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Ethics declarations Ethics approval and consent to participate Animal studies and procedures were carried out in accordance with an approved protocol from the Institutional Animal Care and Use Committee (IACUC) at the Fourth Military Medical University (IACUC-20220505)

Several segments in the ECD of GCGR, the stalk between ECD and transmembrane helix 1 (TM1) of GLP-1R (residues 130135), extracellular loop 1 (ECL1) of GIPR (residues 202207), and intracellular loop 3 (ICL3) of GLP-1R and GIPR (residues 338343 and 330333, respectively) were poorly resolved and not modeled in the structures