{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nThe calcitonin receptor (CALCR) is a prototypical class B G protein–coupled receptor whose structure and ligand‐binding properties have been illuminated by high‐resolution studies. Cryo–electron microscopy and photoaffinity labeling have revealed dynamic rearrangements in its transmembrane and extracellular domains—mapping key residues that directly contact calcitonin and its analogues—and demonstrated that receptor activity–modifying proteins (RAMPs) can allosterically tune the receptor’s peptide selectivity and affinity. These findings underscore a common ligand–binding mode shared among family members."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "7"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nUpon activation, CALCR undergoes tonic internalization and is sorted into recycling vesicles—a process that is influenced by its interactions with cytoskeletal proteins such as filamin. Differential ligand–induced signaling patterns (for example, prolonged cAMP production and β–arrestin recruitment by certain agonists) have been described, and splice variants lacking critical transmembrane or cytoplasmic regions exhibit defective trafficking and even dominant–negative effects. Together, these data highlight that proper receptor localization and conformational integrity are essential for effective CALCR signaling."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "8", "end_ref": "10"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nA wealth of genetic association studies has implicated CALCR polymorphisms in several calcium–regulated and skeletal processes. Variants in the CALCR gene have been correlated with differences in bone mineral density, peak bone mass, osteoporotic fracture risk, and altered osteoclast differentiation in both juvenile and postmenopausal populations. In parallel, associations with conditions such as kidney stone formation, stress fractures, and even altered circulating calcium–regulating hormone levels have been reported across diverse cohorts—underscoring a central role for CALCR genetic variability in bone remodeling and calcium homeostasis."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "11", "end_ref": "29"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its roles in calcium and bone biology, CALCR is emerging as an important player in cancer and immune regulation. Overexpression of CALCR has been observed in aggressive tumors such as prostate and non–small cell lung cancers, where its activation is associated with enhanced cell invasion, disruption of tight junction integrity, and poorer clinical outcomes. Furthermore, CALCR is expressed in lymphocytes, where its engagement elevates intracellular calcium and promotes cell proliferation—suggesting that CALCR signaling may influence both tumor biology and immune cell function."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "30", "end_ref": "34"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nInnovative ligand–engineering approaches have recently been devised whereby fusions of receptor activity–modifying proteins to calcitonin–related peptides generate membrane–anchored ligands that selectively activate CALCR. This autocrine–like mode of signaling holds promise for the development of targeted therapies with minimized off–target effects."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "35"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nFinally, clinical studies have refined the measurement of serum calcitonin and evaluated CALCR genetic markers as potential diagnostic tools in conditions ranging from periodontitis to endemic fluorosis, further illustrating the receptor’s broad diagnostic and therapeutic relevance."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "36"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Kenji Kuwasako, Kazuo Kitamura, Yasuko Nagoshi, et al. 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