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ABSTRACT Photosynthesis is the largest flux of carbon between the atmosphere and Earth's surface and is driven by enzymes that require nitrogen, namely, ribulose‐1,5‐bisphosphate (RuBisCO). Thus, photosynthesis is a key link between the terrestrial carbon and nitrogen cycle, and the representation of this link is critical for coupled carbon‐nitrogen land surface models. Models and observations suggest that soil nitrogen availability can limit plant productivity increases under elevated CO₂. Plants acclimate to elevated CO₂by downregulating RuBisCO and thus nitrogen in leaves, but this acclimation response is not currently included in land surface models. Acclimation of photosynthesis to CO₂can be simulated by the photosynthetic optimality theory in a way that matches observations. Here, we incorporated this theory into the land surface component of the Energy Exascale Earth System Model (ELM). We simulated land surface carbon and nitrogen processes under future elevated CO₂conditions to 2100 using the RCP8.5 high emission scenario. Our simulations showed that when photosynthetic acclimation is considered, photosynthesis increases under future conditions, but maximum RuBisCO carboxylation and thus photosynthetic nitrogen demand decline. We analyzed two simulations that differed as to whether the saved nitrogen could be used in other parts of the plant. The allocation of saved leaf nitrogen to other parts of the plant led to (1) a direct alleviation of plant nitrogen limitation through reduced leaf nitrogen requirements and (2) an indirect reduction in plant nitrogen limitation through an enhancement of root growth that led to increased plant nitrogen uptake. As a result, reallocation of saved leaf nitrogen increased ecosystem carbon stocks by 50.3% in 2100 as compared to a simulation without reallocation of saved leaf nitrogen. These results suggest that land surface models may overestimate future ecosystem nitrogen limitation if they do not incorporate leaf nitrogen savings resulting from photosynthetic acclimation to elevated CO₂. 

Abstract Drought is often thought to reduce ecosystem photosynthesis. However, theory suggests there is potential for increased photosynthesis during meteorological drought, especially in energy-limited ecosystems. Here, we examine the response of photosynthesis (gross primary productivity, GPP) to meteorological drought across the water-energy limitation spectrum. We find a consistent increase in eddy covariance GPP during spring drought in energy-limited ecosystems (83% of the energy-limited sites). Half of spring GPP sensitivity to precipitation was predicted solely from the wetness index (R² = 0.47,p < 0.001), with weaker relationships in summer and fall. Our results suggest GPP increases during spring drought for 55% of vegetated Northern Hemisphere lands ( >30° N). We then compare these results to terrestrial biosphere model outputs and remote sensing products. In contrast to trends detected in eddy covariance data, model mean GPP always declined under spring precipitation deficits after controlling for air temperature and light availability. While remote sensing products captured the observed negative spring GPP sensitivity in energy-limited ecosystems, terrestrial biosphere models proved insufficiently sensitive to spring precipitation deficits. 

Abstract Plants with the C₄photosynthesis pathway typically respond to climate change differently from more common C₃-type plants, due to their distinct anatomical and biochemical characteristics. These different responses are expected to drive changes in global C₄and C₃vegetation distributions. However, current C₄vegetation distribution models may not predict this response as they do not capture multiple interacting factors and often lack observational constraints. Here, we used global observations of plant photosynthetic pathways, satellite remote sensing, and photosynthetic optimality theory to produce an observation-constrained global map of C₄vegetation. We find that global C₄vegetation coverage decreased from 17.7% to 17.1% of the land surface during 2001 to 2019. This was the net result of a reduction in C₄natural grass cover due to elevated CO₂favoring C₃-type photosynthesis, and an increase in C₄crop cover, mainly from corn (maize) expansion. Using an emergent constraint approach, we estimated that C₄vegetation contributed 19.5% of global photosynthetic carbon assimilation, a value within the range of previous estimates (18–23%) but higher than the ensemble mean of dynamic global vegetation models (14 ± 13%; mean ± one standard deviation). Our study sheds insight on the critical and underappreciated role of C₄plants in the contemporary global carbon cycle. 

Abstract Recent years have seen growing appreciation that rapidly intensifying flash droughts are significant climate hazards with major economic and ecological impacts. This has motivated efforts to inventory, monitor, and forecast flash drought events. Here we consider the question of whether the term “flash drought” comprises multiple distinct classes of event, which would imply that understanding and forecasting flash droughts might require more than one framework. To do this, we first extend and evaluate a soil moisture volatility–based flash drought definition that we introduced in previous work and use it to inventory the onset dates and severity of flash droughts across the contiguous United States (CONUS) for the period 1979–2018. Using this inventory, we examine meteorological and land surface conditions associated with flash drought onset and recovery. These same meteorological and land surface conditions are then used to classify the flash droughts based on precursor conditions that may represent predictable drivers of the event. We find that distinct classes of flash drought can be diagnosed in the event inventory. Specifically, we describe three classes of flash drought: “dry and demanding” events for which antecedent evaporative demand is high and soil moisture is low, “evaporative” events with more modest antecedent evaporative demand and soil moisture anomalies, but positive antecedent evaporative anomalies, and “stealth” flash droughts, which are different from the other two classes in that precursor meteorological anomalies are modest relative to the other classes. The three classes exhibit somewhat different geographic and seasonal distributions. We conclude that soil moisture flash droughts are indeed a composite of distinct types of rapidly intensifying droughts, and that flash drought analyses and forecasts would benefit from approaches that recognize the existence of multiple phenomenological pathways. 

Abstract Plants invest a considerable amount of leaf nitrogen in the photosynthetic enzyme ribulose-1,5-bisphosphate carboxylase-oxygenase (RuBisCO), forming a strong coupling of nitrogen and photosynthetic capacity. Variability in the nitrogen-photosynthesis relationship indicates different nitrogen use strategies of plants (i.e., the fraction nitrogen allocated to RuBisCO; fLNR), however, the reason for this remains unclear as widely different nitrogen use strategies are adopted in photosynthesis models. Here, we use a comprehensive database of in situ observations, a remote sensing product of leaf chlorophyll and ancillary climate and soil data, to examine the global distribution in fLNR using a random forest model. We find global fLNR is 18.2 ± 6.2%, with its variation largely driven by negative dependence on leaf mass per area and positive dependence on leaf phosphorus. Some climate and soil factors (i.e., light, atmospheric dryness, soil pH, and sand) have considerable positive influences on fLNR regionally. This study provides insight into the nitrogen-photosynthesis relationship of plants globally and an improved understanding of the global distribution of photosynthetic potential.