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LHCB4 is known to stabilize LHCII attachment to the PSII core dimer (C\n                    <jats:sub>2<\/jats:sub>\n                    ). Here, we show that LHCB4 is essential for the intrinsic stability of the PSII core dimer itself, a fundamental structural unit required for efficient photochemistry. In the\n                    <jats:italic>lhcb4<\/jats:italic>\n                    mutant, clear\u2010native PAGE revealed the absence of C\n                    <jats:sub>2<\/jats:sub>\n                    S\n                    <jats:sub>2<\/jats:sub>\n                    M\n                    <jats:sub>(2)<\/jats:sub>\n                    and only small amounts of C\n                    <jats:sub>2<\/jats:sub>\n                    S\n                    <jats:sub>2<\/jats:sub>\n                    and C\n                    <jats:sub>2<\/jats:sub>\n                    S supercomplexes. Single\u2010particle electron microscopy identified two types of C\n                    <jats:sub>2<\/jats:sub>\n                    S\n                    <jats:sub>(2)<\/jats:sub>\n                    supercomplexes: a canonical configuration with an empty LHCB4 site and a more abundant \u201cpacked\u201d structure, arising from a relative shift of the C(S) monomers. However, most PSII particles were present as monomeric CS complexes with attached LHCB5 antenna. In contrast, PSII analyzed directly in thylakoid membranes retained the canonical C\n                    <jats:sub>2<\/jats:sub>\n                    organization and even formed nonparallel associations of C\n                    <jats:sub>2<\/jats:sub>\n                    S\n                    <jats:sub>2<\/jats:sub>\n                    supercomplexes, indicating that the \u201cpacked\u201d C\n                    <jats:sub>2<\/jats:sub>\n                    S\n                    <jats:sub>2<\/jats:sub>\n                    and CS forms arise during isolation due to reduced PSII dimer stability. The PSII core dimer also readily disintegrates in the mutant lacking all monomeric antenna proteins (NoM), implying that LHCB4 is critical for the stability of the C\n                    <jats:sub>2<\/jats:sub>\n                    dimer itself. Consistently,\n                    <jats:italic>lhcb4<\/jats:italic>\n                    plants exhibited reduced thermal stability of PSII macro\u2010organization and photochemistry in\u00a0vivo. Our findings demonstrate that LHCB4 is not merely a peripheral antenna connector, but also an important structural determinant of PSII core dimer stability, underscoring its essential role in efficient photochemistry and photoprotection.\n                  <\/jats:p>","DOI":"10.1111\/ppl.71144","type":"journal-article","created":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T14:29:20Z","timestamp":1791556160000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Light\u2010Harvesting Protein\n                    <scp>LHCB4<\/scp>\n                    Is Important for Stable Dimeric Organization of Photosystem\n                    <scp>II<\/scp>"],"prefix":"10.1111","volume":"178","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-9156-839X","authenticated-orcid":false,"given":"Tereza","family":"V\u00e1nsk\u00e1","sequence":"first","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3833-6971","authenticated-orcid":false,"given":"Monika","family":"Opat\u00edkov\u00e1","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8211-3348","authenticated-orcid":false,"given":"Roman","family":"Kou\u0159il","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2849-3084","authenticated-orcid":false,"given":"Zuzana","family":"Ku\u010derov\u00e1","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7079-3160","authenticated-orcid":false,"given":"V\u00e1clav","family":"Karlick\u00fd","sequence":"additional","affiliation":[{"name":"Department of Physics, Faculty of Science University of Ostrava  Ostrava Czech Republic"},{"name":"Global Change Research Institute of the Czech Academy of Sciences  Brno Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3547-3110","authenticated-orcid":false,"given":"Iva","family":"Il\u00edkov\u00e1","sequence":"additional","affiliation":[{"name":"Institute of Experimental Botany of the Czech Academy of Sciences, Centre of Plant Structural and Functional Genomics  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0226-6796","authenticated-orcid":false,"given":"Marek","family":"R\u00e1c","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6904-7646","authenticated-orcid":false,"given":"Pavel","family":"Roudnick\u00fd","sequence":"additional","affiliation":[{"name":"Central European Institute of Technology Masaryk University  Brno Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-8226-1577","authenticated-orcid":false,"given":"Vojt\u011bch","family":"Ch\u00fdlek","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8929-8290","authenticated-orcid":false,"given":"Petr","family":"Il\u00edk","sequence":"additional","affiliation":[{"name":"Department of Biophysics, Faculty of Science Palack\u00fd University  Olomouc Czech Republic"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,9]]},"reference":[{"issue":"1","key":"e_1_2_9_2_1","doi-asserted-by":"crossref","first-page":"336","DOI":"10.1111\/ppl.12964","article-title":"Structural and Functional Differentiation of the Light\u2010Harvesting Protein Lhcb4 During Land Plant Diversification","volume":"166","author":"Albanese P.","year":"2019","journal-title":"Physiologia Plantarum"},{"issue":"10","key":"e_1_2_9_3_1","doi-asserted-by":"crossref","first-page":"1651","DOI":"10.1016\/j.bbabio.2016.06.011","article-title":"Dynamic Reorganization of Photosystem II Supercomplexes in Response to 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