems. Resolving variability in canopy conductance, rather than leaf-internal CO2, was more important for modeling Ac at both ecosystems. The PP ecosystem was more sensitive to hydrologic variables that regulate canopy conductance: vapor pressure deficit on weekly time scales and soil moisture on seasonal to interannual time scales. The HW ecosystem was sensitive to water limitation on weekly time scales. A combination of intrinsic drought sensitivity and non-conservative water use at PP was the basis for this response. At both ecosystems, incorporating variability in LAI was required for an accurate spectral representation of modeled NEE. However, nonlinearities imposed by canopy light attenuation were of little importance to spectral fit. The OWT revealed similarities and differences in the scale-wise control of NEE by vegetation with implications for model simplification and improvement.
Assuntos
Dióxido de Carbono/metabolismo , Ecossistema , Modelos Biológicos , Árvores/metabolismo , Pinus taeda/metabolismo , Folhas de Planta/anatomia & histologia , Folhas de Planta/metabolismo , Solo , Análise Espectral , Temperatura , Água/metabolismoRESUMO
Temporal variability in the (13)C of foliage (delta(13)C(F)), soil (delta(13)C(S)) and ecosystem (delta(13)C(R)) respired CO(2) was contrasted between a 17.2-m tall evenly aged loblolly pine forest and a 35-m tall unevenly aged mature second growth mixed broadleaf deciduous forest in North Carolina, USA, over a 2-year period. The two forests are located at the Duke Forest within a kilometer of each other and are subject to identical climate and have similar soil types. The delta(13)C(F), collected just prior to dawn, was primarily controlled by the time-lagged vapor pressure deficit (VPD) in both stands; it was used for calculating the ratio of intercellular to ambient CO(2) ( Ci/ Ca). A remarkable similarity was observed in the relationship between Ci/ Ca and time-lagged VPD in these two forests despite large differences in hydraulic characteristics. This similarity emerged as a result of physiological adjustments that compensated for differences in plant hydraulic characteristics, as predicted by a recently proposed equilibrium hypothesis, and has implications to ecophysiological models. We found that in the broadleaf forest, the delta(13)C of forest floor CO(2) efflux dominated the delta(13)C(R), while in the younger pine forest, the delta(13)C of foliage respired CO(2) dominated delta(13)C(R). This dependence resulted in a more variable delta(13)C(R) in the pine forest when compared to the broadleaf forest due to the larger photosynthetic contribution. Given the sensitivity of the atmospheric inversion models to delta(13)C(R), the results demonstrate that these models could be improved by accounting for stand characteristics, in addition to previously recognized effects of moisture availability, when estimating delta(13)C(R).