Recent advances in structural biology have enabled determination of structures of all 15 class B1 G protein-coupled receptors (GPCRs). While these snapshots provide insights into receptor activation mechanisms, the importance of bound agonist dynamics is underappreciated. Here, we directly observe conformational plasticity in the N-terminus of a class B1 GPCR agonist and provide evidence that these dynamics are vital for activity.1
The glucagon-like peptide 1 receptor (GLP-1R) is an archetypal class B1 GPCR, a critical regulator of glucose homeostasis, and an established target for the treatment of type II diabetes and obesity.2 In recent, full-length structures of the GLP-1R, the critical N-terminus of GLP-1 is rigidly bound in a helical state.3 However, previous studies postulated the existence of a helix-capping motif in class B1 hormones in this region.4 To better understand the determinants of GLP-1R activation by peptide hormones, we systematically probed the N-terminus of GLP-1 and exenatide, a high affinity, high potency, GLP-1R agonist drug, by incorporating single amino acid substitutions in place of a highly conserved glycine residue. We found that disfavoring agonist helicity by incorporation of a single D-alanine residue minimally affects potency but decreases affinity. Conversely, incorporation of another unnatural amino acid, one that readily adopts a helix, abrogates activity but not affinity, yielding a potent inhibitor. These data suggest that the ability of GLP-1 to access non-helical conformations is important for activity. We then used single particle cryo-electron microscopy (cryo-EM) to visualize a potent agonist containing D-alanine bound to GLP-1R/Gs heterotrimer. 3D classification and 3D variance analysis of the cryo-EM data revealed that, unlike GLP-1, the N-terminal region of this analogue transited through multiple conformations while bound to the receptor. Molecular dynamics simulations supported the hypothesis of a mobile, unwinding peptide N-terminus concurrent with outward extracellular loop 3 (ECL3) movement that was also observed in analysis of the cryo-EM data. Collectively, our evidence supports the hypothesis that agonist mobility is important for the activation of the GLP-1R.