{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T09:08:22Z","timestamp":1789981702636,"version":"4.0.1"},"reference-count":57,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,9,20]],"date-time":"2026-09-20T00:00:00Z","timestamp":1789862400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFF1002000"],"award-info":[{"award-number":["2023YFF1002000"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100012166","name":"National Key Research and Development Program of China","doi-asserted-by":"publisher","award":["2023YFD1201500"],"award-info":[{"award-number":["2023YFD1201500"]}],"id":[{"id":"10.13039\/501100012166","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Key Research and Development Project of the Autonomous Region","award":["2025B02008"],"award-info":[{"award-number":["2025B02008"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Antioxidants"],"abstract":"<jats:p>Global climate change increases concurrent waterlogging and heat waves, threatening crop production. Tomato, a widely cultivated vegetable crop, is susceptible to both stresses, but their combined effects on tomato responses remain poorly understood. This study analyzed the anatomical, physiological, and molecular responses of two tomato genotypes, \u2018NT212\u2019 and \u2018NT606\u2019 (sensitive and tolerant to combined waterlogging and heat stresses), under control (C), waterlogging (W), heat stress (H1, H2, and H3) (34\/27 \u00b0C, 38\/31 \u00b0C, 42\/35 \u00b0C), and combined stress (WH1, WH2, and WH3) for 48 h. \u2018NT212\u2019 exhibited only abaxial epidermal peeling under H2, but severe anatomical degradation including abaxial epidermal peeling and sponge parenchyma cell lysis under WH2. The superoxide anion (O2\u2022\u2212) production rate and PsbS and HSP70 expression were significantly higher in \u2018NT212\u2019 under WH2 than W and H2. Moreover, as the stress intensity increased from WH1 to WH2 and WH3, the O2\u2022\u2212 production rate and PsbS and HSP70 expression were significantly increased in \u2018NT212\u2019. In contrast, \u2018NT606\u2019 did not exhibit obvious cell lysis under any stress treatments. The O2\u2022\u2212 production rate was highest in \u2018NT606\u2019 under WH3 among all the treatments. HSP70 expression was significantly upregulated in \u2018NT606\u2019 under H3 and WH3 compared with the other treatments. Consequently, \u2018NT212\u2019 exhibited severe wilting under the H2, H3, and all combined stresses, while \u2018NT606\u2019 showed severe wilting only under WH3. Thus, combined stress caused synergistic damage exceeding individual stress, with injury intensifying as temperature increased. This study provided vital knowledge for breeding climate-resilient tomato cultivars, especially under combined waterlogging and heat stress.<\/jats:p>","DOI":"10.3390\/antiox15091210","type":"journal-article","created":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T08:03:47Z","timestamp":1789977827000},"page":"1210","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Enhanced Levels of Combined Waterlogging and Heat Stress Exacerbate Anatomical Damage, Photosynthetic Impairment, and Oxidative Stress in Tomato"],"prefix":"10.3390","volume":"15","author":[{"given":"Wen","family":"Qin","sequence":"first","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-7351-1533","authenticated-orcid":false,"given":"Yankai","family":"Li","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yang","family":"Sun","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3880-0241","authenticated-orcid":false,"given":"Rosa M.","family":"Rivero","sequence":"additional","affiliation":[{"name":"Center of Edaphology and Applied Biology of Segura (CEBAS-CSIC), Department of Plant Nutrition, Campus Universitario Espinardo, Ed. 25, 30100 Murcia, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0084-3668","authenticated-orcid":false,"given":"Xiaoming","family":"Song","sequence":"additional","affiliation":[{"name":"School of Life Sciences, North China University of Science and Technology, Tangshan 063000, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ron","family":"Mittler","sequence":"additional","affiliation":[{"name":"Division of Plant Science and Technology, College of Agriculture, Food and Natural Resources, Bond Life Sciences Center, University of Missouri, Columbia, MO 65201, USA"},{"name":"Advanced Agriculture Initiative, Tel-Hai University of Kiryat Shmona in the Galilee, Kiryat Shmona 1220800, Israel"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1112-0580","authenticated-orcid":false,"given":"Fangling","family":"Jiang","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhen","family":"Wu","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dong","family":"Xiao","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuedong","family":"Yang","sequence":"additional","affiliation":[{"name":"Horticulture Research Institute, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5755-1258","authenticated-orcid":false,"given":"Rong","family":"Zhou","sequence":"additional","affiliation":[{"name":"Sanya Research Institute, College of Horticulture, Nanjing Agricultural University, Nanjing 210095, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,20]]},"reference":[{"key":"ref_1","unstructured":"Masson-Delmotte, V., Zhai, P., Pirani, A., Connors, S.L., P\u00e9an, C., Berger, S., Caud, N., Chen, Y., Goldfarb, L., and Gomis, M.I. (2023). Weather and Climate Extreme Events in a Changing Climate. Climate Change 2021\u2014The Physical Science Basis: Working Group I Contribution to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change, Cambridge University Press."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1016\/j.tplants.2021.02.011","article-title":"Global warming, climate change, and environmental pollution: Recipe for a multifactorial stress combination disaster","volume":"26","author":"Zandalinas","year":"2021","journal-title":"Trends Plant Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/j.tplants.2005.11.002","article-title":"Abiotic stress, the field environment and stress combination","volume":"11","author":"Mittler","year":"2006","journal-title":"Trends Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1111\/nph.12797","article-title":"Abiotic and biotic stress combinations","volume":"203","author":"Suzuki","year":"2014","journal-title":"New Phytol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.jplph.2014.11.008","article-title":"The interactive effects of simultaneous biotic and abiotic stresses on plants: Mechanistic understanding from drought and pathogen combination","volume":"176","author":"Ramegowda","year":"2015","journal-title":"J. Plant Physiol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/j.pedsph.2022.06.031","article-title":"Stress combination: When two negatives may become antagonistic, synergistic or additive for plants?","volume":"33","author":"Nadeem","year":"2023","journal-title":"Pedosphere"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1106","DOI":"10.1104\/pp.17.01232","article-title":"Signal dynamics and interactions during flooding stress","volume":"176","author":"Sasidharan","year":"2018","journal-title":"Plant Physiol."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Pan, J., Sharif, R., Xu, X., and Chen, X. (2021). Mechanisms of waterlogging tolerance in plants: Research progress and prospects. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.627331"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"2215","DOI":"10.1093\/plphys\/kiad433","article-title":"Rapid systemic responses of Arabidopsis to waterlogging stress","volume":"193","author":"Tukuli","year":"2023","journal-title":"Plant Physiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1016\/S0098-8472(03)00063-7","article-title":"Antioxidative enzymes in barley plants subjected to soil flooding","volume":"51","author":"Yordanova","year":"2004","journal-title":"Environ. Exp. Bot."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"9643","DOI":"10.3390\/ijms14059643","article-title":"Physiological, biochemical, and molecular mechanisms of heat stress tolerance in plants","volume":"14","author":"Hasanuzzaman","year":"2013","journal-title":"Int. J. Mol. Sci."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1007\/s11120-008-9331-0","article-title":"Heat stress: An overview of molecular responses in photosynthesis","volume":"98","author":"Allakhverdiev","year":"2008","journal-title":"Photosynth. Res."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Hu, S., Ding, Y., and Zhu, C. (2020). Sensitivity and responses of chloroplasts to heat stress in plants. Front. Plant Sci., 11.","DOI":"10.3389\/fpls.2020.00375"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"759","DOI":"10.1007\/s11103-005-5750-x","article-title":"The heat stress transcription factor HsfA2 serves as a regulatory amplifier of a subset of genes in the heat stress response in Arabidopsis","volume":"60","author":"Schramm","year":"2006","journal-title":"Plant Mol. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"405","DOI":"10.1111\/pbi.12659","article-title":"The heat shock protein\/chaperone network and multiple stress resistance","volume":"15","author":"Jacob","year":"2017","journal-title":"Plant Biotechnol. J."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"uhaf309","DOI":"10.1093\/hr\/uhaf309","article-title":"Molecular mechanisms and breeding strategies for heat tolerance in vegetable crops under global warming","volume":"13","author":"Li","year":"2026","journal-title":"Hortic. Res."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Ploschuk, R.A., Savin, R., and Slafer, G.A. (2025). Dual stress, equivalent harm? hypothesizing on the type of interactions between waterlogging and high temperature. Front. Plant Sci., 15.","DOI":"10.3389\/fpls.2024.1472665"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1471","DOI":"10.1104\/pp.109.149815","article-title":"The heat-inducible transcription factor HsfA2 enhances anoxia tolerance in Arabidopsis","volume":"152","author":"Banti","year":"2010","journal-title":"Plant Physiol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1111\/j.1469-8137.2009.02830.x","article-title":"Causes and consequences of variation in leaf mass per area (LMA): A meta-analysis","volume":"182","author":"Poorter","year":"2009","journal-title":"New Phytol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.carbpol.2012.05.015","article-title":"Heat stress causes alterations in the cell-wall polymers and anatomy of coffee leaves (Coffea arabica L.)","volume":"93","author":"Lima","year":"2013","journal-title":"Carbohydr. Polym."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1104\/pp.110.165472","article-title":"Leaf functional anatomy in relation to photosynthesis","volume":"155","author":"Terashima","year":"2011","journal-title":"Plant Physiol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1447","DOI":"10.1111\/nph.14496","article-title":"Physiological and structural tradeoffs underlying the leaf economics spectrum","volume":"214","author":"Onoda","year":"2017","journal-title":"New Phytol."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1556","DOI":"10.1104\/pp.114.237107","article-title":"Stomatal size, speed, and responsiveness impact on photosynthesis and water use efficiency","volume":"164","author":"Lawson","year":"2014","journal-title":"Plant Physiol."},{"key":"ref_24","first-page":"20","article-title":"Systemic effects of the vapor pressure deficit on the physiology and productivity of protected vegetables","volume":"3","author":"Yu","year":"2023","journal-title":"Veg. Res."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1002\/pei3.10117","article-title":"Parallels between drought and flooding: An integrated framework for plant eco-physiological responses to water stress","volume":"4","author":"Chen","year":"2023","journal-title":"Plant-Environ. Interact."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Zait, Y., Joseph, A., and Assmann, S.M. (2024). Stomatal responses to VPD utilize guard cell intracellular signaling components. Front. Plant Sci., 15.","DOI":"10.3389\/fpls.2024.1351612"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1171","DOI":"10.1111\/pce.14797","article-title":"Systemic stomatal responses in plants: Coordinating development, stress, and pathogen defense under a changing climate","volume":"47","author":"Zandalinas","year":"2024","journal-title":"Plant Cell Environ."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"20240241","DOI":"10.1098\/rstb.2024.0241","article-title":"The differential transpiration response of plants to stress","volume":"380","author":"Sinha","year":"2025","journal-title":"Philos. Trans. R. Soc. B"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"104","DOI":"10.1016\/j.bbagrm.2011.10.002","article-title":"The plant heat stress transcription factor (Hsf) family: Structure, function and evolution","volume":"1819","author":"Scharf","year":"2012","journal-title":"Biochim. Biophys. Acta Gene Regul. Mech."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.tplants.2016.08.015","article-title":"Transcriptional regulatory network of plant heat stress response","volume":"22","author":"Ohama","year":"2017","journal-title":"Trends Plant Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1016\/j.tplants.2004.03.006","article-title":"Role of plant heat-shock proteins and molecular chaperones in the abiotic stress response","volume":"9","author":"Wang","year":"2004","journal-title":"Trends Plant Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1007\/s10535-009-0090-3","article-title":"Waterlogging induced oxidative stress and antioxidant enzyme activities in pigeon pea","volume":"53","author":"Sairam","year":"2009","journal-title":"Biol. Plant."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1038\/35000131","article-title":"A pigment-binding protein essential for regulation of photosynthetic light harvesting","volume":"403","author":"Li","year":"2000","journal-title":"Nature"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1126\/science.aai8878","article-title":"Improving photosynthesis and crop productivity by accelerating recovery from photoprotection","volume":"354","author":"Kromdijk","year":"2016","journal-title":"Science"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"112435","DOI":"10.1016\/j.scienta.2023.112435","article-title":"Tomato heat tolerance: Progress and prospects","volume":"322","author":"Liu","year":"2023","journal-title":"Sci. Hortic."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"6230","DOI":"10.1093\/jxb\/eraf203","article-title":"An overview of low-oxygen sensing and flooding responses of tomato","volume":"76","author":"Eerdekens","year":"2025","journal-title":"J. Exp. Bot."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Zhou, R., Niu, L., Yin, J., Jiang, F., Wang, Y., Zhao, T., Wu, Z., and Zhu, W. (2023). Differences in physiological responses of two tomato genotypes to combined waterlogging and cadmium stresses. Antioxidants, 12.","DOI":"10.3390\/antiox12061205"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1309","DOI":"10.1016\/j.jplph.2013.04.007","article-title":"Smaller stomata require less severe leaf drying to close: A case study in Rosa hydrida","volume":"170","author":"Giday","year":"2013","journal-title":"J. Plant Physiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1007\/s11738-018-2781-x","article-title":"Protective effect of a natural ally on simultaneous mild heat and salt episodes in maize seedlings","volume":"40","author":"Pehlivan","year":"2018","journal-title":"Acta Physiol. Plant."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1016\/S0254-6299(16)30702-5","article-title":"Leaf anatomy, chloroplast organization and photosynthetic rate of hyperhydric Eucalyptus saligna Sm. material","volume":"59","author":"Jones","year":"1993","journal-title":"S. Afr. J. Bot."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"602","DOI":"10.1111\/j.1365-3040.2007.01757.x","article-title":"Mesophyll conductance to CO2: Current knowledge and future prospects","volume":"31","author":"Flexas","year":"2007","journal-title":"Plant Cell Environ."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.plantsci.2012.05.009","article-title":"Mesophyll diffusion conductance to CO2: An unappreciated central player in photosynthesis","volume":"193\u2013194","author":"Flexas","year":"2012","journal-title":"Plant Sci."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"20203145","DOI":"10.1098\/rspb.2020.3145","article-title":"Maximum CO2 diffusion inside leaves is limited by the scaling of cell size and genome size","volume":"288","author":"Roddy","year":"2021","journal-title":"Proc. R. Soc. B"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1016\/j.plantsci.2016.06.015","article-title":"Controlling stomatal aperture in semi-arid regions\u2014The dilemma of saving water or being cool?","volume":"251","author":"Chaves","year":"2016","journal-title":"Plant Sci."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.plaphy.2010.08.016","article-title":"Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants","volume":"48","author":"Gill","year":"2010","journal-title":"Plant Physiol. Biochem."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Suzuki, N. (2023). Fine tuning of ROS, redox and energy regulatory systems associated with the functions of chloroplasts and mitochondria in plants under heat stress. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24021356"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1111\/j.1365-3040.2005.01327.x","article-title":"Oxidant and antioxidant signalling in plants: A re-evaluation of the concept of oxidative stress in a physiological context","volume":"28","author":"Foyer","year":"2005","journal-title":"Plant Cell Environ."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1016\/j.molcel.2010.10.006","article-title":"The heat shock response: Life on the verge of death","volume":"40","author":"Richter","year":"2010","journal-title":"Mol. Cell"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Li, Y., Jiang, F., He, Z., Liu, Y., Chen, Z., Ottosen, C.O., Mittler, R., Wu, Z., and Zhou, R. (2024). Higher intensity of salt stress accompanied by heat inhibits stomatal conductance and induces ROS accumulation in tomato plants. Antioxidants, 13.","DOI":"10.3390\/antiox13040448"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1007\/BF00232226","article-title":"Environmental scanning electron microscopy of the surface of normal and vitrified leaves of Gypsophila paniculata (Babies Breath) cultured in vitro","volume":"15","author":"Gribble","year":"1996","journal-title":"Plant Cell Rep."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1016\/j.scienta.2012.08.019","article-title":"Anatomical responses to waterlogging in Chrysanthemum zawadskii","volume":"146","author":"Yin","year":"2012","journal-title":"Sci. Hortic."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"2125","DOI":"10.2135\/cropsci2010.10.0571","article-title":"High-temperature stress and soybean leaves: Leaf anatomy and photosynthesis","volume":"51","author":"Djanaguiraman","year":"2011","journal-title":"Crop Sci."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1146\/annurev.pp.33.060182.001533","article-title":"Stomatal conductance and photosynthesis","volume":"33","author":"Farquhar","year":"1982","journal-title":"Annu. Rev. Plant Physiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1053","DOI":"10.1104\/pp.010357","article-title":"Exceptional sensitivity of Rubisco activase to thermal denaturation in vitro and in vivo","volume":"127","author":"Salvucci","year":"2001","journal-title":"Plant Physiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11120-004-6434-0","article-title":"Inhibition of the repair of photosystem II by oxidative stress in cyanobacteria","volume":"84","author":"Nishiyama","year":"2005","journal-title":"Photosynth. Res."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Demmig-Adams, B., Garab, G., Adams, W. (2014). Non-Photochemical Fluorescence Quenching Across Scales: From Chloroplasts to Plants to Communities. Non-Photochemical Quenching and Energy Dissipation in Plants, Algae and Cyanobacteria, Springer.","DOI":"10.1007\/978-94-017-9032-1"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"2017","DOI":"10.1042\/BCJ20190328","article-title":"Less photoprotection can be good in some genetic and environmental contexts","volume":"476","author":"Stewart","year":"2019","journal-title":"Biochem. J."}],"container-title":["Antioxidants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-3921\/15\/9\/1210\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T08:57:08Z","timestamp":1789981028000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-3921\/15\/9\/1210"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,20]]},"references-count":57,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["antiox15091210"],"URL":"https:\/\/doi.org\/10.3390\/antiox15091210","relation":{},"ISSN":["2076-3921"],"issn-type":[{"value":"2076-3921","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,20]]}}}