Differential Responses of Stomata and Photosynthesis to Elevated Temperature in Two Co-occurring Subtropical Forest Tree Species.
ABSTRACT: Global warming could increase leaf transpiration and soil evaporation, which can potentially cause water deficit to plants. As valves, leaf stomata can control plant water loss and carbon gain, particularly under water stress conditions. To investigate the responses of stomata to elevated temperature in Schima superba and Syzygium rehderianum, two co-occurring subtropical forest dominant tree species, functional traits related to gas exchange, stomatal anatomy, and drought resistance were measured under control and warming environment (ca. 2°C higher). We found that leaf water potential at both predawn and midday significantly decreased for the two species grown under warming conditions compared with those grown in the control environment. Warming resulted in significant decrease of stomatal size in S. rehderianum, but had no obvious effect on that of S. superba. By contrast, stomatal density of S. superba significantly decreased under warming conditions, while non-significant change was observed for S. rehderianum. In addition, warming significantly reduced photosynthetic rate, stomatal conductance, and stomatal sensitivity to leaf water potential of S. superba, but had non-significant effects on those of S. rehderianum. Overall, our results demonstrated that, confronting water deficit caused by elevated temperature, the two co-occurring subtropical tree species responded differently through the adjustment of stomatal morphology and photosynthetic function. Consequently, S. rehderianum was able to maintain similar carbon assimilation as under control environment, while S. superba showed a decrease in carbon gain that might bring adverse effect on its dominancy in subtropical forest community under future climate change.
Project description:Ongoing climate warming induced by human activities may have great impacts on trees, yet it remains unresolved how subtropical tree species respond to rising temperature in the field. Here, we used downward translocation to investigate the effects of climate warming on leaf photosynthesis of six common tree species in subtropical China. During the experimental period between 2012 and 2014, the mean average photosynthetic rates (Asat) under saturating light for Schima superba, Machilus breviflora, Pinus massoniana and Ardisia lindleyana in the warm site were7%, 19%, 20% and 29% higher than those in the control site. In contrast, seasonal Asat for Castanopsis hystrix in the warm site were lower compared to the control site. Changes in Asat in response to translocation were mainly associated with those in leaf stomatal conductance (gs) and photosynthetic capacity (RuBP carboxylation, RuBP regeneration capacity). Our results imply that climate warming could have potential impacts on species composition and community structure in subtropical forests.
Project description:The opening and closing of stomata are controlled by the integration of environmental and endogenous signals. Here, we show the effects of combining elevated atmospheric carbon dioxide concentration (<i>eCO</i> <sub>2</sub>; 600 ?mol mol<sup>-1</sup>) and warming (+2°C) on stomatal properties and their consequence to plant function in a <i>Stylosanthes capitata</i> Vogel (C<sub>3</sub>) tropical pasture. The <i>eCO</i> <sub>2</sub> treatment alone reduced stomatal density, stomatal index, and stomatal conductance (<i>g<sub>s</sub></i> ), resulting in reduced transpiration, increased leaf temperature, and leading to maintenance of soil moisture during the growing season. Increased CO<sub>2</sub> concentration inside leaves stimulated photosynthesis, starch content levels, water use efficiency, and PSII photochemistry. Under warming, plants developed leaves with smaller stomata on both leaf surfaces; however, we did not see effects of warming on stomatal conductance, transpiration, or leaf water status. Warming alone enhanced PSII photochemistry and photosynthesis, and likely starch exports from chloroplasts. Under the combination of warming and <i>eCO</i> <sub>2</sub>, leaf temperature was higher than that of leaves from the warming or <i>eCO</i> <sub>2</sub> treatments. Thus, warming counterbalanced the effects of CO<sub>2</sub> on transpiration and soil water content but not on stomatal functioning, which was independent of temperature treatment. Under warming, and in combination with <i>eCO</i> <sub>2</sub>, leaves also produced more carotenoids and a more efficient heat and fluorescence dissipation. Our combined results suggest that control on stomatal opening under <i>eCO</i> <sub>2</sub> was not changed by a warmer environment; however, their combination significantly improved whole-plant functioning.
Project description:Life history and photosynthetic type both affect the economics of leaf physiological function. Annual plants have lower tissue densities and resource-use efficiencies than perennials, while C4 photosynthesis, facilitated in grasses by specific changes in leaf anatomy, improves photosynthetic efficiency and water-use efficiency, especially in hot climates. This study aimed to determine whether C4 photosynthesis affects differences in functional traits between annual and perennial species. We measured 26 traits and characterised niche descriptors for 42 grasses from subtropical China. Differences in the majority of traits were explained by life history. The ranges of annual species (particularly C4 annuals) extended to regions with greater temperature seasonality and lower precipitation, and annuals had less-negative turgor-loss points, higher specific leaf areas, and lower water-use efficiencies, stomatal conductances, and leaf areas per stem area than perennials. Photosynthetic type largely affected leaf physiology as expected, but interacted with life history in determining specific traits. Leaf hydraulic conductance was intermediate in perennials, highest in C4-annuals, and lowest in C3-annuals. Densities of stomata and stem vessels were similar across C3-perennials and C4 species, but stomatal densities were lower and stem vessel densities higher in C3-annuals. Phylogenetic principal component analysis confirmed that in this subtropical environment life history is the predominant axis separating species, and annuals and perennials were more different within C3 than C4 grasses. The interplay between life history and photosynthetic type may be an overlooked factor in shaping the physiological ecology of grasses.
Project description:The mechanism underlying the effect of drought on the photosynthetic traits of leaves in forest ecosystems in subtropical regions is unclear. In this study, three limiting processes (stomatal, mesophyll and biochemical limitations) that control the photosynthetic capacity and three resource use efficiencies (intrinsic water use efficiency (iWUE), nitrogen use efficiency (NUE) and light use efficiency (LUE)), which were characterized as the interactions between photosynthesis and environmental resources, were estimated in two species (Schima superba and Pinus massoniana) under drought conditions. A quantitative limitation analysis demonstrated that the drought-induced limitation of photosynthesis in Schima superba was primarily due to stomatal limitation, whereas for Pinus massoniana, both stomatal and non-stomatal limitations generally exhibited similar magnitudes. Although the mesophyll limitation represented only 1% of the total limitation in Schima superba, it accounted for 24% of the total limitations for Pinus massoniana. Furthermore, a positive relationship between the LUE and NUE and a marginally negative relationship or trade-off between the NUE and iWUE were observed in the control plots. However, drought disrupted the relationships between the resource use efficiencies. Our findings may have important implications for reducing the uncertainties in model simulations and advancing the understanding of the interactions between ecosystem functions and climate change.
Project description:Stomata are simultaneously tasked with permitting the uptake of carbon dioxide for photosynthesis while limiting water loss from the plant. This process is mainly regulated by guard cell control of the stomatal aperture, but recent advancements have highlighted the importance of several genes that control stomatal development. Using targeted genetic manipulations of the stomatal lineage and a combination of gas exchange and microscopy techniques, we show that changes in stomatal development of the epidermal layer lead to coupled changes in the underlying mesophyll tissues. This coordinated response tends to match leaf photosynthetic potential (V<sub>cmax</sub> ) with gas-exchange capacity (g<sub>smax</sub> ), and hence the uptake of carbon dioxide for water lost. We found that different genetic regulators systematically altered tissue coordination in separate ways: the transcription factor SPEECHLESS (SPCH) primarily affected leaf size and thickness, whereas peptides in the EPIDERMAL PATTERNING FACTOR (EPF) family altered cell density in the mesophyll. It was also determined that interlayer coordination required the cell-surface receptor TOO MANY MOUTHS (TMM). These results demonstrate that stomata-specific regulators can alter mesophyll properties, which provides insight into how molecular pathways can organize leaf tissues to coordinate gas exchange and suggests new strategies for improving plant water-use efficiency.
Project description:Stomata, the microvalves on leaf surfaces, exert major influences across scales, from plant growth and productivity to global carbon and water cycling. Stomatal opening enables leaf photosynthesis, and plant growth and water use, whereas plant survival of drought depends on stomatal closure. Here we report that stomatal function is constrained by a safety-efficiency trade-off, such that species with greater stomatal conductance under high water availability (g<sub>max</sub>) show greater sensitivity to closure during leaf dehydration, i.e., a higher leaf water potential at which stomatal conductance is reduced by 50% (?<sub>gs50</sub>). The g<sub>max</sub> - ?<sub>gs50</sub> trade-off and its mechanistic basis is supported by experiments on leaves of California woody species, and in analyses of previous studies of the responses of diverse flowering plant species around the world. Linking the two fundamental key roles of stomata-the enabling of gas exchange, and the first defense against drought-this trade-off constrains the rates of water use and the drought sensitivity of leaves, with potential impacts on ecosystems.
Project description:We examined the warming effects on the stomatal frequency, stomatal aperture size and shape, and their spatial distribution pattern of maize (Zea may L.) leaves using a light microscope, an electron scanning microscope, and geostatistic techniques. A field manipulative experiment was conducted to elevate canopy temperature by 2.08°C, on average. We found that experimental warming had little effect on stomatal density, but significantly increased stomatal index due to the reduction in the number of epidermal cells under the warming treatment. Warming also significantly decreased stomatal aperture length and increased stomatal aperture width. As a result, warming significantly increased the average stomatal aperture area and stomatal aperture circumference. In addition, warming dramatically changed the stomatal spatial distribution pattern with a substantial increase in the average nearest neighbor distance between stomata on both adaxial and abaxial surfaces. The spatial distribution pattern of stomata was scale dependent with regular patterns at small scales and random patterns at larger scales on both leaf surfaces. Warming caused the stomatal distribution to become more regular on both leaf surfaces with smaller L(t) values (Ripley's K-function, L(t) is an expectation of zero for any value of t) in the warming plots than the control plots.
Project description:Quantifying changes in interspecific plant growth and physiology under climate warming will facilitate explanation of the shifts in community structure in subtropical forest. We evaluated the effects of 3 years climate warming (ca. 1°C, 2012-2015) on plant growth and physiological parameters of six subtropical tree species by translocating seedlings and soil from a higher to a lower elevation site. We found that an increase in soil/air temperature had divergent effects on six co-occurring species. Warming increased the biomass of Schima superba and Pinus massoniana, whereas it decreased their specific leaf area and intrinsic water use efficiency compared to other species. Warming decreased the foliar non-structural carbohydrates for all species. Our findings demonstrated that a warmer climate would have species-specific effects on the physiology and growth of subtropical trees, which may cause changes in the competitive balance and composition of these forests.
Project description:Light is one of the factors that can play a role in bacterial infiltration into leafy greens by keeping stomata open and providing photosynthetic products for microorganisms. We model chemotactic transport of bacteria within a leaf tissue in response to photosynthesis occurring within plant mesophyll. The model includes transport of carbon dioxide, oxygen, bicarbonate, sucrose/glucose, bacteria, and autoinducer-2 within the leaf tissue. Biological processes of carbon fixation in chloroplasts, and respiration in mitochondria of the plant cells, as well as motility, chemotaxis, nutrient consumption and communication in the bacterial community are considered. We show that presence of light is enough to boost bacterial chemotaxis through the stomatal opening and toward photosynthetic products within the leaf tissue. Bacterial chemotactic ability is a major player in infiltration, and plant stomatal defense in closing the stomata as a perception of microbe-associated molecular patterns is an effective way to inhibit the infiltration.
Project description:Whereas warming enhances plant nutrient status and photosynthesis in most terrestrial ecosystems, dryland vegetation is vulnerable to the likely increases in evapotranspiration and reductions in soil moisture caused by elevated temperatures. Any warming-induced declines in plant primary production and cover in drylands would increase erosion, land degradation, and desertification. We conducted a four-year manipulative experiment in a semi-arid Mediterranean ecosystem to evaluate the impacts of a ~2°C warming on the photosynthesis, transpiration, leaf nutrient status, chlorophyll content, isotopic composition, biomass growth, and postsummer survival of the native shrub Helianthemum squamatum. We predicted that warmed plants would show reduced photosynthetic activity and growth, primarily due to the greater stomatal limitation imposed by faster and more severe soil drying under warming. On average, warming reduced net photosynthetic rates by 36% across the study period. Despite this strong response, warming did not affect stomatal conductance and transpiration. The reduction of peak photosynthetic rates with warming was more pronounced in a drought year than in years with near-average rainfall (75% and 25-40% reductions relative to controls, respectively), with no indications of photosynthetic acclimation to warming through time. Warmed plants had lower leaf N and P contents, ? (13)C, and sparser and smaller leaves than control plants. Warming reduced shoot dry mass production by 31%. However, warmed plants were able to cope with large reductions in net photosynthesis, leaf area, and shoot biomass production without changes in postsummer survival rates. Our findings highlight the key role of nonstomatal factors (biochemical and/or nutritional) in reducing net carbon assimilation rates and growth under warming, which has important implications for projections of plant carbon balance under the warmer and drier climatic scenario predicted for drylands worldwide. Projected climate warming over the coming decades could reduce net primary production by about one-third in semi-arid gypsum shrublands dominated by H. squamatum.