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The wild-type human CCK1R and CCK2R with C-terminal 15-residue depleted were subcloned into pFastBac vector with an N-terminal FLAG-tag and C-terminal 10× His-tag. A dominant-negative human Gαs (DNGαs) was generated by site-directed mutagenesis as previously described to limit G protein dissociation. Twenty-nine amino acids at the N-terminus of wild-type Gαq were replaced by the corresponding sequence in Gαi1 to facilitate the binding of scFv16. The constructs were cloned into both pcDNA3.1 and pFastBac vectors for functional assays in mammalian cells and protein expression in insect cells, respectively. All modifications of the receptor had no effect on ligand binding and receptor activation.
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- The long history has made the CCK literature comprehensive and at some points also confusing because impure CCK preparations with little attention paid to species differences and to physiological levels were used initially.
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In the CCK2R–Gq complex, Gq undergoes an anticlockwise rotation relative to the receptor compared to Gs (except for EP4R–Gs) (Supplementary Fig. S5a, f). This difference makes the α5 helix of Gq moving closer to the TM2–TM3–TM4 half (Supplementary Fig. S5h, i). It remains mysterious that the CCKRs with highly conserved orthosteric binding pocket display distinct affinity and potency on various endogenous ligands, according to previous homology modeling and mutagenesis studies45.
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.