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Thylakoid stacking promotes the physical separation of Photosystems I and II, thereby ensuring a balanced flow of excitation energy between them. Through a genetic dissection of the light-harvesting complex subfamily Lhcb, we showed that removal of the antenna system serving Photosystem II completely abolished grana formation, whereas deletion of specific Lhcb subgroups reduced stacking by up to 40%. Lhcb5 alone was sufficient to partially restore stacking, whereas exclusive expression of Lhcb2 resulted in grana with an unusually large size, which disassembled upon illumination. Time-resolved fluorescence measurements revealed that the extent of excitation energy spillover from PSII to Photosystem I correlated with the level of Lhcb depletion, rather than with granal architecture. These findings provide genetic evidence for a functional link between Photosystem II antenna composition, lateral heterogeneity, and excitation energy partition between photosystems.<\/jats:p>","DOI":"10.1093\/plcell\/koag256","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T08:10:24Z","timestamp":1788250224000},"source":"Crossref","is-referenced-by-count":0,"title":["Specific light-harvesting complexes mediate grana stacking and prevent energy spillover between photosystems in plant chloroplasts"],"prefix":"10.1093","volume":"38","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6773-4047","authenticated-orcid":false,"given":"Zeno","family":"Guardini","sequence":"first","affiliation":[{"name":"Dipartimento di Biotecnologie, Universit\u00e0 di Verona , Strada Le Grazie 15, Verona 37134 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