{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\n GPR55 is an atypical G protein–coupled receptor that has emerged as a novel cannabinoid receptor because it is activated not only by select exogenous cannabinoids but also by endogenous lipid ligands such as lysophosphatidylinositol (LPI)."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "4"}]}, {"type": "t", "text": " Upon activation, GPR55 couples primarily to G_12/13 and G_q proteins to trigger several downstream signaling cascades—including mobilization of intracellular calcium, activation of RhoA, ERK1/2 and p38 MAPK pathways, and induction of transcription factors such as NFAT and NF-κB."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "5", "end_ref": "7"}]}, {"type": "t", "text": " These signaling events underpin its diverse physiological roles."}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n In the nervous system, for example, GPR55 modulates synaptic transmission and motor coordination by increasing presynaptic release probability."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "8"}]}, {"type": "t", "text": " In immune cells, activation of GPR55 in monocytes and natural killer cells enhances proinflammatory cytokine production and cytotoxic responses, indicating an immunomodulatory function."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "10"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n GPR55 also plays an important role in bone physiology by regulating osteoclast number and function, thereby influencing bone resorption and remodeling."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "11"}]}, {"type": "t", "text": " In metabolic tissues, its expression in pancreatic β‐cells and adipocytes enables GPR55 activation to stimulate calcium‐dependent insulin secretion and modulate lipogenic gene expression, implicating this receptor in glucose homeostasis and obesity."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "12", "end_ref": "14"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n In cancer, aberrant overexpression and signaling via GPR55 have been linked to enhanced proliferation, migration, and tumor aggressiveness in several tumor types including those of the colon, pancreas, prostate, ovary, and skin."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "15", "end_ref": "18"}]}, {"type": "t", "text": " Furthermore, post‐transcriptional regulation of GPR55—such as via microRNAs—can suppress its expression and consequently inhibit cancer cell growth and metastasis."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "19"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n Additional complexity is added by the ability of GPR55 to form heteromers with classical cannabinoid receptors like CB₁ and CB₂, thereby modulating signaling specificity and intensity."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "20"}]}, {"type": "t", "text": " In the gastrointestinal tract, GPR55 is expressed on myenteric neurons where its activation modulates motility by influencing neuronal excitability and muscle contraction."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "22"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n Collectively, studies indicate that GPR55 functions as a multifunctional receptor involved in neuromodulation, immunomodulation, bone remodeling, metabolic regulation, and oncogenesis. 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