Photosynthesis is the fundamental biological process that introduced oxygen into Earth's atmosphere and continues to power life, from the earliest single-celled organisms to entire global ecosystems. Yet, measuring photosynthesis across scales has been challenging because traditional techniques have not transcended scales. The emergence of remote-sensing techniques to measure solar-induced chlorophyll fluorescence (SIF) provides a unique approach to estimate photosynthesis across spatiotemporal scales, representing a new age for optical remote sensing to study photosynthesis and shaping the decades of satellite SIF research. Here, focusing on spatiotemporal scales, we review the mechanisms that drive the relationship between SIF and photosynthesis. Remotely sensed SIF is modulated by biological drivers, environmental drivers, the interaction between biological and environmental drivers, and the viewing geometry. Studying fluorescence at small scales provides the ecophysiological understanding needed to disentangle the biological and environmental drivers of SIF at larger scales. Leveraging progress in satellite SIF, future research should focus on cross-scale mechanistic understanding of the drivers of SIF and using SIF as a metric for plant function beyond photosynthesis.

The model of C3 photosynthesis of Farquhar et al. (1980) integrated knowledge of the functioning of the biochemical components of photosynthetic carbon assimilation in C3 plants. The model linked equations describing activated Rubisco kinetics with those on the stoichiometry of the photosynthetic carbon reduction cycle and the photorespiratory carbon oxidation cycle, particularly on their energetic (electron transport, ATP synthesis and NADPH) requirements. It included temperature dependencies of these processes and combined them with a semi-empirical equation for the dependence of potential electron transport rate on absorbed irradiance. The model aimed to match generalized observations of photosynthetic gas exchange of leaves with predictions from this mathematical summary of photosynthesis. In this model, we introduced the hypothesis that the rate of Rubisco carboxylation could not exceed the capacity for RuP2 regeneration or the enzymatic capacity to consume RuP2, and that the system behaved as a teeter-totter (see-saw) with a sharp transition from one limiting state to the other. We suggested that to model genotypic and phenotypic variations amongst leaves most parameters could be assigned a priori and only the maximum Rubisco activity (Vcmax) and the maximal electron transport rate, Jmax, needed to be varied. This simplicity of use led to the wider spread application and success of the model.

Solar-induced chlorophyll fluorescence (SIF) is widely accepted as a proxy for gross primary productivity (GPP). Among the various SIF measurements, tower-based SIF measurements allow for continuous monitoring of SIF variation at a canopy scale with high temporal resolution, making it suitable for monitoring highly variable plant physiological responses to environmental changes. However, because of the strong and close relationship between SIF and absorbed photosynthetically active radiation (aPAR), it may be difficult to detect the influence of environmental drivers other than light conditions. Among the drivers, atmospheric dryness (vapor pressure deficit, VPD) is projected to increase as drought becomes more frequent and severe in the future, negatively impacting plants. In this study, we evaluated the tower-based high-frequency SIF measurement as a tool for detecting plant response to highly variable VPD. The study was performed in a mixed temperate forest in Virginia, USA, where a 40-m-tall flux tower has been measuring gas and energy exchanges and ancillary environmental drivers, and the Fluospec 2 system has been measuring SIF. We show that a proper definition of light availability to vegetation can reproduce SIF response to changing VPD that is comparable to GPP response as estimated from eddy covariance measurement: GPP decreased with rising VPD regardless of how aPAR was defined, whereas SIF decreased only when aPAR was defined as the PAR absorbed by chlorophyll (aPARchl) or simulated by a model (Soil Canopy Observation, Photochemistry and Energy fluxes, SCOPE). We simulated the effect of VPD on SIF with two different simulation modes of fluorescence emission representing contrasting moisture conditions, ‘Moderate’ and ‘Soil Moisture (SM) Stress’ modes. The decreasing SIF to rising VPD was only found in the SM Stress mode, implying that the SIF-VPD relationship depends on soil moisture conditions. Furthermore, we observed a similar response of SIF to VPD at hourly and daily scales, indicating that satellite measurements can be used to study the effects of environmental drivers other than light conditions. Finally, the definition of aPAR emphasizes the importance of canopy structure research to interpret remote sensing observations properly.

Vegetation 'greenness' characterized by spectral vegetation indices (VIs) is an integrative measure of vegetation leaf abundance, biochemical properties and pigment composition. Surprisingly, satellite observations reveal that several major VIs over the US Corn Belt are higher than those over the Amazon rainforest, despite the forests having a greater leaf area. This contradicting pattern underscores the pressing need to understand the underlying drivers and their impacts to prevent misinterpretations. Here we show that macroscale shadows cast by complex forest structures result in lower greenness measures compared with those cast by structurally simple and homogeneous crops. The shadow-induced contradictory pattern of VIs is inevitable because most Earth-observing satellites do not view the Earth in the solar direction and thus view shadows due to the sun-sensor geometry. The shadow impacts have important implications for the interpretation of VIs and solar-induced chlorophyll fluorescence as measures of global vegetation changes. For instance, a land-conversion process from forests to crops over the Amazon shows notable increases in VIs despite a decrease in leaf area. Our findings highlight the importance of considering shadow impacts to accurately interpret remotely sensed VIs and solar-induced chlorophyll fluorescence for assessing global vegetation and its changes.

Aerosols can affect photosynthesis through radiative perturbations such as scattering and absorbing solar radiation. This biophysical impact has been widely studied using field measurements, but the sign and magnitude at continental scales remain uncertain. Solar-induced fluorescence (SIF), emitted by chlorophyll, strongly correlates with photosynthesis. With recent advancements in Earth observation satellites, we leverage SIF observations from the Tropospheric Monitoring Instrument (TROPOMI) with unprecedented spatial resolution and near-daily global coverage, to investigate the impact of aerosols on photosynthesis. Our analysis reveals that on weekends when there is more plant-available sunlight due to less particulate pollution, 64% of regions across Europe show increased SIF, indicating more photosynthesis. Moreover, we find a widespread negative relationship between SIF and aerosol loading across Europe. This suggests the possible reduction in photosynthesis as aerosol levels increase, particularly in ecosystems limited by light availability. By considering two plausible scenarios of improved air quality-reducing aerosol levels to the weekly minimum 3-d values and levels observed during the COVID-19 period-we estimate a potential of 41 to 50 Mt net additional annual CO2 uptake by terrestrial ecosystems in Europe. This work assesses human impacts on photosynthesis via aerosol pollution at continental scales using satellite observations. Our results highlight i) the use of spatiotemporal variations in satellite SIF to estimate the human impacts on photosynthesis and ii) the potential of reducing particulate pollution to enhance ecosystem productivity.

Solar-induced chlorophyll fluorescence (SIF) shows enormous promise as a proxy for photosynthesis and as a tool for modeling variability in gross primary productivity and net biosphere exchange (NBE). In this study, we explore the skill of SIF and other vegetation indicators in predicting variability in global atmospheric CO2 observations, and thus global variability in NBE. We do so using a 4-year record of CO2 observations from NASA's Orbiting Carbon Observatory 2 satellite and using a geostatistical inverse model. We find that existing SIF products closely correlate with space-time variability in atmospheric CO2 observations, particularly in the extratropics. In the extratropics, all SIF products exhibit greater skill in explaining variability in atmospheric CO2 observations compared to an ensemble of process-based CO2 flux models and other vegetation indicators. With that said, other vegetation indicators, when multiplied by photosynthetically active radiation, yield similar results as SIF and may therefore be an effective structural SIF proxy at regional to global spatial scales. Furthermore, we find that using SIF as a predictor variable in the geostatistical inverse model shifts the seasonal cycle of estimated NBE and yields an earlier end to the growing season relative to other vegetation indicators. These results highlight how SIF can help constrain global-scale variability in NBE.

There remains limited information to characterize the solar-induced chlorophyll fluorescence (SIF)-gross primary production (GPP) relationship in C4 cropping systems. The annual C4 crop corn and perennial C4 crop miscanthus differ in phenology, canopy structure and leaf physiology. Investigating the SIF-GPP relationships in these species could deepen our understanding of SIF-GPP relationships within C4 crops. Using in situ canopy SIF and GPP measurements for both species along with leaf-level measurements, we found considerable differences in the SIF-GPP relationships between corn and miscanthus, with a stronger SIF-GPP relationship and higher slope of SIF-GPP observed in corn compared to miscanthus. These differences were mainly caused by leaf physiology. For miscanthus, high non-photochemical quenching (NPQ) under high light, temperature and water vapor deficit (VPD) conditions caused a large decline of fluorescence yield (ΦF), which further led to a SIF midday depression and weakened the SIF-GPP relationship. The larger slope in corn than miscanthus was mainly due to its higher GPP in mid-summer, largely attributed to the higher leaf photosynthesis and less NPQ. Our results demonstrated variation of the SIF-GPP relationship within C4 crops and highlighted the importance of leaf physiology in determining canopy SIF behaviors and SIF-GPP relationships.

We provide here an overview of the remarkable life and outstanding research of David (Dave) Charles Fork (March 4, 1929–December 13, 2021) in oxygenic photosynthesis. In the words of the late Jack Edgar Myers, he was a top ‘photosynthetiker’. His research dealt with novel findings on light absorption, excitation energy distribution, and redistribution among the two photosystems, electron transfer, and their relation to dynamic membrane change as affected by environmental changes, especially temperature. David was an attentive listener and a creative designer of experiments and instruments, and he was also great fun to work with. He is remembered here by his family, coworkers, and friends from around the world including Australia, France, Germany, Japan, Sweden, Israel, and USA.

Sun-induced chlorophyll fluorescence (SIF) is a promising new tool for remotely estimating photosynthesis. However, the degree to which incoming solar radiation and the structure of the canopy rather than leaf physiology contribute to SIF variations is still not well characterized. Therefore, we investigated relationships between SIF and variables that at least partly capture the canopy structure component of SIF. For this, we relied on high-quality SIF observations from ground-based instruments, high-resolution airborne SIF imagery and the most recent satellite SIF products to cover large ranges in spatial and temporal resolution and diverse ecosystems. We found that the canopy structure-related near-infrared reflectance of vegetation multiplied by incoming sunlight (NIRvP) is a robust proxy for far-red SIF across a wide range of spatial and temporal scales. Our findings indicate that contributions from leaf physiology to SIF variability are small compared to the structure and radiation components. Also, NIRvP captured spatio-temporal patterns of canopy photosynthesis better than SIF, which seems to be mostly due to the greater retrieval noise of SIF. Compared to other relevant structural SIF proxies, NIRvP showed more robust relationships to SIF, especially at the global scale. Our results highlight the promise of using widely available NIRvP data for vegetation monitoring and also indicate the potential of using SIF and NIRvP in combination to extract physiological information from SIF.

Carbonyl sulfide (COS) has emerged as a multi-scale tracer for terrestrial photosynthesis. To infer ecosystem-scale photosynthesis from COS fluxes often requires knowledge of leaf relative uptake (LRU), the concentration-normalized ratio between leaf COS uptake and photosynthesis. However, current mechanistic understanding of LRU variability remains inadequate for deriving robust COS-based estimates of photosynthesis. We derive a set of closed-form equations to describe LRU responses to light, humidity and CO2 based on the Ball-Berry stomatal conductance model and the biochemical model of photosynthesis. This framework reproduces observed LRU responses: decreasing LRU with increasing light or decreasing humidity; it also predicts that LRU increases with ambient CO2 . By fitting the LRU equations to flux measurements on a C3 reed (Typha latifolia), we obtain physiological parameters that control LRU variability, including an estimate of the Ball-Berry slope of 7.1 without using transpiration measurements. Sensitivity tests reveal that LRU is more sensitive to photosynthetic capacity than to the Ball-Berry slope, indicating stomatal response to photosynthesis. This study presents a simple framework for interpreting observed LRU variability and upscaling LRU. The stoma-regulated LRU response to CO2 suggests that COS may offer a unique window into long-term stomatal acclimation to elevated CO2 .

Solar-induced chlorophyll fluorescence (SIF) shows great potential to assess plants physiological state and response to environmental changes. Recently the near-infrared reflectance of vegetation (NIRv) provides a promising way to quantify the confounding effect of canopy structure in SIF, while the difference between SIF and NIRv under varying environmental conditions has not been well explored. Here we developed a simple approach to extract the fluorescence yield (Phi(F)) by the combined use of SIF and the near-infrared radiance of vegetation (NIRvR). The proposed NIRvR approach was evaluated in multiple ways, including with the seasonal leaf-level steady-state fluorescence yield. Results indicate that NIRvR-derived Phi(F) well captured the seasonal variation of the fluorescence yield changes, and achieved similar results with the existing approach. Both SIF and NIRvR were derived from the airborne imaging fluorescence spectrometer HyPlant for three case studies to evaluate the impacts of light adaptation, heat stress and water limitation on Phi(F). For the light adaptation case study, Phi(F) over the low-light adapted sugar beet field was about 1.3 times larger compared to an unaffected reference area while the difference in NIRvR was minimal, which clearly shows the short-term photosynthetic light induction effect and the ability of SIF to detect plant physiological responses. For the heat stress experiment, OF decreased during a natural heatwave in 2015 in the fields of rapeseed from 0.0150 to 0.0130, barley from 0.0152 to 0.0144, and wheat from 0.0146 to 0.0142 which showed signs of senescence, while slightly increased from 0.0125 to 0.0130 in the corn field which was still in growing. At the water-limited sugar beet field, Phi(F) first increased towards solar noon and then slightly decreased during the afternoon over the water-limited areas from 0.017 to 0.021 and 0.020, with high temperature and high light at noon. The advantages to use SIF/NIRvR as a proxy of Phi(F) to detect stress-induced limitations in photosynthesis include that the impacts of canopy structure and sun-sensor geometry on the Phi(F) estimation are explicitly cancelled, and photosynthetically active radiation (PAR) is not required as input. Finally, our approach is directly applicable to satellite-derived estimates of SIF, enabling the study of variations in Phi(F) to detect the effects of abiotic changes and stresses at large scale.

Remote sensing of solar-induced fluorescence (SIF) opens a new window for quantifying a key ecological variable, the terrestrial ecosystem gross primary production (GPP), because of the revealed strong SIF-GPP correlation. However, similar to many other remotely sensed metrics, SIF observations suffer from the sun-sensor geometry effects, which may have important impacts on the SIF-GPP relationship but remain poorly understood. Here we used remotely sensed SIF, globally distributed tower GPP data, and a mechanistic model to provide a systematic analysis. Our results reveal that leaf physiology, canopy structure, and sun-sensor geometries all affect the SIF-GPP relationship. In particular, we found that SIF observations in the sun-tracking hotspot direction can be a better proxy of GPP due to the similar responses of light use efficiency and SIF escaping probability in the hotspot direction to the increasing incoming solar radiation. Such conclusions are supported by a variety of modeling simulations and satellite observations over various plant function types, at different time scales and with satellite observational modes. This study demonstrates the potential and advantage of normalizing SIF observations to the hotspot direction for better global GPP estimations. This study also demonstrates the great potentials of current and future spaceborne sun-tracking satellite missions for a significant improvement in measuring and monitoring, at a wide range of spatial and temporal scales, the changes in terrestrial ecosystem GPP in response to anticipated changes in the Earth's environmental conditions.

Sun-induced chlorophyll fluorescence (SIF) measurements have shown unique potential for quantifying plant physiological stress. However, recent investigations found canopy structure and radiation largely control SIF, and physiological relevance of SIF remains yet to be fully understood. This study aims to evaluate whether the SIF-derived physiological signal improves quantification of crop responses to environmental stresses, by analyzing data at three different spatial scales within the U.S. Corn Belt, i.e. experiment plot, field, and regional scales, where ground-based portable, stationary and space-borne hyperspectral sensing systems are used, respectively. We found that, when controlling for variations in incoming radiation and canopy structure, crop SIF signals can be decomposed into non-physiological (i.e. canopy structure and radiation, 60% ∼ 82%) and physiological information (i.e. physiological SIF yield, ΦF, 17% ∼ 31%), which confirms the contribution of physiological variation to SIF. We further evaluated whether ΦF indicated plant responses under high-temperature and high vapor pressure deficit (VPD) stresses. The plot-scale data showed that ΦF responded to the proxy for physiological stress (partial correlation coefficient, r p= 0.40, p< 0.001) while non-physiological signals of SIF did not respond (p> 0.1). The field-scale ΦF data showed water deficit stress from the comparison between irrigated and rainfed fields, and ΦF was positively correlated with canopy-scale stomatal conductance, a reliable indicator of plant physiological condition (correlation coefficient r= 0.60 and 0.56 for an irrigated and rainfed sites, respectively). The regional-scale data showed ΦF was more strongly correlated spatially with air temperature and VPD (r= 0.23 and 0.39) than SIF (r= 0.11 and 0.34) for the U.S. Corn Belt. The lines of evidence suggested that ΦF reflects crop physiological responses to environmental stresses with greater sensitivity to stress factors than SIF, and the stress quantification capability of ΦF is spatially scalable. Utilizing ΦF for physiological investigations will contribute to improve our understanding of vegetation responses to high-temperature and high-VPD stresses.

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet, Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of "Plant Functional Types" (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, that is, to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages ("Lineage Functional Types") will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Solar-induced chlorophyll fluorescence (SIF) provides us with new opportunities to understand the physiological and structural dynamics of vegetation from leaf to global scales. However, the relationships between SIF and gross primary productivity (GPP) are not fully understood, which is mainly due to the challenges of decoupling structural and physiological factors that control the relationships. Here, we report the results of continuous observations of canopy-level SIF, GPP, absorbed photosynthetically active radiation (APAR), and chlorophyll: carotenoid index (CCI) in a temperate evergreen needleleaf forest. To understand the mechanisms underlying the relationship between GPP and SIF, we investigated the relationships of light use efficiency (LUEp), chlorophyll fluorescence yield (?F), and the fraction of emitted SIF photons escaping from the canopy (fesc) separately. We found a strongly non-linear relationship between GPP and SIF at diurnal and seasonal time scales (R2 = 0.91 with a hyperbolic regression function, daily). GPP saturated with APAR, while SIF did not. Also, there were differential responses of LUEp and ?F to air temperature. While LUEp reached saturation at high air temperatures, ?F did not saturate. We found that the canopy-level chlorophyll: carotenoid index was strongly correlated to canopy-level ?F (R2 = 0.84) implying that ?F could be more closely related to pigment pool changes rather than LUEp. In addition, we found that the fesc contributed to a stronger SIF-GPP relationship by partially capturing the response of LUEp to diffuse light. These findings can help refine physiological and structural links between canopy-level SIF and GPP in evergreen needleleaf forest.

Disentangling the individual contributions from vegetation and soil in measured canopy reflectance is a grand challenge to the remote sensing and ecophysiology communities. Since Solar Induced chlorophyll Fluorescence (SIF) is uniquely emitted from vegetation, it can be used to evaluate how well reflectance-based vegetation indices (VIs) can separate the vegetation and soil components. Due to the residual soil background contributions, Near-infrared (NIR) reflectance of vegetation (NIRv) and Difference Vegetation index (DVI) present offsets when compared to SIF (i.e., the value of NIRv or DVI is non-zero when SIF is zero). In this study, we proposed a simple framework for estimating the true NIR reflectance of vegetation from Hyperspectral measurements (NIRvH) with minimal soil impacts. NIRvH takes advantage of the spectral shape variations in the red-edge region to minimize the soil effects. We evaluated the capability of NIRvH, NIRv and DVI in isolating the true NIR reflectance of vegetation using the data from both the model-based simulations and Hyperspectral Plant imaging spectrometer (HyPlant) measurements. Benchmarked by simultaneously measured SIF, NIRvH has the smallest offset (0–0.037), as compared to an intermediate offset of 0.047–0.062 from NIRv, and the largest offset of 0.089–0.112 from DVI. The magnitude of the offset can vary with different soil reflectance spectra across spatio-temporal scales, which may lead to bias in the downstream NIRv-based photosynthesis estimates. NIRvH and SIF measurements from the same sensor platform avoided complications due to different geometry, footprint and time of observation across sensors when studying the radiative transfer of reflected photons and SIF. In addition, NIRvH was primarily determined by canopy structure rather than chlorophyll content and soil brightness. Our work showcases that NIRvH is promising for retrieving canopy structure parameters such as leaf area index and leaf inclination angle, and for estimating fluorescence yield with current and forthcoming hyperspectral satellite measurements.

High temperature and accompanying high vapor pressure deficit often stress plants without causing distinctive changes in plant canopy structure and consequential spectral signatures. Sun-induced chlorophyll fluorescence (SIF), because of its mechanistic link with photosynthesis, may better detect such stress than remote sensing techniques relying on spectral reflectance signatures of canopy structural changes. However, our understanding about physiological mechanisms of SIF and its unique potential for physiological stress detection remains less clear. In this study, we measured SIF at a high-temperature experiment, Temperature Free-Air Controlled Enhancement, to explore the potential of SIF for physiological investigations. The experiment provided a gradient of soybean canopy temperature with 1.5, 3.0, 4.5, and 6.0°C above the ambient canopy temperature in the open field environments. SIF yield, which is normalized by incident radiation and the fraction of absorbed photosynthetically active radiation, showed a high correlation with photosynthetic light use efficiency (r = 0.89) and captured dynamic plant responses to high-temperature conditions. SIF yield was affected by canopy structural and plant physiological changes associated with high-temperature stress (partial correlation r = 0.60 and -0.23). Near-infrared reflectance of vegetation, only affected by canopy structural changes, was used to minimize the canopy structural impact on SIF yield and to retrieve physiological SIF yield (ΦF ) signals. ΦF further excludes the canopy structural impact than SIF yield and indicates plant physiological variability, and we found that ΦF outperformed SIF yield in responding to physiological stress (r = -0.37). Our findings highlight that ΦF sensitively responded to the physiological downregulation of soybean gross primary productivity under high temperature. ΦF , if reliably derived from satellite SIF, can support monitoring regional crop growth and different ecosystems' vegetation productivity under environmental stress and climate change.

The uptake of carbonyl sulfide (COS) by terrestrial plants is linked to photosynthetic uptake of CO2 as these gases partly share the same uptake pathway. Applying COS as a photosynthesis tracer in models requires an accurate representation of biosphere COS fluxes, but these models have not been extensively evaluated against field observations of COS fluxes. In this paper, the COS flux as simulated by the Simple Biosphere Model, version 4 (SiB4), is updated with the latest mechanistic insights and evaluated with site observations from different biomes: one evergreen needleleaf forest, two deciduous broadleaf forests, three grasslands, and two crop fields spread over Europe and North America. We improved SiB4 in several ways to improve its representation of COS. To account for the effect of atmospheric COS mole fractions on COS biosphere uptake, we replaced the fixed atmospheric COS mole fraction boundary condition originally used in SiB4 with spatially and temporally varying COS mole fraction fields. Seasonal amplitudes of COS mole fractions are ∼50–200 ppt at the investigated sites with a minimum mole fraction in the late growing season. Incorporating seasonal variability into the model reduces COS uptake rates in the late growing season, allowing better agreement with observations. We also replaced the empirical soil COS uptake model in SiB4 with a mechanistic model that represents both uptake and production of COS in soils, which improves the match with observations over agricultural fields and fertilized grassland soils. The improved version of SiB4 was capable of simulating the diurnal and seasonal variation in COS fluxes in the boreal, temperate, and Mediterranean region. Nonetheless, the daytime vegetation COS flux is underestimated on average by 8±27 %, albeit with large variability across sites. On a global scale, our model modifications decreased the modeled COS terrestrial biosphere sink from 922 Gg S yr−1 in the original SiB4 to 753 Gg S yr−1 in the updated version. The largest decrease in fluxes was driven by lower atmospheric COS mole fractions over regions with high productivity, which highlights the importance of accounting for variations in atmospheric COS mole fractions. The change to a different soil model, on the other hand, had a relatively small effect on the global biosphere COS sink. The secondary role of the modeled soil component in the global COS budget supports the use of COS as a global photosynthesis tracer. A more accurate representation of COS uptake in SiB4 should allow for improved application of atmospheric COS as a tracer of local- to global-scale terrestrial photosynthesis.

Here, we describe a model of C 3 , C 3 –C 4 intermediate, and C 4 photosynthesis that is designed to facilitate quantitative analysis of physiological measurements. The model relates the factors limiting electron transport and carbon metabolism, the regulatory processes that coordinate these metabolic domains, and the responses to light, carbon dioxide, and temperature. It has three unique features. First, mechanistic expressions describe how the cytochrome b 6 f complex controls electron transport in mesophyll and bundle sheath chloroplasts. Second, the coupling between the mesophyll and bundle sheath expressions represents how feedback regulation of Cyt b 6 f coordinates electron transport and carbon metabolism. Third, the temperature sensitivity of Cyt b 6 f is differentiated from that of the coupling between NADPH, Fd, and ATP production. Using this model, we present simulations demonstrating that the light dependence of the carbon dioxide compensation point in C 3 –C 4 leaves can be explained by co-occurrence of light saturation in the mesophyll and light limitation in the bundle sheath. We also present inversions demonstrating that population-level variation in the carbon dioxide compensation point in a Type I C 3 –C 4 plant, Flaveria chloraefolia , can be explained by variable allocation of photosynthetic capacity to the bundle sheath. These results suggest that Type I C 3 –C 4 intermediate plants adjust pigment and protein distributions to optimize the glycine shuttle under different light and temperature regimes, and that the malate and aspartate shuttles may have originally functioned to smooth out the energy supply and demand associated with the glycine shuttle. This model has a wide range of potential applications to physiological, ecological, and evolutionary questions.