{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T03:00:58Z","timestamp":1789441258952,"version":"build-2803163510"},"reference-count":34,"publisher":"MDPI AG","issue":"11","license":[{"start":{"date-parts":[[2023,5,26]],"date-time":"2023-05-26T00:00:00Z","timestamp":1685059200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"JSPS KAKENHI","award":["JP20H02891"],"award-info":[{"award-number":["JP20H02891"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>The barley cultivar Sarab 1 (SRB1) can continue photosynthesis despite its low Fe acquisition potential via roots and dramatically reduced amounts of photosystem I (PSI) reaction-center proteins under Fe-deficient conditions. We compared the characteristics of photosynthetic electron transfer (ET), thylakoid ultrastructure, and Fe and protein distribution on thylakoid membranes among barley cultivars. The Fe-deficient SRB1 had a large proportion of functional PSI proteins by avoiding P700 over-reduction. An analysis of the thylakoid ultrastructure clarified that SRB1 had a larger proportion of non-appressed thylakoid membranes than those in another Fe-tolerant cultivar, Ehimehadaka-1 (EHM1). Separating thylakoids by differential centrifugation further revealed that the Fe-deficient SRB1 had increased amounts of low\/light-density thylakoids with increased Fe and light-harvesting complex II (LHCII) than did EHM1. LHCII with uncommon localization probably prevents excessive ET from PSII leading to elevated NPQ and lower PSI photodamage in SRB1 than in EHM1, as supported by increased Y(NPQ) and Y(ND) in the Fe-deficient SRB1. Unlike this strategy, EHM1 may preferentially supply Fe cofactors to PSI, thereby exploiting more surplus reaction center proteins than SRB1 under Fe-deficient conditions. In summary, SRB1 and EHM1 support PSI through different mechanisms during Fe deficiency, suggesting that barley species have multiple strategies for acclimating photosynthetic apparatus to Fe deficiency.<\/jats:p>","DOI":"10.3390\/plants12112111","type":"journal-article","created":{"date-parts":[[2023,5,27]],"date-time":"2023-05-27T16:16:04Z","timestamp":1685204164000},"page":"2111","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":5,"title":["Barley Cultivar Sarab 1 Has a Characteristic Region on the Thylakoid Membrane That Protects Photosystem I under Iron-Deficient Conditions"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5220-4895","authenticated-orcid":false,"given":"Akihiro","family":"Saito","sequence":"first","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kimika","family":"Hoshi","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuna","family":"Wakabayashi","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Takumi","family":"Togashi","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tomoki","family":"Shigematsu","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maya","family":"Katori","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2469-5890","authenticated-orcid":false,"given":"Takuji","family":"Ohyama","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5963-9402","authenticated-orcid":false,"given":"Kyoko","family":"Higuchi","sequence":"additional","affiliation":[{"name":"Laboratory of Biochemistry in Plant Productivity, Department of Agricultural Chemistry, Tokyo University of Agriculture, Setagaya-ku, Tokyo 156-8502, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2023,5,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1080\/01904168909363962","article-title":"Iron regulation of chloroplast photosynthetic function: Mediation of PS I development","volume":"12","author":"Pushnik","year":"1989","journal-title":"J. Plant Nutr."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1111\/j.1399-3054.1995.tb05127.x","article-title":"Correlation of Iron Content, Spectral Forms of chlorophyll and chlorophyll-proteins in Iron Deficient Cucumber (Cucumis sativus)","volume":"93","author":"Fodor","year":"1995","journal-title":"Physiol. Plant."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1023\/A:1006282714942","article-title":"The role of trace metals in photosynthetic electron transport in O2-evolving organisms","volume":"60","author":"Raven","year":"1999","journal-title":"Photosynth. Res."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1016\/j.tplants.2020.03.003","article-title":"Chloroplast transition metal regulation for efficient photosynthesis","volume":"25","author":"Schmidt","year":"2020","journal-title":"Trends Plant Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"505","DOI":"10.1080\/00380768.2022.2106115","article-title":"Elucidation of efficient photosynthesis in plants with limited iron","volume":"68","author":"Higuchi","year":"2022","journal-title":"Soil Sci. Plant Nutr."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Saito, A., Shinjo, S., Ito, D., Doi, Y., Sato, A., Wakabayashi, Y., Honda, J., Arai, Y., Maeda, T., and Ohyama, T. (2021). Enhancement of photosynthetic iron-use efficiency is an important trait of Hordeum vulgare for adaptation of photosystems to iron deficiency. Plants, 10.","DOI":"10.3390\/plants10020234"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Higuchi, K., Kurita, K., Sakai, T., Suzui, N., Sasaki, M., Katori, M., Wakabayashi, Y., Majima, Y., Saito, A., and Ohyama, T. (2022). \u2018Live-Autoradiography\u2019 technique reveals genetic variation in the rate of Fe uptake by barley cultivars. Plants, 11.","DOI":"10.3390\/plants11060817"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1038\/35089104","article-title":"A giant chlorophyll-protein complex induced by iron deficiency in cyanobacteria","volume":"412","author":"Boekema","year":"2001","journal-title":"Nature"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"232","DOI":"10.1038\/s42003-020-0949-6","article-title":"Structure of a cyanobacterial photosystem I surrounded by octadecameric IsiA antenna proteins","volume":"3","author":"Akita","year":"2020","journal-title":"Commun. Biol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"36","DOI":"10.1111\/j.0031-9317.2004.0229.x","article-title":"Adaptation of the photosynthetic electron transport chain in cyanobacteria to iron deficiency: The function of IdiA and IsiA","volume":"120","author":"Michel","year":"2004","journal-title":"Physiol. Plant."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"10305","DOI":"10.1074\/jbc.M511057200","article-title":"A chlorophyll a\/b-binding Protein Homolog That Is Induced by Iron Deficiency Is Associated with Enlarged photosystem I Units in the Eucaryotic Alga Dunaliella salina","volume":"281","author":"Varsano","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"20431","DOI":"10.1074\/jbc.M414486200","article-title":"N-terminal processing of Lhca3 Is a key step in remodeling of the photosystem I-light-harvesting complex under iron deficiency in Chlamydomonas reinhardtii","volume":"280","author":"Naumann","year":"2005","journal-title":"J. Biol. Chem."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"3964","DOI":"10.1002\/pmic.200700407","article-title":"Comparative quantitative proteomics to investigate the remodeling of bioenergetic pathways under iron deficiency in Chlamydomonas reinhardtii","volume":"7","author":"Naumann","year":"2007","journal-title":"Proteomics"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1007\/s11120-018-0580-2","article-title":"Changes in the photosynthetic apparatus and lipid droplet formation in Chlamydomonas reinhardtii under iron deficiency","volume":"139","author":"Devadasu","year":"2019","journal-title":"Photosynth. Res."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"6709","DOI":"10.1093\/emboj\/cdf666","article-title":"Adaptation to Fe-deficiency requires remodeling of the photosynthetic apparatus","volume":"21","author":"Moseley","year":"2002","journal-title":"EMBO J."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"2649","DOI":"10.1105\/tpc.112.098962","article-title":"Fe sparing and Fe recycling contribute to increased superoxide dismutase capacity in iron-starved Chlamydomonas reinhardtii","volume":"24","author":"Page","year":"2012","journal-title":"Plant Cell"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2013","DOI":"10.1093\/pcp\/pcq160","article-title":"Remodeling of the major light-harvesting antenna protein of PSII protects the young leaves of barley (Hordeum vulgare L.) from photoinhibition under prolonged iron deficiency","volume":"51","author":"Saito","year":"2010","journal-title":"Plant Cell Physiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"2042","DOI":"10.1016\/j.febslet.2014.04.031","article-title":"Fe deficiency induces phosphorylation and translocation of Lhcb1 in barley thylakoid membranes","volume":"588","author":"Saito","year":"2014","journal-title":"FEBS Lett."},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Miyake, C. (2020). Molecular mechanism of oxidation of P700 and suppression of ROS Production in photosystem I in response to electron-sink limitations in C3 plants. Antioxidants, 9.","DOI":"10.3390\/antiox9030230"},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Kaiser, E., Walther, D., and Armbruster, U. (2020). Photorespiration enhances acidification of the thylakoid lumen, reduces the plastoquinone pool, and contributes to the oxidation of P700 at a lower partial pressure of CO2 in wheat leaves. Plants, 9.","DOI":"10.3390\/plants9030319"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2954","DOI":"10.1111\/pce.14400","article-title":"Plants acclimate to photosystem I photoinhibition by readjusting the photosynthetic machinery","volume":"45","author":"Aro","year":"2022","journal-title":"Plant Cell Environ."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"3515","DOI":"10.1098\/rstb.2012.0373","article-title":"Towards elucidation of dynamic structural changes of plant thylakoid architecture","volume":"367","author":"Anderson","year":"2012","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1983","DOI":"10.1111\/ppl.13408","article-title":"Growth-phase dependent morphological alteration in higher plant thylakoid is accompanied by changes in both photodamage and repair rates","volume":"172","author":"Nozue","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1617","DOI":"10.3389\/fpls.2018.01617","article-title":"Oxidation of P700 ensures robust photosynthesis","volume":"9","author":"Shimakawa","year":"2018","journal-title":"Front. Plant Sci."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1016\/bs.abr.2020.08.001","article-title":"P700 oxidation suppresses the production of reactive oxygen species in photosystem I","volume":"96","author":"Furutani","year":"2020","journal-title":"Adv. Bot. Res."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1093\/pcp\/pcaa155","article-title":"The mechanism of non-photochemical quenching in plants: Localization and driving forces","volume":"62","author":"Ruban","year":"2021","journal-title":"Plant Cell Physiol."},{"key":"ref_27","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_28","doi-asserted-by":"crossref","first-page":"326","DOI":"10.1080\/23818107.2019.1619195","article-title":"Spatial organization of thylakoid network in higher plants","volume":"166","author":"Kowalewska","year":"2019","journal-title":"Bot. Lett."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e00280","DOI":"10.1002\/pld3.280","article-title":"Measuring the dynamic response of the thylakoid architecture in plant leaves by electron microscopy","volume":"4","author":"Li","year":"2020","journal-title":"Plant Direct"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"791124","DOI":"10.3389\/fpls.2021.791124","article-title":"Photosystem I inhibition, protection and signalling: Knowns and unknowns","volume":"12","author":"Aro","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"971","DOI":"10.1016\/j.tplants.2022.04.006","article-title":"An ancient function of PGR5 in iron delivery?","volume":"27","author":"Leister","year":"2022","journal-title":"Trends Plant Sci."},{"key":"ref_32","first-page":"27","article-title":"Complementary PS II quantum yields calculated from simple fluorescence parameters measured by PAM fluorometry and the Saturation Pulse method","volume":"1","author":"Klughammer","year":"2008","journal-title":"PAM Appl. Notes"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"384","DOI":"10.1016\/S0005-2728(89)80347-0","article-title":"Determination of accurate extinction coefficients and simultaneous-equations for assaying chlorophyll-a and chlorophyll-B extracted with 4 different solvents\u2014Verification of the concentration of chlorophyll standards by atomic-absorption spectroscopy","volume":"975","author":"Porra","year":"1989","journal-title":"Biochim. Biophys. Acta"},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Ohgane, K., and Yoshioka, H. (2019). Quantification of Gel Bands by an ImageJ Macro, Band\/Peak Quantification Tool, Protocols.io.","DOI":"10.17504\/protocols.io.7vghn3w"}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/12\/11\/2111\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,10]],"date-time":"2025-10-10T19:42:41Z","timestamp":1760125361000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/12\/11\/2111"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,5,26]]},"references-count":34,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2023,6]]}},"alternative-id":["plants12112111"],"URL":"https:\/\/doi.org\/10.3390\/plants12112111","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,5,26]]}}}