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We investigate sensing and signaling mechanisms for H + , [Formula: see text] and CO 2  in basilar arteries using out-of-equilibrium solutions. Selectively varying pH o , [[Formula: see text]] o,  or pCO 2 , we find: (a) lowering pH o  attenuates vasoconstriction and vascular smooth muscle cell (VSMC) Ca 2+ -responses whereas raising pH o  augments vasoconstriction independently of VSMC [Ca 2+ ] i , (b) lowering [[Formula: see text]] o  increases arterial agonist-sensitivity of tone development without affecting VSMC [Ca 2+ ] i  but c) no evidence that CO 2  has direct net vasomotor effects. Receptor protein tyrosine phosphatase (RPTP)γ is transcribed in endothelial cells, and direct vasomotor effects of [Formula: see text] are absent in arteries from RPTPγ-knockout mice. At pH o  7.4, selective changes in [[Formula: see text]] o  or pCO 2  have little effect on pH i . At pH o  7.1, decreased [[Formula: see text]] o  or increased pCO 2  causes intracellular acidification, which attenuates vasoconstriction. Under equilibrated conditions, anti-contractile effects of CO 2 /[Formula: see text] are endothelium-dependent and absent in arteries from RPTPγ-knockout mice. With CO 2 /[Formula: see text] present, contractile responses to agonist-stimulation are potentiated in arteries from RPTPγ-knockout compared to wild-type mice, and this difference is larger for respiratory than metabolic acidosis. In conclusion, decreased pH o  and pH i  inhibit vasoconstriction, whereas decreased [[Formula: see text]] o  promotes vasoconstriction through RPTPγ-dependent changes in VSMC Ca 2+ -sensitivity. [Formula: see text] serves dual roles, providing substrate for pH i -regulating membrane transporters and modulating arterial responses to acid–base disturbances.

Extracellular ʜ ᴄ ᴏ3̅ is sensed by mouse cerebral arteries: Regulation of tone by receptor protein tyrosine phosphatase γ