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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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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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For the SR146131–CCK1R–Gs–Nb35 complex, auto-picking-yielded 5,213,984 particle projections were subjected to 3D classification to discard particles in poorly defined classes, producing 898,915 particle projections. The re-picked particles were extracted and subjected to two rounds of 3D classification with four classes; the best-resolved class was selected each time. The resulting set of 606,684 particles was subjected to CTF refinement, Bayesian polishing and 3D auto refinement using the pixel size of 1.014 Å. The final map has an indicated global resolution of 3.0 Å at an FSC of 0.143. For the CCK-8–CCK1R–Gs–Nb35 complex, auto-picking-yielded 5,768,688 particle projections were subjected to 3D classification to discard particles in poorly defined classes, producing 1,608,563 particle projections.
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Extraintestinal Endocrine Cells
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In this paper, we reported four high-resolution cryo-EM structures of CCK1R and CCK2R signaling complexes. 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. The sulfation level of CCK and gastrin is also correlated with sub-cellular and cellular distribution. Interestingly, the non-sulfated CCK is enriched in neuronal cell bodies, whereas the sulfated CCK predominantly distributes in the nerve terminals62. CCKRs differentially recognize the sulfated and non-sulfated ligands enabling subtle and precisely regulated responses to extracellular stimuli.
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