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Keenan, Trevor F.

Publications and source records attributed to Keenan, Trevor F..

At least 37 records · Page 2

No evidence for a negative effect of growing season photosynthesis on leaf senescence timing

The length of the growing season has a large influence on the carbon, water, and energy fluxes of global terrestrial ecosystems. While there has been mounting evidence of an advanced start of the growing season mostly due to elevated spring air temperatures, the mechanisms that control the end of the growing season (EOS) in most ecosystems remain relatively less well understood. Recently, a strong lagged control of EOS by growing season photosynthesis has been proposed, suggesting that more productive growing seasons lead to an earlier EOS. However, this relationship has not been extensively tested with in-situ observations across a variety of ecosystems. In this work, we use observations from 40 eddy-covariance flux tower sites in temperate and boreal ecosystems in the northern hemisphere with more than 10 years of observations (594 site-years), ground observations of phenology, satellite observations from the Moderate Resolution Imaging Spectroradiometer (MODIS), and three leaf senescence models to test the extent of a relationship between growing season photosynthesis and end of season senescence. The results suggest that there is no significant negative relationship between growing season photosynthesis and observed leaf senescence, flux-inferred EOS estimates, or remotely sensed phenological metrics, in most ecosystems. On the contrary, while we found negative effects of summer air temperatures and autumn vapor pressure deficit on EOS, more productive growing seasons were typically related to a later, not earlier, EOS. Our results challenge recent reports of carry-over effects of photosynthesis on EOS timing, and suggest those results may not hold over a large range of ecosystems.

59 BASIC BIOLOGICAL SCIENCES↗

Exacerbated drought impacts on global ecosystems due to structural overshoot

Vegetation dynamics are affected not only by the concurrent climate but also by memory-induced lagged responses. For example, favourable climate in the past could stimulate vegetation growth to surpass the ecosystem carrying capacity, leaving an ecosystem vulnerable to climate stresses. This phenomenon, known as structural overshoot, could potentially contribute to worldwide drought stress and forest mortality but the magnitude of the impact is poorly known due to the dynamic nature of overshoot and complex influencing timescales. Here, we use a dynamic statistical learning approach to identify and characterize ecosystem structural overshoot globally and quantify the associated drought impacts. We find that structural overshoot contributed to around 11% of drought events during 1981-2015 and is often associated with compound extreme drought and heat, causing faster vegetation declines and greater drought impacts compared to non-overshoot related droughts. The fraction of droughts related to overshoot is strongly related to mean annual temperature, with biodiversity, aridity and land cover as secondary factors. These results highlight the large role vegetation dynamics play in drought development and suggest that soil water depletion due to warming-induced future increases in vegetation could cause more frequent and stronger overshoot droughts.

54 ENVIRONMENTAL SCIENCES↗

Global variation in the fraction of leaf nitrogen allocated to photosynthesis

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.

54 ENVIRONMENTAL SCIENCES↗

Eco-evolutionary optimality as a means to improve vegetation and land-surface models

Global vegetation and land-surface models embody interdisciplinary scientific understanding of the behaviour of plants and ecosystems, and are indispensable to project the impacts of environmental change on vegetation and the interactions between vegetation and climate. Furthermore, systematic errors and persistently large differences among carbon and water cycle projections by different models highlight the limitations of current process formulations. In this review, focusing on core plant functions in the terrestrial carbon and water cycles, we show how unifying hypotheses derived from eco-evolutionary optimality (EEO) principles can provide novel, parameter-sparse representations of plant and vegetation processes. We present case studies that demonstrate how EEO generates parsimonious representations of core, leaf-level processes that are individually testable and supported by evidence. EEO approaches to photosynthesis and primary production, dark respiration and stomatal behaviour are ripe for implementation in global models. EEO approaches to other important traits, including the leaf economics spectrum and applications of EEO at the community level are active research areas. Independently tested modules emerging from EEO studies could profitably be integrated into modelling frameworks that account for the multiple time scales on which plants and plant communities adjust to environmental change.

59 BASIC BIOLOGICAL SCIENCES↗

Midwest US Croplands Determine Model Divergence in North American Carbon Fluxes

Large uncertainties in North American terrestrial carbon fluxes hinder regional climate projections. Terrestrial biosphere models (TBMs), the essential tools for understanding continental-scale carbon cycle, diverge on whether temperate forests or croplands dominate carbon uptake in North America. Evidence from novel photosynthetic proxies, such as those based on chlorophyll fluorescence, has cast doubt on the “weak cropland, strong forest” carbon uptake patterns simulated by most TBMs. However, no systematic evaluation of TBMs has yet been attempted to pin down space-time patterns that are most consistent with regional CO 2 observational constraints. Here, we leverage atmospheric CO 2 observations and satellite-observed photosynthetic proxies to understand emergent space-time patterns in North American carbon fluxes from a large suite of TBMs and data-driven models. To do so, we evaluate how well the atmospheric signals resulting from carbon flux estimates reproduce the space-time variability in atmospheric CO 2 , as is observed by a network of continuous-monitoring towers over North America. Models with gross or net carbon fluxes that are consistent with the observed CO 2 variability share a salient feature of growing-season carbon uptake in Midwest US croplands. Conversely, the remaining models place most growing-season uptake in boreal or temperate forests. Differences in model explanatory power depend mainly on the simulated annual cycles of cropland uptake—especially, the timing of peak uptake—rather than the distribution of annual mean fluxes across biomes. Our results suggest that improved model representation of cropland phenology is crucial to robust, policy-relevant estimation of North American carbon exchange.

54 ENVIRONMENTAL SCIENCES↗

AI-Constrained Bottom-Up Ecohydrology and Improved Prediction of Seasonal, Interannual, and Decadal Flood and Drought Risks

Focal Areas: (2) Predictive modeling through the use of AI techniques and AI-derived model components; the use of AI and other tools to design a prediction system composed of a hierarchy of models (3)Insight gleaned from complex data (both observed & simulated) using AI, big data analytics, and other advanced methods, including explainable AI and physics- or knowledge-guided AI

54 ENVIRONMENTAL SCIENCES↗

A trade-off between plant and soil carbon storage under elevated CO 2

Terrestrial ecosystems remove about 30 per cent of the carbon dioxide (CO 2 ) emitted by human activities each year, yet the persistence of this carbon sink depends partly on how plant biomass and soil organic carbon (SOC) stocks respond to future increases in atmospheric CO 2 . Although plant biomass often increases in elevated CO 2 (eCO 2 ) experiments, SOC has been observed to increase, remain unchanged or even decline. The mechanisms that drive this variation across experiments remain poorly understood, creating uncertainty in climate projections. In this study we synthesized data from 108 eCO 2 experiments and found that the effect of eCO 2 on SOC stocks is best explained by a negative relationship with plant biomass: when plant biomass is strongly stimulated by eCO 2 , SOC storage declines; conversely, when biomass is weakly stimulated, SOC storage increases. This trade-off appears to be related to plant nutrient acquisition, in which plants increase their biomass by mining the soil for nutrients, which decreases SOC storage. We found that, overall, SOC stocks increase with eCO 2 in grasslands (8 ± 2 per cent) but not in forests (0 ± 2 per cent), even though plant biomass in grasslands increase less (9 ± 3 per cent) than in forests (23 ± 2 per cent). Ecosystem models do not reproduce this trade-off, which implies that projections of SOC may need to be revised.

54 ENVIRONMENTAL SCIENCES↗

10 New Insights in Climate Science 2020 - a Horizon Scan

We summarize some of the past year’s most important findings within climate change-related research. New research has improved our understanding of Earth’s sensitivity to carbon dioxide, finds that permafrost thaw could release more carbon emissions than expected and that the uptake of carbon in tropical ecosystems is weakening. Adverse impacts on human society include increasing water shortages and impacts on mental health. Options for solutions emerge from rethinking economic models, rights-based litigation, strengthened governance systems and a new social contract. The disruption caused by COVID-19 could be seized as an opportunity for positive change, directing economic stimulus towards sustainable investments.

29 ENERGY PLANNING, POLICY, AND ECONOMY↗

Improving collaborations between empiricists and modelers to advance grassland community dynamics in ecosystem models

Climate change, increasing atmospheric CO 2 , and land use change have altered biogeochemical and hydrologic cycles world-wide, with grassland systems being particularly vulnerable to resulting vegetation shifts (Komatsu et al., 2019). Therefore, incorporating plant community dynamics into ecosystem models is critical for accurate forecasting of ecosystem responses to global change (Levine, 2016). Process-based ecosystem models, which simulate the biogeochemical transfers of mass and energy among biota, the subsurface, and atmosphere, require representation of dynamic composition of organisms within ecosystems. For example, these models simulate leaf and plant-level characteristics, such as electron transport rate and allometry of carbon (C) allocation, to predict how net primary productivity and other ecosystem processes respond to abiotic drivers. These models are particularly useful in scaling from organismal to ecosystem levels but are still underdeveloped in their ability to capture community change, especially in grassland ecosystems. To represent compositional changes, these models must simulate competition, mortality, establishment, and reproduction of plant populations within communities. Yet, current ecosystem modeling approaches to forecast plant community change have derived from studies of forested systems and are either too coarse to capture fine-scale community dynamics (e.g. dynamic global vegetation models (DGVMs)) or too complex to be used at large spatial scales (e.g. forest gap models).

54 ENVIRONMENTAL SCIENCES↗

Evaluating three evapotranspiration estimates from model of different complexity over China using the ILAMB benchmarking system

The land surface models range in complexity of terrestrial evapotranspiration, yet it is unknown how model complexity translates to accuracy of modeled evapotranspiration estimates. Here, we use the International Land Model Benchmarking system to assess ET estimates from three models of varying complexity driven by the same forcing datasets: an earth system model, a terrestrial biosphere model, and a stand-alone ET model. The performance assessment includes both temporal and spatial evaluation, and different plant functional types across China. Our results indicate that the most complex model, an earth system model, performed best against the benchmarking datasets and metrics. Terrestrial biosphere model performed best in simulating inter-annual variability of ET, while earth system model performed best in simulating the seasonal cycle. The more complex models (earth system model and terrestrial biosphere model) perform better in forest, shrub and crop ecosystems, while the simpler model (stand-alone ET model) perform better in grass ecosystems. Our study demonstrates the impact of model complexity on ET estimates and highlights directions for future ET model improvements.

54 ENVIRONMENTAL SCIENCES↗

Integrating the evidence for a terrestrial carbon sink caused by increasing atmospheric CO 2

Atmospheric carbon dioxide concentration ([CO 2 ]) is increasing, which increases leaf-scale photosynthesis and intrinsic water-use efficiency. These direct responses have the potential to increase plant growth, vegetation biomass, and soil organic matter; transferring carbon from the atmosphere into terrestrial ecosystems (a carbon sink). A substantial global terrestrial carbon sink would slow the rate of [CO 2 ] increase and thus climate change. However, ecosystem CO 2 -responses are complex or confounded by concurrent changes in multiple agents of global change and evidence for a [CO 2 ]-driven terrestrial carbon sink can appear contradictory. In this work, we synthesise theory and broad, multi-disciplinary evidence for the effects of increasing [CO 2 ] (iCO 2 ) on the global terrestrial carbon sink. Evidence suggests a substantial increase in global photosynthesis since pre-industry. Established theory, supported by experiments, indicates that iCO 2 is likely responsible for about half of the increase. Global carbon budgeting, atmospheric data, and forest inventories indicate a historical carbon sink, and these apparent iCO 2 -responses are high in comparison with experiments and theory. Plant mortality and soil carbon iCO 2 -responses are highly uncertain. In conclusion, a range of evidence supports a positive terrestrial carbon sink in response to iCO 2 , albeit with uncertain magnitude and strong suggestion of a role for additional agents of global change.

54 ENVIRONMENTAL SCIENCES↗

Carbon budget of the Harvard Forest Long-Term Ecological Research site: pattern, process, and response to global change

How, where, and why carbon (C) moves into and out of an ecosystem through time are long-standing questions in biogeochemistry. Here, we bring together hundreds of thousands of C-cycle observations at the Harvard Forest in central Massachusetts, USA, a mid-latitude landscape dominated by 80–120-yr-old closed-canopy forests. These data answered four questions: (1) where and how much C is presently stored in dominant forest types; (2) what are current rates of C accrual and loss; (3) what biotic and abiotic factors contribute to variability in these rates; and (4) how has climate change affected the forest’s C cycle? Harvard Forest is an active C sink resulting from forest regrowth following land abandonment. Soil and tree biomass comprise nearly equal portions of existing C stocks. Net primary production (NPP) averaged 680–750 g C·m –2 ·yr –1 ; belowground NPP contributed 38–47% of the total, but with large uncertainty. Mineral soil C measured in the same inventory plots in 1992 and 2013 was too heterogeneous to detect change in soil-C pools; however, radiocarbon data suggest a small but persistent sink of 10–30 g C·m –2 ·yr –1 . Net ecosystem production (NEP) in hardwood stands averaged ~300 g C·m –2 ·yr –1 . NEP in hemlock-dominated forests averaged ~450 g C·m –2 ·yr –1 until infestation by the hemlock woolly adelgid turned these stands into a net C source. Since 2000, NPP has increased by 26%. For the period 1992–2015, NEP increased 93%. The increase in mean annual temperature and growing season length alone accounted for ~30% of the increase in productivity. Interannual variations in GPP and NEP were also correlated with increases in red oak biomass, forest leaf area, and canopy-scale light-use efficiency. Compared to long-term global change experiments at the Harvard Forest, the C sink in regrowing biomass equaled or exceeded C cycle modifications imposed by soil warming, N saturation, and hemlock removal. Furthermore, the results of this synthesis and comparison to simulation models suggest that forests across the region are likely to accrue C for decades to come but may be disrupted if the frequency or severity of biotic and abiotic disturbances increases.

54 ENVIRONMENTAL SCIENCES↗

Global evidence for the acclimation of ecosystem photosynthesis to light

Photosynthesis responds quickly to changes in light, increasing with incoming photosynthetic photon flux density (PPFD) until the leaves become light saturated. This instantaneous response to PPFD, which is widely studied and incorporated into models of photosynthesis, is overlaid on non-instantaneous photosynthetic changes resulting from the acclimation of plants to average PPFD over intermediate timescales of a week to months ($\overline{PPFD}$). Such photosynthetic light acclimation is not typically incorporated into models, due to the lack of observational constraints. In this work, we use eddy covariance observations from globally distributed and automated sensor networks, along with photosynthesis estimates from nine terrestrial biosphere models (TBMs), to quantify and assess photosynthetic acclimation to light in natural environments. We also use recent theoretical developments to incorporate light acclimation in a TBM. Our results show widespread light acclimation of ecosystem photosynthesis. On average, a 1 μmol m -2 s -1 increase in $\overline{PPFD}_{10}$ (ten-day average PPFD) leads to a 0.031 ± 0.013 μmol C m -2 s -1 increase in the maximum photosynthetic assimilation rate (A max ), with croplands having stronger acclimation rates than grasslands and forests. Our analysis shows that the TBMs examined either neglect or substantially underestimate light acclimation. By updating a TBM to include photosynthetic acclimation, successfully reproducing the $\overline{PPFD}_{10}$-A max relationship, we provide a robust method for the incorporation of photosynthetic light acclimation in future models.

59 BASIC BIOLOGICAL SCIENCES↗

Mechanisms underlying leaf photosynthetic acclimation to warming and elevated CO 2 as inferred from least‐cost optimality theory

Abstract 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.

Smith, Nicholas G.↗

Partitioning net carbon dioxide fluxes into photosynthesis and respiration using neural networks

The eddy covariance (EC) technique is used to measure the net ecosystem exchange (NEE) of CO 2 between ecosystems and the atmosphere, offering a unique opportunity to study ecosystem responses to climate change. NEE is the difference between the total CO 2 release due to all respiration processes (RECO), and the gross carbon uptake by photosynthesis (GPP). These two gross CO 2 fluxes are derived from EC measurements by applying partitioning methods that rely on physiologically based functional relationships with a limited number of environmental drivers. However, the partitioning methods applied in the global FLUXNET network of EC observations do not account for the multiple co-acting factors that modulate GPP and RECO flux dynamics. To overcome this limitation, we developed a hybrid data-driven approach based on combined neural networks (NN C-part ). NN C-part incorporates process knowledge by introducing a photosynthetic response based on the light-use efficiency (LUE) concept, and uses a comprehensive dataset of soil and micrometeorological variables as fluxes drivers. We applied the method to 36 sites from the FLUXNET2015 dataset and found a high consistency in the results with those derived from other standard partitioning methods for both GPP (R 2 > .94) and RECO (R 2 > .8). High consistency was also found for (a) the diurnal and seasonal patterns of fluxes and (b) the ecosystem functional responses. NNC-part performed more realistic than the traditional methods for predicting additional patterns of gross CO 2 fluxes, such as: (a) the GPP response to VPD, (b) direct effects of air temperature on GPP dynamics, (c) hysteresis in the diel cycle of gross CO 2 fluxes, (d) the sensitivity of LUE to the diffuse to direct radiation ratio, and (e) the post rain respiration pulse after a long dry period. In conclusion, NN C-part is a valid data-driven approach to provide GPP and RECO estimates and complementary to the existing partitioning methods.

54 ENVIRONMENTAL SCIENCES↗

P-model v1.0: an optimality-based light use efficiency model for simulating ecosystem gross primary production

Terrestrial photosynthesis is the basis for vegetation growth and drives the land carbon cycle. Accurately simulating gross primary production (GPP, ecosystem-level apparent photosynthesis) is key for satellite monitoring and Earth system model predictions under climate change. While robust models exist for describing leaf-level photosynthesis, predictions diverge due to uncertain photosynthetic traits and parameters which vary on multiple spatial and temporal scales. Here, we describe and evaluate a GPP (photosynthesis per unit ground area) model, the P-model, that combines the Farquhar–von Caemmerer–Berry model for C 3 photosynthesis with an optimality principle for the carbon assimilation–transpiration trade-off, and predicts a multi-day average light use efficiency (LUE) for any climate and C 3 vegetation type. The model builds on the theory developed in Prentice et al. (2014) and Wang et al. (2017a) and is extended to include low temperature effects on the intrinsic quantum yield and an empirical soil moisture stress factor. The model is forced with site-level data of the fraction of absorbed photosynthetically active radiation (fAPAR) and meteorological data and is evaluated against GPP estimates from a globally distributed network of ecosystem flux measurements. Although the P-model requires relatively few inputs, the R 2 for predicted versus observed GPP based on the full model setup is 0.75 (8 d mean, 126 sites) – similar to comparable satellite-data-driven GPP models but without predefined vegetation-type-specific parameters. The R 2 is reduced to 0.70 when not accounting for the reduction in quantum yield at low temperatures and effects of low soil moisture on LUE. The R 2 for the P-model-predicted LUE is 0.32 (means by site) and 0.48 (means by vegetation type). Applying this model for global-scale simulations yields a total global GPP of 106–122 Pg C yr –1 (mean of 2001–2011), depending on the fAPAR forcing data. The P-model provides a simple but powerful method for predicting – rather than prescribing – light use efficiency and simulating terrestrial photosynthesis across a wide range of conditions. The model is available as an R package (rpmodel).

54 ENVIRONMENTAL SCIENCES↗