Photosynthetica X:X | DOI: 10.32615/ps.2026.012

Drought stress-induced changes in PSII functioning in ecologically contrasting plants: chlorophyll fluorescence study of poikilohydric and homoiohydric species

J. HÁJEK1, A. PUHOVKIN2, 3, 4, 5, J. LANG6, S.I. UL HAQ2, M. BERNATOVÁ1, D. GIORDANO2, J. SEKERÁK JR.2, M. BARTÁK2
1 Department of Forest Protection and Wildlife Management, Faculty of Forestry and Wood Technology, Mendel University, Zemědělská 3, 613 00 Brno, Czech Republic
2 Department of Experimental Biology, Laboratory of Photosynthetic Processes, Faculty of Science, Masaryk University, Kamenice 5, C-13, 625 00 Brno, Czech Republic
3 Institute for Problems of Cryobiology and Cryomedicine, National Academy of Science of Ukraine, Pereyaslavska Str. 23, 61016 Kharkiv, Ukraine
4 National Antarctic Scientific Centre, State Institution, Ministry of Education and Science of Ukraine, Taras Shevchenko Blvd. 16, 01601 Kyiv, Ukraine
5 Department of Geography, Faculty of Science, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic
6 Agricultural Research, Ltd., Zahradní 1, 664 41 Troubsko, Czech Republic

Dehydration reduces photosynthetic performance in leaves and other photosynthetic organs in response to drought. This decline is linked to changes in chlorophyll fluorescence, which reflects reduced photosystem II activity and increased protective non-photochemical quenching (NPQ). This study examined NPQ dynamics during desiccation across diverse species, including lichens, mosses, conifers, crops, and C3 and C4 plants, using the method of NPQ induction and relaxation curves. Gradual dehydration from a full-hydration state (RWC 100%) to complete dryness (RWC 0%) was applied in order to compare poikilohydric and homoiohydric species. NPQmax (light period), NPQrelax (dark period end), and the initial slopes of NPQ rise (α) and decline (β) were evaluated. Relative electron transport rate curves were also assessed. Results showed the above NPQ parameters are highly sensitive to dehydration, suggesting their usefulness - along with standard maximum (FV/FM) and effective quantum yield (ΦPSII) - as early indicators of drought stress in PSII across plant groups.

Additional key words: desiccation; FV, FM; non-photochemical quenching; relative water content.

Received: October 27, 2025; Revised: May 15, 2026; Accepted: May 19, 2026; Prepublished online: June 10, 2026 

Download citation

References

  1. Acebron K., Matsubara S., Jedmowski C. et al.: Diurnal dynamics of nonphotochemical quenching in Arabidopsis npq mutants assessed by solar-induced fluorescence and reflectance measurements in the field. - New Phytol. 229: 2104-2119, 2020. Go to original source...
  2. Banks J.A.: Chlorophyll fluorescence as a tool to identify drought stress in Acer genotypes. - Environ. Exp. Bot. 155: 118-127, 2018. Go to original source...
  3. Barboričová M., Filaček A., Mlynáriková Vysoká D. et al.: Sensitivity of fast chlorophyll fluorescence parameters to combined heat and drought stress in wheat genotypes. - Plant Soil Environ. 68: 309-316, 2022. Go to original source...
  4. Barták M., Hájek J., Halici M.G. et al.: Resistance of primary photosynthesis to photoinhibition in antarctic lichen Xanthoria elegans: photoprotective mechanisms activated during a short period of high light stress. - Plants-Basel 12: 2259, 2023. Go to original source...
  5. Barták M., Hájek J., Orekhova A. et al.: Inhibition of primary photosynthesis in desiccating antarctic lichens differing in their photobionts, thallus morphology, and spectral properties. -Microorganisms 9: 818, 2021. Go to original source...
  6. Barták M., Hazdrová J., Skácelová K., Hájek J.: Dehydration-induced responses of primary photosynthetic processes and spectral reflectance indices in Antarctic Nostoc commune. - Czech Polar Rep. 6: 87-95, 2016. Go to original source...
  7. Barták M., Trnková K., Hansen E.S. et al.: Effect of dehydration on spectral reflectance and photosynthetic efficiency in Umbilicaria arctica and U. hyperborea. - Biol. Plantarum 59: 357-365, 2015. Go to original source...
  8. Beckett R., Minibayeva F., Solhaug K., Roach T.: Photoprotection in lichens: adaptations of photobionts to high light. - Lichenologist 53: 21-33, 2021a. Go to original source...
  9. Beckett R.P., Minibayeva F.V., Mkhize K.W.G.: Shade lichens are characterized by rapid relaxation of non-photochemical quenching on transition to darkness. - Lichenologist 53: 409-414, 2021b. Go to original source...
  10. Calatayud A., Deltoro V.I., Barreno E., del Valle-Tascon S.: Changes in vivo chlorophyll fluorescence quenching in lichen thalli as a function of water content and suggestion of zeaxanthin-associated photoprotection. - Physiol. Plantarum 101: 93-102, 1997. Go to original source...
  11. Carbonera D., Gerotto C., Posocco B. et al.: NPQ activation reduces chlorophyll triplet state formation in the moss Physcomitrella patens. - BBA-Bioenergetics 1817: 1608-1615, 2012. Go to original source...
  12. Cardoso A.A., Brodribb T.J., Lucani C.J. et al.: Coordinated plasticity maintains hydraulic safety in sunflower leaves. - Plant Cell Environ. 41: 2567-2576, 2018. Go to original source...
  13. Dalal V.K., Tripathy B.C.: Water-stress induced downsizing of light-harvesting antenna complex protects developing rice seedlings from photo-oxidative damage. - Sci. Rep.-UK 8: 5955, 2018. Go to original source...
  14. Demmig-Adams B., Adams III W.W.: Photoprotection in an ecological context: the remarkable complexity of thermal energy dissipation. - New Phytol. 172: 11-21, 2006. Go to original source...
  15. Demmig-Adams B., Koh S.-C., Cohu C.M. et al.: Non-photochemical fluorescence quenching in contrasting plant species and environments. - In: Demmig-Adams B., Garab G., Adams W.W., Govindjee (ed.): Non-Photochemical Quenching and Energy Dissipation in Plants, Algae and Cyanobacteria. Advances in Photosynthesis and Respiration. Vol. 40. Pp. 531-552. Springer, Dordrecht 2014. Go to original source...
  16. Evdokimova O.V., Kabashnikova L.F., Savchenko G.E.: Biogenesis of photosynthetic apparatus under the inhibition of energy processes in de-etiolated seedlings of barley (Hordeum vulgare L.). - Biochemistry-Moscow 7: 148-156, 2013. Go to original source...
  17. Flexas J., Medrano H.: Drought-inhibition of photosynthesis in C3 plants: stomatal and non-stomatal limitations revisited. - Ann. Bot.-London 89: 183-189, 2002. Go to original source...
  18. Gerotto C., Alboresi A., Meneghesso A. et al.: Flavodiiron proteins act as safety valve for electrons in Physcomitrella patens. - PNAS 113: 12322-12327, 2016. Go to original source...
  19. Giudici G.N.M.: Combined chlorophyll fluorescence techniques to study environmental impact on the mountain moss Polytrichum commune. - Czech Polar Rep. 11: 161-173, 2021. Go to original source...
  20. Gotarkar D., Doran L., Burns M. et al.: High-throughput analysis of non-photochemical quenching in crops using pulse amplitude modulated chlorophyll fluorometry. - J. Vis. Exp. 185: e63485, 2022. Go to original source...
  21. Granda E., Camarero J.J.: Drought reduces growth and stimulates sugar accumulation: new evidence of environmentally driven non-structural carbohydrate use. - Tree Physiol. 37: 997-1000, 2017. Go to original source...
  22. Hazrati S., Tahmasebi-Sarvestani Z., Modarres-Sanavy S.A.M. et al.: Effects of water stress and light intensity on chlorophyll fluorescence parameters and pigments of Aloe vera L. - Plant Physiol. Biochem. 106: 141-148, 2016. Go to original source...
  23. Heber U., Bilger W., Shuvalov V.A.: Thermal energy dissipation in reaction centres and in the antenna of photosystem II protects desiccated poikilohydric mosses against photo-oxidation. - J. Exp. Bot. 57: 2993-3006, 2006a. Go to original source...
  24. Heber U.: Photoprotection of green plants: a mechanism of ultra-fast thermal energy dissipation in desiccated lichens. - Planta 228: 641-650, 2008. Go to original source...
  25. Heber U., Bilger W., Türk R., Lange O.L.: Photoprotection of reaction centres in photosynthetic organisms: mechanisms of thermal energy dissipation in desiccated thalli of the lichen Lobaria pulmonaria. - New Phytol. 185: 459-470, 2010. Go to original source...
  26. Heber U., Lange O.L., Shuvalov V.A.: Conservation and dissipation of light energy as complementary processes: homoiohydric and poikilohydric autotrophs. - J. Exp. Bot. 57: 1211-1223, 2006b. Go to original source...
  27. Hoegh-Guldberg O., Jacob D., Taylor M. et al.: Impacts of 1.5°C global warming on natural and human systems. - In: Masson-Delmotte V., Zhai P., Pörtner H.-O. et al. (Ed.): Global warming of 1.5°C. An IPCC Special Report on the impacts of global warming of 1.5°C above pre-industrial levels and related global greenhouse gas emission pathways, in the context of strengthening the global response to the threat of climate change, sustainable development, and efforts to eradicate poverty. Pp. 175-312. Cambridge University Press, Cambridge-New York 2018.
  28. Ikeuchi M., Sato F., Endo T.: Allocation of absorbed light energy in photosystem II in NPQ mutants of Arabidopsis. - Plant Cell Physiol. 57: 1484-1494, 2016. Go to original source...
  29. Itam M., Hall D., Kramer D., Merewitz E.: Early detection of Kentucky bluegrass and perennial ryegrass responses to drought stress by measuring chlorophyll fluorescence parameters. - Crop Sci. 64: 1015-1026, 2024. Go to original source...
  30. Kalariya K.A., Singh A.L., Chakraborty K. et al.: Relative water content as an index of permanent wilting in groundnut under progressive water deficit stress. - Electron. J. Environ. Sci. 8: 17-22, 2015.
  31. Khapte P.S., Changan S.S., Kumar P. et al.: Deciphering desiccation tolerance in wild eggplant species: insights from chlorophyll fluorescence dynamics. - BMC Plant Biol. 24: 702, 2024. Go to original source...
  32. Kitao M., Yasuda Y., Kodani E. et al.: Integration of electron flow partitioning improves estimation of photosynthetic rate under various environmental conditions based on chlorophyll fluorescence. - Remote Sens. Environ. 254: 112273, 2021. Go to original source...
  33. Kress E., Jahns P.: The dynamics of energy dissipation and xanthophyll conversion in Arabidopsis indicate an indirect photoprotective role of zeaxanthin in slowly inducible and relaxing components of non-photochemical quenching of excitation energy. - Front. Plant Sci. 8: 2094, 2017. Go to original source...
  34. Lawlor D.W., Cornic G.: Photosynthetic carbon assimilation and associated metabolism in relation to water deficits in higher plants. - Plant Cell Environ. 25: 275-294, 2002. Go to original source...
  35. Lazár D.: Parameters of photosynthetic energy partitioning. - J. Plant Physiol. 175: 131-147, 2015. Go to original source...
  36. le Roux M.-S.L., Burger N.F.V., Vlok M. et al.: EMS derived wheat mutant BIG8-1 (Triticum aestivum L.) - a new drought tolerant mutant wheat line. - Int. J. Mol. Sci. 22: 5314, 2021. Go to original source...
  37. Li X.P., Gilmore A.M., Caffarri S. et al.: Regulation of photosynthetic light harvesting involves intrathylakoid lumen pH sensing by the PsbS protein. - J. Biol. Chem. 279: 22866-22874, 2004. Go to original source...
  38. Li X.-P., Müller-Moulé P., Gilmore A.M., Niyogi K.K.: PsbS-dependent enhancement of feedbackde-excitation protects photosystem II from photoinhibition. - PNAS 99: 15222-15227, 2002. Go to original source...
  39. Liu M., Qi H., Zhang Z.P. et al.: Response of photosynthesis and chlorophyll fluorescence to drought stress in two maize cultivars. - Afr. J. Agr. Res. 7: 4751-4760, 2012. Go to original source...
  40. Lu H.B., Qiao Y.M., Gong X.C. et al.: Influence of drought stress on the photosynthetic characteristics and dry matter accumulation of hybrid millet. - Photosynthetica 53: 306-311, 2015. Go to original source...
  41. Marchin R.M., Backes D., Ossola A. et al.: Extreme heat increases stomatal conductance and drought-induced mortality risk in vulnerable plant species. - Glob. Change Biol. 28: 1133-1146, 2022. Go to original source...
  42. Mena-Petite A., Duñabeitia M.K., Gonzalez-Moro B. et al.: Sequential effects of acidic precipitation and drought on water relations of Pinus radiata seedlings. - J. Plant Physiol. 155: 93-100, 1999. Go to original source...
  43. Meng L.L., Song J.F., Wen J. et al.: Effects of drought stress on fluorescence characteristics of photosystem II in leaves of Plectranthus scutellarioides. - Photosynthetica 54: 414-421, 2016. Go to original source...
  44. Mishra K.B., Rosputimský M., Grieą M., Puhovkin A.: Photoprotective mechanisms activated in Antarctic moss Chorisodontium aciphyllum during desiccation. - Czech Polar Rep. 14: 1-9, 2024. Go to original source...
  45. Mkhize K., Minibayeva F., Beckett R.: Adaptions of photosynthesis in sun and shade in populations of some Afromontane lichens. - Lichenologist 54: 319-329, 2022. Go to original source...
  46. Morales-Sánchez J.Á., Mark K., Souza J.P.S., Niinemets Ü.: Desiccation-rehydration measurements in bryophytes: current status and future insights. - J. Exp. Bot. 73: 4338-4361, 2022. Go to original source...
  47. Murchie E.H., Niyogi K.K.: Manipulation of photoprotection to improve plant photosynthesis. - Plant Physiol. 155: 86-92, 2011. Go to original source...
  48. Nayaka S., Saxena P.: Physiological responses and ecological success of lichen Stereocaulon foliolosum and moss Racomitrium subsecundum growing in same habitat in Himalaya. - Indian J. Fundam. Appl. Life Sci. 4: 167-179, 2014.
  49. Niyogi K.K., Grossman A.R., Björkman O.: Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion. - Plant Cell 10: 1121-1134, 1998. Go to original source...
  50. Orekhova A., Hájek J.: Responses of primary photosynthetic processes to repetitive rehydration differ in two representatives of Svalbard moss flora. - Czech Polar Rep. 12: 103-114, 2022. Go to original source...
  51. Permann C., Pierangelini M., Remias D. et al.: Photophysiological investigations of the temperature stress responses of Zygnema spp. (Zygnematophyceae) from subpolar and polar habitats (Iceland, Svalbard). - Phycologia 61: 299-311, 2022. Go to original source...
  52. Proctor M.C.F., Ligrone R., Duckett J.G.: Desiccation tolerance in the moss Polytrichum formosum: physiological and fine-structural changes during desiccation and recovery. - Ann. Bot.-London 99: 75-93, 2007. Go to original source...
  53. Puhovkin A., Bezsmertna O., Parnikoza I.: Interspecific differences in desiccation tolerance of selected Antarctic lichens: analysis of photosystem II effectivity and quenching mechanisms. - Czech Polar Rep. 12: 31-43, 2022. Go to original source...
  54. Ramakers L.A.I., Harbinson J., Wientjes E., van Amerongen H.: Unravelling the different components of nonphotochemical quenching using a novel analytical pipeline. - New Phytol. 245: 625-636, 2025. Go to original source...
  55. Ruban A.V.: Quantifying the efficiency of photoprotection. - Philos. T. Roy. Soc. B 372: 20160393, 2017. Go to original source...
  56. Sahay S., Grzybowski M., Schnable J.C., Głowacka K. : Genotype-specific nonphotochemical quenching responses to nitrogen deficit are linked to chlorophyll a to b ratios. - J. Plant Physiol. 297: 154261, 2024. Go to original source...
  57. Seleiman M.F., Al-Suhaibani N., Ali N. et al.: Drought stress impacts on plants and different approaches to alleviate its adverse effects. - Plants-Basel 10: 259, 2021. Go to original source...
  58. Singh G.M., Goldberg S., Schaefer D. et al.: Biochemical, gas exchange, and chlorophyll fluorescence analysis of maize genotypes under drought stress reveals important insights into their interaction and homeostasis. - Photosynthetica 60: 376-388, 2022. Go to original source...
  59. Skotnica J., Matouąková M., Nauą J. et al.: Thermoluminescence and fluorescence study of changes in Photosystem II photochemistry in desiccating barley leaves. - Photosynth. Res. 65: 29-40, 2000. Go to original source...
  60. Swoczyna T., Kalaji H.M., Bussotti F. et al.: Environmental stress - what can we learn from chlorophyll a fluorescence analysis in woody plants? A review. - Front. Plant Sci. 13: 1048582, 2022. Go to original source...
  61. ©pundová M., Kučerová Z., Noľková V. et al.: What to choose for estimating leaf water status-spectral reflectance or in vivo chlorophyll fluorescence? - Plant Phenomics 6: 0243, 2024. Go to original source...
  62. Takahashi S., Milward S.E., Fan D.-Y. et al.: How does cyclic electron flow alleviate photoinhibition in Arabidopsis? - Plant Physiol. 149: 1560-1567, 2009. Go to original source...
  63. Tezara W., Mitchell V.J., Driscoll S.D., Lawlor D.W.: Water stress inhibits plant photosynthesis by decreasing coupling factor and ATP. - Nature 401: 914-917, 1999. Go to original source...
  64. Townsend A.J., Saccon F., Giovagnetti V. et al.: The causes of altered chlorophyll fluorescence quenching induction in the Arabidopsis mutant lacking all minor antenna complexes. - BBA-Bioenergetics 1859: 666-675, 2018. Go to original source...
  65. Trueba S., Pan R., Scoffoni C. et al.: Thresholds for leaf damage due to dehydration: declines of hydraulic function, stomatal conductance and cellular integrity precede those for photochemistry. - New Phytol. 223: 134-149, 2019. Go to original source...
  66. Wang B., Yang X., Chen L. et al.: Physiological mechanism of drought-resistant rice coping with drought stress. - J. Plant Growth Regul. 41: 2638-2651, 2022. Go to original source...
  67. Wang Z., Li G., Sun H. et al.: Effects of drought stress on photosynthesis and photosynthetic electron transport chain in young apple tree leaves. - Biol. Open 7: bio035279, 2018. Go to original source...
  68. Woo N.S., Badger M.R., Pogson B.J.: A rapid, non-invasive procedure for quantitative assessment of drought survival using chlorophyll fluorescence. - Plant Methods 4: 27, 2008. Go to original source...
  69. Xu Z.Z., Zhou G.S., Wang Y.L. et al.: Changes in chlorophyll fluorescence in maize plants with imposed rapid dehydration at different leaf ages. - J. Plant Growth Regul. 27: 83-92, 2008. Go to original source...
  70. Xue S., Zang Y., Chen J. et al.: Effects of enhanced UV-B radiation on photosynthetic performance and non-photochemical quenching process of intertidal red macroalgae Neoporphyra haitanensis. - Environ. Exp. Bot. 199: 104888, 2022. Go to original source...
  71. Yamakawa H., Fukushima Y., Itoh S., Heber U.: Three different mechanisms of energy dissipation of a desiccation-tolerant moss serve one common purpose: to protect reaction centres against photo-oxidation. - J. Exp. Bot. 63: 3765-3775, 2012. Go to original source...
  72. Yamakawa H., Itoh S.: Dissipation of excess excitation energy by drought-induced nonphotochemical quenching in two species of drought-tolerant moss: desiccation-induced acceleration of photosystem II fluorescence decay. - Biochemistry 52: 4451-4459, 2013. Go to original source...
  73. Zait Y., Shemer O.E., Cochavi A.: Dynamic responses of chlorophyll fluorescence parameters to drought across diverse plant families. - Physiol. Plantarum 176: e14527, 2024. Go to original source...
  74. Zaks J., Amarnath K., Sylak-Glassman E.J., Fleming G.R.: Models and measurements of energy-dependent quenching. - Photosynth. Res. 116: 389-409, 2013. Go to original source...
  75. Zhang L., Yang C., Liu C.: Revealing the significance of chlorophyll b in the moss Physcomitrium patens by knocking out two functional chlorophyllide a oxygenase. - Photosynth. Res. 158: 171-180, 2023. Go to original source...
  76. Zivcak M., Olsovska K., Brestic M., Slabbert M.M.: Critical temperature derived from the selected chlorophyll a fluorescence parameters of indigenous vegetable species of South Africa treated with high temperature. - In: Photosynthesis Research for Food, Fuel and the Future. Advanced Topics in Science and Technology in China. Pp. 628-632. Springer, Berlin-Heidelberg 2010. Go to original source...
  77. Zlatev Z.: Drought-induced changes in chlorophyll fluorescence of young wheat plants. - Biotechnol. Biotec. Eq. 23: 438-441, 2009. Go to original source...
  78. Zlatev Z., Lidon F.C.: An overview on drought induced changes in plant growth, water relations and photosynthesis. - Emir. J. Food Agric. 24: 57-72, 2012. Go to original source...