Synaptic targets and cellular sources of CB1 cannabinoid receptor and
vesicular glutamate transporter-3 expressing nerve terminals in relation
to GABAergic neurons in the human cerebral cortex
Abstract
Cannabinoid receptor 1 (CB1) regulates synaptic transmission through
presynaptic receptors in nerve terminals, and its physiological roles
are of clinical relevance. The cellular sources and synaptic targets of
CB1-expressing terminals in the human cerebral cortex are undefined. We
demonstrate a variable laminar pattern of CB1-immunorective axons and
electron microscopically show that CB1-positive GABAergic terminals make
type-2 synapses innervating dendritic shafts (69%), dendritic spines
(20%) and somata (11%) in neocortical layers 2-3. Of the
CB1-immunopositive GABAergic terminals, 25% were
vesicular-glutamate-transporter-3 (VGLUT3)-immunoreactive, suggesting
GABAergic/glutamatergic co-transmission on dendritic shafts. In vitro
recorded and labelled VGLUT3 or CB1-positive GABAergic interneurons
expressed cholecystokinin, vasoactive-intestinal-polypeptide and
calretinin, had diverse firing, axons and dendrites, and included
rosehip, neurogliaform and basket cells, but not double bouquet or
axo-axonic cells. CB1-positive interneurons innervated pyramidal cells
and GABAergic interneurons. Most glutamatergic synaptic terminals formed
type-1 synapses and some were positive for CB1 receptor concentrated in
the presynaptic active zone, unlike in GABAergic terminals. From the
sampled VGLUT3-positive terminals, 60% formed type-1 synapses with
dendritic spines (80%) or shafts (20%) and 52% were also positive for
VGLUT1, suggesting intracortical origin. Some VGLUT3-positive terminals
were immunopositive for vesicular-monoamine-transporter-2, suggesting
5-HT/glutamate co-transmission. Overall, the results show that CB1
regulates GABA release mainly to dendritic shafts of both pyramidal
cells and interneurons, and predict CB1-regulated co-release of GABA and
glutamate from single cortical interneurons. We also demonstrate the
co-existence of multiple vesicular glutamate transporters in a select
population of terminals probably originating from cortical neurons and
innervating dendritic spines in the human cerebral cortex.