stomatal regulation Dotaz Zobrazit nápovědu
The initiation of stomata, microscopic valves in the epidermis of higher plants that control of gas exchange, requires a co-ordinated sequence of asymmetric and symmetric divisions, which is under tight environmental and developmental control. Arabidopsis leaves grown under elevated photosynthetic photon flux density have a higher density of stomata. STOMAGEN encodes an epidermal patterning factor produced in the mesophyll, and our observations indicated that elevated photosynthetic irradiation stimulates STOMAGEN expression. Our analysis of gain and loss of function of STOMAGEN further detailed its function as a positive regulator of stomatal formation on both sides of the leaf, not only in terms of stomatal density across the leaf surface but also in terms of their stomatal index. STOMAGEN function was rate limiting for the light response of the stomatal lineage in the adaxial epidermis. Mutants in pathways that regulate stomatal spacing in the epidermis and have elevated stomatal density, such as stomatal density and distribution (sdd1) and too many mouth alleles, displayed elevated STOMAGEN expression, suggesting that STOMAGEN is either under the direct control of these pathways or is indirectly affected by stomatal patterning, suggestive of a feedback mechanism. These observations support a model in which changes in levels of light irradiation are perceived in the mesophyll and control the production of stomata in the epidermis by mesophyll-produced STOMAGEN, and whereby, conversely, stomatal patterning, either directly or indirectly, influences STOMAGEN levels.
- MeSH
- Arabidopsis genetika růst a vývoj metabolismus účinky záření MeSH
- fotosyntéza MeSH
- listy rostlin růst a vývoj metabolismus účinky záření MeSH
- proteiny huseníčku genetika metabolismus MeSH
- průduchy rostlin genetika růst a vývoj metabolismus účinky záření MeSH
- regulace genové exprese u rostlin účinky záření MeSH
- signální transdukce MeSH
- světlo MeSH
- vývojová regulace genové exprese účinky záření MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Stomatal conductance directly modifies plant water relations and photosynthesis. Many environmental factors affecting the stomatal conductance have been intensively studied but temperature has been largely neglected, even though it is one of the fastest changing environmental variables and it is rising due to climate change. In this study, we describe how stomata open when the temperature increases. Stomatal conductance increased by ca 40% in a broadleaf and a coniferous species, poplar (Populus deltoides x nigra) and loblolly pine (Pinus taeda) when temperature was increased by 10 °C, from 30 °C to 40 °C at a constant vapor pressure deficit of 1 kPa. The mechanism of regulating stomatal conductance by temperature was, at least partly, independent of other known mechanisms linked to water status and carbon metabolism. Stomatal conductance increased with rising temperature despite the decrease in leaf water potential, increase in transpiration, increase in intercellular CO2 concentration and was decoupled from photosynthesis. Increase in xylem and mesophyll hydraulic conductance coming from lower water viscosity may to some degree explain temperature dependent opening of stomata. The direct stomatal response to temperature allows plants to benefit from increased evaporative cooling during the heat waves and from lower stomatal limitations to photosynthesis but they may be jeopardized by faster depletion of soil water.
- MeSH
- borovice fyziologie MeSH
- Populus fyziologie MeSH
- průduchy rostlin fyziologie MeSH
- teplota * MeSH
- tlak par MeSH
- voda MeSH
- Publikační typ
- časopisecké články MeSH
Stomatal ontogenesis, patterning, and function are hallmarks of environmental plant adaptation, especially to conditions limiting plant growth, such as elevated temperatures and reduced water availability. The specification and distribution of a stomatal cell lineage and its terminal differentiation into guard cells require a master regulatory protein phosphorylation cascade involving the YODA mitogen-activated protein kinase kinase kinase. YODA signaling results in the activation of MITOGEN-ACTIVATED PROTEIN KINASEs (MPK3 and MPK6), which regulate transcription factors, including SPEECHLESS (SPCH). Here, we report that acute heat stress affects the phosphorylation and deactivation of SPCH and modulates stomatal density. By using complementary molecular, genetic, biochemical, and cell biology approaches, we provide solid evidence that HEAT SHOCK PROTEINS 90 (HSP90s) play a crucial role in transducing heat-stress response through the YODA cascade. Genetic studies revealed that YODA and HSP90.1 are epistatic, and they likely function linearly in the same developmental pathway regulating stomata formation. HSP90s interact with YODA, affect its cellular polarization, and modulate the phosphorylation of downstream targets, such as MPK6 and SPCH, under both normal and heat-stress conditions. Thus, HSP90-mediated specification and differentiation of the stomatal cell lineage couples stomatal development to environmental cues, providing an adaptive heat stress response mechanism in plants.
- MeSH
- Arabidopsis fyziologie MeSH
- buněčná diferenciace MeSH
- buněčné dělení MeSH
- buněčný rodokmen MeSH
- epigeneze genetická MeSH
- fosforylace MeSH
- kotyledon cytologie MeSH
- MAP kinasy kinas (kinas) genetika metabolismus MeSH
- mitogenem aktivované proteinkinasy kinas metabolismus MeSH
- mitogenem aktivované proteinkinasy metabolismus MeSH
- mutace MeSH
- proteiny huseníčku genetika metabolismus MeSH
- proteiny tepelného šoku HSP90 genetika metabolismus MeSH
- průduchy rostlin cytologie růst a vývoj metabolismus MeSH
- reakce na tepelný šok * MeSH
- regulace genové exprese u rostlin MeSH
- signální transdukce MeSH
- transkripční faktory bHLH metabolismus MeSH
- vazba proteinů MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Occurrence of stomata on both leaf surfaces (amphistomaty) promotes higher stomatal conductance and photosynthesis while simultaneously increasing exposure to potential disease agents in black cottonwood (Populus trichocarpa). A genome-wide association study (GWAS) with 2.2M single nucleotide polymorphisms generated through whole-genome sequencing found 280 loci associated with variation in adaxial stomatal traits, implicating genes regulating stomatal development and behavior. Strikingly, numerous loci regulating plant growth and response to biotic and abiotic stresses were also identified. The most significant locus was a poplar homologue of SPEECHLESS (PtSPCH1). Individuals possessing PtSPCH1 alleles associated with greater adaxial stomatal density originated primarily from environments with shorter growing seasons (e.g. northern latitudes, high elevations) or with less precipitation. PtSPCH1 was expressed in developing leaves but not developing stem xylem. In developing leaves, RNA sequencing showed patterns of coordinated expression between PtSPCH1 and other GWAS-identified genes. The breadth of our GWAS results suggests that the evolution of amphistomaty is part of a larger, complex response in plants. Suites of genes underpin this response, retrieved through genetic association to adaxial stomata, and show coordinated expression during development. We propose that the occurrence of amphistomaty in P. trichocarpa involves PtSPCH1 and reflects selection for supporting rapid growth over investment in immunity.
- MeSH
- alely MeSH
- celogenomová asociační studie MeSH
- druhová specificita MeSH
- fenotyp MeSH
- genotyp MeSH
- imunita rostlin genetika MeSH
- jednonukleotidový polymorfismus genetika MeSH
- kvantitativní znak dědičný MeSH
- podnebí MeSH
- Populus genetika růst a vývoj imunologie fyziologie MeSH
- průduchy rostlin genetika fyziologie MeSH
- regulace genové exprese u rostlin MeSH
- rostlinné geny MeSH
- rostlinné proteiny genetika metabolismus MeSH
- rozvržení tělního plánu * MeSH
- vývoj rostlin MeSH
- zeměpis MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Element content and expression of genes of interest on single cell types, such as stomata, provide valuable insights into their specific physiology, improving our understanding of leaf gas exchange regulation. We investigated how far differences in stomatal conductance (gs ) can be ascribed to changes in guard cells functioning in amphistomateous leaves. gs was measured during the day on both leaf sides, on well-watered and drought-stressed trees (two Populus euramericana Moench and two Populus nigra L. genotypes). In parallel, guard cells were dissected for element content and gene expressions analyses. Both were strongly arranged according to genotype, and drought had the lowest impact overall. Normalizing the data by genotype highlighted a structure on the basis of leaf sides and time of day both for element content and gene expression. Guard cells magnesium, phosphorus, and chlorine were the most abundant on the abaxial side in the morning, where gs was at the highest. In contrast, genes encoding H+ -ATPase and aquaporins were usually more abundant in the afternoon, whereas genes encoding Ca2+ -vacuolar antiporters, K+ channels, and ABA-related genes were in general more abundant on the adaxial side. Our work highlights the unique physiology of each leaf side and their analogous rhythmicity through the day.
- MeSH
- genotyp MeSH
- komplementární DNA genetika izolace a purifikace MeSH
- listy rostlin genetika metabolismus MeSH
- mikroanalýza elektronovou sondou MeSH
- období sucha MeSH
- Populus klasifikace genetika metabolismus MeSH
- protonové ATPasy genetika metabolismus MeSH
- průduchy rostlin genetika metabolismus MeSH
- regulace genové exprese u rostlin MeSH
- RNA rostlin genetika izolace a purifikace MeSH
- rostlinné proteiny genetika metabolismus MeSH
- stromy genetika metabolismus MeSH
- transpirace rostlin fyziologie MeSH
- voda fyziologie MeSH
- vývoj rostlin MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Conifers growing at high elevations need to optimize their stomatal conductance (gs ) for maximizing photosynthetic yield while minimizing water loss under less favourable thermal conditions. Yet the ability of high-elevation conifers to adjust their gs sensitivity to environmental drivers remains largely unexplored. We used 4 years of sap flow measurements to elucidate intraspecific and interspecific variability of gs in Larix decidua Mill. and Picea abies (L.) Karst along an elevational gradient and contrasting soil moisture conditions. Site- and species-specific gs response to main environmental drivers were examined, including vapour pressure deficit, air temperature, solar irradiance, and soil water potential. Our results indicate that maximum gs of L. decidua is >2 times higher, shows a more plastic response to temperature, and down-regulates gs stronger during atmospheric drought compared to P. abies. These differences allow L. decidua to exert more efficient water use, adjust to site-specific thermal conditions, and reduce water loss during drought episodes. The stronger plasticity of gs sensitivity to temperature and higher conductance of L. decidua compared to P. abies provide new insights into species-specific water use strategies, which affect species' performance and should be considered when predicting terrestrial water dynamics under future climatic change.
- MeSH
- borovice fyziologie MeSH
- cévnaté rostliny * fyziologie MeSH
- fyziologická adaptace MeSH
- modřín fyziologie MeSH
- období sucha MeSH
- průduchy rostlin fyziologie MeSH
- půda MeSH
- teplota MeSH
- transpirace rostlin fyziologie MeSH
- voda fyziologie MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Guard cells on the leaf epidermis regulate stomatal opening for gas exchange between plants and the atmosphere, allowing a balance between photosynthesis and transpiration. Given that guard cells possess several characteristics of sink tissues, their metabolic activities should largely depend on mesophyll-derived sugars. Early biochemical studies revealed sugar uptake into guard cells. However, the transporters that are involved and their relative contribution to guard cell function are not yet known. Here, we identified the monosaccharide/proton symporters Sugar Transport Protein 1 and 4 (STP1 and STP4) as the major plasma membrane hexose sugar transporters in the guard cells of Arabidopsis thaliana. We show that their combined action is required for glucose import to guard cells, providing carbon sources for starch accumulation and light-induced stomatal opening that are essential for plant growth. These findings highlight mesophyll-derived glucose as an important metabolite connecting stomatal movements with photosynthesis.
Understanding the response of a crop to drought is the first step in the breeding of tolerant genotypes. In our study, two maize (Zea mays L.) genotypes with contrasting sensitivity to dehydration were subjected to moderate drought conditions. The subsequent analysis of their physiological parameters revealed a decreased stomatal conductance accompanied by a slighter decrease in the relative water content in the sensitive genotype. In contrast, the tolerant genotype maintained open stomata and active photosynthesis, even under dehydration conditions. Drought-induced changes in the leaf proteome were analyzed by two independent approaches, 2D gel electrophoresis and iTRAQ analysis, which provided compatible but only partially overlapping results. Drought caused the up-regulation of protective and stress-related proteins (mainly chaperones and dehydrins) in both genotypes. The differences in the levels of various detoxification proteins corresponded well with the observed changes in the activities of antioxidant enzymes. The number and levels of up-regulated protective proteins were generally lower in the sensitive genotype, implying a reduced level of proteosynthesis, which was also indicated by specific changes in the components of the translation machinery. Based on these results, we propose that the hypersensitive early stomatal closure in the sensitive genotype leads to the inhibition of photosynthesis and, subsequently, to a less efficient synthesis of the protective/detoxification proteins that are associated with drought tolerance.
- MeSH
- 2D gelová elektroforéza MeSH
- antioxidancia metabolismus MeSH
- dehydratace MeSH
- fyziologická adaptace MeSH
- genotyp MeSH
- glutathionreduktasa metabolismus MeSH
- katalasa metabolismus MeSH
- kukuřice setá enzymologie genetika fyziologie MeSH
- období sucha MeSH
- proteomika MeSH
- průduchy rostlin fyziologie MeSH
- superoxiddismutasa metabolismus MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Responses to drought, heat, and combined stress were compared in tobacco (Nicotiana tabacum L.) plants ectopically expressing the cytokinin oxidase/dehydrogenase CKX1 gene of Arabidopsis thaliana L. under the control of either the predominantly root-expressed WRKY6 promoter or the constitutive 35S promoter, and in the wild type. WRKY6:CKX1 plants exhibited high CKX activity in the roots under control conditions. Under stress, the activity of the WRKY6 promoter was down-regulated and the concomitantly reduced cytokinin degradation coincided with raised bioactive cytokinin levels during the early phase of the stress response, which might contribute to enhanced stress tolerance of this genotype. Constitutive expression of CKX1 resulted in an enlarged root system, a stunted, dwarf shoot phenotype, and a low basal level of expression of the dehydration marker gene ERD10B. The high drought tolerance of this genotype was associated with a relatively moderate drop in leaf water potential and a significant decrease in leaf osmotic potential. Basal expression of the proline biosynthetic gene P5CSA was raised. Both wild-type and WRKY6:CKX1 plants responded to heat stress by transient elevation of stomatal conductance, which correlated with an enhanced abscisic acid catabolism. 35S:CKX1 transgenic plants exhibited a small and delayed stomatal response. Nevertheless, they maintained a lower leaf temperature than the other genotypes. Heat shock applied to drought-stressed plants exaggerated the negative stress effects, probably due to the additional water loss caused by a transient stimulation of transpiration. The results indicate that modulation of cytokinin levels may positively affect plant responses to abiotic stress through a variety of physiological mechanisms.
- MeSH
- Arabidopsis enzymologie genetika MeSH
- cytokininy metabolismus MeSH
- exprese genu MeSH
- geneticky modifikované rostliny chemie genetika fyziologie MeSH
- období sucha MeSH
- oxidoreduktasy genetika metabolismus MeSH
- promotorové oblasti (genetika) MeSH
- proteiny huseníčku genetika metabolismus MeSH
- regulace genové exprese u rostlin MeSH
- tabák chemie genetika fyziologie MeSH
- vysoká teplota MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Cytokinins (CKs) are phytohormones regulating plant growth and development as well as response to the environment. In order to evaluate their function in heat stress (HS) responses, the effect of CK elevation was determined during three types of HS - targeted to shoots, targeted to roots and applied to the whole plant. The early (30min) and longer term (3h) responses were followed at the hormonal, transcriptomic and proteomic levels in Arabidopsis transformants with dexamethasone-inducible expression of the CK biosynthetic gene isopentenyltransferase (ipt) and the corresponding wild-type (Col-0). Combination of hormonal and phenotypic analyses showed transient up-regulation of the CK/abscisic acid ratio, which controls stomatal aperture, to be more pronounced in the transformant. HS responses of the root proteome and Rubisco-immunodepleted leaf proteome were followed using 2-D gel electrophoresis and MALDI-TOF/TOF. More than 100 HS-responsive proteins were detected, most of them being modulated by CK increase. Proteome and transcriptome analyses demonstrated that CKs have longer term positive effects on the stress-related proteins and transcripts, as well as on the photosynthesis-related ones. Transient accumulation of CKs and stimulation of their signal transduction in tissue(s) not exposed to HS indicate that they are involved in plant stress responses.
- MeSH
- alkyltransferasy a aryltransferasy fyziologie MeSH
- Arabidopsis účinky léků metabolismus fyziologie MeSH
- cytokininy fyziologie MeSH
- dexamethason farmakologie MeSH
- kořeny rostlin metabolismus fyziologie MeSH
- kyselina abscisová fyziologie MeSH
- proteomika MeSH
- reakce na tepelný šok fyziologie MeSH
- regulace genové exprese u rostlin fyziologie MeSH
- regulátory růstu rostlin fyziologie MeSH
- signální transdukce účinky léků fyziologie MeSH
- spektrometrie hmotnostní - ionizace laserem za účasti matrice MeSH
- stanovení celkové genové exprese MeSH
- výhonky rostlin metabolismus fyziologie MeSH
- vysoká teplota MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH