Many photosynthetic organisms globally, including crops, forests and algae, must grow in environments where the availability of light energy fluctuates dramatically. How photosynthesis maintains high efficiency despite such fluctuations in its energy source remains poorly understood. Here we show that Arabidopsis thaliana K+ efflux antiporter (KEA3) is critical for high photosynthetic efficiency under fluctuating light. On a shift from dark to low light, or high to low light, kea3 mutants show prolonged dissipation of absorbed light energy as heat. KEA3 localizes to the thylakoid membrane, and allows proton efflux from the thylakoid lumen by proton/potassium antiport. KEA3's activity accelerates the downregulation of pH-dependent energy dissipation after transitions to low light, leading to faster recovery of high photosystem II quantum efficiency and increased CO2 assimilation. Our results reveal a mechanism that increases the efficiency of photosynthesis under fluctuating light.

Earth's atmospheric CO2 concentration (c(a)) for the Phanerozoic Eon is estimated from proxies and geochemical carbon cycle models. Most estimates come with large, sometimes unbounded uncertainty. Here, we calculate tightly constrained estimates of c(a) using a universal equation for leaf gas exchange, with key variables obtained directly from the carbon isotope composition and stomatal anatomy of fossil leaves. Our new estimates, validated against ice cores and direct measurements of c(a), are less than 1000 ppm for most of the Phanerozoic, from the Devonian to the present, coincident with the appearance and global proliferation of forests. Uncertainties, obtained from Monte Carlo simulations, are typically less than for c(a) estimates from other approaches. These results provide critical new empirical support for the emerging view that large (similar to 2000-3000 ppm), long-term swings in c(a) do not characterize the post-Devonian and that Earth's long-term climate sensitivity to c(a) is greater than originally thought.

We have extended a conventional photosynthesis model to simulate field and laboratory measurements of chlorophyll fluorescence at the leaf scale. The fluorescence paramaterization is based on a close nonlinear relationship between the relative light saturation of photosynthesis and nonradiative energy dissipation in plants of different species. This relationship diverged only among examined data sets under stressed (strongly light saturated) conditions, possibly caused by differences in xanthophyll pigment concentrations. The relationship was quantified after analyzing data sets of pulse amplitude modulated measurements of chlorophyll fluorescence and gas exchange of leaves of different species exposed to different levels of light, CO2, temperature, nitrogen fertilization treatments, and drought. We used this relationship in a photosynthesis model. The coupled model enabled us to quantify the relationships between steady state chlorophyll fluorescence yield, electron transport rate, and photosynthesis in leaves under different environmental conditions.

Carbonyl sulfide (COS) has recently emerged as an atmospheric tracer of gross primary production. All modeling studies of COS air-monitoring data rely on a climatological anthropogenic inventory that does not reflect present conditions or support interpretation of ice core and firn trends. Here we develop a global anthropogenic inventory for the years 1850 to 2013 based on new emission measurements and material-specific data. By applying methods from a recent regional inventory to global data, we find that the anthropogenic source is similar in magnitude to the plant sink, confounding carbon cycle applications. However, a material-specific approach results in a current anthropogenic source that is only one third of plant uptake and is concentrated in Asia, supporting carbon cycle applications of global air-monitoring data. Furthermore, changes in the anthropogenic source alone cannot explain the century-scale mixing ratio growth, which suggests that ice and firn data may provide the first global history of gross primary production.

Regional gross primary productivity (GPP) estimates are crucial to estimating carbon-climate feedbacks but are highly uncertain with existing methods. An emerging approach uses atmospheric carbonyl sulphide (COS) as a tracer for carbon dioxide: COS plant uptake is simulated by scaling GPP. A critical parameter for this method is leaf-scale relative uptake (LRU). Plant chamber and eddy covariance studies find a narrow range of LRU values but some atmospheric modelling studies assign values well outside this range. Here we study this discrepancy by conducting new regional chemical transport simulations for North America using the underlying data from previous studies. We find the wide range of ecosystem model GPP estimates can explain the discrepancy in LRU values. We also find that COS concentration uncertainty is more sensitive to GPP uncertainty than to LRU parameter uncertainty. These results support the COS tracer technique as a useful approach for constraining GPP estimates.

The response of tropical forests to droughts is highly uncertain(1). During the dry season, canopy photosynthesis of some tropical forests can decline, whereas in others it can be maintained at the same or a higher level than during the wet season(2). However, it remains uncertain to what extent water availability is responsible for productivity declines of tropical forests during the dry season(2,3). Here we use global satellite observations of two independent measures of vegetation photosynthetic properties (enhanced vegetation index from 2002 to 2012 and solar-induced chlorophyll fluorescence from 2007 to 2012) to investigate links between hydroclimate and tropical forest productivity. We find that above an annual rainfall threshold of approximately 2,000mmyr(-1), the evergreen state is sustained during the dry season in tropical rainforests worldwide, whereas below that threshold, this is not the case. Through awater-budget analysis of precipitation, potential evapotranspiration and satellite measurements of water storage change, we demonstrate that this threshold determines whether the supply of seasonally redistributed subsurface water storage from the wet season can satisfy plant water demands in the subsequent dry season. We conclude that water availability exerts a first-order control on vegetation seasonality in tropical forests globally. Our framework can also help identify where tropical forests may be vulnerable or resilient to future hydroclimatic changes.

We examine satellite-based measurements of solar-induced chlorophyll fluorescence (SIF) over the region impacted by the Russian drought and heat wave of 2010. Like the popular Normalized Difference Vegetation Index (NDVI) that has been used for decades to measure photosynthetic capacity, SIF measurements are sensitive to the fraction of absorbed photosynthetically-active radiation (fPAR). However, in addition, SIF is sensitive to PAR as well as the fluorescence yield that is related to the photosynthetic yield. Both SIF and NDVI from satellite data show drought-related declines early in the growing season in 2010 as compared to other years between 2007 and 2013 for areas dominated by crops and grasslands. This suggests an early manifestation of the dry conditions on fPAR. Using MODIS fPAR retrievals, we computed SIF/APAR which is related to light use efficiencies (LUEs) for fluorescence and photosynthesis. We found drought-related losses in fluorescence efficiency for all areas including those dominated by mixed forests. Unlike croplands and grasslands, areas of mixed forest did not show significant drought-related declines in fPAR. We also simulated SIF and Gross Primary Productivity (GPP) using a global land surface model driven by observation-based meteorological fields. The model provides a reasonable simulation of the drought and heat impacts on SIF in terms of the spatial extents of anomalies, but some differences were found in timing of the peak drought response between modeled and observed SIP. Model data also suggested that drought-related declines in LUE for fluorescence and photosynthesis in areas of mixed forest produce losses in SIF and GPP. SIF and GPP losses due to drought in croplands and grasslands result from both LUE and fPAR reductions. The combination of SIF and NDVI or fPAR data is shown to be an important source of information for evaluating model performance. (C) 2015 Elsevier Inc All rights reserved.

The projected responses of forest ecosystems to warming and drying associated with twenty-first-century climate change vary widely from resiliency to widespread tree mortality(1-3). Current vegetation models lack the ability to account for mortality of overstorey trees during extreme drought owing to uncertainties in mechanisms and thresholds causing mortality(4,5). Here we assess the causes of tree mortality, using field measurements of branch hydraulic conductivity during ongoing mortality in Populus tremuloides in the southwestern United States and a detailed plant hydraulics model. We identify a lethal plant water stress threshold that corresponds with a loss of vascular transport capacity from air entry into the xylem. We then use this hydraulic-based threshold to simulate forest dieback during historical drought, and compare predictions against three independent mortality data sets. The hydraulic threshold predicted with 75% accuracy regional patterns of tree mortality as found in field plots and mortality maps derived from Landsat imagery. In a high-emissions scenario, climate models project that drought stress will exceed the observed mortality threshold in the southwestern United States by the 2050s. Our approach provides a powerful and tractable way of incorporating tree mortality into vegetation models to resolve uncertainty over the fate of forest ecosystems in a changing climate.

Several studies have shown that satellite retrievals of solar-induced chlorophyll fluorescence (SIF) provide useful information on terrestrial photosynthesis or gross primary production (GPP). Here, we have incorporated equations coupling SIF to photosynthesis in a land surface model, the National Center for Atmospheric Research Community Land Model version 4 (NCAR CLM4), and have demonstrated its use as a diagnostic tool for evaluating the calculation of photosynthesis, a key process in a land surface model that strongly influences the carbon, water, and energy cycles. By comparing forward simulations of SIF, essentially as a byproduct of photosynthesis, in CLM4 with observations of actual SIF, it is possible to check whether the model is accurately representing photosynthesis and the processes coupled to it. We provide some background on how SIF is coupled to photosynthesis, describe how SIF was incorporated into CLM4, and demonstrate that our simulated relationship between SIF and GPP values are reasonable when compared with satellite (Greenhouse gases Observing SATellite; GOSAT) and in situ flux-tower measurements. CLM4 overestimates SIF in tropical forests, and we show that this error can be corrected by adjusting the maximum carboxylation rate (V-max) specified for tropical forests in CLM4. Our study confirms that SIF has the potential to improve photosynthesis simulation and thereby can play a critical role in improving land surface and carbon cycle models.

Carbonyl sulfide (OCS), the most abundant sulfur gas in the atmosphere, has a summer minimum associated with uptake by vegetation and soils, closely correlated with CO2. We report the first direct measurements to our knowledge of the ecosystem flux of OCS throughout an annual cycle, at a mixed temperate forest. The forest took up OCS during most of the growing season with an overall uptake of 1.36 +/- 0.01 mol OCS per ha (43.5 +/- 0.5 g S per ha, 95% confidence intervals) for the year. Daytime fluxes accounted for 72% of total uptake. Both soils and incompletely closed stomata in the canopy contributed to nighttime fluxes. Unexpected net OCS emission occurred during the warmest weeks in summer. Many requirements necessary to use fluxes of OCS as a simple estimate of photosynthesis were not met because OCS fluxes did not have a constant relationship with photosynthesis throughout an entire day or over the entire year. However, OCS fluxes provide a direct measure of ecosystem-scale stomatal conductance and mesophyll function, without relying on measures of soil evaporation or leaf temperature, and reveal previously unseen heterogeneity of forest canopy processes. Observations of OCS flux provide powerful, independent means to test and refine land surface and carbon cycle models at the ecosystem scale.