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Cholecystokinin (CCK) receptor family belongs to class-A sevenfold transmembrane G protein-coupled receptors, and is divided into CCK1 receptor (CCK1R) and CCK2 receptor (CCK2R)1. CCK1R is mainly distributed in the gastrointestinal tract2,3,4, peripheral nervous system5,6, and some regions of the brain, e.g., the area postrema, the nucleus tractus solitarius, and the hypothalamus7,8,9,10. Hence, CCK receptors (CCKRs) regulate a variety of physiological functions including digestion, satiety, emotion regulation, pain sensation, and memory process18,19,20,21,22,23,24,25. Besides, CCKRs are expressed in the brain–gut axis which is a region critical for the transmission of information between gut and brain including satiation signals. For these functions, there are already some specific drugs designed for CCK1R, such as ceruletide, or for CCK2R, including proglumide and pentagastrin. Although all of them have been used for gastrointestinal diseases, they and some drugs under clinical trial also have the potential to treat CNS diseases, including pain and anxiety26,27.
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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).
There are species differences, so it is now assumed that CCK in man stimulates pancreatic enzyme secretion through a cholinergic pathway that is less significant in rodents (82–84). The homologous bioactive sequences of peptide systems belonging to the cholecystokinin (CCK) family (upper panel). CCK and the antral hormone, gastrin, are the only mammalian members of the family. Caerulein and phyllocaerulein are identified from frogskin extracts. Cionin is a neuropeptide isolated from the central ganglion of the protochord, ciona intestinalis.