Comparing ecosystem gaseous elemental mercury fluxes over a deciduous and coniferous forest.

Sources of neurotoxic mercury in forests are dominated by atmospheric gaseous elemental mercury (GEM) deposition, but a dearth of direct GEM exchange measurements causes major uncertainties about processes that determine GEM sinks. Here we present three years of forest-level GEM deposition measurements in a coniferous forest and a deciduous forest in northeastern USA, along with flux partitioning into canopy and forest floor contributions. Annual GEM deposition is 13.4 ± 0.80 µg m-2 (coniferous forest) and 25.1 ± 2.4 µg m-2 (deciduous forest) dominating mercury inputs (62 and 76% of total deposition). GEM uptake dominates in daytime during active vegetation periods and correlates with CO2 assimilation, attributable to plant stomatal uptake of mercury. Non-stomatal GEM deposition occurs in the coniferous canopy during nights and to the forest floor in the deciduous forest and accounts for 24 and 39% of GEM deposition, respectively. Our study shows that GEM deposition includes various pathways and is highly ecosystem-specific, which complicates global constraints of terrestrial GEM sinks.

Informing Nature-based Climate Solutions for the United States with the best-available science.

Nature-based Climate Solutions (NbCS) are managed alterations to ecosystems designed to increase carbon sequestration or reduce greenhouse gas emissions. While they have growing public and private support, the realizable benefits and unintended consequences of NbCS are not well understood. At regional scales where policy decisions are often made, NbCS benefits are estimated from soil and tree survey data that can miss important carbon sources and sinks within an ecosystem, and do not reveal the biophysical impacts of NbCS for local water and energy cycles. The only direct observations of ecosystem-scale carbon fluxes, for example, by eddy covariance flux towers, have not yet been systematically assessed for what they can tell us about NbCS potentials, and state-of-the-art remote sensing products and land-surface models are not yet being widely used to inform NbCS policymaking or implementation. As a result, there is a critical mismatch between the point- and tree-scale data most often used to assess NbCS benefits and impacts, the ecosystem and landscape scales where NbCS projects are implemented, and the regional to continental scales most relevant to policymaking. Here, we propose a research agenda to confront these gaps using data and tools that have long been used to understand the mechanisms driving ecosystem carbon and energy cycling, but have not yet been widely applied to NbCS. We outline steps for creating robust NbCS assessments at both local to regional scales that are informed by ecosystem-scale observations, and which consider concurrent biophysical impacts, future climate feedbacks, and the need for equitable and inclusive NbCS implementation strategies. We contend that these research goals can largely be accomplished by shifting the scales at which pre-existing tools are applied and blended together, although we also highlight some opportunities for more radical shifts in approach.

Contrasting responses of woody and grassland ecosystems to increased CO2 as water supply varies.

Experiments show that elevated atmospheric CO2 (eCO2) often enhances plant photosynthesis and productivity, yet this effect varies substantially and may be climate sensitive. Understanding if, where and how water supply regulates CO2 enhancement is critical for projecting terrestrial responses to increasing atmospheric CO2 and climate change. Here, using data from 14 long-term ecosystem-scale CO2 experiments, we show that the eCO2 enhancement of annual aboveground net primary productivity is sensitive to annual precipitation and that this sensitivity differs between woody and grassland ecosystems. During wetter years, CO2 enhancement increases in woody ecosystems but declines in grass-dominated systems. Consistent with this difference, woody ecosystems can increase leaf area index in wetter years more effectively under eCO2 than can grassland ecosystems. Overall, and across different precipitation regimes, woody systems had markedly stronger CO2 enhancement (24%) than grasslands (13%). We developed an empirical relationship to quantify aboveground net primary productivity enhancement on the basis of changes in leaf area index, providing a new approach for evaluating eCO2 impacts on the productivity of terrestrial ecosystems.

Opportunities for forest sector emissions reductions: a state-level analysis.

The forest sector can play a significant role in climate change mitigation. We evaluated forest sector carbon trends and potential mitigation scenarios in Vermont using a systems-based modeling framework that accounts for net emissions from all forest sector components. These components comprise (1) the forest ecosystem, including land-use change, (2) harvested wood products (HWP), and (3) substitution effects associated with using renewable wood-based products and fuels in place of more emission-intensive materials and fossil fuel-based energy. We assessed baseline carbon trends from 1995 through 2050 using a business as usual (BAU) scenario. Emission reductions associated with different forest management and HWP scenarios were evaluated relative to the BAU scenario from 2020 to 2050. We estimated uncertainty for each forest sector component and used a Monte Carlo approach to estimate the distribution of cumulative total mitigation for each scenario relative to baseline. Our analysis indicates that the strength of the forest sector carbon sink in Vermont has been declining and will continue to decline over coming decades under the BAU scenario. However, several scenarios evaluated here could be effective in reducing emissions and enhancing carbon uptake. Shifting HWP to longer-lived commodities resulted in a 14% reduction in net cumulative emissions by 2050, the largest reduction of all scenarios. A scenario that combined extending harvest rotations, utilizing additional harvest residues for bioenergy, and increasing forest productivity resulted in a 12% reduction in net cumulative emissions. Shifting commodities from pulp and paper to bioenergy showed a 7.3% reduction in emissions. In contrast, shortening rotations to increase harvests for bioenergy use resulted in a 5.5% increase in emissions. In summary, model simulations suggest that net emissions could be reduced by up to 14% relative to BAU, depending on the management and HWP-use scenario. Combining multiple scenarios could further enhance reductions. However, realizing the full climate mitigation potential of these forests may be challenging due to socioeconomic barriers to implementation, as well as alternative management objectives that must be considered along with carbon sequestration.

Seasonal variation in the canopy color of temperate evergreen conifer forests.

Evergreen conifer forests are the most prevalent land cover type in North America. Seasonal changes in the color of evergreen forest canopies have been documented with near-surface remote sensing, but the physiological mechanisms underlying these changes, and the implications for photosynthetic uptake, have not been fully elucidated. Here, we integrate on-the-ground phenological observations, leaf-level physiological measurements, near surface hyperspectral remote sensing and digital camera imagery, tower-based CO2 flux measurements, and a predictive model to simulate seasonal canopy color dynamics. We show that seasonal changes in canopy color occur independently of new leaf production, but track changes in chlorophyll fluorescence, the photochemical reflectance index, and leaf pigmentation. We demonstrate that at winter-dormant sites, seasonal changes in canopy color can be used to predict the onset of canopy-level photosynthesis in spring, and its cessation in autumn. Finally, we parameterize a simple temperature-based model to predict the seasonal cycle of canopy greenness, and we show that the model successfully simulates interannual variation in the timing of changes in canopy color. These results provide mechanistic insight into the factors driving seasonal changes in evergreen canopy color and provide opportunities to monitor and model seasonal variation in photosynthetic activity using color-based vegetation indices.

Canopy photosynthetic capacity drives contrasting age dynamics of resource use efficiencies between mature temperate evergreen and deciduous forests.

Forest resource use efficiencies (RUEs) can vary with tree age, but the nature of these trends and their underlying mechanisms are not well understood. Understanding the age dynamics of forest RUEs and their drivers is vital for assessing the trade-offs between forest functions and resource consumption, making rational management policy, and projecting ecosystem carbon dynamics. Here we used the FLUXNET2015 and AmeriFlux datasets and published literature to explore the age-dependent variability of forest light use efficiency (LUE) and inherent water use efficiency as well as their main regulatory variables in temperate regions. Our results showed that evergreen forest RUEs initially increased before reaching the mature stage (i.e., around 90 years old), and then gradually declined; in contrast, RUEs continuously increased with age for mature deciduous forests. Changing canopy photosynthetic capacity (Amax) was the primary cause of age-related changes in RUEs across temperate forest sites. More importantly, soil nitrogen (N) increased in mature deciduous forests through time but decreased in older evergreen forests. The age-dependent changes in soil N were closely linked with the age dynamics of Amax for mature temperate forests. Additionally, soil nutrient conditions played a greater role in deciduous forest RUEs than evergreen forest RUEs. This study highlights the importance of stand age and forest type on temperate forest RUEs over the long term.

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.

Six years of ecosystem-atmosphere greenhouse gas fluxes measured in a sub-boreal forest.

Carbon dioxide (CO2), methane (CH4), and nitrous oxide (N2O) are the greenhouse gases largely responsible for anthropogenic climate change. Natural plant and microbial metabolic processes play a major role in the global atmospheric budget of each. We have been studying ecosystem-atmosphere trace gas exchange at a sub-boreal forest in the northeastern United States for over two decades. Historically our emphasis was on turbulent fluxes of CO2 and water vapor. In 2012 we embarked on an expanded campaign to also measure CH4 and N2O. Here we present continuous tower-based measurements of the ecosystem-atmosphere exchange of CO2 and CH4, recorded over the period 2012-2018 and reported at a 30-minute time step. Additionally, we describe a five-year (2012-2016) dataset of chamber-based measurements of soil fluxes of CO2, CH4, and N2O (2013-2016 only), conducted each year from May to November. These data can be used for process studies, for biogeochemical and land surface model validation and benchmarking, and for regional-to-global upscaling and budgeting analyses.

Solar-induced chlorophyll fluorescence exhibits a universal relationship with gross primary productivity across a wide variety of biomes.

In our recent study in Global Change Biology (Li et al., ), we examined the relationship between solar-induced chlorophyll fluorescence (SIF) measured from the Orbiting Carbon Observatory-2 (OCO-2) and gross primary productivity (GPP) derived from eddy covariance flux towers across the globe, and we discovered that there is a nearly universal relationship between SIF and GPP across a wide variety of biomes. This finding reveals the tremendous potential of SIF for accurately mapping terrestrial photosynthesis globally.

Solar-induced chlorophyll fluorescence is strongly correlated with terrestrial photosynthesis for a wide variety of biomes: First global analysis based on OCO-2 and flux tower observations.

Solar-induced chlorophyll fluorescence (SIF) has been increasingly used as a proxy for terrestrial gross primary productivity (GPP). Previous work mainly evaluated the relationship between satellite-observed SIF and gridded GPP products both based on coarse spatial resolutions. Finer resolution SIF (1.3 km × 2.25 km) measured from the Orbiting Carbon Observatory-2 (OCO-2) provides the first opportunity to examine the SIF-GPP relationship at the ecosystem scale using flux tower GPP data. However, it remains unclear how strong the relationship is for each biome and whether a robust, universal relationship exists across a variety of biomes. Here we conducted the first global analysis of the relationship between OCO-2 SIF and tower GPP for a total of 64 flux sites across the globe encompassing eight major biomes. OCO-2 SIF showed strong correlations with tower GPP at both midday and daily timescales, with the strongest relationship observed for daily SIF at the 757 nm (R2  = 0.72, p < 0.0001). Strong linear relationships between SIF and GPP were consistently found for all biomes (R2  = 0.57-0.79, p < 0.0001) except evergreen broadleaf forests (R2  = 0.16, p < 0.05) at the daily timescale. A higher slope was found for C4 grasslands and croplands than for C3 ecosystems. The generally consistent slope of the relationship among biomes suggests a nearly universal rather than biome-specific SIF-GPP relationship, and this finding is an important distinction and simplification compared to previous results. SIF was mainly driven by absorbed photosynthetically active radiation and was also influenced by environmental stresses (temperature and water stresses) that determine photosynthetic light use efficiency. OCO-2 SIF generally had a better performance for predicting GPP than satellite-derived vegetation indices and a light use efficiency model. The universal SIF-GPP relationship can potentially lead to more accurate GPP estimates regionally or globally. Our findings revealed the remarkable ability of finer resolution SIF observations from OCO-2 and other new or future missions (e.g., TROPOMI, FLEX) for estimating terrestrial photosynthesis across a wide variety of biomes and identified their potential and limitations for ecosystem functioning and carbon cycle studies.

Quantifying climate-growth relationships at the stand level in a mature mixed-species conifer forest.

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year-to-year variability in growth. Numerous dendrochronological (tree-ring) studies have identified climate factors that influence year-to-year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand-level (as opposed to species-level) growth. We argue that stand-level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed-species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand-level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year-to-year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand-level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species- and canopy-position level. Our climate models were better fit to stand-level biomass increment than to species-level or canopy-position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand-level growth varies less from to year-to-year than species-level or canopy-position growth responses. By assessing stand-level growth response to climate, we provide an alternative perspective on climate-growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.

A remotely sensed pigment index reveals photosynthetic phenology in evergreen conifers.

In evergreen conifers, where the foliage amount changes little with season, accurate detection of the underlying "photosynthetic phenology" from satellite remote sensing has been difficult, presenting challenges for global models of ecosystem carbon uptake. Here, we report a close correspondence between seasonally changing foliar pigment levels, expressed as chlorophyll/carotenoid ratios, and evergreen photosynthetic activity, leading to a "chlorophyll/carotenoid index" (CCI) that tracks evergreen photosynthesis at multiple spatial scales. When calculated from NASA's Moderate Resolution Imaging Spectroradiometer satellite sensor, the CCI closely follows the seasonal patterns of daily gross primary productivity of evergreen conifer stands measured by eddy covariance. This discovery provides a way of monitoring evergreen photosynthetic activity from optical remote sensing, and indicates an important regulatory role for carotenoid pigments in evergreen photosynthesis. Improved methods of monitoring photosynthesis from space can improve our understanding of the global carbon budget in a warming world of changing vegetation phenology.

Greenness indices from digital cameras predict the timing and seasonal dynamics of canopy-scale photosynthesis.

The proliferation of digital cameras co-located with eddy covariance instrumentation provides new opportunities to better understand the relationship between canopy phenology and the seasonality of canopy photosynthesis. In this paper we analyze the abilities and limitations of canopy color metrics measured by digital repeat photography to track seasonal canopy development and photosynthesis, determine phenological transition dates, and estimate intra-annual and interannual variability in canopy photosynthesis. We used 59 site-years of camera imagery and net ecosystem exchange measurements from 17 towers spanning three plant functional types (deciduous broadleaf forest, evergreen needleleaf forest, and grassland/crops) to derive color indices and estimate gross primary productivity (GPP). GPP was strongly correlated with greenness derived from camera imagery in all three plant functional types. Specifically, the beginning of the photosynthetic period in deciduous broadleaf forest and grassland/crops and the end of the photosynthetic period in grassland/crops were both correlated with changes in greenness; changes in redness were correlated with the end of the photosynthetic period in deciduous broadleaf forest. However, it was not possible to accurately identify the beginning or ending of the photosynthetic period using camera greenness in evergreen needleleaf forest. At deciduous broadleaf sites, anomalies in integrated greenness and total GPP were significantly correlated up to 60 days after the mean onset date for the start of spring. More generally, results from this work demonstrate that digital repeat photography can be used to quantify both the duration of the photosynthetically active period as well as total GPP in deciduous broadleaf forest and grassland/crops, but that new and different approaches are required before comparable results can be achieved in evergreen needleleaf forest.

Increase in forest water-use efficiency as atmospheric carbon dioxide concentrations rise.

Terrestrial plants remove CO2 from the atmosphere through photosynthesis, a process that is accompanied by the loss of water vapour from leaves. The ratio of water loss to carbon gain, or water-use efficiency, is a key characteristic of ecosystem function that is central to the global cycles of water, energy and carbon. Here we analyse direct, long-term measurements of whole-ecosystem carbon and water exchange. We find a substantial increase in water-use efficiency in temperate and boreal forests of the Northern Hemisphere over the past two decades. We systematically assess various competing hypotheses to explain this trend, and find that the observed increase is most consistent with a strong CO2 fertilization effect. The results suggest a partial closure of stomata-small pores on the leaf surface that regulate gas exchange-to maintain a near-constant concentration of CO2 inside the leaf even under continually increasing atmospheric CO2 levels. The observed increase in forest water-use efficiency is larger than that predicted by existing theory and 13 terrestrial biosphere models. The increase is associated with trends of increasing ecosystem-level photosynthesis and net carbon uptake, and decreasing evapotranspiration. Our findings suggest a shift in the carbon- and water-based economics of terrestrial vegetation, which may require a reassessment of the role of stomatal control in regulating interactions between forests and climate change, and a re-evaluation of coupled vegetation-climate models.

Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees.

Nonstructural carbohydrate reserves support tree metabolism and growth when current photosynthates are insufficient, offering resilience in times of stress. We monitored stemwood nonstructural carbohydrate (starch and sugars) concentrations of the dominant tree species at three sites in the northeastern United States. We estimated the mean age of the starch and sugars in a subset of trees using the radiocarbon ((14) C) bomb spike. With these data, we then tested different carbon (C) allocation schemes in a process-based model of forest C cycling. We found that the nonstructural carbohydrates are both highly dynamic and about a decade old. Seasonal dynamics in starch (two to four times higher in the growing season, lower in the dormant season) mirrored those of sugars. Radiocarbon-based estimates indicated that the mean age of the starch and sugars in red maple (Acer rubrum) was 7-14 yr. A two-pool (fast and slow cycling reserves) model structure gave reasonable estimates of the size and mean residence time of the total NSC pool, and greatly improved model predictions of interannual variability in woody biomass increment, compared with zero- or one-pool structures used in the majority of existing models. This highlights the importance of nonstructural carbohydrates in the context of forest ecosystem carbon cycling.

Thermal optimality of net ecosystem exchange of carbon dioxide and underlying mechanisms.

• It is well established that individual organisms can acclimate and adapt to temperature to optimize their functioning. However, thermal optimization of ecosystems, as an assemblage of organisms, has not been examined at broad spatial and temporal scales. • Here, we compiled data from 169 globally distributed sites of eddy covariance and quantified the temperature response functions of net ecosystem exchange (NEE), an ecosystem-level property, to determine whether NEE shows thermal optimality and to explore the underlying mechanisms. • We found that the temperature response of NEE followed a peak curve, with the optimum temperature (corresponding to the maximum magnitude of NEE) being positively correlated with annual mean temperature over years and across sites. Shifts of the optimum temperature of NEE were mostly a result of temperature acclimation of gross primary productivity (upward shift of optimum temperature) rather than changes in the temperature sensitivity of ecosystem respiration. • Ecosystem-level thermal optimality is a newly revealed ecosystem property, presumably reflecting associated evolutionary adaptation of organisms within ecosystems, and has the potential to significantly regulate ecosystem-climate change feedbacks. The thermal optimality of NEE has implications for understanding fundamental properties of ecosystems in changing environments and benchmarking global models.

Variation in foliar nitrogen and albedo in response to nitrogen fertilization and elevated CO2.

Foliar nitrogen has been shown to be positively correlated with midsummer canopy albedo and canopy near infrared (NIR) reflectance over a broad range of plant functional types (e.g., forests, grasslands, and agricultural lands). To date, the mechanism(s) driving the nitrogen­albedo relationship have not been established, and it is unknown whether factors affecting nitrogen availability will also influence albedo. To address these questions, we examined variation in foliar nitrogen in relation to leaf spectral properties, leaf mass per unit area, and leaf water content for three deciduous species subjected to either nitrogen (Harvard Forest, MA, and Oak Ridge, TN) or CO(2) fertilization (Oak Ridge, TN). At Oak Ridge, we also obtained canopy reflectance data from the airborne visible/infrared imaging spectrometer (AVIRIS) to examine whether canopy-level spectral responses were consistent with leaf-level results. At the leaf level, results showed no differences in reflectance or transmittance between CO(2) or nitrogen treatments, despite significant changes in foliar nitrogen. Contrary to our expectations, there was a significant, but negative, relationship between foliar nitrogen and leaf albedo, a relationship that held for both full spectrum leaf albedo as well as leaf albedo in the NIR region alone. In contrast, remote sensing data indicated an increase in canopy NIR reflectance with nitrogen fertilization. Collectively, these results suggest that altered nitrogen availability can affect canopy albedo, albeit by mechanisms that involve canopy-level processes rather than changes in leaf-level reflectance.