Knowledge of the ability of plants to respond to climate change via phenotypic plasticity or genetic adaptation in ecophysiological traits and of the link of these traits to fitness is still limited. We studied the clonal grass Festuca rubra from 11 localities representing factorially crossed gradients of temperature and precipitation and cultivated them in growth chambers simulating temperature and moisture regime in the four extreme localities. We measured net photosynthetic rate, Fv /Fm , specific leaf area, osmotic potential and stomatal density and length and tested their relationship to proxies of fitness. We found strong phenotypic plasticity in photosynthetic traits and genetic differentiation in stomatal traits. The effects of temperature and moisture interacted (either as conditions of origin or growth chambers), as were effects of growth and origin. The relationships between the ecophysiological and fitness-related traits were significant but weak. Phenotypic plasticity and genetic differentiation of the species indicate the potential ability of F. rubra to adapt to novel climatic conditions. The most important challenge for the plants seems to be increasing moisture exposing plants to hypoxia. However, the plants have the potential to respond to increased moisture by changes in stomatal size and density and adjustments of osmotic potential. Changes in ecophysiological traits translate into variation in plant fitness, but the selection on the traits is relatively weak and depends on actual conditions. Despite the selection, the plants do not show strong local adaptation and local adaptation is thus likely not restricting species ability to adjust to novel conditions.

Department of Botany Faculty of Science Charles University Prague Czech Republic

Faculty of Science University of South Bohemia České Budějovice Czech Republic

Institute of Botany Czech Academy of Sciences Prague Czech Republic

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Acuna-Rodriguez, I.S., Torres-Diaz, C., Hereme, R. & Molina-Montenegro, M.A. (2017) Asymmetric responses to simulated global warming by populations of Colobanthus quitensis along a latitudinal gradient. PeerJ, 5, 3718.

Adams, W.W., Stewart, J.J., Cohu, C.M., Muller, O. & Demmig-Adams, B. (2016) Habitat temperature and precipitation of Arabidopsis thaliana ecotypes determine the response of foliar vasculature, photosynthesis, and transpiration to growth temperature. Frontiers in Plant Science, 7, 1026.

Agrawal, A.A., Erwin, A.C. & Cook, S.C. (2008) Natural selection on and predicted responses of ecophysiological traits of swamp milkweed (Asclepias incarnata). Journal of Ecology, 96, 536-542.

Aitken, S.N. & Whitlock, M.C. (2013) Assisted gene flow to facilitate local adaptation to climate change. Annual Review of Ecology, Evolution, and Systematics, 44, 367-388.

Anderson, J.T. & Gezon, Z.J. (2015) Plasticity in functional traits in the context of climate change: a case study of the subalpine forb Boechera stricta (Brassicaceae). Global Change Biology, 21, 1689-1703.

Ashraf, M. & Harris, P.J.C. (2013) Photosynthesis under stressful environments: an overview. Photosynthetica, 51, 163-190.

Aspinwall, M.J., Varhammar, A., Blackman, C.J., Tjoelker, M.G., Ahrens, C., Byrne, M. et al. (2017) Adaptation and acclimation both influence photosynthetic and respiratory temperature responses in Corymbia calophylla. Tree Physiology, 37, 1095-1112.

Avolio, M.L. & Smith, M.D. (2013) Intra-specific responses of a dominant C-4 grass to altered precipitation patterns. Plant Ecology, 214, 1377-1389.

Azhar, A., Sathornkich, J., Rattanawong, R. & Kasemsap, P. (2014) Responses of chlorophyll fluorescence, stomatal conductance, and net photosynthesis rates of four rubber (Hevea brasiliensis) genotypes to drought. Advanced Materials Research, 844, 11-14.

Bartlett, M.K., Zhang, Y., Kreidler, N., Sun, S.W., Ardy, R., Cao, K.F. et al. (2014) Global analysis of plasticity in turgor loss point, a key drought tolerance trait. Ecology Letters, 17, 1580-1590.

Baslam, M., Mitsui, T., Hodges, M., Priesack, E., Herritt, M.T., Aranjuelo, I. et al. (2020) Photosynthesis in a changing global climate: scaling up and scaling down in crops. Frontiers of Plant Science, 11, 882.

Bates, D., Machler, M., Bolker, B.M. & Walker, S.C. (2015) Fitting linear mixed-effects models using lme4. Journal of Statistical Software, 67, 1-48.

Beck, E.H., Fettig, S., Knake, C., Hartig, K. & Bhattarai, T. (2007) Specific and unspecific responses of plants to cold and drought stress. Journal of Biosciences, 32, 501-510.

Becklin, K.M., Anderson, J.T., Gerhart, L.M., Wadgymar, S.M., Wessinger, C.A. & Ward, J.K. (2016) Examining plant physiological responses to climate change through an evolutionary lens. Plant Physiology, 172, 635-649.

Benomar, L., Lamhamedi, M.S., Rainville, A., Beaulieu, J., Bousquet, J. & Margolis, H.A. (2016) Genetic adaptation vs. ecophysiological plasticity of photosynthetic-related traits in young Picea glauca trees along a regional climatic gradient. Frontiers in Plant Science, 7, 48.

Blokhina, O., Virolainen, E. & Fagerstedt, K.V. (2003) Antioxidants, oxidative damage and oxygen deprivation stress: a review. Annals of Botany, 91(2), 179-194.

Blum, A. (2017) Osmotic adjustment is a prime drought stress adaptive engine in support of plant production. Plant, Cell and Environment, 40, 4-10.

Bolker, B.M., Brooks, M.E., Clark, C.J., Geange, S.W., Poulsen, J.R., Stevens, M.H.H. et al. (2009) Generalized linear mixed models: a practical guide for ecology and evolution. TREE, 24, 127-135.

Bussotti, F., Pancrazi, M., Matteucci, G. & Gerosa, G. (2005) Leaf morphology and chemistry in Fagus sylvatica (beech) trees as affected by site factors and ozone: results from CONECOFOR permanent monitoring plots in Italy. Tree Physiology, 25, 211-219.

Butof, A., von Riedmatten, L.R., Dormann, C.F., Scherer-Lorenzen, M., Welk, E. & Bruelheide, H. (2012) The responses of grassland plants to experimentally simulated climate change depend on land use and region. Global Change Biology, 18, 127-137.

Carlson, J.E., Adams, C.A. & Holsinger, K.E. (2016) Intraspecific variation in stomatal traits, leaf traits and physiology reflects adaptation along aridity gradients in a South African shrub. Annals of Botany, 117, 195-207.

Crawley, M. (2012) The R book, 2nd edition. Chichester, UK: John Wiley & Sons.

Cunningham, S.A., Summerhayes, B. & Westoby, M. (1999) Evolutionary divergences in leaf structure and chemistry, comparing rainfall and soil nutrient gradients. Ecological Monographs, 69, 569-588.

Davis, M.B. & Shaw, R.G. (2001) Range shifts and adaptive responses to quaternary climate change. Science, 292, 673-679.

De Frenne, P., Brunet, J., Shevtsova, A., Kolb, A., Graae, B.J., Chabrerie, O. et al. (2011) Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient. Global Change Biology, 17, 3240-3253.

Donovan, L.A., Dudley, S.A., Rosenthal, D.M. & Ludwig, F. (2007) Phenotypic selection on leaf water use efficiency and related ecophysiological traits for natural populations of desert sunflowers. Oecologia, 152, 13-25.

Dostálek, T., Rokaya, M.B. & Münzbergová, Z. (2020) Plant palatability and trait responses to experimental warming. Scientific Reports, 10, 10526.

Dudley, S.A. (1996a) Differing selection on plant physiological traits in response to environmental water availability: a test of adaptive hypotheses. Evolution, 50, 92-102.

Dudley, S.A. (1996b) The response to differing selection on plant physiological traits: evidence for local adaptation. Evolution, 50, 103-110.

Franks, P.J., Drake, P.L. & Beerling, D.J. (2009) Plasticity in maximum stomatal conductance constrained by negative correlation between stomatal size and density: an analysis using Eucalyptus globulus. Plant, Cell and Environment, 32, 1737-1748.

Franks, S.J., Weber, J.J. & Aitken, S.N. (2014) Evolutionary and plastic responses to climate change in terrestrial plant populations. Evolutionary Applications, 7, 123-139.

Gamon, J.A., Field, C.B., Goulden, M.L., Griffin, K.L., Hartley, A.E., Joel, G. et al. (1995) Relationships between NDVI, canopy structure, and photosynthesis in 3 Californian vegetation types. Ecological Applications, 5, 28-41.

Garnier, E., Shipley, B., Roumet, C. & Laurent, G. (2001) A standardized protocol for the determination of specific leaf area and leaf dry matter content. Functional Ecology, 15(5), 688-695.

Ghalambor, C.K., McKay, J.K., Carroll, S.P. & Reznick, D.N. (2007) Adaptive versus non-adaptive phenotypic plasticity and the potential for contemporary adaptation in new environments. Functional Ecology, 21, 394-407.

Gienapp, P., Teplitsky, C., Alho, J.S., Mills, J.A. & Merila, J. (2008) Climate change and evolution: disentangling environmental and genetic responses. Molecular Ecology, 17(1), 167-178.

Gitz, D.C. & Baker, J.T. (2009) Methods for creating stomatal impressions directly onto archivable slides. Agronomy Journal, 101, 232-236.

Giuliani, A.L., Kelly, E.F. & Knapp, A.K. (2014) Geographic variation in growth and phenology of two dominant central US grasses: consequences for climate change. Journal of Plant Ecology, 7, 211-221.

Givnish, T.J. & Montgomery, R.A. (2014) Common-garden studies on adaptive radiation of photosynthetic physiology among Hawaiian lobeliads. Proceedings of the Royal Society B: Biological Sciences, 281(1779), 20132944.

Gobel, L., Coners, H., Hertel, D., Willinghofer, S. & Leuschner, C. (2019) The role of low soil temperature for photosynthesis and stomatal conductance of three graminoids from different elevations. Frontiers in Plant Science, 10, 9.

Gonzalo-Turpin, H. & Hazard, L. (2009) Local adaptation occurs along altitudinal gradient despite the existence of gene flow in the alpine plant species Festuca eskia. Journal of Ecology, 97, 742-751.

Gower, S.T., McMurtrie, R.E. & Murty, D. (1996) Aboveground net primary production decline with stand age: potential causes. TREE, 11, 378-382.

Hamann, E., Kesselring, H., Armbruster, G.F.J., Scheepens, J.F. & Stocklin, J. (2016) Evidence of local adaptation to fine- and coarse-grained environmental variability in Poa alpina in the Swiss Alps. Journal of Ecology, 104, 1627-1637.

Hamann, E., Kesselring, H. & Stocklin, J. (2018) Plant responses to simulated warming and drought: a comparative study of functional plasticity between congeneric mid and high elevation species. Journal of Plant Ecology, 11, 364-374.

Hare, P.D., Cress, W.A. & Van Staden, J. (1998) Dissecting the roles of osmolyte accumulation during stress. Plant, Cell and Environment, 21, 535-553.

Hemrová, L., Knappová, J. & Münzbergová, Z. (2016) Assessment of habitat suitability is affected by plant-soil feedback: comparison of field and garden experiment. PLoS One, 11, e0157800.

Henn, J.J., Buzzard, V., Enquist, B.J., Halbritter, A.H., Klanderuds, K., Maitner, B.S. et al. (2018) Intraspecific trait variation and phenotypic plasticity mediate alpine plant species response to climate change. Frontiers in Plant Science, 9, 11.

Heschel, M.S., Sultan, S.E., Glover, S. & Sloan, D. (2004) Population differentiation and plastic responses to drought stress in the generalist annual Polygonum persicaria. International Journal of Plant Sciences, 165, 817-824.

Huntley, B. (1991) How plants respond to climate change-migration rates, individualism and the consequences for plant-communities. Annals of Botany, 67, 15-22.

Inouye, D.W. (2008) Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers. Ecology, 89, 353-362.

IPCC. (2014) Climate change 2014: synthesis report, Vol. 2016. Geneva, Switzerland: IPCC.

Jiao, X.R., Korup, K., Andersen, M.N., Petersen, K.K., Prade, T., Jezowski, S. et al. (2016) Low-temperature leaf photosynthesis of a Miscanthus germplasm collection correlates positively to shoot growth rate and specific leaf area. Annals of Botany, 117, 1229-1239.

Jump, A.S. & Penuelas, J. (2005) Running to stand still: adaptation and the response of plants to rapid climate change. Ecology Letters, 8, 1010-1020.

Kawasaki, S.I., Tominaga, J., Yabuta, S., Watanabe, K., Jaiphong, T., Ueno, M. et al. (2015) Responses of growth, photosynthesis, and associated components to hypoxia at different light intensities in red leaf lettuce. Horticultural Science, 193, 330-336.

Kawecki, T.J. & Ebert, D. (2004) Conceptual issues in local adaptation. Ecology Letters, 7, 1225-1241.

Kingsolver, J.G., Diamond, S.E., Siepielski, A.M. & Carlson, S.M. (2012) Synthetic analyses of phenotypic selection in natural populations: lessons, limitations and future directions. Evolutionary Ecology, 26, 1101-1118.

Kingsolver, J.G., Hoekstra, H.E., Hoekstra, J.M., Berrigan, D., Vignieri, S.N., Hill, C.E. et al. (2001) The strength of phenotypic selection in natural populations. The American Naturalist, 157, 245-261.

Klanderud, K., Vandvik, V. & Goldberg, D. (2015) The importance of biotic vs. abiotic drivers of local plant community composition along regional bioclimatic gradients. PLoS One, 10, e0130205.

Knutzen, F., Meier, I.C. & Leuschner, C. (2015) Does reduced precipitation trigger physiological and morphological drought adaptations in European beech (Fagus sylvatica L.)? Comparing provenances across a precipitation gradient. Tree Physiology, 35, 949-963.

Lande, R. & Arnold, S.J. (1983) The measurement of selection on correlated characters. Evolution, 37, 1210-1226.

Lu, X.M., Zhou, G.S., Wang, Y.H. & Song, X.L. (2016) Effects of changing precipitation and warming on functional traits of zonal Stipa plants from inner Mongolian grassland. Journal of Meteorological Research, 30, 412-425.

Majekova, M., Martinkova, J. & Hajek, T. (2019) Grassland plants show no relationship between leaf drought tolerance and soil moisture affinity, but rapidly adjust to changes in soil moisture. Functional Ecology, 33, 774-785.

Maricle, B.R. & Adler, P.B. (2011) Effects of precipitation on photosynthesis and water potential in Andropogon gerardii and Schizachyrium scoparium in a southern mixed grass prairie. Environmental and Experimental Botany, 72, 223-231.

Matesanz, S., Ramos-Munoz, M., Blanco-Sanchez, M. & Escudero, A. (2020) High differentiation in functional traits but similar phenotypic plasticity in populations of a soil specialist along a climatic gradient. Annals of Botany, 125, 969-980.

Maxwell, K. & Johnson, G.N. (2000) Chlorophyll fluorescence-a practical guide. Journal of Experimental Botany, 51, 659-668.

Meineri, E., Skarpaas, O., Spindelbock, J., Bargmann, T. & Vandvik, V. (2014) Direct and size-dependent effects of climate on flowering performance in alpine and lowland herbaceous species. Journal of Vegetation Science, 25, 275-286.

Meineri, E., Skarpaas, O. & Vandvik, V. (2012) Modeling alpine plant distributions at the landscape scale: do biotic interactions matter? Ecological Modelling, 231, 1-10.

Meineri, E., Spindelbock, J. & Vandvik, V. (2013) Seedling emergence responds to both seed source and recruitment site climates: a climate change experiment combining transplant and gradient approaches. Plant Ecology, 214, 607-619.

Molina-Montenegro, M.A., Salgado-Luarte, C., Oses, R. & Torres-Diaz, C. (2013) Is physiological performance a good predictor for fitness? Insights from an invasive plant species. PLoS One, 8, 9.

Montoro, A. & Sadras, V. (2014) Elevated temperature increased stomata size and leaf photosynthesis in shiraz grapevine. VII International Symposium on Irrigation of Horticultural Crops 1038: 443-447

Moran, E.V., Hartig, F. & Bell, D.M. (2016) Intraspecific trait variation across scales: implications for understanding global change responses. Global Change Biology, 22, 137-150.

Münzbergová, Z., Hadincová, V., Skálová, H. & Vandvik, V. (2017) Genetic differentiation and plasticity interact along temperature and precipitation gradients to determine plant performance under climate change. Journal of Ecology, 105, 1358-1373.

Münzbergová, Z., Kosová, V., Schnáblová, R., Rokaya, M., Synková, H., Haisel, D. et al. (2020) Plant origin, but not phylogeny, drive species ecophysiological response to projected climate. Frontiers in Plant Science, 11, 400.

Münzbergová, Z., Latzel, V., Šurinová, M. & Hadincová, V. (2019) DNA methylation as a possible mechanism affecting ability of natural populations to adapt to changing climate. Oikos, 128, 124-134.

Münzbergová, Z., Vandvik, V. & Hadincová, V. (2021) Evolutionary rescue as a mechanism allowing a clonal grass to adapt to novel climates. Frontiers in Plant Science, 12, 659479.

Nicotra, A.B., Atkin, O.K., Bonser, S.P., Davidson, A.M., Finnegan, E.J., Mathesius, U. et al. (2010) Plant phenotypic plasticity in a changing climate. Trends in Plant Science, 15, 684-692.

Niu, S.L., Li, L.H., Jiang, G.M., Gao, L.M., Li, Y.G., Peng, Y. et al. (2004) Gas exchange and chlorophyll fluorescence response to simulated rainfall in Hedysarum fruticosum var. mongolicumi. Photosynthetica, 42, 1-6.

Ocheltree, T.W., Nippert, J.B. & Prasad, P.V.V. (2014) Stomatal responses to changes in vapor pressure deficit reflect tissue-specific differences in hydraulic conductance. Plant, Cell and Environment, 37, 132-139.

Ougham, H.J., Morris, P. & Thomas, H. (2005) The colors of autumn leaves as symptoms of cellular recycling and defenses against environmental stresses. Current Topics in Developmental Biology, 66, 135-160.

Pavlíková, Z., Holá, D., Vlasáková, B., Procházka, T. & Münzbergová, Z. (2017) Physiological and fitness differences between cytotypes vary with stress in a grassland perennial herb. PLoS One, 12(11), e0188795.

Peng, S.B., Krieg, D.R. & Girma, F.S. (1991) Leaf photosynthetic rate is correlated with biomass and grain production in grain Sorghum lines. Photosynthesis Research, 28, 1-7.

Poorter, H., Niinemets, U., Poorter, L., Wright, I.J. & Villar, R. (2009) Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. The New Phytologist, 182(3), 565-588.

Pratt, J.D. & Mooney, K.A. (2013) Clinical adaptation and adaptive plasticity in Artemisia californica: implications for the response of a foundation species to predicted climate change. Global Change Biology, 19, 2454-2466.

Prud'homme, G.O., Lamhamedi, M.S., Benomar, L., Rainville, A., DeBlois, J., Bousquet, J. et al. (2018) Ecophysiology and growth of white spruce seedlings from various seed sources along a climatic gradient support the need for assisted migration. Frontiers in Plant Science, 8, 2214.

Psidova, E., Ditmarova, L., Jamnicka, G., Kurjak, D., Majerova, J., Czajkowski, T. et al. (2015) Photosynthetic response of beech seedlings of different origin to water deficit. Photosynthetica, 53, 187-194.

Quezada, I.M., Saldana, A. & Gianoli, E. (2017) Divergent patterns of selection on Crassulacean acid metabolism photosynthesis in contrasting environments. International Journal of Plant Sciences, 178, 398-405.

R Development Core Team. (2011) A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing.

Ramirez-Valiente, J.A., Center, A., Sparks, J.P., Sparks, K.L., Etterson, J.R., Longwell, T. et al. (2017) Population-level differentiation in growth rates and leaf traits in seedlings of the neotropical live oak Quercus oleoides grown under natural and manipulated precipitation regimes. Frontiers of Plant Science, 8, 585.

Ramirez-Valiente, J.A., Sanchez-Gomez, D., Aranda, I. & Valladares, F. (2010) Phenotypic plasticity and local adaptation in leaf ecophysiological traits of 13 contrasting cork oak populations under different water availabilities. Tree Physiology, 30, 618-627.

Reinhardt, K., Castanha, C., Germino, M.J. & Kueppers, L.M. (2011) Ecophysiological variation in two provenances of Pinus flexilis seedlings across an elevation gradient from forest to alpine. Tree Physiology, 31, 615-625.

Rice, W.R. (1989) Analyzing tables of statistical tests. Evolution, 43, 223-225.

Robakowski, P., Li, Y. & Reich, P.B. (2012) Local ecotypic and species range-related adaptation influence photosynthetic temperature optima in deciduous broadleaved trees. Plant Ecology, 213, 113-125.

Saldana, A., Lusk, C.H., Gonzales, W.L. & Gianoli, E. (2007) Natural selection on ecophysiological traits of a fern species in a temperate rainforest. Evolutionary Ecology, 21, 651-662.

Shaw, R.G. & Etterson, J.R. (2012) Rapid climate change and the rate of adaptation: insight from experimental quantitative genetics. The New Phytologist, 195, 752-765.

Shi, F.S., Wu, Y., Wu, N. & Luo, P. (2010) Different growth and physiological responses to experimental warming of two dominant plant species Elymus nutans and Potentilla anserina in an alpine meadow of the eastern Tibetan Plateau. Photosynthetica, 48, 437-445.

Song, X., Wang, Y. & Lv, X. (2016) Responses of plant biomass, photosynthesis and lipid peroxidation to warming and precipitation change in two dominant species (Stipa grandis and Leymus chinensis) from North China grasslands. Ecology and Evolution, 6, 1871-1882.

Souther, S., Lechowicz, M.J. & McGraw, J.B. (2012) Experimental test for adaptive differentiation of ginseng populations reveals complex response to temperature. Annals of Botany, 110, 829-837.

Stojanova, B., Kolaříková, V., Šurinová, M., Klápště, J., Hadincová, V. & Münzbergová, Z. (2019) Evolutionary potential of a widespread clonal grass under changing climate. Journal of Evolutionary Biology, 32, 1057-1068.

Stojanova, B., Šurinová, M., Klápště, J., Koláříková, V., Hadincová, V. & Münzbergová, Z. (2018) Adaptive differentiation of Festuca rubra along a climate gradient revealed by molecular markers and quantitative traits. PLoS One, 13(4), e0194670.

Sultan, S.E., Horgan-Kobelski, T., Nichols, L.M., Riggs, C.E. & Waples, R.K. (2013) A resurrection study reveals rapid adaptive evolution within populations of an invasive plant. Evolutionary Applications, 6, 266-278.

Šurinová, M., Hadincová, V., Vandvik, V. & Münzbergová, Z. (2019) Temperature and precipitation, but not geographic distance, explain genetic relatedness among populations in the perennial grass Festuca rubra. Journal of Plant Ecology, 12, 730-741.

Taylor, S.H., Franks, P.J., Hulme, S.P., Spriggs, E., Christin, P.A., Edwards, E.J. et al. (2012) Photosynthetic pathway and ecological adaptation explain stomatal trait diversity amongst grasses. The New Phytologist, 193, 387-396.

Thompson, J.D. (1991) Phenotypic plasticity as a component of evolutionary change. TREE, 6, 246-249.

Turner, N.C. (2018) Turgor maintenance by osmotic adjustment: 40 years of progress. Journal of Experimental Botany, 69, 3223-3233.

Valladares, F., Matesanz, S., Guilhaumon, F., Araujo, M.B., Balaguer, L., Benito-Garzon, M. et al. (2014) The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change. Ecology Letters, 17, 1351-1364.

Vandvik, V., Klanderud, K., Meineri, E., Maren, I.E. & Topper, J. (2016) Seed banks are biodiversity reservoirs: species-area relationships above versus below ground. Oikos, 125, 218-228.

Vandvik, V., Klanderud, K., Skarpaas, O., Telford, R., Halbritter, A. & Goldberg, D. (2020) Biotic rescaling reveals importance of species interactions for variation in biodiversity responses to climate change. PNAS, 17, 22858-22865.

Vasseur, F., Sartori, K., Baron, E., Fort, F., Kazakou, E., Segrestin, J. et al. (2018) Climate as a driver of adaptive variations in ecological strategies in Arabidopsis thaliana. Annals of Botany, 122, 935-945.

Veselá, A., Dostálek, T., Rokaya, M.B. & Münzbergová, Z. (2020) Seed mass and plant home site environment interact to determine alpine species germination patterns along an elevation gradient. Alpine Botany, 130, 101-113.

Vitasse, Y., Lenz, A., Kollas, C., Randin, C.F., Hoch, G. & Korner, C. (2014) Genetic vs. non-genetic responses of leaf morphology and growth to elevation in temperate tree species. Functional Ecology, 28, 243-252.

Vogan, P.J. & Maherali, H. (2014) Increased photosynthetic capacity as a mechanism of drought adaptation in C3 plants. International Journal of Plant Sciences, 175, 1033-1041.

Walisch, T.J., Colling, G., Bodenseh, M. & Matthies, D. (2015) Divergent selection along climatic gradients in a rare central European endemic species, Saxifraga sponhemica. Annals of Botany, 115, 1177-1190.

Wellstein, C., Poschlod, P., Gohlke, A., Chelli, S., Chelli, S., Campetella, G. et al. (2017) Effects of extreme drought on specific leaf area of grassland species: a meta-analysis of experimental studies in temperate and sub-Mediterranean systems. Global Change Biology, 23, 2473-2481.

Wild, J., Kopecky, M., Macek, M., Sanda, M., Jankovec, J. & Haase, T. (2019) Climate at ecologically relevant scales: a new temperature and soil moisture logger for long-term microclimate measurement. Agricultural and Forest Meteorology, 268, 40-47.

Williams, D.G., Mack, R.N. & Black, R.A. (1995) Ecophysiology of introduced Pennisetum setaceum on Hawaii-the role of phenotypic plasticity. Ecology, 76, 1569-1580.

Woodward, F.I., Lake, J.A. & Quick, W.P. (2002) Stomatal development and CO2: ecological consequences. The New Phytologist, 153, 477-484.

Wu, J.S., Wurst, S. & Zhang, X.Z. (2016) Plant functional trait diversity regulates the nonlinear response of productivity to regional climate change in Tibetan alpine grasslands. Scientific Reports, 6, 35649.

Xu, P.B., Zhang, X.W., Zhao, C.M., Chen, L.T., Gao, X.L., Yao, B.Q. et al. (2014) Foliar responses of Abies fargesii Franch. To altitude in the Tabai mountain, China. Polish Journal of Ecology, 62, 481-492.

Xu, Z. & Zhou, G. (2008) Responses of leaf stomatal density to water status and its relationship with photosynthesis in a grass. Journal of Experimental Botany, 59, 3317-3325.

Yamori, W., Hikosaka, K. & Way, D.A. (2014) Temperature response of photosynthesis in C-3, C-4, and CAM plants: temperature acclimation and temperature adaptation. Photosynthesis Research, 119, 101-117.

Yan, W.M., Zhong, Y.Q.W. & Shangguan, Z.P. (2017) Contrasting responses of leaf stomatal characteristics to climate change: a considerable challenge to predict carbon and water cycles. Global Change Biology, 23, 3781-3793.

Yeh, C.H., Kaplinsky, N.J., Hu, C. & Charng, Y.Y. (2012) Some like it hot, some like it warm: phenotyping to explore thermotolerance diversity. Plant Science, 195, 10-23.

Younginger, B.S., Sirova, D., Cruzan, M.B. & Ballhorn, D.J. (2017) Is biomass a reliable estimate of plant fitness? Applications in Plant Sciences, 5(2), 1600094.

Zhang, L., Niu, H., Wang, S., Zhu, X., Luo, C., Li, Y. et al. (2012) Gene or environment? Species-specific control of stomatal density and length. Ecology and Evolution, 2, 1065-1070.

Zhang, Y.Q., Lin, Q., Liu, J.B., Zhang, H.S. & Zhao, C.X. (2011) Effects of drought stress on photosynthetic characteristics and yield of different fertilizer and water types of wheat. Journal of Triticeae Crops, 31, 724-730.

Zuniga-Feest, A., Bustos-Salazar, A., Alves, F., Martinez, V. & Smith-Ramirez, C. (2017) Physiological and morphological responses to permanent and intermittent waterlogging in seedlings of four evergreen trees of temperate swamp forests. Tree Physiology, 37, 779-789.

Effect of DNA methylation, modified by 5-azaC, on ecophysiological responses of a clonal plant to changing climate

Rhizome trait scaling relationships are modulated by growth conditions and are linked to plant fitness