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The article "Stomata: custodians of leaf gaseous exchange" published in the Journal of Experimental Botany discusses the role of stomata in regulating gaseous exchange in plants. Stomata, surrounded by guard cells, modulate CO2 uptake for photosynthesis while minimizing water loss. The article explores the impact of stomatal conductance on photosynthesis, water use efficiency, and plant performance, emphasizing the potential for crop improvement through enhanced photosynthesis and water use efficiency. It also delves into the manipulation of stomatal anatomical features to improve water use efficiency and plant resilience to climate-induced stresses. [Extracted from the article]

Artificial intelligence and machine learning (AI/ML) can be used to automatically analyze large image datasets. One valuable application of this approach is estimation of plant trait data contained within images. Here we review 39 papers that describe the development and/or application of such models for estimation of stomatal traits from epidermal micrographs. In doing so, we hope to provide plant biologists with a foundational understanding of AI/ML and summarize the current capabilities and limitations of published tools. While most models show human-level performance for stomatal density (SD) quantification at superhuman speed, they are often likely to be limited in how broadly they can be applied across phenotypic diversity associated with genetic, environmental, or developmental variation. Other models can make predictions across greater phenotypic diversity and/or additional stomatal/epidermal traits, but require significantly greater time investment to generate ground-truth data. We discuss the challenges and opportunities presented by AI/ML-enabled computer vision analysis, and make recommendations for future work to advance accelerated stomatal phenotyping. [ABSTRACT FROM AUTHOR]

Enhancing crop water use efficiency (WUE) is a key target trait for climatic resilience and expanding cultivation on marginal lands. Engineering lower stomatal density to reduce stomatal conductance (g s) has improved WUE in multiple C3 crop species. However, reducing g s in C3 species often reduces photosynthetic carbon gain. A different response is expected in C4 plants because they possess specialized anatomy and biochemistry which concentrates CO2 at the site of fixation. This modifies the relationship of photosynthesis (A N) with intracellular CO2 concentration (c i), such that photosynthesis is CO2 saturated and reductions in g s are unlikely to limit A N. To test this hypothesis, genetic strategies were investigated to reduce stomatal density in the C4 crop sorghum. Constitutive expression of a synthetic epidermal patterning factor (EPF) transgenic allele in sorghum led to reduced stomatal densities, reduced g s, reduced plant water use, and avoidance of stress during a period of water deprivation. In addition, moderate reduction in stomatal density did not increase stomatal limitation to A N. However, these positive outcomes were associated with negative pleiotropic effects on reproductive development and photosynthetic capacity. Avoiding pleiotropy by targeting expression of the transgene to specific tissues could provide a pathway to improved agronomic outcomes. [ABSTRACT FROM AUTHOR]

Stomata regulate CO2 and water vapor exchange between leaves and the atmosphere. Stomata are a target for engineering to improve crop intrinsic water use efficiency (iWUE). One example is by expressing genes that lower stomatal density (SD) and reduce stomatal conductance (g sw). However, the quantitative relationship between reduced SD, g sw, and the mechanisms underlying it is poorly understood. We addressed this knowledge gap using low-SD sugarcane (Saccharum spp. hybrid) as a case study alongside a meta-analysis of data from 10 species. Transgenic expression of EPIDERMAL PATTERNING FACTOR 2 from Sorghum bicolor (SbEPF2) in sugarcane reduced SD by 26–38% but did not affect g sw compared with the wild type. Further, no changes occurred in stomatal complex size or proxies for photosynthetic capacity. Measurements of gas exchange at low CO2 concentrations that promote complete stomatal opening to normalize aperture size between genotypes were combined with modeling of maximum g sw from anatomical data. These data suggest that increased stomatal aperture is the only possible explanation for maintaining g sw when SD is reduced. Meta-analysis across C3 dicots, C3 monocots, and C4 monocots revealed that engineered reductions in SD are strongly correlated with lower g sw (r 2=0.60–0.98), but this response is damped relative to the change in anatomy. [ABSTRACT FROM AUTHOR]

Gas exchange measurements enable mechanistic insights into the processes that underpin carbon and water fluxes in plant leaves which in turn inform understanding of related processes at a range of scales from individual cells to entire ecosytems. Given the importance of photosynthesis for the global climate discussion it is important to (a) foster a basic understanding of the fundamental principles underpinning the experimental methods used by the broad community, and (b) ensure best practice and correct data interpretation within the research community. In this review, we outline the biochemical and biophysical parameters of photosynthesis that can be investigated with gas exchange measurements and we provide step‐by‐step guidance on how to reliably measure them. We advise on best practices for using gas exchange equipment and highlight potential pitfalls in experimental design and data interpretation. The Supporting Information contains exemplary data sets, experimental protocols and data‐modelling routines. This review is a community effort to equip both the experimental researcher and the data modeller with a solid understanding of the theoretical basis of gas‐exchange measurements, the rationale behind different experimental protocols and the approaches to data interpretation. Summary statement: This review outlines the fundamental principles underpinning photosynthetic gas exchange and gives guidance on best practice for obtaining and correctly interpreting measurements. It is meant to provide a solid foundation in gas exchange techniques to the novice and act as a point of reference for the experienced researcher. [ABSTRACT FROM AUTHOR]

The deleterious effects of ozone (O3) pollution on crop physiology, yield, and productivity are widely acknowledged. It has also been assumed that C4 crops with a carbon concentrating mechanism and greater water use efficiency are less sensitive to O3 pollution than C3 crops. This assumption has not been widely tested. Therefore, we compiled 46 journal articles and unpublished datasets that reported leaf photosynthetic and biochemical traits, plant biomass, and yield in five C3 crops (chickpea, rice, snap bean, soybean, and wheat) and four C4 crops (sorghum, maize, Miscanthus × giganteus, and switchgrass) grown under ambient and elevated O3 concentration ([O3]) in the field at free-air O3 concentration enrichment (O3-FACE) facilities over the past 20 y. When normalized by O3 exposure, C3 and C4 crops showed a similar response of leaf photosynthesis, but the reduction in chlorophyll content, fluorescence, and yield was greater in C3 crops compared with C4 crops. Additionally, inbred and hybrid lines of rice and maize showed different sensitivities to O3 exposure. This study quantitatively demonstrates that C4 crops respond less to elevated [O3] than C3 crops. This understanding could help maintain cropland productivity in an increasingly polluted atmosphere. [ABSTRACT FROM AUTHOR]

The deleterious effects of ozone (O3) pollution on crop physiology, yield, and productivity are widely acknowledged. It has also been assumed that C4 crops with a carbon concentrating mechanism and greater water use efficiency are less sensitive to O3 pollution than C3 crops. This assumption has not been widely tested. Therefore, we compiled 46 journal articles and unpublished datasets that reported leaf photosynthetic and biochemical traits, plant biomass, and yield in five C3 crops (chickpea, rice, snap bean, soybean, and wheat) and four C4 crops (sorghum, maize, Miscanthus × giganteus, and switchgrass) grown under ambient and elevated O3 concentration ([O3]) in the field at free-air O3 concentration enrichment (O3-FACE) facilities over the past 20 y. When normalized by O3 exposure, C3 and C4 crops showed a similar response of leaf photosynthesis, but the reduction in chlorophyll content, fluorescence, and yield was greater in C3 crops compared with C4 crops. Additionally, inbred and hybrid lines of rice and maize showed different sensitivities to O3 exposure. This study quantitatively demonstrates that C4 crops respond less to elevated [O3] than C3 crops. This understanding could help maintain cropland productivity in an increasingly polluted atmosphere. [ABSTRACT FROM AUTHOR]

Dietary deficiencies of zinc and iron are a substantial global public health problem. An estimated two billion people suffer these deficiencies (1), causing a loss of 63 million life-years annually (2,3). Most of these people depend on [C.sub.3] grains and legumes as their primary dietary source of zinc and iron. Here we report that [C.sub.3] grains and legumes have lower concentrations of zinc and iron when grown under field conditions at the elevated atmospheric C[O.sub.2] concentration predicted for the middle of this century. [C.sub.3] crops other than legumes also have lower concentrations of protein, whereas [C.sub.4] crops seem to be less affected. Differences between cultivars of a single crop suggest that breeding for decreased sensitivity to atmospheric C[O.sub.2] concentration could partly address these new challenges to global health., In the 1990s, several investigators found that elevated atmospheric C[O.sub.2] concentration (hereafter abbreviated to [C[O.sub.2]]) decreased the concentrations of zinc, iron and protein in grains of wheat (4-7), barley (5) [...]

Increased nutrient inputs due to anthropogenic activity are expected to increase primary productivity across terrestrial ecosystems, but changes in allocation aboveground versus belowground with nutrient addition have different implications for soil carbon (C) storage. Thus, given that roots are major contributors to soil C storage, understanding belowground net primary productivity (BNPP) and biomass responses to changes in nutrient availability is essential to predicting carbon–climate feedbacks in the context of interacting global environmental changes. To address this knowledge gap, we tested whether a decade of nitrogen (N) and phosphorus (P) fertilization consistently influenced aboveground and belowground biomass and productivity at nine grassland sites spanning a wide range of climatic and edaphic conditions in the continental United States. Fertilization effects were strong aboveground, with both N and P addition stimulating aboveground biomass at nearly all sites (by 30% and 36%, respectively, on average). P addition consistently increased root production (by 15% on average), whereas other belowground responses to fertilization were more variable, ranging from positive to negative across sites. Site‐specific responses to P were not predicted by the measured covariates. Atmospheric N deposition mediated the effect of N fertilization on root biomass and turnover. Specifically, atmospheric N deposition was positively correlated with root turnover rates, and this relationship was amplified with N addition. Nitrogen addition increased root biomass at sites with low N deposition but decreased it at sites with high N deposition. Overall, these results suggest that the effects of nutrient supply on belowground plant properties are context dependent, particularly with regard to background N supply rates, demonstrating that site conditions must be considered when predicting how grassland ecosystems will respond to increased nutrient loading from anthropogenic activity. [ABSTRACT FROM AUTHOR]

The leaf economics spectrum (LES) describes multivariate correlations in leaf structural, physiological and chemical traits, originally based on diverse C3 species grown under natural ecosystems. However, the specific contribution of C4 species to the global LES is studied less widely. C4 species have a CO2 concentrating mechanism which drives high rates of photosynthesis and improves resource use efficiency, thus potentially pushing them towards the edge of the LES. Here, we measured foliage morphology, structure, photosynthesis, and nutrient content for hundreds of genotypes of the C4 grass Miscanthus× giganteus grown in two common gardens over two seasons. We show substantial trait variations across M.× giganteus genotypes and robust genotypic trait relationships. Compared to the global LES, M.× giganteus genotypes had higher photosynthetic rates, lower stomatal conductance, and less nitrogen content, indicating greater water and photosynthetic nitrogen use efficiency in the C4 species. Additionally, tetraploid genotypes produced thicker leaves with greater leaf mass per area and lower leaf density than triploid genotypes. By expanding the LES relationships across C3 species to include C4 crops, these findings highlight that M.× giganteus occupies the boundary of the global LES and suggest the potential for ploidy to alter LES traits. SUMMARY STATEMENT: There is substantial variation in leaf‐level photosynthesis, N and P content, and leaf mass per unit area among Miscanthus× giganteus genotypes that contributes significantly to deviation in global leaf economics spectrum (LES) trait relationships. C4 species occupy a distinct niche of the LES. [ABSTRACT FROM AUTHOR]

Miscanthus is one of the most promising perennial crops for bioenergy production, with high yield potential and a low environmental footprint. The increasing interest in this crop requires accelerated selection and the development of new screening techniques. New analytical methods that are more accurate and less labor-intensive are needed to better characterize the effects of genetics and the environment on key traits under field conditions. We used persistent multispectral and photogrammetric UAV time-series imagery collected 10 times over the season, together with ground-truth data for thousands of Miscanthus genotypes, to determine the flowering time, culm length, and biomass yield traits. We compared the performance of convolutional neural network (CNN) architectures that used image data from single dates (2D-spatial) versus the integration of multiple dates by 3D-spatiotemporal architectures. The ability of UAV-based remote sensing to rapidly and non-destructively assess large-scale genetic variation in flowering time, height, and biomass production was improved through the use of 3D-spatiotemporal CNN architectures versus 2D-spatial CNN architectures. The performance gains of the best 3D-spatiotemporal analyses compared to the best 2D-spatial architectures manifested in up to 23% improvements in R2, 17% reductions in RMSE, and 20% reductions in MAE. The integration of photogrammetric and spectral features with 3D architectures was crucial to the improved assessment of all traits. In conclusion, our findings demonstrate that the integration of high-spatiotemporal-resolution UAV imagery with 3D-CNNs enables more accurate monitoring of the dynamics of key phenological and yield-related crop traits. This is especially valuable in highly productive, perennial grass crops such as Miscanthus, where in-field phenotyping is especially challenging and traditionally limits the rate of crop improvement through breeding. [ABSTRACT FROM AUTHOR]

Stomata regulate leaf CO2 assimilation (A) and water loss. The Ball–Berry and Medlyn models predict stomatal conductance (gs) with a slope parameter (m or g1) that reflects the sensitivity of gs to A, atmospheric CO2 and humidity, and is inversely related to water use efficiency (WUE). This study addressed knowledge gaps about what the values of m and g1 are in C4 crops under field conditions, as well as how they vary among genotypes and with drought stress. Four inbred maize genotypes were unexpectedly consistent in how m and g1 decreased as water supply decreased. This was despite genotypic variation in stomatal patterning, A and gs. m and g1 were strongly correlated with soil water content, moderately correlated with predawn leaf water potential (Ψpd), but not correlated with midday leaf water potential (Ψmd). This implied that m and g1 respond to long‐term water supply more than short‐term drought stress. The conserved nature of m and g1 across anatomically diverse genotypes and water supplies suggests there is flexibility in structure‐function relationships underpinning WUE. This evidence can guide the simulation of maize gs across a range of water supply in the primary maize growing region and inform efforts to improve WUE. Summary Statement: Parameter values for models simulating stomatal conductance were unexpectedly consistent for anatomically and physiologically diverse genotypes of the model C4 crop maize when they were grown across a range of water supplies in the field. [ABSTRACT FROM AUTHOR]

Stomata play a central role in surface–atmosphere exchange by controlling the flux of water and CO2 between the leaf and the atmosphere. Representation of stomatal conductance (gsw) is therefore an essential component of models that seek to simulate water and CO2 exchange in plants and ecosystems. For given environmental conditions at the leaf surface (CO2 concentration and vapor pressure deficit or relative humidity), models typically assume a linear relationship between gsw and photosynthetic CO2 assimilation (A). However, measurement of leaf‐level gsw response curves to changes in A are rare, particularly in the tropics, resulting in only limited data to evaluate this key assumption. Here, we measured the response of gsw and A to irradiance in six tropical species at different leaf phenological stages. We showed that the relationship between gsw and A was not linear, challenging the key assumption upon which optimality theory is based—that the marginal cost of water gain is constant. Our data showed that increasing A resulted in a small increase in gsw at low irradiance, but a much larger increase at high irradiance. We reformulated the popular Unified Stomatal Optimization (USO) model to account for this phenomenon and to enable consistent estimation of the key conductance parameters g0 and g1. Our modification of the USO model improved the goodness‐of‐fit and reduced bias, enabling robust estimation of conductance parameters at any irradiance. In addition, our modification revealed previously undetectable relationships between the stomatal slope parameter g1 and other leaf traits. We also observed nonlinear behavior between A and gsw in independent data sets that included data collected from attached and detached leaves, and from plants grown at elevated CO2 concentration. We propose that this empirical modification of the USO model can improve the measurement of gsw parameters and the estimation of plant and ecosystem‐scale water and CO2 fluxes. [ABSTRACT FROM AUTHOR]

Previous studies have found that maximum quantum yield of CO2 assimilation (Φ CO2,max,app) declines in lower canopies of maize and miscanthus, a maladaptive response to self-shading. These observations were limited to single genotypes, leaving it unclear whether the maladaptive shade response is a general property of this C4 grass tribe, the Andropogoneae. We explored the generality of this maladaptation by testing the hypothesis that erect leaf forms (erectophiles), which allow more light into the lower canopy, suffer less of a decline in photosynthetic efficiency than drooping leaf (planophile) forms. On average, Φ CO2,max,app declined 27% in lower canopy leaves across 35 accessions, but the decline was over twice as great in planophiles than in erectophiles. The loss of photosynthetic efficiency involved a decoupling between electron transport and assimilation. This was not associated with increased bundle sheath leakage, based on 13C measurements. In both planophiles and erectophiles, shaded leaves had greater leaf absorptivity and lower activities of key C4 enzymes than sun leaves. The erectophile form is considered more productive because it allows a more effective distribution of light through the canopy to support photosynthesis. We show that in sorghum, it provides a second benefit, maintenance of higher Φ CO2,max,app to support efficient use of that light resource. [ABSTRACT FROM AUTHOR]