Disentangling the role of photosynthesis and stomatal conductance on rising forest water-use efficiency.

Multiple lines of evidence suggest that plant water-use efficiency (WUE)-the ratio of carbon assimilation to water loss-has increased in recent decades. Although rising atmospheric CO2 has been proposed as the principal cause, the underlying physiological mechanisms are still being debated, and implications for the global water cycle remain uncertain. Here, we addressed this gap using 30-y tree ring records of carbon and oxygen isotope measurements and basal area increment from 12 species in 8 North American mature temperate forests. Our goal was to separate the contributions of enhanced photosynthesis and reduced stomatal conductance to WUE trends and to assess consistency between multiple commonly used methods for estimating WUE. Our results show that tree ring-derived estimates of increases in WUE are consistent with estimates from atmospheric measurements and predictions based on an optimal balancing of carbon gains and water costs, but are lower than those based on ecosystem-scale flux observations. Although both physiological mechanisms contributed to rising WUE, enhanced photosynthesis was widespread, while reductions in stomatal conductance were modest and restricted to species that experienced moisture limitations. This finding challenges the hypothesis that rising WUE in forests is primarily the result of widespread, CO2-induced reductions in stomatal conductance.

Montane forest productivity across a semiarid climatic gradient.

High-elevation montane forests are disproportionately important to carbon sequestration in semiarid climates where low elevations are dry and characterized by low carbon density ecosystems. However, these ecosystems are increasingly threatened by climate change with seasonal implications for photosynthesis and forest growth. As a result, we leveraged eddy covariance data from six evergreen conifer forest sites in the semiarid western United States to extrapolate the status of carbon sequestration within a framework of projected warming and drying. At colder locations, the seasonal evolution of gross primary productivity (GPP) was characterized by a single broad maximum during the summer that corresponded to snow melt-derived moisture and a transition from winter dormancy to spring activity. Conversely, winter dormancy was transient at warmer locations, and GPP was responsive to both winter and summer precipitation such that two distinct GPP maxima were separated by a period of foresummer drought. This resulted in a predictable sequence of primary limiting factors to GPP beginning with air temperature in winter and proceeding to moisture and leaf area during the summer. Due to counteracting winter (positive) and summer (negative) GPP responses to warming, leaf area index and moisture availability were the best predictors of annual GPP differences across sites. Overall, mean annual GPP was greatest at the warmest site due to persistent vegetation photosynthetic activity throughout the winter. These results indicate that the trajectory of this region's carbon sequestration will be sensitive to reduced or delayed summer precipitation, especially if coupled to snow drought and earlier soil moisture recession, but summer precipitation changes remain highly uncertain. Given the demonstrated potential for seasonally offsetting responses to warming, we project that decadal semiarid montane forest carbon sequestration will remain relatively stable in the absence of severe disturbance.

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO2 sink. However, such analyses are poorly constrained by measured CO2 exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm-2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (Reco ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and Reco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO2 exchange may be up to 3-5 times larger than current estimates.

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Recovery of ponderosa pine ecosystem carbon and water fluxes from thinning and stand-replacing fire.

Carbon uptake by forests is a major sink in the global carbon cycle, helping buffer the rising concentration of CO2 in the atmosphere, yet the potential for future carbon uptake by forests is uncertain. Climate warming and drought can reduce forest carbon uptake by reducing photosynthesis, increasing respiration, and by increasing the frequency and intensity of wildfires, leading to large releases of stored carbon. Five years of eddy covariance measurements in a ponderosa pine (Pinus ponderosa)-dominated ecosystem in northern Arizona showed that an intense wildfire that converted forest into sparse grassland shifted site carbon balance from sink to source for at least 15 years after burning. In contrast, recovery of carbon sink strength after thinning, a management practice used to reduce the likelihood of intense wildfires, was rapid. Comparisons between an undisturbed-control site and an experimentally thinned site showed that thinning reduced carbon sink strength only for the first two posttreatment years. In the third and fourth posttreatment years, annual carbon sink strength of the thinned site was higher than the undisturbed site because thinning reduced aridity and drought limitation to carbon uptake. As a result, annual maximum gross primary production occurred when temperature was 3 °C higher at the thinned site compared with the undisturbed site. The severe fire consistently reduced annual evapotranspiration (range of 12-30%), whereas effects of thinning were smaller and transient, and could not be detected in the fourth year after thinning. Our results show large and persistent effects of intense fire and minor and short-lived effects of thinning on southwestern ponderosa pine ecosystem carbon and water exchanges.

Management Impacts on Carbon Dynamics in a Sierra Nevada Mixed Conifer Forest.

Forest ecosystems can act as sinks of carbon and thus mitigate anthropogenic carbon emissions. When forests are actively managed, treatments can alter forests carbon dynamics, reducing their sink strength and switching them from sinks to sources of carbon. These effects are generally characterized by fast temporal dynamics. Hence this study monitored for over a decade the impacts of management practices commonly used to reduce fire hazards on the carbon dynamics of mixed-conifer forests in the Sierra Nevada, California, USA. Soil CO2 efflux, carbon pools (i.e. soil carbon, litter, fine roots, tree biomass), and radial tree growth were compared among un-manipulated controls, prescribed fire, thinning, thinning followed by fire, and two clear-cut harvested sites. Soil CO2 efflux was reduced by both fire and harvesting (ca. 15%). Soil carbon content (upper 15 cm) was not significantly changed by harvest or fire treatments. Fine root biomass was reduced by clear-cut harvest (60-70%) but not by fire, and the litter layer was reduced 80% by clear-cut harvest and 40% by fire. Thinning effects on tree growth and biomass were concentrated in the first year after treatments, whereas fire effects persisted over the seven-year post-treatment period. Over this period, tree radial growth was increased (25%) by thinning and reduced (12%) by fire. After seven years, tree biomass returned to pre-treatment levels in both fire and thinning treatments; however, biomass and productivity decreased 30%-40% compared to controls when thinning was combined with fire. The clear-cut treatment had the strongest impact, reducing ecosystem carbon stocks and delaying the capacity for carbon uptake. We conclude that post-treatment carbon dynamics and ecosystem recovery time varied with intensity and type of treatments. Consequently, management practices can be selected to minimize ecosystem carbon losses while increasing future carbon uptake, resilience to high severity fire, and climate related stresses.

Soil Respiration in European Grasslands in Relation to Climate and Assimilate Supply.

Soil respiration constitutes the second largest flux of carbon (C) between terrestrial ecosystems and the atmosphere. This study provides a synthesis of soil respiration (R(s)) in 20 European grasslands across a climatic transect, including ten meadows, eight pastures and two unmanaged grasslands. Maximum rates of R(s) (R(s(max) )), R(s) at a reference soil temperature (10°C; R(s(10) )) and annual R(s) (estimated for 13 sites) ranged from 1.9 to 15.9 µmol CO(2) m(-2) s(-1), 0.3 to 5.5 µmol CO(2) m(-2) s(-1) and 58 to 1988 g C m(-2) y(-1), respectively. Values obtained for Central European mountain meadows are amongst the highest so far reported for any type of ecosystem. Across all sites R(s(max) ) was closely related to R(s(10) ).Assimilate supply affected R(s) at timescales from daily (but not necessarily diurnal) to annual. Reductions of assimilate supply by removal of aboveground biomass through grazing and cutting resulted in a rapid and a significant decrease of R(s). Temperature-independent seasonal fluctuations of R(s) of an intensively managed pasture were closely related to changes in leaf area index (LAI). Across sites R(s(10) ) increased with mean annual soil temperature (MAT), LAI and gross primary productivity (GPP), indicating that assimilate supply overrides potential acclimation to prevailing temperatures. Also annual R(s) was closely related to LAI and GPP. Because the latter two parameters were coupled to MAT, temperature was a suitable surrogate for deriving estimates of annual R(s) across the grasslands studied. These findings contribute to our understanding of regional patterns of soil C fluxes and highlight the importance of assimilate supply for soil CO(2) emissions at various timescales.