{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T11:23:08Z","timestamp":1787743388816,"version":"build-2784847793"},"reference-count":100,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T00:00:00Z","timestamp":1787702400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Cyanobacteria are promising platforms for bioenergy, carbon sequestration, and bioproduct synthesis, but their photosynthetic efficiency is limited by suboptimal light utilization, electron transport constraints, and environmental stress. Titanium dioxide nanoparticles (n-TiO2) have emerged as powerful photocatalytic materials that can enhance light absorption, modulate electron transport, and influence the redox balance in biological systems. This review advances the concept of photocatalytic-biological coupling, in which n-TiO2 functions as artificial light amplifiers that augment cyanobacterial photosynthesis. Current evidence on the physicochemical properties of n-TiO2, their interactions with cyanobacterial cells, and their effects on photosystems, electron transport chains, and downstream metabolic processes is examined. Particular emphasis is placed on the integration of photophysical and biological mechanisms, including reactive oxygen species (ROS)-mediated signaling, proton motive force (PMF) enhancement, and adenosine triphosphate (ATP) synthesis. Emerging approaches, including nano\u2013bio interface engineering, environmental biotechnology applications, and artificial intelligence-guided optimization, are highlighted. By bridging photophysics, cellular bioenergetics, and computational design within a unified mechanistic framework, this review establishes the scientific foundation needed to translate photocatalytic\u2013biological coupling into scalable and biotechnologically deployable nano-enabled photosynthetic systems.<\/jats:p>","DOI":"10.3390\/molecules31172983","type":"journal-article","created":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T10:59:23Z","timestamp":1787741963000},"page":"2983","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Titanium Dioxide Nanoparticle-Driven Metabolic and Molecular Reprogramming in Cyanobacteria"],"prefix":"10.3390","volume":"31","author":[{"given":"Shyama Malika","family":"Malwalage","sequence":"first","affiliation":[{"name":"Bioenvironmental Sciences Program, Morgan State University, Baltimore, MD 21251, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mst","family":"Sayadujjhara","sequence":"additional","affiliation":[{"name":"Bioenvironmental Sciences Program, Morgan State University, Baltimore, MD 21251, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0096-569X","authenticated-orcid":false,"given":"Viji","family":"Sitther","sequence":"additional","affiliation":[{"name":"Department of Biology, School of Computer, Mathematical and Natural Sciences, Morgan State University, Baltimore, MD 21251, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1440","DOI":"10.1111\/nph.16249","article-title":"On the origin of oxygenic photosynthesis and Cyanobacteria","volume":"225","author":"Cardona","year":"2020","journal-title":"New Phytol."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Agarwal, P., Soni, R., Kaur, P., Madan, A., Mishra, R., Pandey, J., Singh, S., and Singh, G. (2022). Cyanobacteria as a promising alternative for sustainable environment: Synthesis of biofuel and biodegradable plastics. Front. Microbiol., 13.","DOI":"10.3389\/fmicb.2022.939347"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1016\/j.biotechadv.2019.02.010","article-title":"Phycobiliproteins from cyanobacteria: Chemistry and biotechnological applications","volume":"37","author":"Pagels","year":"2019","journal-title":"Biotechnol. Adv."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"e2021523118","DOI":"10.1073\/pnas.2021523118","article-title":"Improved photosynthetic capacity and photosystem I oxidation via heterologous metabolism engineering in cyanobacteria","volume":"118","author":"Torrado","year":"2021","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1016\/j.plantsci.2009.06.005","article-title":"Solar energy conversion efficiencies in photosynthesis: Minimizing the chlorophyll antennae to maximize efficiency","volume":"177","author":"Melis","year":"2009","journal-title":"Plant Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1111\/ppl.13404","article-title":"Regulatory electron transport pathways of photosynthesis in cyanobacteria and microalgae: Recent advances and biotechnological prospects","volume":"173","author":"Nikkanen","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Stadnichuk, I.N., and Kusnetsov, V.V. (2023). Phycobilisomes and phycobiliproteins in the pigment apparatus of oxygenic photosynthetics: From cyanobacteria to tertiary endosymbiosis. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms24032290"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1007\/s11120-018-0561-5","article-title":"Blue light reduces photosynthetic efficiency of cyanobacteria through an imbalance between photosystems I and II","volume":"138","author":"Luimstra","year":"2018","journal-title":"Photosynth. Res."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Wyatt, L., Gichuki, S., Yalcin, Y.S., and Sitther, V. (2023). Impact of ascorbic acid on zero-valent iron nanoparticle and UV-B mediated stress in the cyanobacterium, Fremyella diplosiphon. Microorganisms, 11.","DOI":"10.3390\/microorganisms11051245"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1016\/j.bbabio.2013.11.017","article-title":"Co-existence of photosynthetic and respiratory activities in cyanobacterial thylakoid membranes","volume":"1837","author":"Mullineaux","year":"2014","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Bioenerg."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Rathore, C., Yadav, V.K., Gacem, A., AbdelRahim, S.K., Verma, R.K., Chundawat, R.S., Gnanamoorthy, G., Yadav, K.K., Choudhary, N., and Sahoo, D.K. (2023). Microbial synthesis of titanium dioxide nanoparticles and their importance in wastewater treatment and antimicrobial activities: A review. Front. Microbiol., 14.","DOI":"10.3389\/fmicb.2023.1270245"},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Wang, Q., Li, Y., Huang, F., Song, S., Ai, G., Xin, X., Zhao, B., Zheng, Y., and Zhang, Z. (2023). Recent advances in g-C3N4-based materials and their application in energy and environmental sustainability. Molecules, 28.","DOI":"10.3390\/molecules28010432"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"19413","DOI":"10.1021\/acsomega.3c00496","article-title":"Biofunctionalized CdS quantum dots: A case study on nanomaterial toxicity in the photocatalytic wastewater treatment process","volume":"8","author":"Shivaji","year":"2023","journal-title":"ACS Omega"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Li, P., Xia, Y., Song, K., and Liu, D. (2024). The impact of nanomaterials on photosynthesis and antioxidant mechanisms in gramineae plants: Research progress and future prospects. Plants, 13.","DOI":"10.3390\/plants13070984"},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Rudi, L., Cepoi, L., Chiriac, T., and Djur, S. (2025). Interactions between potentially toxic nanoparticles (Cu, CuO, ZnO, and TiO2) and the cyanobacterium Arthrospira platensis: Biological adaptations to xenobiotics. Nanomaterials, 15.","DOI":"10.3390\/nano15010046"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"e14519","DOI":"10.1111\/1751-7915.14519","article-title":"Limiting factors in the operation of photosystems I and II in cyanobacteria","volume":"17","author":"Grettenberger","year":"2024","journal-title":"Microb. Biotechnol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1111\/j.1399-3054.2011.01457.x","article-title":"Protein synthesis is the primary target of reactive oxygen species in the photoinhibition of photosystem II","volume":"142","author":"Nishiyama","year":"2011","journal-title":"Physiol. Plant."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1016\/j.tplants.2019.09.013","article-title":"Changing color for photoprotection: The orange carotenoid protein","volume":"25","author":"Muzzopappa","year":"2020","journal-title":"Trends Plant Sci."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"3476","DOI":"10.1093\/jxb\/erad118","article-title":"Electron transport in cyanobacterial thylakoid membranes: Are cyanobacteria simple models for photosynthetic organisms?","volume":"74","author":"Shimakawa","year":"2023","journal-title":"J. Exp. Bot."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"414","DOI":"10.1016\/j.bbabio.2006.11.019","article-title":"Photoinhibition of photosystem II under environmental stress","volume":"1767","author":"Murata","year":"2007","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Bioenerg."},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Singh, V.K., Jha, S., Rana, P., Mishra, S., Kumari, N., Singh, S.C., Anand, S., Upadhye, V., and Sinha, R.P. (2023). Resilience and mitigation strategies of cyanobacteria under ultraviolet radiation stress. Int. J. Mol. Sci., 24.","DOI":"10.3390\/ijms241512381"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"koae323","DOI":"10.1093\/plcell\/koae323","article-title":"Temperature-driven changes in membrane fluidity differentially impact FILAMENTATION TEMPERATURE-SENSITIVE H2-mediated photosystem II repair","volume":"37","author":"Zhang","year":"2025","journal-title":"Plant Cell"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.bbrc.2023.06.051","article-title":"Titanium dioxide nanoparticles enhance photocurrent generation of cyanobacteria","volume":"672","author":"Li","year":"2023","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"9919","DOI":"10.1021\/cr5001892","article-title":"Understanding TiO2 photocatalysis: Mechanisms and materials","volume":"114","author":"Schneider","year":"2014","journal-title":"Chem. Rev."},{"key":"ref_25","doi-asserted-by":"crossref","unstructured":"Lettieri, S., Pavone, M., Fioravanti, A., Santamaria Amato, L., and Maddalena, P. (2021). Charge carrier processes and optical properties in TiO2 and TiO2-based heterojunction photocatalysts: A review. Materials, 14.","DOI":"10.3390\/ma14071645"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.cattod.2018.10.053","article-title":"Charge carrier trapping, recombination and transfer during TiO2 photocatalysis: An overview","volume":"335","author":"Qian","year":"2019","journal-title":"Catal. Today"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"4043","DOI":"10.1038\/srep04043","article-title":"Why is anatase a better photocatalyst than rutile? Model studies on epitaxial TiO2 films","volume":"4","author":"Luttrell","year":"2014","journal-title":"Sci. Rep."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"11302","DOI":"10.1021\/acs.chemrev.7b00161","article-title":"Generation and detection of reactive oxygen species in photocatalysis","volume":"117","author":"Nosaka","year":"2017","journal-title":"Chem. Rev."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"4545","DOI":"10.1021\/jp0273934","article-title":"Explaining the enhanced photocatalytic activity of Degussa P25 mixed-phase TiO2 using EPR","volume":"107","author":"Hurum","year":"2003","journal-title":"J. Phys. Chem. B"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1016\/j.jphotochem.2010.07.024","article-title":"What is Degussa (Evonik) P25? Crystalline composition analysis, reconstruction from isolated pure particles and photocatalytic activity test","volume":"216","author":"Ohtani","year":"2010","journal-title":"J. Photochem. Photobiol. A Chem."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2644","DOI":"10.1021\/acscatal.7b00131","article-title":"Trapping-induced enhancement of photocatalytic activity on brookite TiO2 powders: Comparison with anatase and rutile TiO2 powders","volume":"7","author":"Vequizo","year":"2017","journal-title":"ACS Catal."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"24538","DOI":"10.1021\/acs.jpcc.5b09236","article-title":"Distinctive behavior of photogenerated electrons and holes in anatase and rutile TiO2 powders","volume":"119","author":"Yamakata","year":"2015","journal-title":"J. Phys. Chem. C"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1038\/nature06964","article-title":"Anatase TiO2 single crystals with a large percentage of reactive facets","volume":"453","author":"Yang","year":"2008","journal-title":"Nature"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"4085","DOI":"10.1021\/cm200597m","article-title":"Anatase TiO2 with dominant high-energy {001} facets: Synthesis, properties, and applications","volume":"23","author":"Liu","year":"2011","journal-title":"Chem. Mater."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Gonzalo, S., Llaneza, V., Pulido-Reyes, G., Fern\u00e1ndez-Pi\u00f1as, F., Bonzongo, J.C., Leganes, F., Rosal, R., Garc\u00eda-Calvo, E., and Rodea-Palomares, I. (2014). A colloidal singularity reveals the crucial role of colloidal stability for nanomaterials in vitro toxicity testing: nZVI-microalgae colloidal system as a case study. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0109645"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"120990","DOI":"10.1016\/j.watres.2023.120990","article-title":"Formation of extracellular polymeric substances corona on TiO2 nanoparticles: Roles of crystalline phase and exposed facets","volume":"249","author":"Du","year":"2024","journal-title":"Water Res."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"4333","DOI":"10.1021\/acsomega.1c06112","article-title":"Visible-light-driven carbon-doped TiO2-based nanocatalysts for enhanced activity toward microbes and removal of dye","volume":"7","author":"Ghumro","year":"2022","journal-title":"ACS Omega"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1016\/j.surfrep.2008.10.001","article-title":"TiO2 photocatalysis and related surface phenomena","volume":"63","author":"Fujishima","year":"2008","journal-title":"Surf. Sci. Rep."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.jphotochemrev.2015.08.003","article-title":"Visible-light activation of TiO2 photocatalysts: Advances in theory and experiments","volume":"25","author":"Etacheri","year":"2015","journal-title":"J. Photochem. Photobiol. C Photochem. Rev."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1077","DOI":"10.1007\/s00253-015-7144-6","article-title":"Application of light-emitting diodes (LEDs) in cultivation of phototrophic microalgae: Current state and perspectives","volume":"100","author":"Glemser","year":"2016","journal-title":"Appl. Microbiol. Biotechnol."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Miller, R.J., Bennett, S., Keller, A.A., Pease, S., and Lenihan, H.S. (2012). TiO2 nanoparticles are phototoxic to marine phytoplankton. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0030321"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Liu, Q., Li, X., Chen, Y., Zhang, X., Gao, B., Ma, M., Yang, H., Yuan, S., and Zhang, Q. (2025). Strategies for Regulating Reactive Oxygen Species in Carbon Nitride-Based Photocatalysis. Molecules, 30.","DOI":"10.3390\/molecules30173586"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1126\/science.1061051","article-title":"Visible-light photocatalysis in nitrogen-doped titanium oxides","volume":"293","author":"Asahi","year":"2001","journal-title":"Science"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"7530","DOI":"10.1021\/es0709122","article-title":"Mesoporous nitrogen-doped TiO2 for the photocatalytic destruction of the cyanobacterial toxin microcystin-LR under visible light irradiation","volume":"41","author":"Choi","year":"2007","journal-title":"Environ. Sci. Technol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"961437","DOI":"10.1155\/2014\/961437","article-title":"Influence of TiO2 nanoparticles on growth and phenolic compounds production in photosynthetic microorganisms","volume":"2014","author":"Comotto","year":"2014","journal-title":"Sci. World J."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1007\/s00284-025-04629-9","article-title":"Strain-specific impact of titanium dioxide nanoparticles on Fremyella diplosiphon physiological and metabolic responses","volume":"83","author":"Sayadujjhara","year":"2025","journal-title":"Curr. Microbiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"8522","DOI":"10.1021\/acs.jafc.8b01522","article-title":"Phycobiliproteins production enhancement and lipidomic alteration by titanium dioxide nanoparticles in Synechocystis sp. PCC 6803 culture","volume":"66","author":"Zahra","year":"2018","journal-title":"J. Agric. Food Chem."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"116508","DOI":"10.1016\/j.envpol.2021.116508","article-title":"Toxic and protective mechanisms of cyanobacterium Synechocystis sp. in response to titanium dioxide nanoparticles","volume":"274","author":"Xu","year":"2021","journal-title":"Environ. Pollut."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Guti\u00e9rrez, G.G., Rivas-Vald\u00e9s, F., Benavente, B.P., Olivares, R., Hepp, M.I., Barra, R.O., and Urrutia, R. (2025). Toxicological responses of photosynthetic genes in Chlorella vulgaris exposed to environmentally relevant concentrations of TiO2 nanoparticles. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms262110271"},{"key":"ref_50","doi-asserted-by":"crossref","unstructured":"Binh, C.T.T., Peterson, C.G., Tong, T., Gray, K.A., Gaillard, J.F., and Kelly, J.J. (2015). Comparing acute effects of a nano-TiO2 pigment on cosmopolitan freshwater phototrophic microbes using high-throughput screening. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0125613"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"6814","DOI":"10.1021\/acs.jpclett.8b02892","article-title":"How to correctly determine the band gap energy of modified semiconductor photocatalysts based on UV\u2013Vis spectra","volume":"9","author":"Pacia","year":"2018","journal-title":"J. Phys. Chem. Lett."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"258","DOI":"10.1111\/j.1574-6976.2008.00134.x","article-title":"Oxidative stress in cyanobacteria","volume":"33","author":"Latifi","year":"2009","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Khedr, T.M., El-Sheikh, S.M., Ismail, A.A., Kowalska, E., and Bahnemann, D.W. (2019). Photodegradation of Microcystin-LR Using Visible Light-Activated C\/N-co-Modified Mesoporous TiO2 Photocatalyst. Materials, 12.","DOI":"10.3390\/ma12071027"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1038\/s41580-020-0230-3","article-title":"Reactive oxygen species (ROS) as pleiotropic physiological signalling agents","volume":"21","author":"Sies","year":"2020","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"14036","DOI":"10.1002\/anie.201406625","article-title":"Molecular-level understanding of the photocatalytic activity difference between anatase and rutile nanoparticles","volume":"53","author":"Kim","year":"2014","journal-title":"Angew. Chem. Int. Ed."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1016\/j.apcata.2007.03.015","article-title":"Photocatalytic reactivity for O2\u00b7\u2212 and \u00b7OH radical formation in anatase and rutile TiO2 suspension as the effect of H2O2 addition","volume":"325","author":"Hirakawa","year":"2007","journal-title":"Appl. Catal. A Gen."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1021\/la025785w","article-title":"OH surface density of SiO2 and TiO2 by thermogravimetric analysis","volume":"19","author":"Mueller","year":"2003","journal-title":"Langmuir"},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"\u00d6nal Acet, B., G\u00fcl, D., Stauber, R.H., Odaba\u015f\u0131, M., and Acet, \u00d6. (2024). A review for uncovering the \u201cprotein\u2013nanoparticle alliance\u201d: Implications of the protein corona for biomedical applications. Nanomaterials, 14.","DOI":"10.3390\/nano14100823"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"e1788","DOI":"10.1002\/wnan.1788","article-title":"Influence of surface chemistry and morphology of nanoparticles on protein corona formation","volume":"14","author":"Bilardo","year":"2022","journal-title":"Wiley Interdiscip. Rev. Nanomed. Nanobiotechnol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"12258","DOI":"10.1021\/acs.est.6b03684","article-title":"Interactions between algal extracellular polymeric substances and commercial TiO2 nanoparticles in aqueous media","volume":"50","author":"Adeleye","year":"2016","journal-title":"Environ. Sci. Technol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1111\/j.1574-6976.2009.00183.x","article-title":"Complexity of cyanobacterial exopolysaccharides: Composition, structures, inducing factors and putative genes involved in their biosynthesis and assembly","volume":"33","author":"Pereira","year":"2009","journal-title":"FEMS Microbiol. Rev."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1002\/etc.445","article-title":"Impact of nano titanium dioxide exposure on cellular structure of Anabaena variabilis and evidence of internalization","volume":"30","author":"Cherchi","year":"2011","journal-title":"Environ. Toxicol. Chem."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1016\/j.ecoenv.2018.06.027","article-title":"Effect and mechanism of TiO2 nanoparticles on the photosynthesis of Chlorella pyrenoidosa","volume":"161","author":"Middepogu","year":"2018","journal-title":"Ecotoxicol. Environ. Saf."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"256","DOI":"10.1016\/j.bbabio.2015.11.010","article-title":"Distribution and dynamics of electron transport complexes in cyanobacterial thylakoid membranes","volume":"1857","author":"Liu","year":"2016","journal-title":"Biochim. Biophys. Acta (BBA)\u2014Bioenerg."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"5873","DOI":"10.1038\/s41598-023-32466-y","article-title":"Plants exposed to titanium dioxide nanoparticles acquired contrasting photosynthetic and morphological strategies depending on the growing light intensity: A case study in radish","volume":"13","author":"Vatankhah","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1385\/BTER:111:1:239","article-title":"Mechanism of nano-anatase TiO2 on promoting photosynthetic carbon reaction of spinach","volume":"111","author":"Gao","year":"2006","journal-title":"Biol. Trace Elem. Res."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1007\/s12011-007-0047-3","article-title":"Effects of nano-anatase TiO2 on photosynthesis of spinach chloroplasts under different light illumination","volume":"119","author":"Lei","year":"2007","journal-title":"Biol. Trace Elem. Res."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"4742","DOI":"10.1038\/s41467-021-24396-y","article-title":"Timing the evolution of antioxidant enzymes in cyanobacteria","volume":"12","author":"Boden","year":"2021","journal-title":"Nat. Commun."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Qiao, Y., Wang, W., and Lu, X. (2020). High light induced alka(e)ne biodegradation for lipid and redox homeostasis in cyanobacteria. Front. Microbiol., 11.","DOI":"10.3389\/fmicb.2020.01659"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"114293","DOI":"10.1016\/j.jphotochem.2022.114293","article-title":"Photocatalytic hydrogen peroxide production in natural seawater by AgQDs(0D)\/Bi2O3(3D) hybrid structure for in situ bacterial inactivation","volume":"435","author":"Zhang","year":"2023","journal-title":"J. Photochem. Photobiol. A Chem."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1146\/annurev-biochem-061516-045037","article-title":"Oxidative stress","volume":"86","author":"Sies","year":"2017","journal-title":"Annu. Rev. Biochem."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Yi, S., Guo, X., Lou, W., Mao, S., Luan, G., and Lu, X. (2024). Structure, regulation, and significance of cyanobacterial and chloroplast adenosine triphosphate synthase in the adaptability of oxygenic photosynthetic organisms. Microorganisms, 12.","DOI":"10.3390\/microorganisms12050940"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"631","DOI":"10.1146\/annurev-biochem-060614-034124","article-title":"ATP synthase","volume":"84","author":"Junge","year":"2015","journal-title":"Annu. Rev. Biochem."},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Davis, G.A., and Kramer, D.M. (2020). Optimization of ATP synthase c-rings for oxygenic photosynthesis. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01778"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.cub.2021.10.051","article-title":"Atp\u0398 is an inhibitor of F0F1 ATP synthase to arrest ATP hydrolysis during low-energy conditions in cyanobacteria","volume":"32","author":"Song","year":"2022","journal-title":"Curr. Biol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"116263","DOI":"10.1016\/j.envres.2023.116263","article-title":"High efficiency lipid production, biochar yield and chlorophyll a content of Chlorella sp. microalgae exposed to seawater and TiO2 nanoparticles","volume":"232","author":"Shanmuganathan","year":"2023","journal-title":"Environ. Res."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Wang, S., Alenius, H., El-Nezami, H., and Karisola, P. (2022). A new look at the effects of engineered ZnO and TiO2 nanoparticles: Evidence from transcriptomics studies. Nanomaterials, 12.","DOI":"10.3390\/nano12081247"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"1909","DOI":"10.1021\/pr4000043","article-title":"Global phosphoproteomic analysis reveals diverse functions of serine\/threonine\/tyrosine phosphorylation in the model cyanobacterium Synechococcus sp. strain PCC 7002","volume":"12","author":"Yang","year":"2013","journal-title":"J. Proteome Res."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Babele, P.K., Kumar, J., and Chaturvedi, V. (2019). Proteomic de-regulation in cyanobacteria in response to abiotic stresses. Front. Microbiol., 10.","DOI":"10.3389\/fmicb.2019.01315"},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Satti, S.H., Raja, N.I., Ikram, M., Oraby, H.F., Mashwani, Z.U.R., Mohamed, A.H., Singh, A., and Omar, A.A. (2022). Plant-based titanium dioxide nanoparticles trigger biochemical and proteome modifications in Triticum aestivum L. under biotic stress of Puccinia striiformis. Molecules, 27.","DOI":"10.3390\/molecules27134274"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"2922","DOI":"10.1039\/D2EN00260D","article-title":"Metabolic alterations in alga Chlamydomonas reinhardtii exposed to nTiO2 materials","volume":"9","author":"Liu","year":"2022","journal-title":"Environ. Sci. Nano"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1038\/s41586-024-07487-w","article-title":"Accurate structure prediction of biomolecular interactions with AlphaFold 3","volume":"630","author":"Abramson","year":"2024","journal-title":"Nature"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"126007","DOI":"10.1016\/j.biortech.2021.126007","article-title":"Metabolic systems biology and multi-omics of cyanobacteria: Perspectives and future directions","volume":"343","author":"Pathania","year":"2022","journal-title":"Bioresour. Technol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1038\/s41586-021-03819-2","article-title":"Highly accurate protein structure prediction with AlphaFold","volume":"596","author":"Jumper","year":"2021","journal-title":"Nature"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"D368","DOI":"10.1093\/nar\/gkad1011","article-title":"AlphaFold Protein Structure Database in 2024: Providing structure coverage for over 214 million protein sequences","volume":"52","author":"Varadi","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"W324","DOI":"10.1093\/nar\/gkae300","article-title":"SwissDock 2024: Major enhancements for small-molecule docking with Attracting Cavities and AutoDock Vina","volume":"52","author":"Bugnon","year":"2024","journal-title":"Nucleic Acids Res."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"3891","DOI":"10.1021\/acs.jcim.1c00203","article-title":"AutoDock Vina 1.2.0: New docking methods, expanded force field, and Python bindings","volume":"61","author":"Eberhardt","year":"2021","journal-title":"J. Chem. Inf. Model."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"939","DOI":"10.3762\/bjnano.14.77","article-title":"Prediction of cytotoxicity of heavy metals adsorbed on nano-TiO2 with periodic table descriptors using machine learning approaches","volume":"14","author":"Roy","year":"2023","journal-title":"Beilstein J. Nanotechnol."},{"key":"ref_89","doi-asserted-by":"crossref","unstructured":"Ji, Z., and Yin, Z. (2025). Machine learning-driven prediction of reactive oxygen species dynamics for assessing nanomaterials\u2019 cytotoxicity. Biomimetics, 10.","DOI":"10.3390\/biomimetics10110718"},{"key":"ref_90","doi-asserted-by":"crossref","unstructured":"Khan, M.F., and Khan, M.T. (2026). AI-driven enzyme engineering: Emerging models and next-generation biotechnological applications. Molecules, 31.","DOI":"10.3390\/molecules31010045"},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"G\u00e9rin, S., Mathy, G., and Franck, F. (2014). Modeling the dependence of respiration and photosynthesis upon light, acetate, carbon dioxide, nitrate and ammonium in Chlamydomonas reinhardtii using design of experiments and multiple regression. BMC Syst. Biol., 8.","DOI":"10.1186\/s12918-014-0096-0"},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Ritz, C., Baty, F., Streibig, J.C., and Gerhard, D. (2015). Dose-response analysis using R. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0146021"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v012.i05","article-title":"Bioassay analysis using R","volume":"12","author":"Ritz","year":"2005","journal-title":"J. Stat. Softw."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"101818","DOI":"10.1016\/j.isci.2020.101818","article-title":"A hybrid flux balance analysis and machine learning pipeline elucidates metabolic adaptation in cyanobacteria","volume":"23","author":"Vijayakumar","year":"2020","journal-title":"iScience"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"6757","DOI":"10.1021\/es901102n","article-title":"Titanium nanomaterial removal and release from wastewater treatment plants","volume":"43","author":"Kiser","year":"2009","journal-title":"Environ. Sci. Technol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1039\/c1em10017c","article-title":"Occurrence and removal of titanium at full-scale wastewater treatment plants: Implications for TiO2 nanomaterials","volume":"13","author":"Westerhoff","year":"2011","journal-title":"J. Environ. Monit."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"6705","DOI":"10.1021\/es204530n","article-title":"Development of environmental fate models for engineered nanoparticles\u2014A case study of TiO2 nanoparticles in the Rhine River","volume":"46","author":"Praetorius","year":"2012","journal-title":"Environ. Sci. Technol."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.scitotenv.2015.03.043","article-title":"Impact of TiO2 nanoparticles on freshwater bacteria from three Swedish lakes","volume":"535","author":"Farkas","year":"2015","journal-title":"Sci. Total Environ."},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Saccardo, A., Bezzo, F., and Sforza, E. (2022). Microalgae growth in ultra-thin steady-state continuous photobioreactors: Assessing self-shading effects. Front. Bioeng. Biotechnol., 10.","DOI":"10.3389\/fbioe.2022.977429"},{"key":"ref_100","doi-asserted-by":"crossref","unstructured":"Bockenstedt, J., Vidwans, N.A., Gentry, T., and Vaddiraju, S. (2021). Catalyst recovery, regeneration and reuse during large-scale disinfection of water using photocatalysis. Water, 13.","DOI":"10.3390\/w13192623"}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/17\/2983\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T10:59:38Z","timestamp":1787741978000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/31\/17\/2983"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,26]]},"references-count":100,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["molecules31172983"],"URL":"https:\/\/doi.org\/10.3390\/molecules31172983","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,26]]}}}