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leukocyte Ig-like receptors (LILR) LILRB1 and LILRB2 are immune checkpoint receptors that regulate a wide range of physiological responses by binding to diverse ligands, including HLA-G. HLA-G is exclusively expressed in the placenta, some immunoregulatory cells, and tumors and has several unique isoforms. However, the recognition of HLA-G isoforms by LILRs is poorly understood. In this study, we characterized LILR binding to the \u03b22-microglobulin (\u03b22m)-free HLA-G1 isoform, which is synthesized by placental trophoblast cells and tends to dimerize and multimerize. The multimerized \u03b22m-free HLA-G1 dimer lacked detectable affinity for LILRB1, but bound strongly to LILRB2. We also determined the crystal structure of the LILRB1 and HLA-G1 complex, which adopted the typical structure of a classical HLA class I complex. LILRB1 exhibits flexible binding modes with the \u03b13 domain, but maintains tight contacts with \u03b22m, thus accounting for \u03b22m-dependent binding. Notably, both LILRB1 and B2 are oriented at suitable angles to permit efficient signaling upon complex formation with HLA-G1 dimers. These structural and functional features of ligand recognition by LILRs provide novel insights into their important roles in the biological regulations.","bibjson":{"author":[{"initials":"K","lastname":"Kuroki","name":"Kuroki K"},{"initials":"H","lastname":"Matsubara","name":"Matsubara H"},{"initials":"R","lastname":"Kanda","name":"Kanda R"},{"initials":"N","lastname":"Miyashita","name":"Miyashita N"},{"initials":"M","lastname":"Shiroishi","name":"Shiroishi M"},{"initials":"Y","lastname":"Fukunaga","name":"Fukunaga Y"},{"initials":"J","lastname":"Kamishikiryo","name":"Kamishikiryo J"},{"initials":"A","lastname":"Fukunaga","name":"Fukunaga A"},{"initials":"H","lastname":"Fukuhara","name":"Fukuhara H"},{"initials":"K","lastname":"Hirose","name":"Hirose K"},{"initials":"JS","lastname":"Hunt","name":"Hunt JS"},{"initials":"Y","lastname":"Sugita","name":"Sugita Y"},{"initials":"S","lastname":"Kita","name":"Kita S"},{"initials":"T","lastname":"Ose","name":"Ose T"},{"initials":"K","lastname":"Maenaka","name":"Maenaka K"}],"identifier":[{"id":"10.4049/jimmunol.1900562","type":"doi"},{"id":"31694909","type":"pubmed"}],"issue":[null],"journal":{"iso_abbreviation":"J. 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