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At least 163 records · Page 9

Reducing Long‐Standing Surface Ozone Overestimation in Earth System Modeling by High‐Resolution Simulation and Dry Deposition Improvement

The overestimation of surface ozone concentration in low‐resolution global atmospheric chemistry and climate models has been a long‐standing issue. We first update the ozone dry deposition scheme in both high‐ (0.25°) and low‐resolution (1°) Community Earth System Model (CESM) version 1.3 runs, by adding the effects of leaf area index and correcting the sunlit and shaded fractions of stomatal resistances. With this update, 5‐year‐long summer simulations (2015–2019) using the low‐resolution CESM still exhibit substantial ozone overestimation (by 6.0–16.2 ppbv) over the U.S., Europe, eastern China, and ozone pollution hotspots. The ozone dry deposition scheme is further improved by adjusting the leaf cuticle conductance, reducing the mean ozone bias by 19%, and increasing the model resolution further reduces the ozone overestimation by 43%. We elucidate the mechanism by which model grid spacing influences simulated ozone, revealing distinctive pathways in urban versus rural areas. In rural areas, grid spacing mainly affects daytime ozone levels, where additional NO x emissions from nearby urban areas result in an ozone boost and overestimation in low‐resolution simulations. In contrast, over urban areas, daytime ozone overestimation follows a similar mechanism due to the influence of volatile organic compounds from surrounding rural areas. However, nighttime ozone overestimation is closely linked to weakened NO titration owing to the redistribution of urban NO x to rural areas. Additionally, stratosphere‐troposphere exchange may also contribute to reducing ozone bias in high‐resolution simulations, warranting further investigation. This optimized high‐resolution CESM may enhance understanding of ozone formation mechanisms, sources, and changes in a warming climate.

54 ENVIRONMENTAL SCIENCES↗

Explicit Consideration of Plant Xylem Hydraulic Transport Improves the Simulation of Crop Response to Atmospheric Dryness in the U.S. Corn Belt

Abstract Atmospheric dryness (i.e., high vapor pressure deficit, VPD), together with soil moisture stress, limits plant photosynthesis and threatens ecosystem functioning. Regions where rainfall and soil moisture are relatively sufficient, such as the rainfed part of the U.S. Corn Belt, are especially prone to high VPD stress. With globally projected rising VPD under climate change, it is crucial to understand, simulate, and manage its negative impacts on agricultural ecosystems. However, most existing models simulating crop response to VPD are highly empirical and insufficient in capturing plant response to high VPD, and improved modeling approaches are urgently required. In this study, by leveraging recent advances in plant hydraulic theory, we demonstrate that the VPD constraints in the widely used coupled photosynthesis‐stomatal conductance models alone are inadequate to fully capture VPD stress effects. Incorporating plant xylem hydraulic transport significantly improves the simulation of transpiration under high VPD, even when soil moisture is sufficient. Our results indicate that the limited water transport capability from the plant root to the leaf stoma could be a major mechanism of plant response to high VPD stress. We then introduce a Demand‐side Hydraulic Limitation Factor (DHLF) that simplifies the xylem and the leaf segments of the plant hydraulic model to only one parameter yet captures the effect of plant hydraulic transport on transpiration response to high VPD with similar accuracy. We expect the improved understanding and modeling of crop response to high VPD to help contribute to better management and adaptation of agricultural systems in a changing climate.

Environmental Sciences & Ecology↗

A Vertically Resolved Canopy Improves Chemical Transport Model Predictions of Ozone Deposition to North Temperate Forests

Abstract Dry deposition is the second largest tropospheric ozone (O 3 ) sink and occurs through stomatal and nonstomatal pathways. Current O 3 uptake predictions are limited by the simplistic big‐leaf schemes commonly used in chemical transport models (CTMs) to parameterize deposition. Such schemes fail to reproduce observed O 3 fluxes over terrestrial ecosystems, highlighting the need for more realistic treatment of surface‐atmosphere exchange in CTMs. We address this need by linking a resolved canopy model (1D Multi‐Layer Canopy CHemistry and Exchange Model, MLC‐CHEM) to the GEOS‐Chem CTM and use this new framework to simulate O 3 fluxes over three north temperate forests. We compare results with in situ measurements from four field studies and with standalone, observationally constrained MLC‐CHEM runs to test current knowledge of O 3 deposition and its drivers. We show that GEOS‐Chem overpredicts observed O 3 fluxes across all four studies by up to 2×, whereas the resolved‐canopy models capture observed diel profiles of O 3 deposition and in‐canopy concentrations to within 10%. Relative humidity and solar irradiance are strong O 3 flux drivers over these forests, and uncertainties in those fields provide the largest remaining source of model deposition biases. Flux partitioning analysis shows that: (a) nonstomatal loss accounts for 60% of O 3 deposition on average; (b) in‐canopy chemistry makes only a small contribution to total O 3 fluxes; and (c) the CTM big‐leaf treatment overestimates O 3 ‐driven stomatal loss and plant phytotoxicity in these temperate forests by up to 7×. Results motivate the application of fully online vertically explicit canopy schemes in CTMs for improved O 3 predictions.

Vermeuel, Michael P. [Department of Soil, Water, a↗

Plant‐Induced Changes Mediate Belowground Carbon Cycling in an Experimentally Warmed Peatland

Warming and elevated atmospheric CO 2 profoundly impact peatland ecosystems, particularly through changes in plant species composition. Plants regulate the initial input of organic compounds to peatland belowground systems, controlling the availability of electron donors and electron acceptors that fuel microbially mediated organic matter decomposition to CO 2 and CH 4 . However, explicit links between porewater CO 2 and CH 4 dynamics and plant-derived chemical compounds remain relatively undefined. In a whole ecosystem warming experiment, we investigated how warming affects plant leaf chemical composition and species assemblages, and how the alteration of leaf-derived organic compounds supplied to the subsurface impacts belowground CO 2 and CH 4 production. While earlier studies at our site found no temperature-dependent changes in CH 4 production pathways, our extended timeseries has revealed increased acetoclastic methanogenesis at higher temperatures in certain peat depths, correlated with elevated porewater phenolics. These changes appear driven by the observed increased plant productivity and altered vegetation inputs, which accelerate decomposition and fuel CH 4 production through enhanced substrate availability. In conclusion, we observed warming-induced changes in molecular composition both between and within plant species, suggesting that plant-mediated controls on belowground carbon processing are more complex than previously recognized.

Wilson, Rachel M. [Florida State Univ., Tallahasse↗

Experimental Soil Warming Impacts Soil Moisture and Plant Water Stress and Thereby Ecosystem Carbon Dynamics

Experimental soil heating experiments have found a consistent increase in soil-surface CO 2 emissions ( F s ), but inconsistent soil organic carbon (SOC) responses. Interpretation of heating effects is complicated by spatial heterogeneity and soil moisture, nitrogen availability, and microbial and plant responses. Here we applied a mechanistic ecosystem model to interpret heating impacts on a California forest subjected to 1 m deep, 4°C heating. The model accurately simulated control-plot CO 2 fluxes, SOC stocks, fine root biomass, soil moisture, and soil temperature, and the observed increases in F s and decreases in fine root biomass. We show that a complex suite of interactions can lead to a consistent increase in F s (~17%) over the 5-year study period, with very small changes in SOC stocks (<1%). Modeled increases in leaf water stress from soil drying reduced GPP and NPP. The resulting reduction in leaf and fine root allocation increased fine root litter inputs to the soil and reduced root exudation. Soil heating led to about a 50% larger increase in root autotrophic respiration than in heterotrophic respiration, with the heating effect on both these fluxes decreasing over the simulation period. Increased heterotrophic respiration led to increased soil N availability and plant N uptake. These heating responses are mechanistically linked, of magnitudes that can affect ecosystem dynamics, and long-term observations of them are rarely made. Therefore, we conclude that a coupled observational and mechanistic modeling framework is needed to interpret manipulation experiments, and to improve projections of climate change impacts on terrestrial ecosystem carbon dynamics.

54 ENVIRONMENTAL SCIENCES↗

Investigating the Global Biogeophysical Impact of Area and Mass Based Wood Harvest in a Vegetation Demography Model

Wood harvesting alters land surface properties and energy redistribution, but there is a lack of studies estimating these changes on a global scale. We coupled a vegetation demographic model, the Functionally Assembled Terrestrial Ecosystem Simulator, with the E3SM land model to perform offline model simulation to investigate the land biogeophysical responses, including canopy coverage, leaf area index, albedo, surface roughness length, and energy fluxes, to historical wood harvest on the global scale. In this study, we found 50% less harvested carbon (C) when choosing the area-based harvest rate as driving data that has not been spatially harmonized, compared to reharmonized mass-based harvesting. By considering the uncertainty from reconstruction of historical wood harvest time series and the choice of wood harvest approach in the model, continuous wood harvest (1850–2015) results in 5%–10% of canopy coverage loss, contributing 0.5%–1% increase of albedo over disturbed land, which is much stronger than a non-demographic land surface model. Changes in energy flux from the wood harvest are negligible (<1%), but the responses of land surface properties vary (up to 30%) due to differences in model structure between the single canopy, sun-shade leaf model and vegetation demographic model.

Shu, Shijie [Lawrence Berkeley National Laboratory↗

Evidence for widespread thermal acclimation of canopy photosynthesis

Plants acclimate to temperature by adjusting their photosynthetic capacity over weeks to months. However, most evidence for photosynthetic acclimation derives from leaf-scale experiments. Here we address the scarcity of evidence for canopy-scale photosynthetic acclimation by examining the correlation between maximum photosynthetic rates (A max,2,000 ) and growth temperature $(\overline{T_{air}})$ across a range of concurrent temperatures and canopy foliage quantity, using data from >200 eddy covariance sites. We detect widespread thermal acclimation of canopy-scale photosynthesis, demonstrated by enhanced A max,2,000 under higher $\overline{T_{air}}$, across flux sites with adequate water availability. A 14-day period is identified as the most relevant timescale for acclimation across all sites, with a range of 12-25 days for different plant functional types. The mean apparent thermal acclimation rate across all ecosystems is 0.41 (-0.38-1.04 for 5th-95th percentile range) µmol m -2 s -1 °C -1 , with croplands showing the largest acclimation rates and grasslands the lowest. Incorporating an optimality-based prediction of leaf photosynthetic capacities into a biochemical photosynthesis model is shown to improve the representation of thermal acclimation. Our results underscore the critical need for enhanced understanding and modelling of canopy-scale photosynthetic capacity to accurately predict plant responses to warmer growing seasons.

59 BASIC BIOLOGICAL SCIENCES↗

The ‘photosynthetic C 1 pathway’ links carbon assimilation and growth in California poplar

Although primarily studied in relation to photorespiration, serine metabolism in chloroplasts may play a key role in plant CO 2 fertilization responses by linking CO 2 assimilation with growth. Here, we show that the phosphorylated serine pathway is part of a 'photosynthetic C 1 pathway' and demonstrate its high activity in foliage of a C 3 tree where it rapidly integrates photosynthesis and C 1 metabolism contributing to new biomass via methyl transfer reactions, imparting a large natural 13 C-depleted signature. Using 13 CO 2 -labelling, we show that leaf serine, the S-methyl group of leaf methionine, pectin methyl esters, and the associated methanol released during cell wall expansion during growth, are directly produced from photosynthetically-linked C 1 metabolism, within minutes of light exposure. We speculate that the photosynthetic C 1 pathway is highly conserved across the photosynthetic tree of life, is responsible for synthesis of the greenhouse gas methane, and may have evolved with oxygenic photosynthesis by providing a mechanism of directly linking carbon and ammonia assimilation with growth. Although the rise in atmospheric CO 2 inhibits major metabolic pathways like photorespiration, our results suggest that the photosynthetic C 1 pathway may accelerate and represents a missing link between enhanced photosynthesis and plant growth rates during CO 2 fertilization under a changing climate.

59 BASIC BIOLOGICAL SCIENCES↗

Forest residue harvest optimization: spanning the bridge between plant biology and biorefinery performance

Forestry residues have immense potential as alternative feedstocks to petroleum, yet their inherent complexity remains a major challenge to widespread use. Pairing the temporal rhythms of plant biology with biorefinery performance is critical to industrial-scale biorefinery development. Here, we provide the first report of a techno-economic analysis (TEA) and life cycle assessment (LCA) for a model integrated reductive catalytic fractionation (RCF)–molten salt hydrolysis process for forestry residues varying in tree part, species, and phenophase. All forestry residues resulted in net-negative greenhouse gas (GHG) emissions vs. comparable petroleum feedstocks, with GHG emissions potentially reduced >4.0× through composition-based feedstock selection (e.g., harvesting American beech bark in spring vs. summer). Moreover, American beech twigs/branchlets and bark in leafed and emergence phenophases, respectively, had 7.9× lower predicted phenolic minimum selling prices (MSPs) vs. other feedstocks and MSPs within the current global phenolic market range. Hemicellulose content and RCF yield emerged as key parameters impacting GHG emissions and biorefinery revenue, identifying hardwood twigs/branchlets in the leafed phenophase as optimal biofeedstocks. Biorefinery expenses were dominated by purchased equipment, raw materials, and utility costs, highlighting essential areas for future study. Notably, RCF reactor pressures drove 85–90% of equipment costs, but sensitivity analysis revealed that decreasing the pressure 20% could reduce the phenol MSP 4-fold. Structural carbohydrate dynamics were also investigated using a two-step acid hydrolysis method to resolve tissue- and species-level patterns in biomass composition throughout the year to enable harvest optimization based on TEA/LCA findings. Ultimately, elucidating the impact of biofeedstock dynamics on biorefinery performance enables harvest optimization, informed engineering design, and progress towards an expanded bioeconomy.

Shapiro, Alison J. [University of Delaware, Newark↗

Cooling outweighs warming across phenological transitions in the Northern Hemisphere

Vegetation phenology, i.e., seasonal biological events such as leaf-out and leaf-fall, regulates local climate through biophysical processes like evapotranspiration (ET) and albedo. However, the net surface temperature impact of these processes—whether ET cooling or albedo-induced warming predominates—and how the dominance changes across phenological transitions and regions remains poorly understood. Here, we investigated the effects of vegetation foliage on daytime land surface temperature (LST) following six phenological transitions, spanning from the start of season to end of season, in deciduous and mixed forests across the mid- to high-latitude Northern Hemisphere during 2013–2021 using multiple satellite products and ground observations. We quantified vegetation effect as the difference between observed LST and LST estimates from the Annual Temperature Cycle (ATC) model, representing a no-foliage scenario. We found that vegetation-induced cooling consistently outweighs warming following all phenological transitions except for the end of the season. Cooling intensity increased with vegetation greenness, ranging from 1.0 ± 0.5 °C (mean ± 0.15 SD) in 59% of forests after the start of the season (SOS) to 6.1 ± 0.8 °C in 89% of forests following the onset of maturity, before declining toward the end of the season. Over half of the regions experiencing cooling showed intensification of surface cooling with climate warming, suggesting an amplified vegetation-mediated cooling under future climate change. The findings provide a more precise understanding of the role of vegetation in modulating climate at the intraseasonal scale, highlighting the importance of integrating phenological impacts into climate adaptation strategies and Earth system modeling.

albedo↗

Divergent trait controls on soluble sugars and starch underlie global strategies of tree carbohydrate storage

Nonstructural carbohydrate (NSC) stores buffer tree metabolism, osmotic regulation, and defense, thereby mediating tolerance and survival under climate extremes. Yet, the functional and evolutionary determinants of interspecific variation in NSC remain elusive, limiting understanding and prediction of forest carbon allocation and mortality under global change. Here, we present a cross-species synthesis of NSC concentrations across multiple organs for 281 woody species from 102 mixed forest communities worldwide, where we quantified species-specific deviations from community means to disentangle intrinsic trait effects from environmental and methodological variation. We found phylogenetic signals in NSC deviations, with coniferous gymnosperms and evergreen species consistently maintaining lower stem soluble sugars and starch concentrations than co-occurring angiosperms and deciduous species, respectively. A global pattern emerged where greater stomatal sensitivity to leaf water potential was associated with declines in the relative concentrations of both sugars and starch. In contrast, xylem hydraulic safety traits showed weak and organ-dependent relationships with NSC concentrations. Sugars increased with photosynthetic capacity and declined with wood density, whereas starch showed the reverse pattern, which aligned with the distinct functional-metabolic roles of sugars and starch. By integrating trait-based ecology with a community-centered framework, our study provides global evidence that stomatal regulation, photosynthetic capacity, specific leaf area, and wood density jointly govern interspecific NSC variation, through contrasting effects on sugars and starch. These are among the most broadly measured traits globally, thus the emergent carbohydrate–trait relationships can have broad applications toward understanding and predicting forest growth and survival under climate change.

tropic system↗

Changes in high-latitude surface energy balance driven by snowpack and vegetation dynamics under warmer climate

With rapid climate warming, expected changes in snowpack and vegetation will alter the seasonal surface albedo of high-latitude ecosystems. The extent to which these albedo changes may affect surface energy balances and thus soil temperatures is uncertain, but represents a potentially important component of ecosystem feedbacks to climate change. Here, we apply a well-tested process-rich ecosystem model, ecosys , to examine changes in seasonal surface albedo and soil temperature driven by climate-induced snowpack and vegetation changes across Alaska under a warmer twenty-first century climate. Under the Representative Concentration Pathway 8.5 climate change scenario, the modeled changes in surface albedo exhibited large seasonal and spatial variations. We found spring albedo decreases driven by increases in snow-free periods (>20 d) and an extended growing season length that resulted in greater gains in leaf area index (LAI) in most parts of Alaska. In contrast, we modeled increases in summer and winter albedo (despite modeled increases in LAI) across much of the boreal forest due to an increased proportion of aspen, which has a higher leaf albedo than the currently dominant black spruce. Modeled latent heat fluxes generally increase across the twenty-first century, particularly during the spring and summer. Overall, climate warming and changes in surface energy fluxes resulted in a 3.5 ± 0.50 °C increase in spatial- and annual-averaged top 10 cm surface soil temperatures across Alaskan ecosystems by the year 2100, with larger increases in tundra than boreal forest regions. We conclude that under warmer climates, seasonal variations in albedo and surface energy fluxes are particularly pronounced during the spring and summer, driven by changes in snowpack and vegetation dynamics.

54 ENVIRONMENTAL SCIENCES↗

Thermal stress in degraded forests in the Brazilian Amazon Arc of Deforestation

Understanding thermal stress in tropical forests has taken on new urgency in light of accelerating climate change and expansion of deforestation and forest degradation. Degraded tropical forests in particular may be approaching critical temperature thresholds even more rapidly than intact forests, with implications for tree survival and ecosystem recovery. We investigate thermal stress in degraded tropical forests within the Brazilian Amazon Arc of Deforestation. Using land surface temperature data from the ECOsystem Spaceborne Thermal Radiometer Experiment on the international Space Station (ECOSTRESS), we compared canopy temperatures of intact, selectively logged, and burned forests in Feliz Natal, Mato Grosso, Brazil. Upper canopy temperatures in previously burned forests were 4.1% higher (mean = 36.5 °C) and 50.9% more variable compared to intact and logged forests, which showed remarkably similar temperature distributions (means of 34.9 °C and 35.1 °C, respectively). Modeled leaf temperature distributions based on canopy temperature measurements from one of the warmest days in a 2 year dry season record indicated that 87% of leaves in the warmest burned forest patches exceeded the temperature threshold where respiration surpasses photosynthesis, compared to approximately 72%–74% in intact and logged forests, respectively, in the same time period. After controlling for environmental factors, burned forests were predicted to be 2.6 °C warmer on average [1.39 °C–3.96 °C, 95% credible interval] than intact forests across a 5–40 m canopy height range. Burned forests showed modest thermal recovery over time, with temperatures decreasing by approximately 1.2 °C over a 30 year recovery period. In contrast, logged forests showed minimal thermal differences from intact forests (−0.353 on average, [−1.125–0.274, 95% credible interval]) and negligible change (0.15 °C) over the same timeframe. While the absolute probabilities of exceeding damaging thermal thresholds remain low across all forest types under current climate conditions, the probability of leaves exceeding temperatures that cause permanent leaf damage was ten times higher in burned forests, with implications for the future of burned forest regeneration in water-limited regions of the Amazon basin. In particular, the combination of higher mean temperatures, greater temperature variability, and more frequent exposure to damaging thermal thresholds implies that burned tropical forests will experience substantially higher mortality rates and slower biomass recovery compared to intact and selectively logged forests, especially in water-limited regions where trees cannot rely on evaporative cooling to moderate canopy temperatures.

Brazilian Amazon Arc of Deforestation↗

Observations and modeling reveal that heatwaves reduce photosynthesis, plant carbon reserves, and net carbon uptake

Heatwaves threaten ecosystem carbon balances, yet the mechanisms driving short-term carbon flux responses remain poorly understood. Here, integrating high-frequency eddy covariance (EC) data from 140 global flux tower sites (872 site-years) with detailed process-based modeling, we examine ecosystem responses during and immediately after heatwaves. We show that heatwaves caused a −40% (range [−29%, −128%]) reduction in net ecosystem productivity (NEP) compared to pre-heatwave values, with this reduction persisting over the following two weeks (−38% range [+3%, −154%]). We attributed NEP decreases to photosynthesis decreases more than to ecosystem respiration (RE) increases. Forest sites had greater NEP decreases during heatwaves than non-forest sites, but remained carbon sinks afterwards, indicating resilience. Our modeling analysis of extreme heatwaves at selected EC sites shows that decreased photosynthesis, increased maintenance respiration, and decreased plant non-structural carbon reserves during heatwaves drive carbon cycle changes that persist for weeks. Consistent with phenocam observations, we modeled a reduction in leaf area index caused by reduced non-structural carbon reserves, leading to early leaf senescence and longer-term impacts. Ongoing increases in heatwaves are therefore likely to reduce NEP across a range of ecosystems, exacerbating carbon cycle feedback.

carbon cycle↗

Identification of candidate host-specificity genes in Exserohilum turcicum using comparative genomics and transcriptomics

Abstract Exserohilum turcicum causes northern corn leaf blight and sorghum leaf blight. While the same species cause disease in both crops, the strains are host-specific. Here, we report the sequence and de novo annotated assemblies of one sorghum- and one maize-specific E. turcicum strain. The strains were sequenced using the PacBio Sequel II system. The total genome length for both assemblies was between 44 and 45 Mb with N50 of ∼2.5 Mb. Ninety-eight percent of the Benchmarking Universal Single-Copy Orthologs (BUSCO) for both assemblies had complete status. The estimated number of genes was 11,762 and 12,029 in the sorghum- and maize-specific isolates, respectively. Funannotate, EffectorP, SignalP, and transcriptome data were used to create functional annotation of each genome. The whole-genome comparison identified ten large-scale inversions and three translocations between the maize- and sorghum-specific strains, along with homologous genes and gene duplications. RNA was sequenced from the maize- and sorghum-specific isolate 10 days post-inoculation in maize and sorghum and from axenic cultures. Gene expression data from planta and axenic growth experiments were compared for each strain. Candidate host-specificity genes were identified by combining results from whole-genome comparison, synteny analysis, gene annotations, and transcriptome data. Overall, this study identified several candidate host-specificity genes that provide insights into E. turcicum interaction with its hosts.

Krone, Mara J. (ORCID:0000000159006624)↗

The Ecosystem as Super-Organ/ism, Revisited: Scaling Hydraulics to Forests under Climate Change

Synopsis Classic debates in community ecology focused on the complexities of considering an ecosystem as a super-organ or organism. New consideration of such perspectives could clarify mechanisms underlying the dynamics of forest carbon dioxide (CO2) uptake and water vapor loss, important for predicting and managing the future of Earth’s ecosystems and climate system. Here, we provide a rubric for considering ecosystem traits as aggregated, systemic, or emergent, i.e., representing the ecosystem as an aggregate of its individuals or as a metaphorical or literal super-organ or organism. We review recent approaches to scaling-up plant water relations (hydraulics) concepts developed for organs and organisms to enable and interpret measurements at ecosystem-level. We focus on three community-scale versions of water relations traits that have potential to provide mechanistic insight into climate change responses of forest CO2 and H2O gas exchange and productivity: leaf water potential (Ψcanopy), pressure volume curves (eco-PV), and hydraulic conductance (Keco). These analyses can reveal additional ecosystem-scale parameters analogous to those typically quantified for leaves or plants (e.g., wilting point and hydraulic vulnerability) that may act as thresholds in forest responses to drought, including growth cessation, mortality, and flammability. We unite these concepts in a novel framework to predict Ψcanopy and its approaching of critical thresholds during drought, using measurements of Keco and eco-PV curves. We thus delineate how the extension of water relations concepts from organ- and organism-scales can reveal the hydraulic constraints on the interaction of vegetation and climate and provide new mechanistic understanding and prediction of forest water use and productivity.

Zoology↗

A family of α/β hydrolases removes phytol from chlorophyll metabolites for tocopherol biosynthesis in Arabidopsis

Tocopherol synthesis requires phytyl diphosphate derived from phytol esterified to chlorophyll metabolites. The >600-member Arabidopsis thaliana α/β hydrolase (ABH) gene family contains 4 members that can release phytol from chlorophyll metabolites in vitro; however, only pheophytinase (PPH) affects tocopherol synthesis when mutated, reducing seed tocopherols by 5%. We report the biochemical analysis of 2 previously uncharacterized ABHs, chlorophyll dephytylase 2 (CLD2) and CLD3, and their respective mutants singly and in combinations with pph and cld1 alleles. While all CLDs localized to the thylakoid and could hydrolyze phytol from chlorophylls and Pheophytin a in vitro, CLD3 had the highest in vitro activity and the largest effect on tocopherol synthesis in vivo. The 3 CLDs acted cooperatively to provide phytol for 31% of tocopherols synthesized in light-grown leaf tissue. Dark-induced leaf senescence assays showed PPH is required for 18% of the tocopherols synthesized. Though the cld123 triple mutant had no impact on dark-induced tocopherol content, cld123 in the pph background reduced tocopherol levels by an additional 18%. In seeds, pph and cld123 each reduced tocopherol content by 5% and by 15% in the cld123pph quadruple mutant. VTE7 ( ViTamin E7 ) is an envelope-localized ABH that specifically affects chlorophyll biosynthetic intermediates in vivo and is required for 55% of seed tocopherol synthesis. The introduction of cld123pph into the vte7 background further reduced seed tocopherol levels to 23% of that of the wild type. Our findings demonstrate that phytol provision for tocopherol biosynthesis and homeostasis is a complex process involving the coordinated spatiotemporal expression of multiple ABH family members.

arabidopsis↗

CO2 response screen in grass Brachypodium reveals the key role of a MAP kinase in CO2-triggered stomatal closure

Abstract Plants respond to increased CO2 concentrations through stomatal closure, which can contribute to increased water use efficiency. Grasses display faster stomatal responses than eudicots due to dumbbell-shaped guard cells flanked by subsidiary cells working in opposition. However, forward genetic screening for stomatal CO2 signal transduction mutants in grasses has yet to be reported. The grass model Brachypodium distachyon is closely related to agronomically important cereal crops, sharing largely collinear genomes. To gain insights into CO2 control mechanisms of stomatal movements in grasses, we developed an unbiased forward genetic screen with an EMS-mutagenized B. distachyon M5 generation population using infrared imaging to identify plants with altered leaf temperatures at elevated CO2. Among isolated mutants, a “chill1” mutant exhibited cooler leaf temperatures than wild-type Bd21-3 parent control plants after exposure to increased CO2. chill1 plants showed strongly impaired high CO2-induced stomatal closure despite retaining a robust abscisic acid-induced stomatal closing response. Through bulked segregant whole-genome sequencing analyses followed by analyses of further backcrossed F4 generation plants and generation and characterization of sodium azide and CRISPR-cas9 mutants, chill1 was mapped to a protein kinase, Mitogen-Activated Protein Kinase 5 (BdMPK5). The chill1 mutation impaired BdMPK5 protein-mediated CO2/HCO3− sensing together with the High Temperature 1 (HT1) Raf-like kinase in vitro. Furthermore, AlphaFold2-directed structural modeling predicted that the identified BdMPK5-D90N chill1 mutant residue is located at the interface of BdMPK5 with the BdHT1 Raf-like kinase. BdMPK5 is a key signaling component that mediates CO2-induced stomatal movements and is proposed to function as a component of the primary CO2 sensor in grasses.

Lopez, Bryn N. K. (ORCID:0009000937288216)↗