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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 CO 2 . Plants acclimate to elevated CO 2 by downregulating RuBisCO and thus nitrogen in leaves, but this acclimation response is not currently included in land surface models. Acclimation of photosynthesis to CO 2 can 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 CO 2 conditions 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 CO 2 ., (© 2024 John Wiley & Sons Ltd.)

Global warming has been linked to declines in tree growth. However, it is unclear how the asymmetry in daytime and nighttime warming influences this response. Here, we use 2947 residual tree-ring width chronologies covering 32 species at 2493 sites, between 1901 and 2018, across the Northern Hemisphere, to analyze the effects of daytime and nighttime temperatures, precipitation, and drought stress on the radial growth of trees. We show that drought stress was primarily triggered by daytime rather than nighttime warming. The radial growth of trees was more sensitive to drought stress in warm regions than in cold regions, especially for angiosperms. Our study provides robust evidence that daytime warming is the primary driver of the observed declines in forest productivity related to drought stress and that daytime and nighttime warming should be considered separately when modelling forest-climate interactions and feedbacks in a future, warmer world., (© 2022 John Wiley & Sons Ltd.)

Terrestrial ecosystems regulate Earth's climate through water, energy, and biogeochemical transformations. Despite a key role in regulating the Earth system, terrestrial ecology has historically been underrepresented in the Earth system models (ESMs) that are used to understand and project global environmental change. Ecology and Earth system modeling must be integrated for scientists to fully comprehend the role of ecological systems in driving and responding to global change. Ecological insights can improve ESM realism and reduce process uncertainty, while ESMs offer ecologists an opportunity to broadly test ecological theory and increase the impact of their work by scaling concepts through time and space. Despite this mutualism, meaningfully integrating the two remains a persistent challenge, in part because of logistical obstacles in translating processes into mathematical formulas and identifying ways to integrate new theories and code into large, complex model structures. To help overcome this interdisciplinary challenge, we present a framework consisting of a series of interconnected stages for integrating a new ecological process or insight into an ESM. First, we highlight the multiple ways that ecological observations and modeling iteratively strengthen one another, dispelling the illusion that the ecologist's role ends with initial provision of data. Second, we show that many valuable insights, products, and theoretical developments are produced through sustained interdisciplinary collaborations between empiricists and modelers, regardless of eventual inclusion of a process in an ESM. Finally, we provide concrete actions and resources to facilitate learning and collaboration at every stage of data-model integration. This framework will create synergies that will transform our understanding of ecology within the Earth system, ultimately improving our understanding of global environmental change, and broadening the impact of ecological research., (© 2021 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Earlier spring phenological events have been widely reported in plants under global warming. Recent studies reported a slowdown in the warming-induced advanced spring phenology in temperate regions. However, previous research mainly focused on daily mean temperature, thus neglecting the asymmetric phenological responses to daytime and nighttime temperature. Using long-term records of leaf unfolding in eight deciduous species at 1300 sites across central Europe, we assessed and compared the effects of daytime temperature, nighttime temperature, and photoperiod on leaf unfolding during 1951-1980 and 1981-2013. Although leaf unfolding was advanced by daytime warming during 1951-2013, the advancing responses of leaf unfolding significantly decreased from 1951-1980 to 1981-2013 due to a lower accumulation of chilling units by daytime warming. Nighttime warming delayed leaf unfolding during 1951-1980 but advanced it during 1981-2013 due to a higher accumulation of chilling units by nighttime warming. In contrast, critical daylength and plasticity of leaf unfolding dates remained unchanged between 1951 and 2013. Our study provided evidence that daytime warming instead of nighttime warming accounts for the slowdown in the advancing spring phenology and implied that nighttime warming-induced earlier spring phenology may be buffering the slowdown of the advanced spring phenology by daytime warming. The response of spring phenology to nighttime temperature may override that to daytime temperature under the actual trends in global warming., (© 2021 John Wiley & Sons Ltd.)

The mechanisms responsible for photosynthetic acclimation are not well understood, effectively limiting predictability under future conditions. Least-cost optimality theory can be used to predict the acclimation of photosynthetic capacity based on the assumption that plants maximize carbon uptake while minimizing the associated costs. Here, we use this theory as a null model in combination with multiple datasets of C 3 plant photosynthetic traits to elucidate the mechanisms underlying photosynthetic acclimation to elevated temperature and carbon dioxide (CO 2 ). The model-data comparison showed that leaves decrease the ratio of the maximum rate of electron transport to the maximum rate of Rubisco carboxylation (J max /V cmax ) under higher temperatures. The comparison also indicated that resources used for Rubisco and electron transport are reduced under both elevated temperature and CO 2 . Finally, our analysis suggested that plants underinvest in electron transport relative to carboxylation under elevated CO 2 , limiting potential leaf-level photosynthesis under future CO 2 concentrations. Altogether, our results show that acclimation to temperature and CO 2 is primarily related to resource conservation at the leaf level. Under future, warmer, high CO 2 conditions, plants are therefore likely to use less nutrients for leaf-level photosynthesis, which may impact whole-plant to ecosystem functioning., (© 2020 John Wiley & Sons Ltd.)

Plant respiration is an important contributor to the proposed positive global carbon-cycle feedback to climate change. However, as a major component, leaf mitochondrial ('dark') respiration (R d ) differs among species adapted to contrasting environments and is known to acclimate to sustained changes in temperature. No accepted theory explains these phenomena or predicts its magnitude. Here we propose that the acclimation of R d follows an optimal behaviour related to the need to maintain long-term average photosynthetic capacity (V cmax ) so that available environmental resources can be most efficiently used for photosynthesis. To test this hypothesis, we extend photosynthetic co-ordination theory to predict the acclimation of R d to growth temperature via a link to V cmax , and compare predictions to a global set of measurements from 112 sites spanning all terrestrial biomes. This extended co-ordination theory predicts that field-measured R d and V cmax accessed at growth temperature (R d,tg and V cmax,tg ) should increase by 3.7% and 5.5% per degree increase in growth temperature. These acclimated responses to growth temperature are less steep than the corresponding instantaneous responses, which increase 8.1% and 9.9% per degree of measurement temperature for R d and V cmax respectively. Data-fitted responses proof indistinguishable from the values predicted by our theory, and smaller than the instantaneous responses. Theory and data are also shown to agree that the basal rates of both R d and V cmax assessed at 25°C (R d,25 and V cmax,25 ) decline by ~4.4% per degree increase in growth temperature. These results provide a parsimonious general theory for R d acclimation to temperature that is simpler-and potentially more reliable-than the plant functional type-based leaf respiration schemes currently employed in most ecosystem and land-surface models., (© 2019 John Wiley & Sons Ltd.)

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects. We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives., (© 2019 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

While temperature responses of photosynthesis and plant respiration are known to acclimate over time in many species, few studies have been designed to directly compare process-level differences in acclimation capacity among plant types. We assessed short-term (7 day) temperature acclimation of the maximum rate of Rubisco carboxylation (V cmax ), the maximum rate of electron transport (J max ), the maximum rate of phosphoenolpyruvate carboxylase carboxylation (V pmax ), and foliar dark respiration (R d ) in 22 plant species that varied in lifespan (annual and perennial), photosynthetic pathway (C 3 and C 4 ), and climate of origin (tropical and nontropical) grown under fertilized, well-watered conditions. In general, acclimation to warmer temperatures increased the rate of each process. The relative increase in different photosynthetic processes varied by plant type, with C 3 species tending to preferentially accelerate CO 2 -limited photosynthetic processes and respiration and C 4 species tending to preferentially accelerate light-limited photosynthetic processes under warmer conditions. R d acclimation to warmer temperatures caused a reduction in temperature sensitivity that resulted in slower rates at high leaf temperatures. R d acclimation was similar across plant types. These results suggest that temperature acclimation of the biochemical processes that underlie plant carbon exchange is common across different plant types, but that acclimation to warmer temperatures tends to have a relatively greater positive effect on the processes most limiting to carbon assimilation, which differ by plant type. The acclimation responses observed here suggest that warmer conditions should lead to increased rates of carbon assimilation when water and nutrients are not limiting., (© 2017 John Wiley & Sons Ltd.)

To realistically simulate climate feedbacks from the land surface to the atmosphere, models must replicate the responses of plants to environmental changes. Several processes, operating at various scales, cause the responses of photosynthesis and plant respiration to temperature and CO2 to change over time of exposure to new or changing environmental conditions. Here, we review the latest empirical evidence that short-term responses of plant carbon exchange rates to temperature and CO2 are modified by plant photosynthetic and respiratory acclimation as well as biogeochemical feedbacks. We assess the frequency with which these responses have been incorporated into vegetation models, and highlight recently designed algorithms that can facilitate their incorporation. Few models currently include representations of the long-term plant responses that have been recorded by empirical studies, likely because these responses are still poorly understood at scales relevant for models. Studies show that, at a regional scale, simulated carbon flux between the atmosphere and vegetation can dramatically differ between versions of models that do and do not include acclimation. However, the realism of these results is difficult to evaluate, as algorithm development is still in an early stage, and a limited number of data are available. We provide a series of recommendations that suggest how a combination of empirical and modeling studies can produce mechanistic algorithms that will realistically simulate longer term responses within global-scale models., (© 2012 Blackwell Publishing Ltd.)