{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:17:03Z","timestamp":1788221823660,"version":"build-2803163510"},"reference-count":69,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,6,15]],"date-time":"2026-06-15T00:00:00Z","timestamp":1781481600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"vor","delay-in-days":78,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100004281","name":"National Science Centre of Poland","doi-asserted-by":"crossref","award":["OPUS 2019\/35\/B\/NZ9\/01567"],"award-info":[{"award-number":["OPUS 2019\/35\/B\/NZ9\/01567"]}],"id":[{"id":"10.13039\/501100004281","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100004281","name":"National Science Centre of Poland","doi-asserted-by":"crossref","award":["OPUS 2019\/35\/B\/NZ9\/01567"],"award-info":[{"award-number":["OPUS 2019\/35\/B\/NZ9\/01567"]}],"id":[{"id":"10.13039\/501100004281","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100004281","name":"National Science Centre of Poland","doi-asserted-by":"crossref","award":["OPUS 2019\/35\/B\/NZ9\/01567"],"award-info":[{"award-number":["OPUS 2019\/35\/B\/NZ9\/01567"]}],"id":[{"id":"10.13039\/501100004281","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    The chemically diverse group of nonsteroidal anti-inflammatory drugs (NSAIDs) is a significant source of aquatic pollution, inducing oxidative stress in algae. The aim of this study was to evaluate the antioxidant response of\n                    <jats:italic>Chlamydomonas reinhardtii<\/jats:italic>\n                    exposed to four NSAIDs differing in toxicity and structure: flufenamic acid (FFA), nabumetone (NBT), ibuprofen (IBU), and naproxen (NPX). This study demonstrated that all pharmaceuticals significantly increased H\u2082O\u2082 production, confirming redox imbalance in cells. The antioxidant defense also showed compartment- and compound-specific signatures depending on the NSAID toxicity level. Less toxic IBU and NPX induced coordinated SOD isoforms and catalase activation, while more toxic FFA and NBT triggered chloroplast-targeted H\u2082O\u2082 scavenging via APX pathways. Notably,\n                    <jats:italic>MSD3<\/jats:italic>\n                    transcript levels increased in all treatments, indicating its potential as an NSAID stress biomarker. IBR analysis demonstrated that antioxidant efficiency decreased with increasing NSAID toxicity. These findings demonstrate that NSAID toxicity shapes compartment- and isoform-specific antioxidant strategies in\n                    <jats:italic>C. reinhardtii.<\/jats:italic>\n                    We believe that future studies with a broader range of NSAIDs would enable us to investigate subcellular redox dynamics using compartment-specific ROS reporters and to identify NSAID-sensitive biomarkers for aquatic ecotoxicology monitoring.\n                  <\/jats:p>","DOI":"10.1038\/s41598-026-57688-8","type":"journal-article","created":{"date-parts":[[2026,6,15]],"date-time":"2026-06-15T05:05:16Z","timestamp":1781499916000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Differential antioxidant pathways displayed by Chlamydomonas reinhardtii exposed to selected NSAIDs"],"prefix":"10.1038","volume":"16","author":[{"given":"Dominika","family":"Kapu\u015bci\u0144ska","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Monika","family":"Hejna","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wojciech","family":"Pokora","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna","family":"Aksmann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,15]]},"reference":[{"key":"57688_CR1","doi-asserted-by":"publisher","first-page":"757","DOI":"10.1289\/ehp.5947","volume":"111","author":"CG Daughton","year":"2003","unstructured":"Daughton, C. G. Cradle-to-cradle stewardship of drugs for minimizing their environmental disposition while promoting human health. I. Rationale for and avenues toward a green pharmacy. Environ. Health Perspect. 111, 757 (2003).","journal-title":"Environ. Health Perspect."},{"key":"57688_CR2","doi-asserted-by":"publisher","first-page":"7717","DOI":"10.3390\/ijerph19137717","volume":"19","author":"M Hejna","year":"2022","unstructured":"Hejna, M., Kapu\u015bci\u0144ska, D. & Aksmann, A. Pharmaceuticals in the aquatic environment: a review on eco-toxicology and the remediation potential of algae. Int. J. Environ. Res. Public. Health. 19, 7717 (2022).","journal-title":"Int. J. Environ. Res. Public. Health"},{"key":"57688_CR3","doi-asserted-by":"publisher","first-page":"115891","DOI":"10.1016\/j.envpol.2020.115891","volume":"273","author":"K \u015awiacka","year":"2021","unstructured":"\u015awiacka, K., Michnowska, A., Maculewicz, J., Caban, M. & Smolarz, K. Toxic effects of NSAIDs in non-target species: A review from the perspective of the aquatic environment. Environ. Pollut. 273, 115891 (2021).","journal-title":"Environ. Pollut"},{"key":"57688_CR4","doi-asserted-by":"publisher","first-page":"140043","DOI":"10.1016\/j.scitotenv.2020.140043","volume":"740","author":"M Parolini","year":"2020","unstructured":"Parolini, M. Toxicity of the non-steroidal anti-inflammatory drugs (NSAIDs) acetylsalicylic acid, paracetamol, diclofenac, ibuprofen and naproxen towards freshwater invertebrates: A review. Sci. Total Environ. 740, 140043 (2020).","journal-title":"Sci. Total Environ."},{"key":"57688_CR5","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1021\/acs.jafc.1c06499","volume":"70","author":"LM Madikizela","year":"2022","unstructured":"Madikizela, L. M., Botha, T. L., Kamika, I. & Msagati, T. A. M. Uptake, occurrence, and effects of nonsteroidal anti-inflammatory drugs and analgesics in plants and edible crops. J. Agric. Food Chem. 70, 34\u201345 (2022).","journal-title":"J. Agric. Food Chem."},{"key":"57688_CR6","doi-asserted-by":"publisher","first-page":"136176","DOI":"10.1016\/j.scitotenv.2019.136176","volume":"707","author":"H Wang","year":"2020","unstructured":"Wang, H. et al. Photosynthetic toxicity of non-steroidal anti-inflammatory drugs (NSAIDs) on green algae Scenedesmus obliquus. Sci. Total Environ. 707, 136176 (2020).","journal-title":"Sci. Total Environ."},{"key":"57688_CR7","doi-asserted-by":"publisher","first-page":"523","DOI":"10.1111\/tpj.12780","volume":"82","author":"MA Scranton","year":"2015","unstructured":"Scranton, M. A., Ostrand, J. T., Fields, F. J. & Mayfield, S. P. Chlamydomonas as a model for biofuels and bio-products production. Plant. J. 82, 523\u2013531 (2015).","journal-title":"Plant. J."},{"key":"57688_CR8","doi-asserted-by":"publisher","first-page":"108015","DOI":"10.1016\/j.biombioe.2025.108015","volume":"200","author":"A Singh","year":"2025","unstructured":"Singh, A. et al. An exploration of lipid remodeling by microalga Chlamydomonas reinhardtii in autotrophic and mixotrophic cultivation for bioenergy prospects. Biomass Bioenerg. 200, 108015 (2025).","journal-title":"Biomass Bioenerg"},{"key":"57688_CR9","doi-asserted-by":"publisher","first-page":"122256","DOI":"10.1016\/j.renene.2024.122256","volume":"240","author":"A Singh","year":"2025","unstructured":"Singh, A., Singh, S., Singh, S., Prasad, N. & Asthana, R. K. Harnessing bioenergy potential of Chlamydomonas reinhardtii: a comprehensive characterization and valorization of biomass towards energy conversion under circular bioeconomy. Renew. Energy. 240, 122256 (2025).","journal-title":"Renew. Energy"},{"key":"57688_CR10","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1016\/j.nbt.2026.04.005","volume":"93","author":"JW Ahn","year":"2026","unstructured":"Ahn, J. W. & Choi, J. il. Recent advances in Chlamydomonas reinhardtii as a green factory for sustainable biofuels and bioproduct production. N Biotechnol. 93, 362\u2013377 (2026).","journal-title":"N Biotechnol."},{"key":"57688_CR11","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1016\/S0147-6513(03)00141-6","volume":"59","author":"M Cleuvers","year":"2004","unstructured":"Cleuvers, M. Mixture toxicity of the anti-inflammatory drugs diclofenac, ibuprofen, naproxen, and acetylsalicylic acid. Ecotoxicol. Environ. Saf. 59, 309\u2013315 (2004).","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"57688_CR12","doi-asserted-by":"publisher","first-page":"989","DOI":"10.1016\/j.chemosphere.2018.06.156","volume":"209","author":"M Majewska","year":"2018","unstructured":"Majewska, M., Harshkova, D., Gu\u015bciora, M. & Aksmann, A. Phytotoxic activity of diclofenac: Evaluation using a model green alga Chlamydomonas reinhardtii with atrazine as a reference substance. Chemosphere 209, 989\u2013997 (2018).","journal-title":"Chemosphere"},{"key":"57688_CR13","doi-asserted-by":"publisher","first-page":"140853","DOI":"10.1016\/j.chemosphere.2023.140853","volume":"349","author":"D Kapu\u015bci\u0144ska","year":"2024","unstructured":"Kapu\u015bci\u0144ska, D., Narajczyk, M., Liakh, I., Wielgomas, B. & Aksmann, A. Nabumetone and flufenamic acid pose a serious risk to aquatic plants: a study with Chlamydomonas reinhardtii as a model organism. Chemosphere 349, 140853 (2024).","journal-title":"Chemosphere"},{"key":"57688_CR14","doi-asserted-by":"publisher","first-page":"106455","DOI":"10.1016\/j.aquatox.2023.106455","volume":"257","author":"M Seoane","year":"2023","unstructured":"Seoane, M., Conde-P\u00e9rez, K., Esperanza, M., Cid, \u00c1. & Rioboo, C. Unravelling joint cytotoxicity of ibuprofen and oxytetracycline on Chlamydomonas reinhardtii using a programmed cell death-related biomarkers panel. Aquat. Toxicol. 257, 106455 (2023).","journal-title":"Aquat. Toxicol."},{"key":"57688_CR15","doi-asserted-by":"publisher","first-page":"164","DOI":"10.1016\/j.biortech.2017.04.018","volume":"238","author":"T Ding","year":"2017","unstructured":"Ding, T. et al. Biodegradation of naproxen by freshwater algae Cymbella sp. and Scenedesmus quadricauda and the comparative toxicity. Bioresour Technol. 238, 164\u2013173 (2017).","journal-title":"Bioresour Technol."},{"key":"57688_CR16","doi-asserted-by":"publisher","first-page":"743","DOI":"10.1016\/S0006-2952(98)00330-X","volume":"57","author":"R Moreno-S\u00e1nchez","year":"1999","unstructured":"Moreno-S\u00e1nchez, R. et al. Inhibition and uncoupling of oxidative phosphorylation by nonsteroidal anti-inflammatory drugs. Biochem. Pharmacol. 57, 743\u2013752 (1999).","journal-title":"Biochem. Pharmacol."},{"key":"57688_CR17","doi-asserted-by":"publisher","first-page":"111630","DOI":"10.1016\/j.ecoenv.2020.111630","volume":"208","author":"M Majewska","year":"2021","unstructured":"Majewska, M. et al. Does diclofenac act like a photosynthetic herbicide on green algae? Chlamydomonas reinhardtii synchronous culture-based study with atrazine as reference. Ecotoxicol. Environ. Saf. 208, 111630 (2021).","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"57688_CR18","doi-asserted-by":"publisher","first-page":"111954","DOI":"10.1016\/j.ecoenv.2021.111954","volume":"211","author":"X Zhong","year":"2021","unstructured":"Zhong, X., Zhu, Y., Wang, Y., Zhao, Q. & Huang, H. Effects of three antibiotics on growth and antioxidant response of Chlorella pyrenoidosa and Anabaena cylindrica. Ecotoxicol. Environ. Saf. 211, 111954 (2021).","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"57688_CR19","doi-asserted-by":"publisher","first-page":"1056","DOI":"10.1111\/j.1365-3040.2005.01327.x","volume":"28","author":"CH Foyer","year":"2005","unstructured":"Foyer, C. H. & Noctor, G. Oxidant and antioxidant signalling in plants: a re-evaluation of the concept of oxidative stress in a physiological context. Plant. Cell. Environ. 28, 1056\u20131071 (2005).","journal-title":"Plant. Cell. Environ."},{"key":"57688_CR20","doi-asserted-by":"publisher","first-page":"687","DOI":"10.1093\/plphys\/kiab355","volume":"187","author":"S Wakao","year":"2021","unstructured":"Wakao, S. & Niyogi, K. K. Chlamydomonas as a model for reactive oxygen species signaling and thiol redox regulation in the green lineage. Plant. Physiol. 187, 687\u2013698 (2021).","journal-title":"Plant. Physiol."},{"key":"57688_CR21","doi-asserted-by":"crossref","unstructured":"Pokora, W., Tu\u0142odziecki, S., Dettlaff-Pokora, A. & Aksmann, A. Cross talk between hydrogen peroxide and nitric oxide in the unicellular green algae cell cycle: how does it work? Cells 11, (2022).","DOI":"10.3390\/cells11152425"},{"key":"57688_CR22","doi-asserted-by":"publisher","first-page":"124619","DOI":"10.1016\/j.jhazmat.2020.124619","volume":"410","author":"X Xin","year":"2021","unstructured":"Xin, X., Huang, G. & Zhang, B. Review of aquatic toxicity of pharmaceuticals and personal care products to algae. J. Hazard. Mater. 410, 124619 (2021).","journal-title":"J. Hazard. Mater."},{"key":"57688_CR23","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1104\/pp.106.088609","volume":"143","author":"MD Allen","year":"2007","unstructured":"Allen, M. D. et al. Manganese deficiency in Chlamydomonas results in loss of photosystem II and MnSOD function, sensitivity to peroxides, and secondary phosphorus and iron deficiency. Plant. Physiol. 143, 263\u2013277 (2007).","journal-title":"Plant. Physiol."},{"key":"57688_CR24","doi-asserted-by":"crossref","unstructured":"Rajput, V. D. et al. Recent Developments in enzymatic antioxidant defence mechanism in plants with special reference to abiotic stress. Biology 10, 10 (2021).","DOI":"10.3390\/biology10040267"},{"key":"57688_CR25","doi-asserted-by":"publisher","first-page":"31","DOI":"10.1016\/j.ecoenv.2014.08.005","volume":"110","author":"A Aksmann","year":"2014","unstructured":"Aksmann, A. et al. Time-dependent changes in antioxidative enzyme expression and photosynthetic activity of Chlamydomonas reinhardtii cells under acute exposure to cadmium and anthracene. Ecotoxicol. Environ. Saf. 110, 31\u201340 (2014).","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"57688_CR26","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-020-70247-z","volume":"10","author":"EYH Kuo","year":"2020","unstructured":"Kuo, E. Y. H., Cai, M. S. & Lee, T. M. Ascorbate peroxidase 4 plays a role in the tolerance of Chlamydomonas reinhardtii to photo-oxidative stress. Sci. Rep. 10, 1\u201312 (2020).","journal-title":"Sci. Rep."},{"key":"57688_CR27","doi-asserted-by":"publisher","first-page":"277","DOI":"10.3390\/antiox10020277","volume":"10","author":"S Sachdev","year":"2021","unstructured":"Sachdev, S., Ansari, S. A., Ansari, M. I., Fujita, M. & Hasanuzzaman, M. Abiotic stress and reactive oxygen species: generation, signaling, and defense mechanisms. Antioxidants 10, 277 (2021).","journal-title":"Antioxidants"},{"key":"57688_CR28","doi-asserted-by":"publisher","first-page":"101586","DOI":"10.1016\/j.eti.2021.101586","volume":"23","author":"M Gomaa","year":"2021","unstructured":"Gomaa, M., Zien-Elabdeen, A., Hifney, A. F. & Adam, M. S. Phycotoxicity of antibiotics and non-steroidal anti-inflammatory drugs to green algae Chlorella sp. and Desmodesmus spinosus: assessment of combined toxicity by Box\u2013Behnken experimental design. Environ. Technol. Innov. 23, 101586 (2021).","journal-title":"Environ. Technol. Innov."},{"key":"57688_CR29","doi-asserted-by":"crossref","unstructured":"Harshkova, D., Zielinska, E., Narajczyk, M., Kapusta, M. & Aksmann, A. Mitochondria dysfunction is one of the causes of diclofenac toxicity in the green alga Chlamydomonas reinhardtii. PeerJ 12, (2024).","DOI":"10.7717\/peerj.18005"},{"key":"57688_CR30","unstructured":"Berwal, M. K., Ram, C. & Superoxide Dismutase A stable biochemical marker for abiotic stress tolerance in higher plants. Abiotic and Biotic Stress in Plants (2018)."},{"key":"57688_CR31","doi-asserted-by":"publisher","first-page":"477","DOI":"10.3390\/horticulturae11050477","volume":"11","author":"MJ Rao","year":"2025","unstructured":"Rao, M. J. et al. Antioxidant Defense System in Plants: Reactive Oxygen Species production, signaling, and scavenging during abiotic stress-induced oxidative damage. Horticulturae 11, 477 (2025).","journal-title":"Horticulturae"},{"key":"57688_CR32","doi-asserted-by":"publisher","first-page":"974","DOI":"10.1111\/tpj.13053","volume":"84","author":"IK Blaby","year":"2015","unstructured":"Blaby, I. K. et al. Genome-wide analysis on Chlamydomonas reinhardtii reveals the impact of hydrogen peroxide on protein stress responses and overlap with other stress transcriptomes. Plant. J. 84, 974\u2013988 (2015).","journal-title":"Plant. J."},{"key":"57688_CR33","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1016\/j.aquatox.2017.03.007","volume":"186","author":"PK Pandey","year":"2017","unstructured":"Pandey, P. K., Ajima, M. N. O., Kumar, K., Poojary, N. & Kumar, S. Evaluation of DNA damage and physiological responses in Nile tilapia, Oreochromis niloticus (Linnaeus, 1758) exposed to sub-lethal diclofenac (DCF). Aquat. Toxicol. 186, 205\u2013214 (2017).","journal-title":"Aquat. Toxicol."},{"key":"57688_CR34","doi-asserted-by":"publisher","first-page":"108172","DOI":"10.1016\/j.ecolind.2021.108172","volume":"131","author":"SA Mirzaee","year":"2021","unstructured":"Mirzaee, S. A. et al. The possible oxidative stress and DNA damage induced in diclofenac-exposed non-target organisms in the aquatic environment: a systematic review. Ecol. Indic. 131, 108172 (2021).","journal-title":"Ecol. Indic."},{"key":"57688_CR35","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1007\/s11270-025-08758-6","volume":"237","author":"N Touzout","year":"2025","unstructured":"Touzout, N. et al. Understanding the ecotoxicity of diclofenac on the freshwater cyanobacterium arthrospira platensis through growth kinetics, primary biocomponents, oxidative metabolism, and molecular docking. Water Air Soil. Pollut. 237, 119 (2025).","journal-title":"Water Air Soil. Pollut"},{"key":"57688_CR36","doi-asserted-by":"crossref","unstructured":"Kato, N., Nelson, G. & Lauersen, K. J. Subcellular localizations of catalase and exogenously added fatty acid in Chlamydomonas reinhardtii. Cells 10, (2021)","DOI":"10.3390\/cells10081940"},{"key":"57688_CR37","first-page":"1377","volume":"57","author":"T Maruta","year":"2016","unstructured":"Maruta, T., Sawa, Y., Shigeoka, S. & Ishikawa, T. Diversity and evolution of ascorbate peroxidase functions in chloroplasts: more than just a classical antioxidant enzyme? Plant. Cell. Physiol. 57, 1377\u20131386 (2016).","journal-title":"Plant. Cell. Physiol."},{"key":"57688_CR38","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1111\/j.1399-3054.1996.tb00472.x","volume":"97","author":"F Lopez","year":"1996","unstructured":"Lopez, F., Vansuyt, G., Casse-Delbart, F. & Fourcroy, P. Ascorbate peroxidase activity, not the mRNA level, is enhanced in salt-stressed Raphanus sativus plants. Physiol. Plant. 97, 13\u201320 (1996).","journal-title":"Physiol. Plant."},{"key":"57688_CR39","unstructured":"Caccamo, A. The role of ascorbate peroxidase 2 and protein cysteine modifications in the green alga Chlamydomonas reinhardtii. (2024). https:\/\/researchportal.vub.be\/en\/publications\/the-role-of-ascorbate-peroxidase-2-and-protein-cysteine-modificat\/"},{"key":"57688_CR40","doi-asserted-by":"publisher","first-page":"2700","DOI":"10.1093\/jxb\/erae061","volume":"75","author":"K Yoshimura","year":"2024","unstructured":"Yoshimura, K. & Ishikawa, T. Physiological function and regulation of ascorbate peroxidase isoforms. J. Exp. Bot. 75, 2700\u20132715 (2024).","journal-title":"J. Exp. Bot."},{"key":"57688_CR41","doi-asserted-by":"publisher","first-page":"102789","DOI":"10.1016\/j.redox.2023.102789","volume":"64","author":"S Li","year":"2023","unstructured":"Li, S. Novel insight into functions of ascorbate peroxidase in higher plants: more than a simple antioxidant enzyme. Redox Biol. 64, 102789 (2023).","journal-title":"Redox Biol."},{"key":"57688_CR42","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1042\/bj3480321","volume":"348","author":"ANP Hiner","year":"2000","unstructured":"Hiner, A. N. P. et al. Kinetic study of the inactivation of ascorbate peroxidase by hydrogen peroxide. Biochem. J. 348, 321 (2000).","journal-title":"Biochem. J."},{"key":"57688_CR43","doi-asserted-by":"publisher","first-page":"1027730","DOI":"10.3389\/fpls.2021.678111","volume":"12","author":"S Kaur","year":"2021","unstructured":"Kaur, S. et al. Regulation of dual activity of ascorbate peroxidase 1 from Arabidopsis thaliana by conformational changes and posttranslational modifications. Front. Plant. Sci. 12, 1027730 (2021).","journal-title":"Front. Plant. Sci."},{"key":"57688_CR44","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1016\/S0168-9452(03)00164-X","volume":"165","author":"R Madhusudhan","year":"2003","unstructured":"Madhusudhan, R., Ishikawa, T., Sawa, Y., Shigeoka, S. & Shibata, H. Post-transcriptional regulation of ascorbate peroxidase during light adaptation of Euglena gracilis. Plant. Sci. 165, 233\u2013238 (2003).","journal-title":"Plant. Sci."},{"key":"57688_CR45","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1080\/09670262.2019.1576062","volume":"54","author":"K Mukai","year":"2019","unstructured":"Mukai, K. et al. Effects of light and hydrogen peroxide on gene expression of newly identified antioxidant enzymes in the harmful algal bloom species Chattonella marina. Eur. J. Phycol. 54, 393\u2013403 (2019).","journal-title":"Eur. J. Phycol."},{"key":"57688_CR46","doi-asserted-by":"publisher","first-page":"672672","DOI":"10.3389\/fpls.2021.672672","volume":"12","author":"Y Zhang","year":"2021","unstructured":"Zhang, Y., Wang, L. F., Li, T. T. & Liu, W. C. Mutual promotion of LAP2 and CAT2 synergistically regulates plant salt and osmotic stress tolerance. Front. Plant. Sci. 12, 672672 (2021).","journal-title":"Front. Plant. Sci."},{"key":"57688_CR47","doi-asserted-by":"publisher","first-page":"70118","DOI":"10.1111\/ppl.70118","volume":"177","author":"A Jim\u00e9nez","year":"2025","unstructured":"Jim\u00e9nez, A., Mart\u00ed, M. C. & Sevilla, F. Oxidative post-translational modifications of plant antioxidant systems under environmental stress. Physiol. Plant. 177, 70118 (2025).","journal-title":"Physiol. Plant."},{"key":"57688_CR48","doi-asserted-by":"publisher","first-page":"1027730","DOI":"10.3389\/fpls.2022.1027730","volume":"13","author":"JM Mart\u00ed-Guill\u00e9n","year":"2022","unstructured":"Mart\u00ed-Guill\u00e9n, J. M., Pardo-Hern\u00e1ndez, M., Mart\u00ednez-Lorente, S. E., Almagro, L. & Rivero, R. M. Redox post-translational modifications and their interplay in plant abiotic stress tolerance. Front. Plant. Sci. 13, 1027730 (2022).","journal-title":"Front. Plant. Sci."},{"key":"57688_CR49","unstructured":"NCBI. (2025). https:\/\/pubchem.ncbi.nlm.nih.gov\/compound\/4409"},{"key":"57688_CR50","unstructured":"Situmeang, D. & Suwitono, M. R. Predicting pollutant toxicity of over-the-counter (OTC) pain killers (Analgesic) pharmaceutical drug. in 11th International Scholars ConferenceUniversitas Advent Indonesia, Indonesia, (2024)."},{"key":"57688_CR51","doi-asserted-by":"publisher","first-page":"10921","DOI":"10.1021\/jp805614y","volume":"112","author":"KAK Musa","year":"2008","unstructured":"Musa, K. A. K. & Eriksson, L. A. Theoretical study of the phototoxicity of naproxen and the active form of nabumetone. J. Phys. Chem. A. 112, 10921\u201310930 (2008).","journal-title":"J. Phys. Chem. A"},{"key":"57688_CR52","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1007\/s00244-004-2297-9","volume":"47","author":"A Aksmann","year":"2004","unstructured":"Aksmann, A. & Tukaj, Z. The effect of anthracene and phenanthrene on the growth, photosynthesis, and SOD activity of the green alga Scenedesmus armatus depends on the PAR irradiance and CO2 level. Arch. Environ. Contam. Toxicol. 47, 177\u2013184 (2004).","journal-title":"Arch. Environ. Contam. Toxicol."},{"key":"57688_CR53","doi-asserted-by":"publisher","first-page":"26","DOI":"10.1016\/j.chemosphere.2008.09.064","volume":"74","author":"A Aksmann","year":"2008","unstructured":"Aksmann, A. & Tukaj, Z. Intact anthracene inhibits photosynthesis in algal cells: a fluorescence induction study on Chlamydomonas reinhardtii cw92 strain. Chemosphere 74, 26\u201332 (2008).","journal-title":"Chemosphere"},{"key":"57688_CR54","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1111\/pre.12147","volume":"64","author":"A Aksmann","year":"2016","unstructured":"Aksmann, A., Pokora, W., Ba\u015bcik-Remisiewicz, A., Dettlaff-Pokora, A. & Tukaj, Z. High hydrogen peroxide production and antioxidative enzymes expression in the Chlamydomonas reinhardtii cia3 mutant with an increased tolerance to cadmium and anthracene. Phycological Res. 64, 300\u2013311 (2016).","journal-title":"Phycological Res."},{"key":"57688_CR55","doi-asserted-by":"crossref","unstructured":"Harris, E. H. The Chlamydomonas sourcebook: introduction to Chlamydomonas and its laboratory use: Volume 1 (Academic, 2009).","DOI":"10.1016\/B978-0-12-370873-1.00001-0"},{"key":"57688_CR56","doi-asserted-by":"publisher","first-page":"1356","DOI":"10.1111\/jpy.13524","volume":"60","author":"M Hejna","year":"2024","unstructured":"Hejna, M., Kapu\u015bci\u0144ska, D. & Aksmann, A. A sensitive and reliable method for the quantitative determination of hydrogen peroxide produced by microalgae cells. J. Phycol. 60, 1356\u20131370 (2024).","journal-title":"J. Phycol."},{"key":"57688_CR57","doi-asserted-by":"publisher","first-page":"248","DOI":"10.1016\/0003-2697(76)90527-3","volume":"72","author":"MM Bradford","year":"1976","unstructured":"Bradford, M. M. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Anal. Biochem. 72, 248\u2013254 (1976).","journal-title":"Anal. Biochem."},{"key":"57688_CR58","doi-asserted-by":"publisher","first-page":"404","DOI":"10.1111\/j.1749-6632.1964.tb14213.x","volume":"121","author":"BJ Davis","year":"1964","unstructured":"Davis, B. J. Disc electrophoresis \u2013 II method and application to human serum proteins. Ann. N Y Acad. Sci. 121, 404\u2013427 (1964).","journal-title":"Ann. N Y Acad. Sci."},{"key":"57688_CR59","doi-asserted-by":"publisher","first-page":"323","DOI":"10.1016\/j.ecoenv.2005.12.001","volume":"65","author":"Z Tukaj","year":"2006","unstructured":"Tukaj, Z. & Pokora, W. Individual and combined effect of anthracene, cadmium, and chloridazone on growth and activity of SOD izoformes in three Scenedesmus species. Ecotoxicol. Environ. Saf. 65, 323\u2013331 (2006).","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"57688_CR60","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1016\/0003-2697(71)90370-8","volume":"44","author":"C Beauchamp","year":"1971","unstructured":"Beauchamp, C. & Fridovich, I. Superoxide dismutase: improved assays and an assay applicable to acrylamide gels. Anal. Biochem. 44, 276\u2013287 (1971).","journal-title":"Anal. Biochem."},{"key":"57688_CR61","doi-asserted-by":"publisher","first-page":"867","DOI":"10.1093\/oxfordjournals.pcp.a076232","volume":"22","author":"Y Nakano","year":"1981","unstructured":"Nakano, Y. & Asada, K. Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts. Plant. Cell. Physiol. 22, 867\u2013880 (1981).","journal-title":"Plant. Cell. Physiol."},{"key":"57688_CR62","doi-asserted-by":"crossref","unstructured":"Aebi, H. & Catalase in vitro. in Methods in Enzymology 105. 121\u2013126Academic Press, (1984).","DOI":"10.1016\/S0076-6879(84)05016-3"},{"key":"57688_CR63","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1016\/j.jplph.2016.10.008","volume":"208","author":"W Pokora","year":"2017","unstructured":"Pokora, W. et al. Changes in nitric oxide\/hydrogen peroxide content and cell cycle progression: Study with synchronized cultures of green alga Chlamydomonas reinhardtii. J. Plant. Physiol. 208, 84\u201393 (2017).","journal-title":"J. Plant. Physiol."},{"key":"57688_CR64","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1016\/j.jplph.2018.07.015","volume":"230","author":"W Pokora","year":"2018","unstructured":"Pokora, W., Aksmann, A., Ba\u015bcik-Remisiewicz, A., Dettlaff-Pokora, A. & Tukaj, Z. Exogenously applied hydrogen peroxide modifies the course of the Chlamydomonas reinhardtii cell cycle. J. Plant. Physiol. 230, 61\u201372 (2018).","journal-title":"J. Plant. Physiol."},{"key":"57688_CR65","doi-asserted-by":"publisher","first-page":"e45","DOI":"10.1093\/nar\/29.9.e45","volume":"29","author":"MW Pfaffl","year":"2001","unstructured":"Pfaffl, M. W. A new mathematical model for relative quantification in real-time RT\u2013PCR. Nucleic Acids Res. 29, e45 (2001).","journal-title":"Nucleic Acids Res."},{"key":"57688_CR66","doi-asserted-by":"publisher","first-page":"D523","DOI":"10.1093\/nar\/gkac1052","volume":"51","author":"A Bateman","year":"2023","unstructured":"Bateman, A. et al. UniProt: the universal protein knowledgebase in 2023. Nucleic Acids Res. 51, D523\u2013D531 (2023).","journal-title":"Nucleic Acids Res."},{"key":"57688_CR67","doi-asserted-by":"publisher","first-page":"W215","DOI":"10.1093\/nar\/gkae237","volume":"52","author":"MT \u00d8dum","year":"2024","unstructured":"\u00d8dum, M. T. et al. DeepLoc 2.1: multi-label membrane protein type prediction using protein language models. Nucleic Acids Res. 52, W215\u2013W220 (2024).","journal-title":"Nucleic Acids Res."},{"key":"57688_CR68","doi-asserted-by":"publisher","first-page":"3625","DOI":"10.1093\/molbev\/mss178","volume":"29","author":"M Tardif","year":"2012","unstructured":"Tardif, M. et al. PredAlgo: a new subcellular localization prediction tool dedicated to green algae. Mol. Biol. Evol. 29, 3625\u20133639 (2012).","journal-title":"Mol. Biol. Evol."},{"key":"57688_CR69","doi-asserted-by":"publisher","first-page":"1316","DOI":"10.1002\/etc.5620210629","volume":"21","author":"B Beliaeff","year":"2002","unstructured":"Beliaeff, B. & Burgeot, T. Integrated biomarker response: a useful tool for ecological risk assessment. Environ. Toxicol. Chem. 21, 1316\u20131322 (2002).","journal-title":"Environ. Toxicol. Chem."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57688-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57688-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57688-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:02:42Z","timestamp":1788220962000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-57688-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,15]]},"references-count":69,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["57688"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-57688-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,15]]},"assertion":[{"value":"2 April 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"9 June 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"15 June 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"This study does not contain any individual personal data including any individual\u2019s name, images, or videos.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"27289"}}