A kiszárítás intenzitásának meghatározó szerepe a II-es fotokémiai rendszer helyreállásában fényen és sötétben: az abszcizinsav-kezelés és a xantofill-ciklus gátlásának hatása a Porella platyphylla (L.) Pfeiff. májmohában
Absztrakt
A poikilohidrikus leveles májmoha Porella platyphylla (L.) Pfeiff. kiszáradásra adott fotoszintetikus válaszai kulcsfontosságúak a faj túlélési stratégiájának megértésében. Vizsgálatunk célja az volt, hogy meghatározzuk:
(i) az egyhetes, eltérő relatív páratartalmon végzett kiszárítás milyen mértékben befolyásolja a PSII működésének újranedvesítés utáni helyreállását fényen és sötétben; valamint (ii) az abszcizinsav (ABA) milyen időléptékben járul hozzá a korai fényvédelem stabilizálásához. A regenerációt a rehidratációt követő 1., 24. és 48. órában klorofill-fluoreszcencia paraméterek segítségével követtük nyomon, és a zeaxantin-függő energiadisszipáció szerepét ditiotreitol (DTT) alkalmazásával vizsgáltuk. Eredményeink három eltérő helyreállási tartományt különítettek el. A mérsékelt kiszáradást túlélő minták (32–76% RH), valamint a
természetes kiszáradásnak kitett minták (gyors vízvesztés laboratóriumi levegőn, ~35% RH) optimális kvantumhatásfokuk (Fv/Fm) és effektív kvantumhatásfokuk (ΦPSII) értékeit fényben és sötétben egyaránt szinte teljes mértékben helyreállították 24–48 órán belül. Ezzel szemben az 5% RH mellett történő kiszáradás irreverzibilis PSII-károsodást okozott, és sem fényben, sem sötétben nem következett be regeneráció. A DTT jelentősen csökkentette a nem-fotokémiai kioltást (NPQ), megerősítve a zeaxantin-függő, energiadependens kioltás (qE) központi szerepét a fényvédelemben, továbbá kimutatva egy DTT-re nem érzékeny NPQ frakció fennmaradását is. Vizsgálatunk egyik kulcsfontosságú eredménye, hogy az ABA-kezelés már 1 órán belül jelentősen stabilizálta a PSII működését mind fényben, mind sötétben, ami a kezeletlen mintákhoz képest magasabb Fv/Fm- és ΦPSII-értékekben nyilvánult meg. Ez a hatás tartósnak bizonyult, és a 24–48
órás helyreállási szakasz során is megfigyelhető maradt, összhangban a mérsékelt kiszáradást elszenvedő minták újranedvesedési mintázatával. Eredményeink új, integrált megközelítést nyújtanak a P. platyphylla kiszáradási–újranedvesedési ciklusai során működő regenerációs mechanizmusok megértéséhez, kiemelve a korai ABA-jelátvitel és a qE-domináns NPQ meghatározó szerepét a PSII mérsékelt dehidratációt követő helyreállásában.
Hivatkozások
Bartels D., Giarola V., Chandler J. 2025: Unravelling the molecular network of desiccation tolerance in resurrection plants started with the model plant Craterostigma plantagineum. Planta 262(2): 37. https://doi.org/10.1007/s00425-025-04752-8
Beckett R. P., Csintalan Zs., Tuba Z. 2000: ABA treatment increases both the desiccation tolerance of photosynthesis, and nonphotochemical quenching in the moss Atrichum undulatum. Plant Ecology 151(1): 65–71. https://doi.org/10.1023/A:1026546707649
Bukhov N., Heber U., Wiese C., Shuvalov V. A. 2001: Energy dissipation in photosynthesis: Does the quenching of chlorophyll fluorescence originate from antenna complexes of photosystem II or from the reaction center? Planta 212: 749–758. https://doi.org/10.1007/s004250000486
Ćosić M., Vujičić M. M., Sa bovljević M. S., Sabovljević A. D. 2020: Effects of ABA and NaCl on physiological responses in selected bryophyte species. Botany 98(11): 639–650. https://doi.org/10.1139/cjb-2020-0041
Demming-Adams B., Adams W. W. 1996: The role of xanthophyll cycle carotenoids in the protection of hotosynthesis. Trends in Plant Science 1(1): 21–26. https://doi.org/10.1016/S1360-1385(96)80019-7
Dinakar C., Bartels D. 2013: Desiccation tolerance in resurrection plants: new insights from transcriptome, proteome and metabolome analysis. Frontiers in Plant Science 4: 482. https://doi.org/10.3389/fpls.2013.00482
Esteban R., Pollastri S., Brilli F., García-Plazaola J. I., Odriozola I., Michelozzi M., Loreto F. 2024: Dehydration and rehydration differently affect photosynthesis and volatile monoterpenes in bryophytes with contrasting ecological traits. Physiologia Plantarum 176(3): e14395. https://doi.org/10.1111/ppl.14395
Fernández-Marín B., Balaguer L., Esteban R., Becerril J. M., García-Plazaola J. I. 2009: Dark induction of the photoprotective xanthophyll cycle in response to dehydration. Journal of Plant Physiology 166(16): 1734–1744. https://doi.org/10.1016/j.jplph.2009.04.019
Fernández-Marín B., Kranner I., Sebastián M. S., Artetxe U., Laza J. M., Vilas J. L., Pritchard H. W., Nadajaran J., Míguez F., Becerril J. M., García-Plazaola J. I. 2013: Evidence for the absence of enzymatic reactions in the glassy state. A case study of xanthophyll cycle pigments in the desiccation-tolerant moss Syntrichia ruralis. Journal of Experimental Botany 64(10): 3033–3043. https://doi.org/10.1093/jxb/ert145
Fernández-Marín B., Míguez F., Becerril J. M., García-Plazaola J. I. 2011: Activation of violaxanthin cycle in darkness is a common response to different abiotic stresses: a case study in Pelvetia canaliculata. BMC Plant Biology 11(1): 181. https://doi.org/10.1186/1471-2229-11-181
Gao B., Li X., Liang Y., Chen M., Liu H., Liu Y., Wang J., Zhang J., Oliver M. J., Zhang D. 2024: Drying without dying: A genome database for desiccation-tolerant plants and evolution of desiccation tolerance. Plant Physiology 194(4): 2249–2262. https://doi.org/10.1093/plphys/kiad672
Heber U. 2008: Photoprotection of green plants: a mechanism of ultra-fast thermal energy dissipation in desiccated lichens. Planta 228: 641–650. https://doi.org/10.1007/s00425-008-0766-5
Heber U., Bilger W., Shuvalov V. A. 2006: Thermal energy dissipation in reaction centres and in the antenna of photosystem II protects desiccated poikilohydric mosses against photooxidation. Journal of Experimental Botany 57(12): 2993–3006. https://doi.org/10.1093/jxb/erl058
Horton P., Ruban A. 2005: Molecular design of the photosystem II light-harvesting antenna: photosynthesis and photoprotection. Journal of Experimental Botany 56(411): 365–373. https://doi.org/10.1093/jxb/eri023
Jabłońska E., Kotowski W., Soudzilovskaia N. A. 2023: Desiccation avoidance and hummock formation traits of rich fen bryophytes. Wetlands 43(3): 21. https://doi.org/10.1007/s13157-023-01669-4
Jahan A., Yamazaki Y., Islam M., Ghosh T. K., Yoshimura N., Kato H., Ishizaki K., Shinozawa A., Sakata Y., Takezawa D. 2022: Differential regulations of abscisic acid-induced desiccation tolerance and vegetative dormancy by group B3 Raf kinases in liverworts. Frontiers in Plant Science 13: 952820. https://doi.org/10.3389/fpls.2022.952820
Lu D., Zhang Y., Zhang A., Lu C. 2022: Non-photochemical quenching: from light perception to photoprotective gene expression. International Journal of Molecular Sciences 23(2): 687. https://doi.org/10.3390/ijms23020687
Marschall M., Beckett R. P. 2005: Photosynthetic responses in the inducible mechanisms of desiccation tolerance of a liverwort and a moss. Acta Biologica Szegediensis 49(1–2): 155–156.
Marschall M., Borbély P. 2011: Photosynthetic responses of the desiccation intolerant Sphagnum angustifolium in relation to increasing its desiccation tolerance by exogenous ABA. Acta Biologica Szegediensis 55(1): 119–121.
Marschall M., Proctor M. C. F. 1999: Desiccation tolerance and recovery of the leafy liverwort Porella platyphylla (L.) Pfeiff.: chlorophyll-fluorescence measurements. Journal of Bryology 21(4): 257–262. https://doi.org/10.1179/jbr.1999.21.4.257
Marschall M., Proctor M. C. F., Smirnoff N. 1998: Carbohydrate composition and invertase activity of the leafy liverwort Porella platyphylla. The New Phytologist 138(2): 343–353. https://doi.org/10.1046/j.1469-8137.1998.00102.x
Marschall M., Sütő Sz. 2022: Effects of desiccation rate and inhibition of protein synthesis and the violaxanthin cycle on the rewetting recovery of Porella platyphylla. The Bryologist 125(4): 602–625. https://doi.org/10.1639/0007-2745-125.4.602
Mayaba N., Beckett R. P., Csintalan Zs., Tuba Z. 2001: ABA increases the desiccation tolerance of photosynthesis in the Afromontane understorey moss Atrichum androgynum. Annals of Botany 88(6): 1093–1100. https://doi.org/10.1006/anbo.2001.1552
Morales-Sánchez J. Á., Mark K., Souza J. P. S., Niinement Ü. 2022: Desiccation–rehydration measurements in bryophytes: current status and future insights. Journal of Experimental Botany 73(13): 4338–4361. https://doi.org/10.1093/jxb/erac172
Nava-Nolazco R. M., Ríos-Melendez S., Galván-Gordillo S. V., Martínez-Navarro A. C., Sánchez-Pérez M., Chavez-Santoscoy R. A., Bibbins-Martinez M., Maldonado-Mendoza I. E., Arroyo-Becerra A., Villalobos-López M. A. 2025: Selection and validation of reference genes for RT-qPCR in protonemal tissue of the desiccation-tolerant moss Pseudocrossidium replicatum under multiple abiotic stress conditions. Plants 14(12): 1752. https://doi.org/10.3390/plants14121752
Ndhlovu N. T., Zulu S. B., Khuzwayo T. N., Minibayeva F. V., Beckett R. P. 2025: Short-term effects of combined light and chilling stress on non-photochemical quenching in bryophytes. Journal of Bryology 47(4): 247–256. https://doi.org/10.1080/03736687.2025.2554445
Nibau C., Van de Koot W., Spiliotis D., Williams K., Kramaric T., Beckmann M., Mur L., Hiwatashi Y., Doonan J. H. 2022: Molecular and physiological responses to desiccation indicate the abscisic acid pathway is conserved in the peat moss, Sphagnum. Journal of Experimental Botany 73(13): 4576–4591. https://doi.org/10.1093/jxb/erac133
Niu Q., Zhao L., Dang Y., Guo R., Deng K., Xiao L. 2025: Bryophyte ELIPs as evolutionary guardians: engineering chloroplast resilience and ABA‐enhanced drought tolerance in crops. Plant, Cell & Environment 48(11): 7948–7960. https://doi.org/10.1111/pce.70094
Oliver M. J., Farrant J. M., Hilhorst H. W. M., Mundree S., Williams B., Bewley J. D. 2020: Desiccation tolerance: avoiding cellular damage during drying and rehydration. Annual Review of Plant Biology 71(1): 435–460. https://doi.org/10.1146/annurev-arplant-071219-105542
Peng Y., Ma T., Wang X., Zhang M., Xu Y., Wei J., Wei S., Li J. 2023: Proteomic and transcriptomic responses of the desiccation-tolerant moss Racomitrium canescens in the rapid rehydration processes. Genes 14(2): 390. https://doi.org/10.3390/genes14020390
Perera-Castro A. V., Flexas J. 2022: Desiccation tolerance in bryophytes relates to elasticity but is independent of cell wall thickness and photosynthesis. Physiologia Plantarum 174(2): e13661. https://doi.org/10.1111/ppl.13661
Pressel S., Duckett J. G. 2010: Cytological insights into the desiccation biology of a model system: moss protonemata. New Phytologist 185(4): 944–963. https://doi.org/10.1111/j.1469-8137.2009.03148.x
Proctor M. C. 2000a: The bryophyte paradox: tolerance of desiccation, evasion of drought. Plant Ecology 151: 41–49. https://doi.org/10.1023/A:1026517920852
Proctor M. C. F. 2000b: Physiological ecology. In: Shaw A. J., Goffinet B. (eds) Bryophyte biology. Cambridge University Press, Cambridge, pp. 225–247. http://dx.doi.org/10.1017/CBO9781139171304.009
Proctor M. 2001: Patterns of desiccation tolerance and recovery in bryophytes. Plant Growth Regulation 35: 147–156. https://doi.org/10.1023/A:1014429720821
Proctor M. C. F., Smirnoff N. 2000: Rapid recovery of photosystems on rewetting desiccation‐tolerant mosses: chlorophyll fluorescence and inhibitor experiments. Journal of Experimental Botany 51(351): 1695–1704. https://doi.org/10.1093/jexbot/51.351.1695
R Core Team 2025: R : A language and environment for statistical computing. R Foundation for Statistical Computing, https://www.r-project.org/
Rathnayake K. N., Nelson S., Seeve C., Oliver M. J., Koster K. L. 2019: Acclimation and endogenous abscisic acid in the moss Physcomitrella patens during acquisition of desiccation tolerance. Physiologia Plantarum 167(3): 317–329. https://doi.org/10.1111/ppl.12892
Stark L. R. 2017: Ecology of desiccation tolerance in bryophytes: A conceptual framework and methodology. The Bryologist 120(2): 130–165. https://doi.org/10.1639/0007-2745-120.2.130
Stark L. R., Greenwood J. L., Brinda J. C. 2022: How to dry a bryophyte: A review and experimental test of four methods to induce desiccation tolerance. The Bryologist 125: 1–22. https://doi.org/10.1639/0007-2745-125.1.001
Stark L. R., Greenwood J. L., Brinda J. C., Oliver M. J. 2014: Physiological history may mask the inherent inducible desiccation tolerance strategy of the desert moss Crossidium crassinerve. Plant Biology 16(5): 935–946. https://doi.org/10.1111/plb.12140
Sun Y., Pri-Tal O., Michaeli D., Mosquna A. 2020: Evolution of abscisic acid signaling module and its perception. Frontiers in Plant Science 11: 934. https://doi.org/10.3389/fpls.2020.00934
Takezawa D., Komatsu K., Sakata Y. 2011: ABA in bryophytes: how a universal growth regulator in life became a plant hormone? Journal of Plant Research 124(4): 437–453. https://doi.org/10.1007/s10265-011-0410-5
Verhoeven A. S., Berkowitz J. M., Walton B. N., Berube B. K., Willour J. J., Polich S. B. 2021: Is zeaxanthin needed for desiccation tolerance? Sustained forms of thermal dissipation in tolerant versus sensitive bryophytes. Physiologia Plantarum 171(3): 453–467. https://doi.org/10.1111/ppl.13263
Winter K., Königer M. 1989: Dithiothreitol, an inhibitor of violaxanthin de-epoxidation, increases the susceptibility of leaves of Nerium oleander L. to photoinhibition of photosynthesis. Planta 180(1): 24–31. https://doi.org/10.1007/BF02411406
Wood A. J. 2007: The nature and distribution of vegetative desiccation-tolerance in hornworts, liverworts and mosses. The Bryologist 110(2): 163–177. https://doi.org/10.1639/0007-2745(2007)110[163:ienfib]2.0.co;2
Xiao L., Zhao L., Cao X., Li X., Niu Q., Dang Y., Oliver M. J., He Y. 2026: Surviving dehydration: shutdown–restart dynamics in Physcomitrium patens avoids the metabolic collapse seen in Arabidopsis. New Phytologist 249(3): 1283–1302. https://doi.org/10.1111/nph.70703
Xiao L., Yobi A., Koster K. L., He Y., Oliver M. J. 2018: Desiccation tolerance in Physcomitrella patens: rate of dehydration and the involvement of endogenous abscisic acid (ABA). Plant, Cell & Environment 41(1): 275–284. https://doi.org/10.1111/pce.13096
Yang R., Li X., Yang Q., Zhao M., Bai W., Liang Y., Liu X., Gao B., Zhang D. 2023: Transcriptional profiling analysis providing insights into desiccation tolerance mechanisms of the desert moss Syntrichia caninervis. Frontiers in Plant Science 14: 1127541. https://doi.org/10.3389/fpls.2023.1127541
Zimran G., Shpilman M., Hobson E., Kamisugi Y., Baichman-Kass A., Zhang H., RuizPartida R., González-Bermúdez M. R., Azar M., Feuer E., Gal M., Lozano-Juste J., de Vries J., Cuming A. C., Mosquna A., Sun Y. 2025: Abscisic acid receptors functionally converge across 500 million years of land plant evolution. Current Biology 35(4): 818–830. https://doi.org/10.1016/j.cub.2024.12.043
Zuo G. 2025: Non‐photochemical quenching (NPQ) in photoprotection: insights into NPQ levels required to avoid photoinactivation and photoinhibition. New Phytologist 246(5): 1967–1974. https://doi.org/10.1111/nph.70121








