Abstract： The cortical grey matter of mammals has a specific cyto-architecture defined by the process of "tiling" in which protoplasmic astrocytes parcellate the nervous tissue into spatially segregated territorial domains. Within these domains astrocytes integrate neuronal structures, microglial cells and neighboring capillaries into the neuro-gliovascular unit or neurogliovascular unit (NGVU) (Verkhratsky and Nedergaard, 2018; Sweeney et al., 2019). Signaling within NGVU is of fundamental importance for sustaining brain function, while disruptions of these signaling pathways define many aspects of neuropathology.