Structural basis of diadinoxanthin-Chl a/b–binding proteins in the photosystem I supercomplex of Euglena gracilis

Euglena gracilis is a phototrophic flagellate that has evolved through secondary endosymbiotic events and belongs to green-lineage organisms. We report a unique structure of photosystem I-light harvesting complex I (PSI-LHCI) supercomplex from E. gracilis at a 2.23-angstrom resolution by cryo-electron microscopy. The supercomplex is composed of 8 core subunits and 16 LHCIs and exhibits distinctive structural features compared to its counterparts in green algae and plants. The LHCI subunits encircle the core complex, forming a two-layered arrangement that comprises six pairs of tightly packed heterodimers. Specifically, the 16 LHCIs consist of 4 diadinoxanthin-chlorophyll (Chl) a/b-binding antennae (Lhcbm) and 12 diadinoxanthin-Chl a-binding antennae (Lhca), and they exhibit characteristic pigment compositions combining features of green- and red-lineage organisms. These findings provide a robust structural foundation for elucidating the mechanisms of light harvesting and energy transfer as well as insights into the evolutionary changes of green-lineage PSI-LHCI.