The unique binding mode of gastrin in CCK2R permits its selective activation of CCK2R, which is also known as the gastrin receptor. Last, the ECL2 lid possesses a key and exclusive residue R197ECL2 in CCK1R, leading to a positively charged sub-pocket for TYS. Cholecystokinin (CCK) was discovered in 1928 in jejunal extracts as a gallbladder contraction factor. https://cck444.com/ It was later shown to be member of a peptide family, which are all ligands for the CCK1 and CCK2 receptors.
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CCK peptides release insulin and glucagon more potently in man and pig than in dog and rat (51, 85–87). The difference is partly due to neurons in pancreatic islets that release CCK-8 and CCK-5 in man and pig (51), whereas rat and dog islets have no such innervation (50, 51). Moreover, islet cells in man and pig also express the CCK2 receptor abundantly (73), whereas rat islet cells express mainly the CCK1 receptor (88). While the interest in the effect of CCK on the exocrine pancreas was for many years restricted to enzyme secretion, it is now well established that CCK also stimulates fluid and bicarbonate secretion.
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A small fragment (SmBiT) was N-terminally fused to a C68-mutated Gγ2 subunit with a 15-amino acid flexible linker. HEK293T cells were seeded in a six-well plate and allowed to grow to 80% confluence before transfection. For measuring Gs signaling, a plasmid mixture consisting of 200 ng LgBiT-inserted Gαs subunit, 1000 ng Gβ1, 1000 ng C68S-mutant SmBiT-fused Gγ2 (C68S), 400 ng test GPCR in 200 μL of Opti-MEM (Gibco) was transiently transfected with PEI. After 1-day transfection, transfected cells were plated onto a 96-well plate treated by cell adherent reagent (Applygen).
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Conceived and supervised the project and wrote the manuscript with inputs from all co-authors. CCK1R and CCK2R Gs-mediated Gs-cAMP accumulation assays were performed using HEK293T cells (ATCCCRL-11268) transiently expressing human CCK1R and the cAMP biosensor GloSensor-22F (Promega). Cells were seeded (4 × 103 cells, 40 μL/well) into 384-well culture plates and incubated for 24 h at 37 °C in 5% CO2. Next day, the culture medium was removed and the equilibration medium containing 4% (v/v) dilution of the GloSensor™ cAMP reagent stock solution was added to each well.
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- The CCK2R–Gq structure is similar to other reported class-A GPCR–Gq complexes regarding the overall conformation of TMD and the positioning of TM6 (Supplementary Fig. S5d).
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- Sulfation of CCK and gastrin in the body is highly regulated by sulfotransferase (SULT) enzymes with tissue-specific expression patterns, e.g., SULT4 is predominantly expressed in the brain61.
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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. In addition, most assays for measurement of CCK in plasma and elsewhere lacked specificity and sensitivity (10–12). The cellular effects of CCK peptides are mediated via two receptors (66, 67). CCK1 receptors have been found also in the anterior pituitary, the myenteric plexus, and areas of the midbrain (69, 70). The CCK1 receptor binds with high affinity CCK peptides that are amidated and sulfated, whereas the affinity for non-sulfated CCK peptides and gastrins is negligible. In the basal state, the concentration of CCK in plasma is around 1 pmol/l, but often less.
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).