Maximum leaf conductance driven by CO2 effects on stomatal size and density over geologic time.
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ABSTRACT:
SUBMITTER: Franks PJ
PROVIDER: S-EPMC2693183 | biostudies-literature |
REPOSITORIES: biostudies-literature
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Project description:A principle response of C3 plants to increasing concentrations of atmospheric CO(2) (CO(2)) is to reduce transpirational water loss by decreasing stomatal conductance (g(s)) and simultaneously increase assimilation rates. Via this adaptation, vegetation has the ability to alter hydrology and climate. Therefore, it is important to determine the adaptation of vegetation to the expected anthropogenic rise in CO(2). Short-term stomatal opening-closing responses of vegetation to increasing CO(2) are described by free-air carbon enrichments growth experiments, and evolutionary adaptations are known from the geological record. However, to date the effects of decadal to centennial CO(2) perturbations on stomatal conductance are still largely unknown. Here we reconstruct a 34% (±12%) reduction in maximum stomatal conductance (g(smax)) per 100 ppm CO(2) increase as a result of the adaptation in stomatal density (D) and pore size at maximal stomatal opening (a(max)) of nine common species from Florida over the past 150 y. The species-specific g(smax) values are determined by different evolutionary development, whereby the angiosperms sampled generally have numerous small stomata and high g(smax), and the conifers and fern have few large stomata and lower g(smax). Although angiosperms and conifers use different D and a(max) adaptation strategies, our data show a coherent response in g(smax) to CO(2) rise of the past century. Understanding these adaptations of C3 plants to rising CO(2) after decadal to centennial environmental changes is essential for quantification of plant physiological forcing at timescales relevant for global warming, and they are likely to continue until the limits of their phenotypic plasticity are reached.
| S-EPMC3054011 | biostudies-literature
Project description:Leaf gas exchange is influenced by stomatal size, density, distribution between the leaf adaxial and abaxial sides, as well as by pore dimensions. This study aims to quantify which of these traits mainly underlie genetic differences in operating stomatal conductance (gs) and addresses possible links between anatomical traits and regulation of pore width.Stomatal responsiveness to desiccation, gs-related anatomical traits of each leaf side and estimated gs (based on these traits) were determined for 54 introgression lines (ILs) generated by introgressing segments of Solanum pennelli into the S. lycopersicum 'M82'. A quantitative trait locus (QTL) analysis for stomatal traits was also performed.A wide genetic variation in stomatal responsiveness to desiccation was observed, a large part of which was explained by stomatal length. Operating gs ranged over a factor of five between ILs. The pore area per stomatal area varied 8-fold among ILs (2-16 %), and was the main determinant of differences in operating gs between ILs. Operating gs was primarily positioned on the abaxial surface (60-83 %), due to higher abaxial stomatal density and, secondarily, to larger abaxial pore area. An analysis revealed 64 QTLs for stomatal traits in the ILs, most of which were in the direction of S. pennellii.The data indicate that operating and maximum gs of non-stressed leaves maintained under stable conditions deviate considerably (by 45-91 %), because stomatal size inadequately reflects operating pore area (R(2)?=?0·46). Furthermore, it was found that variation between ILs in both stomatal sensitivity to desiccation and operating gs is associated with features of individual stoma. In contrast, genotypic variation in gs partitioning depends on the distribution of stomata between the leaf adaxial and abaxial epidermis.
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Project description:Background and aimsStomatal density (SD) generally decreases with rising atmospheric CO2 concentration, Ca. However, SD is also affected by light, air humidity and drought, all under systemic signalling from older leaves. This makes our understanding of how Ca controls SD incomplete. This study tested the hypotheses that SD is affected by the internal CO2 concentration of the leaf, Ci, rather than Ca, and that cotyledons, as the first plant assimilation organs, lack the systemic signal.MethodsSunflower (Helianthus annuus), beech (Fagus sylvatica), arabidopsis (Arabidopsis thaliana) and garden cress (Lepidium sativum) were grown under contrasting environmental conditions that affected Ci while Ca was kept constant. The SD, pavement cell density (PCD) and stomatal index (SI) responses to Ci in cotyledons and the first leaves of garden cress were compared. (13)C abundance (?(13)C) in leaf dry matter was used to estimate the effective Ci during leaf development. The SD was estimated from leaf imprints.Key resultsSD correlated negatively with Ci in leaves of all four species and under three different treatments (irradiance, abscisic acid and osmotic stress). PCD in arabidopsis and garden cress responded similarly, so that SI was largely unaffected. However, SD and PCD of cotyledons were insensitive to Ci, indicating an essential role for systemic signalling.ConclusionsIt is proposed that Ci or a Ci-linked factor plays an important role in modulating SD and PCD during epidermis development and leaf expansion. The absence of a Ci-SD relationship in the cotyledons of garden cress indicates the key role of lower-insertion CO2 assimilation organs in signal perception and its long-distance transport.
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Project description:Background and Aims:Studies have indicated that plant stomatal conductance (gs) decreases in response to elevated atmospheric CO2, a phenomenon of significance for the global hydrological cycle. However, gs increases across certain CO2 ranges have been predicted by optimization models. The aim of this work was to demonstrate that under certain environmental conditions, gs can increase in response to elevated CO2. Methods:Using (1) an extensive, up-to-date synthesis of gs responses in free air CO2 enrichment (FACE)experiments, (2) in situ measurements across four biomes showing dynamic gs responses to a CO2 rise of ~50 ppm (characterizing the change in this greenhouse gas over the past three decades) and (3) a photosynthesis-stomatal conductance model, it is demonstrated that gs can in some cases increase in response to increasing atmospheric CO2. Key Results:Field observations are corroborated by an extensive synthesis of gs responses in FACE experiments showing that 11.8 % of gs responses under experimentally elevated CO2 are positive. They are further supported by a strong data-model fit (r2 = 0.607) using a stomatal optimization model applied to the field gs dataset. A parameter space identified in the Farquhar-Ball-Berry photosynthesis-stomatal conductance model confirms field observations of increasing gs under elevated CO2 in hot dry conditions. Contrary to the general assumption, positive gs responses to elevated CO2, although relatively rare, are a feature of woody taxa adapted to warm, low-humidity conditions, and this response is also demonstrated in global simulations using the Community Land Model (CLM4). Conclusions:The results contradict the over-simplistic notion that global vegetation always responds with decreasing gs to elevated CO2, a finding that has important implications for predicting future vegetation feedbacks on the hydrological cycle at the regional level.
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Project description:We employed a detailed whole leaf hydraulic model to study the local operation of three stomatal conductance models distributed on the scale of a whole leaf. We quantified the behavior of these models by examining the leaf-area distributions of photosynthesis, transpiration, stomatal conductance, and guard cell turgor pressure. We gauged the models' local responses to changes in environmental conditions of carbon dioxide concentration, relative humidity, and light irradiance. We found that a stomatal conductance model that includes mechanical processes dependent on local variables predicts a spatial variation of physiological activity across the leaf: the leaf functions of photosynthesis and transpiration are not uniformly operative even when external conditions are uniform. The gradient pattern of hydraulic pressure which is needed to produce transpiration from the whole leaf is derived from the gradient patterns of turgor pressures of guard cells and epidermal cells and consequently leads to nonuniform spatial distribution patterns of transpiration and photosynthesis via the mechanical stomatal model. Our simulation experiments, comparing the predictions of two versions of a mechanical stomatal conductance model, suggest that leaves exhibit a more complex spatial distribution pattern of both photosynthesis and transpiration rate and more complex dependencies on environmental conditions when a non-linear relationship between the stomatal aperture and guard cell and epidermal cell turgor pressures is implemented. Our model studies offer a deeper understanding of the mechanism of stomatal conductance and point to possible future experimental measurements seeking to quantify the spatial distributions of several physiological activities taking place over a whole leaf.
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Project description:The impact of global warming on transpiration and photosynthesis would compromise plant fitness, impacting on crop yields and ecosystem functioning. In this frame, we explored the performance of a set of Arabidopsis mutants carrying partial or total loss-of-function alleles of stomatal development genes and displaying distinct stomatal abundances. Using microscopy and non-invasive imaging techniques on this genotype collection, we examined anatomical leaf and stomatal traits, plant growth and development, and physiological performance at optimal (22°C) and supra-optimal (30°C) temperatures. All genotypes showed thermomorphogenetic responses but no signs of heat stress. Data analysis singled out an extremely low stomatal abundance mutant, spch-5. At 22°C, spch-5 had lower transpiration and warmer leaves than the wild type. However, at 30°C, this mutant developed larger stomata and thinner leaves, paralleled by a notable cooling capacity, similar to that of the wild type. Despite their low stomatal density (SD), spch-5 plants grown at 30°C showed no photosynthesis or growth penalties. The behavior of spch-5 at supra-optimal temperature exemplifies how the effect of very low stomatal numbers can be counteracted by a combination of larger stomata and thinner leaves. Furthermore, it provides a novel strategy for coping with high growth temperatures.
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