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Other constructs including the full-length and various site-directed mutated human CCK1R and CCK2R were cloned into the pcDNA3.1 vector for cAMP accumulation and NanoBiT–G protein dissociation assay. The intestinal hormone and neuromodulator cholecystokinin (CCK) receptors CCK1R and CCK2R act as a signaling hub in brain–gut axis, mediating digestion, emotion, and memory regulation. CCK receptors exhibit distinct preferences for ligands in different posttranslational modification (PTM) states. They exist in multiple molecular forms, sharing the same evolutionarily conserved pentapeptide motif, which comprises the minimal sequence required for biological activity and receptor activation (Fig. 1a).

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Although CCK-8 is located in a similar orthosteric pocket in CCK1R and CCK2R, the binding modes significantly differ (Fig. 2d, e). N982.61 and R197ECL2 form hydrogen bonds with the sulfonic acid group of TYS, vital for CCK-8 binding to CCK1R. The importance of the hydrogen bonds was reflected in our mutagenesis studies, wherein R197ECL2A decreased the potency of CCK-8 by over 100-fold and N982.61A abolished the efficacy (Fig. 2g, h; Supplementary Tables S2, S3). The outward movement of TM6 is a hallmark of receptor activation and the various extent of the TM6 openness relative to other TMs was proposed to be the determinants of G protein subtype specificity47,48. Surprisingly, structural superimposition of the CCK1R–Gs complex onto the well-studied Gs-coupled GPCR β2AR reveals striking differences regarding the position of TM6, and TM7–Helix8 hinge. CCK1R TM6 locates between the positions of β2AR TM6 in the inactive and active states (Fig. 5c).

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  • But since CCK receptors also have such widespread expression (66, 67, 70–73, 94, 95), there is both room and need for delineation of the roles of CCK released from the “new” sites.
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Moreover, CCK-8 (sulfated as well as non-sulfated) has been reported to exert a wide specter of stimulation and inhibition on lymphocytes, macrophages, and cytokine release, with ensuing anti-inflammatory effects (108–111). The field is complex due to the many players; but the clinical impact of CCK in inflammatory diseases and endotoxin shock may be significant. The distal part of the gut is as mentioned abundantly innervated with CCK neurons (42, 90). It is therefore likely that an increase of intestinal motor activity by exogenous CCK (91) reflects neuronal control of intestinal muscles by CCK peptide transmission. Neuronal CCK acts both indirectly via acetylcholine release from postganglionic parasympathetic nerves and directly on muscle cells (49). The observation that CCK peptides stimulate intestinal blood flow is in harmony with the occurrence of CCK nerve terminals around blood vessels in the basal lamina propria and the submucosa (42).