GPCRs (green) are bound by an orthosteric agonist (gold ball) and an allosteric modulator (blue triangle). Binding of the agonist to the receptor results in signaling through multiple pathways - labeled by ‘a’, ‘b’ and ‘c’ (arrow thickness and pathway labels indicate the strength of the signaling outcome). Dose-response curves (below) show the effect of agonist on a given signaling outcome in the presence of increasing allosteric modulator concentrations, indicated by increasing blue color saturation. Newest among the classes of allosteric modulators are the BAM and ago-BAM. BAMs are pure modulators that lack intrinsic activity (τ=0) and may or may not alter the binding affinity (α) of the agonist. Characteristically, BAMs enhance signaling (β>1) through some pathways but exert no effect (β=1) or antagonize signaling (β<1) through other pathway subsets. Ago-BAMs act as biasing modulators to exert pathway-specific effects on agonist signaling and act alone as biased allosteric agonists, that is, τ is greater than 0 for some pathways, but not others. (A) This hypothetical BAM potentiates agonist-induced signaling through pathway ‘a’ while antagonizing agonist-induced signaling through pathways ‘b’ and ‘c’. (B) This hypothetical ago-BAM stimulates signaling through pathway ‘a’ when applied alone but has no effect on pathways ‘b’ or ‘c’. In combination with an orthosteric agonist, this ago-BAM enhances agonist signaling efficacy through pathway ‘a’ while antagonizing signaling through ‘b’ and ‘c’. (C-E) Application of Eq. 2, a symmetrical version of the extended operational model, to the agonist ago-BAM pair of neurotensin (NTS) and SBI-553 at the NTSR1. (C) NTS alone acts as a balanced, full orthosteric agonist, activating both Gαq- and β-arrestin-mediated cellular responses. (D) SBI-553 alone acts as a biased allosteric agonist, activating β-arrestin- but not Gαq-mediated cellular responses. (E) SBI-553 confers β-arrestin bias to NTS by concurrently enhancing NTS-induced β-arrestin recruitment and antagonizing NTS-induced Gαq activation. The qualitative behavior depicted here has been directly observed with the action on SBI-553 on NTSR1 [40, 41].