6-Benzylaminopurine Alleviates the Impact of Cu2+ Toxicity on Photosynthetic Performance of Ricinus communis L. Seedlings

Copper (Cu) is an essential element involved in various metabolic processes in plants, but at concentrations above the threshold level, it becomes a potential stress factor. The effects of two different cytokinins, kinetin (KIN) and 6-benzylaminopurine (BAP), on chlorophyll a fluorescence parameters, stomatal responses and antioxidation mechanisms in castor (Ricinus communis L.) under Cu2+ toxicity was investigated. Ricinus communis plants were exposed to 80 and 160 μM CuSO4 added to the growth medium. Foliar spraying of 15 μM KIN and BAP was carried out on these seedlings. The application of these cytokinins enhanced the tissue water status, chlorophyll contents, stomatal opening and photosynthetic efficiency in the castor plants subjected to Cu2+ stress. The fluorescence parameters, such as Fm, Fv/Fo, Sm, photochemical and non-photochemical quantum yields, energy absorbed, energy trapped and electron transport per cross-sections, were more efficiently modulated by BAP application than KIN under Cu2+ toxicity. There was also effective alleviation of reactive oxygen species by enzymatic and non-enzymatic antioxidation systems, reducing the membrane lipid peroxidation, which brought about a relative enhancement in the membrane stability index. Of the various treatments, 80 µM CuSO4 + BAP recorded the highest increase in photosynthetic efficiency compared to other cytokinin treatments. Therefore, it can be concluded that BAP could effectively alleviate the detrimental effects of Cu2+toxicity in cotyledonary leaves of R. communis by effectively modulating stomatal responses and antioxidation mechanisms, thereby enhancing the photosynthetic apparatus’ functioning.

Stomatal Responses to Light, CO2, and Mesophyll Tissue in Vicia faba and Kalanchoë fedtschenkoi

The responses of stomatal aperture to light intensity and CO2 concentration were studied in both Vicia faba (C3) and Kalanchoë fedtschenkoi (Crassulacean acid metabolism; CAM), in material sampled from both light and dark periods. Direct comparison was made between intact leaf segments, epidermises grafted onto exposed mesophyll, and isolated epidermal peels, including transplantations between species and between diel periods. We reported the stomatal opening in response to darkness in isolated CAM peels from the light period, but not from the dark. Furthermore, we showed that C3 mesophyll has stimulated CAM stomata in transplanted peels to behave as C3 in response to light and CO2. By using peels and mesophyll from plants sampled in the dark and the light period, we provided clear evidence that CAM stomata behaved differently from C3. This might be linked to stored metabolites/ions and signalling pathway components within the guard cells, and/or a mesophyll-derived signal. Overall, our results provided evidence for both the involvement of guard cell metabolism and mesophyll signals in stomatal responses in both C3 and CAM species.

Physiological and Yield Responses of Spring Wheat Cultivars under Realistic and Acute Levels of Ozone

Tropospheric ozone (O3) is widely recognized as the cause of substantial yield and quality reduction in crops. Most of the previous studies focused on the exposure of wheat cultivars to elevated O3 levels. Our main objectives were to: (i) investigate the consistency of wheat cultivars’ physiological responses across two different realistic O3 levels; and (ii) compare these physiological responses with those under short acute O3 exposure. Three commercially available hard spring wheat cultivars bred under semiarid and Eastern Mediterranean conditions were exposed to two different O3 levels during two consecutive seasons (2016–2018)—36 and 71 ppbv 7 h mean O3 mixing ratios in open-top chambers. The results were compared to those following short acute O3 exposure (102.8 ppbv, 7 h mean for 10 days) in a greenhouse. Non-stomatal responses were significantly more pronounced than stomatal responses in all cultivars under different levels of O3. The specific cultivar was observed as the most O3-tolerant under all experiments. The fact that the same cultivar was found remarkably tolerant to the local semiarid ambient conditions according to other studies and to O3 exposure based on the present study supports a link between cultivar resistance to drought conditions and O3.

Concentration matters: different stomatal CO2-responses at sub-ambient and above-ambient CO2 levels

Stomatal pores, formed of paired guard cells, mediate CO2 uptake for photosynthesis and water loss via transpiration in plants. Globally rising atmospheric CO2 concentration triggers stomatal closure, contributing to increased leaf temperature and reduced nutrient uptake due to lower transpiration rate. Hence, it is important to understand the signalling pathways that control elevated CO2-induced stomatal closure to identify targets for breeding climate-ready crops. CO2-induced stomatal closure can be studied by increasing CO2 concentration from ambient to above-ambient concentrations, or elevation of CO2 levels from sub-ambient to above-ambient. Previous experiments comparing ferns with angiosperms suggested that stomatal responses to CO2 may be different, when changing CO2 levels in the sub-ambient or above-ambient ranges. Here, we set out to test this by comparing CO2-induced stomatal closure in key guard cell signalling mutants in response to CO2 elevation from 100 to 400 ppm or 400 to 800 ppm. We show that signalling components that contribute to CO2-induced stomatal closure are different in the sub-ambient and above-ambient CO2 levels, with guard cell slow-type anion channel SLAC1 involved mainly in above-ambient CO2-induced stomatal closure.

Conditional Stomatal Closure in a Fern Shares Molecular Features with Flowering Plant Active Stomatal Responses

Stomata evolved as plants transitioned from water to land, enabling carbon dioxide uptake and water loss to be controlled. In flowering plants, the most recently divergent land plant lineage, stomatal pores actively close in response to drought. In this response, the phytohormone abscisic acid (ABA) triggers signalling cascades that lead to ion and water loss in the guard cells of the stomatal complex, causing a reduction in turgor and pore closure. Whether this stimulus-response coupling pathway acts in other major land plant lineages is unclear, with some investigations reporting that stomatal closure involves ABA but others concluding that closure is passive. Here we show that in the model fern Ceratopteris richardii active stomatal closure is conditional on sensitisation by pre-exposure to either low humidity or exogenous ABA and is promoted by ABA. RNA-seq analysis and de novo transcriptome assembly reconstructed the protein coding complement of the C. richardii genome with coverage comparable to other plant models, enabling transcriptional signatures of stomatal sensitisation and closure to be identified. In both cases, changes in abundance of homologs of ABA, Ca2+ and ROS-related signalling components were observed, suggesting that the closure response pathway is conserved in ferns and flowering plants. These signatures further suggested that sensitisation is achieved by lowering the threshold required for a subsequent closure-inducing signal to trigger a response. We conclude that the canonical signalling network for active stomatal closure functioned in at least a rudimentary form in the stomata of the last common ancestor of ferns and flowering plants.Significance StatementStomata are valve-like pores that control the uptake of CO2 and the loss of water vapour in almost all land plants. In flowering plants, stomatal opening and closure is actively regulated by a stimulus-response coupling network. Whether active stomatal responses are present in other land plant lineages such as ferns has been hotly debated. Here we show that stomatal responses in the fern Ceratopteris richardii are active but depend on their past growth environment, and demonstrate that fern stomatal closure and sensitisation are associated with the altered expression of genes whose homologs function in the canonical stomatal regulatory network of flowering plants. Genetic pathways for active stomatal regulation therefore most likely evolved before the divergence of ferns and flowering plants.

Winter wheat and maize under varying soil moisture: from leaf to canopy

<p>Drought is one of the most detrimental factors limiting crop growth and production of important staple crops such as winter wheat and maize. For both crops, stomatal regulation and change of canopy structure responses to water stress can be observed. A substantial range of stomatal behavior in regulating water loss was recently reported while the crop growth and morphological responses to drought stress depend on the intensity and duration of the imposed stress. Insights into the responses from leaf to the canopy are important for crop modeling and soil-vegetation-atmosphere models (SVAT). Stomatal responses and effects of soil water deficit on the dynamic change of canopy photosynthesis and transpiration, and seasonal crop growth of winter wheat and maize are investigated based on data collected from field-grown conditions with varying soil moisture treatments (sheltered, rainfed, irrigated) in 2016, 2017, and 2018. A reduction of leaf net photosynthesis (An), stomatal conductance (Gs), transpiration (E), and leaf water potential (LWP) was observed in the sheltered plot as compared to the rainfed and irrigated plots in winter wheat in 2016, indicating anisohydric stomatal responses. Maize showed seasonal isohydric behaviour with the minimum LWP from -1.5 to -2 MPa in 2017 and -2 to -2.7 MPa in the extremely hot and dry year in 2018. Crop growth (biomass, leaf area index, and yield) was substantially reduced under drought conditions, particularly for maize in 2018. Leaf water use efficiency (An/E) and crop WUE (total dry biomass/canopy transpiration) were not significantly different among treatments in both crops. The reduction of tiller number (in winter wheat) and leaf-rolling and plant size (in maize) resulted in a reduction of canopy transpiration, assimilation rate, and thus biomass. The seasonal isohydry in maize and the seasonal variability of LWP in winter wheat suggest a possibility to use the same critical LWP thresholds for maize and wheat to simulate the stomatal control in process-based crop and SVAT models. The canopy response such as dynamically reducing leaf area under water stress adds complexity in simulating gas exchange and crop growth rate that needs adequate consideration in the current modeling approaches.</p>

Stomatal responses to carbon dioxide and light require abscisic acid catabolism in Arabidopsis

In plants, stomata control water loss and CO 2 uptake. The aperture and density of stomatal pores, and hence the exchange of gases between the plant and the atmosphere, are controlled by internal factors such as the plant hormone abscisic acid (ABA) and external signals including light and CO 2 . In this study, we examine the importance of ABA catabolism in the stomatal responses to CO 2 and light. By using the ABA 8′-hydroxylase-deficient Arabidopsis thaliana double mutant cyp707a1 cyp707a3 , which is unable to break down and instead accumulates high levels of ABA, we reveal the importance of the control of ABA concentration in mediating stomatal responses to CO 2 and light. Intriguingly, our experiments suggest that endogenously produced ABA is unable to close stomata in the absence of CO 2 . Furthermore, we show that when plants are grown in short day conditions ABA breakdown is required for the modulation of both elevated [CO 2 ]-induced stomatal closure and elevated [CO 2 ]-induced reductions in leaf stomatal density. ABA catabolism is also required for the stomatal density response to light intensity, and for the full range of light-induced stomatal opening, suggesting that ABA catabolism is critical for the integration of stomatal responses to a range of environmental stimuli.