{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,2]],"date-time":"2026-04-02T09:42:39Z","timestamp":1775122959781,"version":"3.50.1"},"reference-count":109,"publisher":"MDPI AG","issue":"20","license":[{"start":{"date-parts":[[2020,10,11]],"date-time":"2020-10-11T00:00:00Z","timestamp":1602374400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["PTDC\/BIA-FBT\/30979\/2017"],"award-info":[{"award-number":["PTDC\/BIA-FBT\/30979\/2017"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["UIDP\/50017\/2020+UIDB\/50017\/2020"],"award-info":[{"award-number":["UIDP\/50017\/2020+UIDB\/50017\/2020"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001871","name":"Funda\u00e7\u00e3o para a Ci\u00eancia e a Tecnologia","doi-asserted-by":"publisher","award":["CEECIND\/01434\/2018"],"award-info":[{"award-number":["CEECIND\/01434\/2018"]}],"id":[{"id":"10.13039\/501100001871","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Light is a paramount parameter driving photosynthesis. However, excessive irradiance leads to the formation of reactive oxygen species that cause cell damage and hamper the growth of photosynthetic organisms. Xanthophylls are key pigments involved in the photoprotective response of plants and algae to excessive light. Of particular relevance is the operation of xanthophyll cycles (XC) leading to the formation of de-epoxidized molecules with energy dissipating capacities. Neoxanthin, found in plants and algae in two different isomeric forms, is involved in the light stress response at different levels. This xanthophyll is not directly involved in XCs and the molecular mechanisms behind its photoprotective activity are yet to be fully resolved. This review comprehensively addresses the photoprotective role of 9\u2032-cis-neoxanthin, the most abundant neoxanthin isomer, and one of the major xanthophyll components in plants\u2019 photosystems. The light-dependent accumulation of all-trans-neoxanthin in photosynthetic cells was identified exclusively in algae of the order Bryopsidales (Chlorophyta), that lack a functional XC. A putative photoprotective model involving all-trans-neoxanthin is discussed.<\/jats:p>","DOI":"10.3390\/molecules25204617","type":"journal-article","created":{"date-parts":[[2020,10,17]],"date-time":"2020-10-17T07:23:22Z","timestamp":1602919402000},"page":"4617","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":44,"title":["Photoprotective Role of Neoxanthin in Plants and Algae"],"prefix":"10.3390","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4234-7830","authenticated-orcid":false,"given":"Chiara","family":"Giossi","sequence":"first","affiliation":[{"name":"CESAM\u2013Centre for Environmental and Marine Studies, Department of Biology, University of Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal"}]},{"given":"Paulo","family":"Cartaxana","sequence":"additional","affiliation":[{"name":"CESAM\u2013Centre for Environmental and Marine Studies, Department of Biology, University of Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4775-8161","authenticated-orcid":false,"given":"S\u00f3nia","family":"Cruz","sequence":"additional","affiliation":[{"name":"CESAM\u2013Centre for Environmental and Marine Studies, Department of Biology, University of Aveiro, Campus de Santiago, 3810-193 Aveiro, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2020,10,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"347","DOI":"10.1093\/jxb\/eri041","article-title":"Dynamics of photosystem II: A proteomic approach to thylakoid protein complexes","volume":"56","author":"Aro","year":"2004","journal-title":"J. Exp. Bot."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.tplants.2010.10.001","article-title":"Photoprotection in plants: A new light on photosystem II damage","volume":"16","author":"Takahashi","year":"2011","journal-title":"Trends Plant Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/S1360-1385(99)01504-6","article-title":"The effect of water motion on short-term rates of photosynthesis by marine phytoplankton","volume":"5","author":"MacIntyre","year":"2000","journal-title":"Trends Plant Sci."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1126\/science.1072359","article-title":"Rapid regulation of light harvesting and plant fitness in the field","volume":"297","author":"Agren","year":"2002","journal-title":"Science"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"468","DOI":"10.1016\/j.bbabio.2015.02.008","article-title":"Diverse mechanisms for photoprotection in photosynthesis. Dynamic regulation of photosystem II excitation in response to rapid environmental change","volume":"1847","author":"Derks","year":"2015","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.jplph.2014.03.004","article-title":"Biodiversity of NPQ","volume":"172","author":"Goss","year":"2015","journal-title":"J. Plant Physiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1903","DOI":"10.1104\/pp.15.01935","article-title":"Nonphotochemical chlorophyll fluorescence quenching: Mechanism and effectiveness in protecting plants from photodamage","volume":"170","author":"Ruban","year":"2016","journal-title":"Plant Physiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/0005-2728(90)90088-L","article-title":"Carotenoids and photoprotection in plants: A role for the xanthophyll zeaxanthin","volume":"1020","year":"1990","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1007\/BF02183041","article-title":"Photophysics of the carotenoids associated with the xanthophyll cycle in photosynthesis","volume":"41","author":"Frank","year":"1994","journal-title":"Photosynth. Res."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/S1360-1385(96)80019-7","article-title":"The role of xanthophyll cycle carotenoids in the protection of photosynthesis","volume":"1","author":"Adams","year":"1996","journal-title":"Trends Plant Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1179\/174329211X13020951739938","article-title":"Xanthophyll cycle\u2013a mechanism protecting plants against oxidative stress","volume":"16","author":"Latowski","year":"2011","journal-title":"Redox Rep."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1055\/s-2004-820884","article-title":"The xanthophyll cycle in green algae (Chlorophyta): Its role in the photosynthetic apparatus","volume":"6","author":"Torzillo","year":"2004","journal-title":"Plant Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1038\/nature08587","article-title":"An ancient light-harvesting protein is critical for the regulation of algal photosynthesis","volume":"462","author":"Peers","year":"2009","journal-title":"Nature"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"879","DOI":"10.1111\/j.1365-3040.2005.01467.x","article-title":"Photoprotection mutants of Arabidopsis thaliana acclimate to high light by increasing photosynthesis and specific antioxidants","volume":"29","author":"Golan","year":"2006","journal-title":"Plant Cell Environ."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1048","DOI":"10.1105\/tpc.106.049114","article-title":"The Arabidopsis aba4-1 mutant reveals a specific function for neoxanthin in protection against photooxidative stress","volume":"19","author":"Cazzaniga","year":"2007","journal-title":"Plant Cell"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1506","DOI":"10.1104\/pp.107.108480","article-title":"Zeaxanthin has enhanced antioxidant capacity with respect to all other xanthophylls in Arabidopsis leaves and functions independent of binding to PSII antennae","volume":"145","author":"Havaux","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1132","DOI":"10.1111\/nph.14435","article-title":"Photoprotection in a monophyletic branch of chlorophyte algae is independent of energy-dependent quenching (qE)","volume":"214","author":"Christa","year":"2017","journal-title":"New Phytol."},{"key":"ref_18","unstructured":"Strain, H.H. (1938). Leaf Xanthophylls, Carnegie Institute Publ."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1038\/188491b0","article-title":"The epoxide nature of the carotenoid, neoxanthin","volume":"188","author":"Goldsmith","year":"1960","journal-title":"Nature"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1256","DOI":"10.1039\/J39690001256","article-title":"Carotenoids and related compounds. Part XXI. Structure of neoxanthin (foliaxanthin)","volume":"1","author":"Cholnoky","year":"1969","journal-title":"J. Chem. Soc. C"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"773","DOI":"10.1002\/hlca.19920750314","article-title":"Synthese von (6R, all-E)-neoxanthin und verwandten allen-carotinoiden","volume":"75","author":"Baumeler","year":"1992","journal-title":"Helv. Chim. Acta"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1637","DOI":"10.1002\/hlca.19900730608","article-title":"Neoflor und 6-epineoflor aus bl\u00fcten von trollius europaeus; hochfeld-1H-NMR-spektren von neoxanthin und (9\u2032Z)-neoxanthin","volume":"73","author":"Eugster","year":"1990","journal-title":"Helv. Chim. Acta"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1002\/hlca.19940770405","article-title":"(6R,9\u2032Z)-neoxanthin: Synthese, schmelzverhalten, spektren und konformationsberechnungen","volume":"77","author":"Baumeler","year":"1994","journal-title":"Helv. Chim. Acta"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Roy, S., Llewellyn, C., Egeland, E.S., and Johnsen, G. (2011). Phytoplanton Pigments-Characterization, Chemotaxonomy and Applications in Oceanography Pigments, Cambridge University Press.","DOI":"10.1017\/CBO9780511732263"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"363","DOI":"10.4319\/lom.2007.5.363","article-title":"HPLC determination of phytoplankton and microphytobenthos pigments: Comparing resolution and sensitivity of a C18 and a C8 method","volume":"5","author":"Mendes","year":"2007","journal-title":"Limnol. Oceanogr. Methods"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"5180","DOI":"10.1021\/jf300332m","article-title":"Thermal degradation kinetics of neoxanthin, violaxanthin, and antheraxanthin in virgin olive oils","volume":"60","year":"2012","journal-title":"J. Agric. Food Chem."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"24349","DOI":"10.1074\/jbc.271.40.24349","article-title":"Cloning and functional analysis of the \u03b2-carotene hydroxylase of Arabidopsis thaliana","volume":"271","author":"Sun","year":"1996","journal-title":"J. Biol. Chem."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1007\/BF00239993","article-title":"The role of cis-carotenoids in abscisic acid biosynthesis","volume":"182","author":"Parry","year":"1990","journal-title":"Planta"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"6346","DOI":"10.1046\/j.1432-1327.2000.01722.x","article-title":"Identification of neoxanthin synthase as a carotenoid cyclase paralog","volume":"267","author":"Bouvier","year":"2000","journal-title":"Eur. J. Biochem."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1111\/j.1365-313X.2007.03094.x","article-title":"The Arabidopsis ABA-deficient mutant aba4 demonstrates that the major route for stress-induced ABA accumulation is via neoxanthin isomers","volume":"50","author":"North","year":"2007","journal-title":"Plant J."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/S1369-5266(00)00163-1","article-title":"Carotenoid biosynthesis in flowering plants","volume":"4","author":"Hirschberg","year":"2001","journal-title":"Curr. Opin. Plant Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1016\/S0014-5793(00)02193-1","article-title":"Identification of a novel gene coding for neoxanthin synthase from Solanum tuberosum","volume":"485","author":"Hugueney","year":"2000","journal-title":"FEBS Lett."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"11102","DOI":"10.1073\/pnas.190177497","article-title":"An alternative pathway to \u03b2-carotene formation in plant chromoplasts discovered by map-based cloning of Beta and old-gold color mutations in tomato","volume":"97","author":"Ronen","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1146\/annurev.genet.38.072902.092328","article-title":"Chlamydomonas reinhardtii in the landscape of pigments","volume":"38","author":"Grossman","year":"2004","journal-title":"Annu. Rev. Genet."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1104\/pp.104.056069","article-title":"Genome-based examination of chlorophyll and carotenoid biosynthesis in Chlamydomonas reinhardtii","volume":"138","author":"Lohr","year":"2005","journal-title":"Plant Physiol."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"80","DOI":"10.1111\/tpj.12451","article-title":"The tomato mutation nxd1 reveals a gene necessary for neoxanthin biosynthesis and demonstrates that violaxanthin is a sufficient precursor for abscisic acid biosynthesis","volume":"78","author":"Neuman","year":"2014","journal-title":"Plant J."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"1145","DOI":"10.1093\/jexbot\/52.359.1145","article-title":"The pathway of biosynthesis of abscisic acid in vascular plants: A review of the present state of knowledge of ABA biosynthesis","volume":"52","author":"Milborrow","year":"2001","journal-title":"J. Exp. Bot."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1591","DOI":"10.1104\/pp.102.017921","article-title":"Elucidation of the indirect pathway of abscisic acid biosynthesis by mutants, genes, and enzymes","volume":"131","author":"Schwartz","year":"2003","journal-title":"Plant Physiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1146\/annurev.arplant.56.032604.144046","article-title":"Abscisic acid biosynthesis and catabolism","volume":"56","author":"Nambara","year":"2005","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Li, J., Li, C., and Smith, S.M. (2017). Abscisic acid. Hormone Metabolism and Signaling in Plants, Academic Press.","DOI":"10.1016\/B978-0-12-811562-6.00005-0"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"816","DOI":"10.1104\/pp.108.116749","article-title":"Evidence for abscisic acid biosynthesis in Cuscuta reflexa, a parasitic plant lacking neoxanthin","volume":"147","author":"Qin","year":"2008","journal-title":"Plant Physiol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"2859","DOI":"10.1016\/0031-9422(80)85056-4","article-title":"Formation of neoxanthin, diadinoxanthin and peridinin from [14C]zeaxanthin by a cell-free system from Amphidinium carterae","volume":"21","author":"Swift","year":"1980","journal-title":"Phytochemistry"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2261","DOI":"10.1016\/0031-9422(74)85038-7","article-title":"Carotenoids of the dinophyceae","volume":"13","author":"Johansen","year":"1974","journal-title":"Phytochemistry"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"968","DOI":"10.1093\/oxfordjournals.pcp.a029461","article-title":"Distribution and geometric isomerism of neoxanthin in oxygenic phototrophs: 9\u2032-cis, a sole molecular form","volume":"39","author":"Takaichi","year":"1998","journal-title":"Plant Cell Physiol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"9330","DOI":"10.1021\/jp001752i","article-title":"Identifying the pathways of energy transfer between carotenoids and chlorophylls in LHCII and CP29. A multicolor, femtosecond pump\u2212probe study","volume":"104","author":"Gradinaru","year":"2000","journal-title":"J. Phys. Chem. B"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1016\/S0006-3495(01)76069-9","article-title":"Carotenoid-to-chlorophyll energy transfer in recombinant major light-harvesting complex (LHCII) of higher plants. I. Femtosecond transient absorption measurements","volume":"80","author":"Croce","year":"2001","journal-title":"Biophys. J."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"443","DOI":"10.1016\/S0305-1978(99)00078-2","article-title":"Geometrical E\/Z isomers of (6R)- and (6S)-neoxanthin and biological implications","volume":"28","author":"Strand","year":"2000","journal-title":"Biochem. Syst. Ecol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"29613","DOI":"10.1074\/jbc.274.42.29613","article-title":"Carotenoid-binding sites of the major light-harvesting complex II of higher plants","volume":"274","author":"Croce","year":"1999","journal-title":"J. Biol. Chem."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0014-5793(99)00907-2","article-title":"The neoxanthin binding site of the major light harvesting complex (LHCII) from higher plants","volume":"456","author":"Croce","year":"1999","journal-title":"FEBS Lett."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1038\/nature02373","article-title":"Crystal structure of spinach major light-harvesting complex at 2.72 \u00c5 resolution","volume":"428","author":"Liu","year":"2004","journal-title":"Nature"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1038\/sj.emboj.7600585","article-title":"Mechanisms of photoprotection and nonphotochemical quenching in pea light-harvesting complex at 2.5 \u212b resolution","volume":"24","author":"Standfuss","year":"2005","journal-title":"EMBO J."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"570","DOI":"10.1046\/j.1529-8817.2003.02098.x","article-title":"Photosynthetic pigment composition in the primitive green alga Mesostigma viride (Prasinophyceae): Phylogenetic and evolutionary implications","volume":"39","author":"Yoshii","year":"2003","journal-title":"J. Phycol."},{"key":"ref_53","doi-asserted-by":"crossref","unstructured":"Brodie, J., and Lewis, J. (2007). In the shadow of giants: Systematics of the charophyte green algae. Unravelling the Algae: The Past, Present, and Future of Algal Systematics (The Systematic Association Special Volume Series), CRC Press.","DOI":"10.1201\/9780849379901"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"1872","DOI":"10.1126\/science.276.5320.1872","article-title":"Specific oxidative cleavage of carotenoids by VP14 of maize","volume":"276","author":"Schwartz","year":"1997","journal-title":"Science"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1007\/s11120-014-0011-y","article-title":"Light-dependent conformational change of neoxanthin in a siphonous green alga, Codium intricatum, revealed by Raman spectroscopy","volume":"121","author":"Uragami","year":"2014","journal-title":"Photosynth. Res."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"jeb180463","DOI":"10.1242\/jeb.180463","article-title":"Kleptoplast photoacclimation state modulates the photobehaviour of the solar-powered sea slug Elysia viridis","volume":"221","author":"Cartaxana","year":"2018","journal-title":"J. Exp. Biol."},{"key":"ref_57","unstructured":"Giossi, C. (2020). Photoacclimation and Photoprotection Strategies in Siphonous Green Algae of the Order Bryopsidales (Codium tomentosum and Bryopsis plumosa). [Master\u2019s Thesis, University of Bologna]."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"1220","DOI":"10.1021\/bi001160q","article-title":"Photochemical behavior of xanthophylls in the recombinant photosystem II antenna complex, CP26","volume":"40","author":"Frank","year":"2001","journal-title":"Biochemistry"},{"key":"ref_59","first-page":"253","article-title":"Purification and characterization of light-harvesting chlorophyll a\/b-protein complexes of photosystem II from the green alga, Bryopsis maxima","volume":"31","author":"Nakayama","year":"1990","journal-title":"Plant Cell Physiol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1104\/pp.80.1.231","article-title":"Chlorophyll-protein complexes from Euglena gracilis and mutants deficient in chlorophyll b","volume":"80","author":"Cunningham","year":"1986","journal-title":"Plant Physiol."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1007\/BF00035234","article-title":"Energy transfer and pigment composition in three chlorophyll b-containing light-harvesting complexes isolated from Mantoniella squamata (Prasinophyceae), Chlorella fusca (Chlorophyceae) and Sinapis alba","volume":"13","author":"Wilhelm","year":"1987","journal-title":"Photosynth. Res."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"753","DOI":"10.1016\/j.bbabio.2009.03.012","article-title":"Crystallisation, structure and function of plant light-harvesting Complex II","volume":"1787","author":"Barros","year":"2009","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"616","DOI":"10.1046\/j.1432-1327.2000.01060.x","article-title":"Carotenoid binding sites in LHCIIb","volume":"267","author":"Hobe","year":"2000","journal-title":"Eur. J. Biochem."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"35924","DOI":"10.1074\/jbc.M105199200","article-title":"The major antenna complex of Photosystem II has a xanthophyll binding site not involved in light harvesting","volume":"276","author":"Caffarri","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"25156","DOI":"10.1074\/jbc.M604828200","article-title":"Assembly of the major light-harvesting chlorophyll-a\/b complex: Thermodynamics and kinetics of neoxanthin binding","volume":"281","author":"Hobe","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"4877","DOI":"10.1074\/jbc.M114.607770","article-title":"Pigment interactions in light-harvesting complex II in different molecular environments","volume":"290","author":"Akhtar","year":"2015","journal-title":"J. Biol. Chem."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"1490","DOI":"10.1016\/j.bbabio.2016.06.001","article-title":"Probing the pigment binding sites in LHCII with resonance Raman spectroscopy: The effect of mutations at S123","volume":"1857","author":"Kish","year":"2016","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"4704","DOI":"10.1016\/j.febslet.2007.08.066","article-title":"A specific binding site for neoxanthin in the monomeric antenna proteins CP26 and CP29 of Photosystem II","volume":"581","author":"Caffarri","year":"2007","journal-title":"FEBS Lett."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1135","DOI":"10.1073\/pnas.96.3.1135","article-title":"Unusual carotenoid composition and a new type of xanthophyll cycle in plants","volume":"96","author":"Bungard","year":"1999","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"5898","DOI":"10.1021\/acs.jpclett.7b02634","article-title":"Influence of the carotenoid composition on the conformational dynamics of photosynthetic light-harvesting complexes","volume":"8","author":"Tutkus","year":"2017","journal-title":"J. Phys. Chem. Lett."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1038\/nature18020","article-title":"Structure of spinach photosystem II-LHCII supercomplex at 3.2 \u00c5 resolution","volume":"534","author":"Wei","year":"2016","journal-title":"Nature"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1093\/jxb\/erh237","article-title":"Singlet oxygen production in photosynthesis","volume":"56","year":"2004","journal-title":"J. Exp. Bot."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"836","DOI":"10.1063\/1.1699044","article-title":"A theory of sensitized luminescence in solids","volume":"21","author":"Dexter","year":"1953","journal-title":"J. Chem. Phys."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"3143","DOI":"10.1016\/S0006-3495(98)77756-2","article-title":"Rate of carotenoid triplet formation in solubilized light-harvesting complex II (LHCII) from spinach","volume":"75","author":"Irrgang","year":"1998","journal-title":"Biophys. J."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/j.tplants.2009.01.008","article-title":"Singlet oxygen in plants: Production, detoxification and signaling","volume":"14","author":"Havaux","year":"2009","journal-title":"Trends Plant Sci."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Dall\u2019Osto, L., Lico, C., Alric, J., Giuliano, G., Havaux, M., and Bassi, R. (2006). Lutein is needed for efficient chlorophyll triplet quenching in the major LHCII antenna complex of higher plants and effective photoprotection in vivo under strong light. BMC Plant Biol., 6.","DOI":"10.1186\/1471-2229-6-32"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.bbabio.2011.04.012","article-title":"The role of the xanthophyll cycle and of lutein in photoprotection of photosystem II","volume":"1817","author":"Jahns","year":"2012","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"12208","DOI":"10.1021\/bi9711689","article-title":"Xanthophylls in light-harvesting complex II of higher plants: Light harvesting and triplet quenching","volume":"36","author":"Peterman","year":"1997","journal-title":"Biochemistry"},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"6184","DOI":"10.1074\/jbc.M708961200","article-title":"Photoprotection in the antenna complexes of photosystem II: Role of individual xanthophylls in chlorophyll triplet quenching","volume":"283","author":"Mozzo","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"11337","DOI":"10.1021\/jp402977y","article-title":"Chlorophyll triplet quenching and photoprotection in the higher plant monomeric antenna protein Lhcb5","volume":"117","author":"Ballottari","year":"2013","journal-title":"J. Phys. Chem. B"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.jplph.2013.11.008","article-title":"The inter-monomer interface of the major light-harvesting chlorophyll a\/b complexes of photosystem II (LHCII) influences the chlorophyll triplet distribution","volume":"171","author":"Zhang","year":"2014","journal-title":"J. Plant Physiol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"13517","DOI":"10.1021\/jp106323e","article-title":"Structure-based identification of energy dinks in plant light-harvesting complex II","volume":"114","author":"Madjet","year":"2010","journal-title":"J. Phys. Chem. B"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1146\/annurev.arplant.50.1.601","article-title":"The water-water cycle in chloroplasts: Scavenging of active oxygens and dissipation of excess photons","volume":"50","author":"Asada","year":"1999","journal-title":"Annu. Rev. Plant Physiol. Plant Mol. Biol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1237","DOI":"10.1016\/j.bbabio.2011.06.011","article-title":"Light-induced isomerization of the LHCII-bound xanthophyll neoxanthin: Possible implications for photoprotection in plants","volume":"1807","author":"Zubik","year":"2011","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"7","DOI":"10.1093\/jxb\/eru400","article-title":"Evolution under the sun: Optimizing light harvesting in photosynthesis","volume":"66","author":"Ruban","year":"2015","journal-title":"J. Exp. Bot."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1038\/nature06262","article-title":"Identification of a mechanism of photoprotective energy dissipation in higher plants","volume":"450","author":"Ruban","year":"2007","journal-title":"Nature"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"29505","DOI":"10.1074\/jbc.M802438200","article-title":"Induction of efficient energy dissipation in the isolated light-harvesting complex of photosystem II in the absence of protein aggregation","volume":"283","author":"Ilioaia","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1016\/j.tplants.2004.06.006","article-title":"Insights into the molecular dynamics of plant light-harvesting proteins in vivo","volume":"9","author":"Robert","year":"2004","journal-title":"Trends Plant Sci."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"17020","DOI":"10.1074\/jbc.M109.077750","article-title":"Efficient light harvesting by photosystem II requires an optimized protein packing density in grana thylakoids","volume":"285","author":"Haferkamp","year":"2010","journal-title":"J. Biol. Chem."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1038\/nature03795","article-title":"Molecular basis of photoprotection and control of photosynthetic light-harvesting","volume":"436","author":"Pascal","year":"2005","journal-title":"Nature"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"12253","DOI":"10.1039\/c3cp51925b","article-title":"Switching light harvesting complex II into photoprotective state involves the lumen-facing apoprotein loop","volume":"15","author":"Belgio","year":"2013","journal-title":"Phys. Chem. Chem. Phys."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"12508","DOI":"10.1021\/jp804571k","article-title":"Theoretical investigation of the role of strongly coupled chlorophyll dimers in photoprotection of LHCII","volume":"112","author":"Duffy","year":"2008","journal-title":"J. Phys. Chem. B"},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"26007","DOI":"10.1074\/jbc.M115.669804","article-title":"Modeling of the N-terminal section and the lumenal loop of trimeric light harvesting complex II (LHCII) by using EPR","volume":"290","author":"Fehr","year":"2015","journal-title":"J. Biol. Chem."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"86","DOI":"10.1104\/pp.17.00029","article-title":"9-cis-neoxanthin in light harvesting complexes of photosystem II regulates the binding of violaxanthin and xanthophyll cycle","volume":"174","author":"Wang","year":"2017","journal-title":"Plant Physiol."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1515\/botm.1977.20.7.433","article-title":"A carotenoid characteristic of chlorophycean seaweeds living in deep coastal waters","volume":"20","author":"Yokohama","year":"1977","journal-title":"Bot. Mar."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1016\/0005-2728(83)90096-8","article-title":"Chlorophyll-protein complexes of a Codium species, including a light-harvesting siphonaxanthin-Chlorophylla ab-protein complex, an evolutionary relic of some Chlorophyta","volume":"724","author":"Anderson","year":"1983","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/0005-2728(85)90090-8","article-title":"Chlorophyll-protein complexes of a marine green alga, Codium species (Siphonales)","volume":"806","author":"Anderson","year":"1985","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1007\/s11120-013-9808-3","article-title":"Spectral and functional studies on siphonaxanthin-type light-harvesting complex of photosystem II from Bryopsis corticulans","volume":"117","author":"Wang","year":"2013","journal-title":"Photosynth. Res."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1111\/j.1440-1835.2006.00430.x","article-title":"Diversity and evolution of photosynthetic antenna systems in green plants","volume":"54","author":"Yoshii","year":"2006","journal-title":"Phycol. Res."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1007\/s11120-014-0039-z","article-title":"Isolation and characterization of a PSI\u2013LHCI super-complex and its sub-complexes from a siphonaceous marine green alga, Bryopsis Corticulans","volume":"123","author":"Qin","year":"2015","journal-title":"Photosynth. Res."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1111\/j.1365-3040.1996.tb00419.x","article-title":"Photoinhibition of photosynthesis on a coral reef","volume":"19","author":"Franklin","year":"1996","journal-title":"Plant Cell Environ."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"113","DOI":"10.3354\/meps271113","article-title":"Photosynthetic plasticity of an invasive variety of Caulerpa racemosa in a coastal Mediterranean area: Light harvesting capacity and seasonal acclimation","volume":"271","author":"Raniello","year":"2004","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"7904","DOI":"10.1038\/srep07904","article-title":"Photoprotection in sequestered plastids of sea slugs and respective algal sources","volume":"5","author":"Cruz","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"1293","DOI":"10.1007\/s00425-018-2854-5","article-title":"A siphonous morphology affects light-harvesting modulation in the intertidal green macroalga Bryopsis corticulans (Ulvophyceae)","volume":"247","author":"Giovagnetti","year":"2018","journal-title":"Planta"},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.1016\/j.bbabio.2009.05.012","article-title":"Ultrafast fluorescence study on the location and mechanism of non-photochemical quenching in diatoms","volume":"1787","author":"Miloslavina","year":"2009","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"3625","DOI":"10.1016\/j.febslet.2008.09.044","article-title":"Far-red fluorescence: A direct spectroscopic marker for LHCII oligomer formation in non-photochemical quenching","volume":"582","author":"Miloslavina","year":"2008","journal-title":"FEBS Lett."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.bbabio.2013.09.001","article-title":"Stark fluorescence spectroscopy reveals two emitting sites in the dissipative state of FCP antennas","volume":"1837","author":"Wahadoszamen","year":"2014","journal-title":"Biochim. Biophys. Acta Bioenerg."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1016\/j.cplett.2009.10.085","article-title":"Identification of two quenching sites active in the regulation of photosynthetic light-harvesting studied by time-resolved fluorescence","volume":"483","author":"Holzwarth","year":"2009","journal-title":"Chem. Phys. Lett."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"900","DOI":"10.1016\/j.bbabio.2015.01.006","article-title":"Photosystem II repair in plant chloroplasts\u2014Regulation, assisting proteins and shared components with photosystem II biogenesis","volume":"1847","author":"Suorsa","year":"2015","journal-title":"Biochim. Biophys. Acta Bioenerg."}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/20\/4617\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T10:19:16Z","timestamp":1760177956000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/25\/20\/4617"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,10,11]]},"references-count":109,"journal-issue":{"issue":"20","published-online":{"date-parts":[[2020,10]]}},"alternative-id":["molecules25204617"],"URL":"https:\/\/doi.org\/10.3390\/molecules25204617","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,10,11]]}}}