{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\n ACKR3 (also known as CXCR7 or RDC1) is an atypical chemokine receptor that binds with high affinity to CXCL12 and CXCL11. Unlike classical G‐protein–coupled receptors that mediate rapid Ca²⁺ mobilization and chemotaxis, ACKR3 often acts as a “scavenger” receptor capable of internalizing and degrading its ligands, thereby modulating local chemokine gradients and indirectly regulating CXCR4‐dependent responses."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "5"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n In developmental contexts, ACKR3 plays critical roles in cardiac, neuronal, and vascular formation. It is expressed on a variety of cells—including activated endothelial cells, fetal liver cells, and migrating cortical interneurons—and contributes to processes such as interneuron migration and laminar positioning in the brain."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "6", "end_ref": "8"}]}, {"type": "t", "text": " Moreover, ACKR3 contributes to cardiovascular development, as evidenced by defects in CXCR7‐deficient mice that include myocardial degeneration and fibrosis."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "9"}]}, {"type": "t", "text": "\n "}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n In the context of cancer, ACKR3 is emerging as a key regulator of tumor progression. It is expressed by various tumor cell types—including breast, lung, astroglial, hepatocellular, and esophageal cancers—and promotes tumor cell survival, adhesion, and growth. In many models, ACKR3 enhances angiogenesis and is associated with resistance to drug‐induced apoptosis, increased metastasis, and poorer clinical outcomes. In addition, when co‐expressed with CXCR4, ACKR3 modulates the balance of intracellular signaling by preferentially recruiting β‐arrestin and attenuating canonical G₍i₎–mediated pathways, thereby affecting directional migration and chemotactic responses."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "10", "end_ref": "17"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n ACKR3 also influences the tumor microenvironment by regulating transendothelial migration of cancer cells, modulating integrin rearrangement, and affecting the recruitment of pro‐tumorigenic immune cells. Its expression is upregulated by inflammatory stimuli and hypoxia and can be induced downstream of cytokine signaling (for example via IL‑6 in esophageal cancer), linking the receptor to chemoresistance and aggressive disease phenotypes."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "18", "end_ref": "23"}]}, {"type": "t", "text": "\n "}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n Interestingly, despite its inability to trigger classical chemotactic signaling, ACKR3 actively engages intracellular pathways by recruiting adapter proteins such as β‐arrestin and, in some cell types (e.g., astrocytes), by activating pertussis toxin–sensitive G(i/o) proteins. This ligand‑biased signaling demonstrates that ACKR3 not only modulates chemokine availability but also transduces distinct intracellular signals that can influence processes ranging from cell migration to survival."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "24"}]}, {"type": "t", "text": "\n "}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n Beyond its roles in development and cancer, ACKR3 functions are also implicated in regulating immune cell behavior. For instance, in rheumatoid arthritis, altered promoter methylation of CXCL12 combined with activation via CXCR7 on synovial fibroblasts contributes to enhanced expression of matrix metalloproteinases and joint destruction."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "25"}]}, {"type": "t", "text": " In addition, in breast cancer, differential regulation of ACKR3 (versus CXCR4) by estrogens affects cell growth and tumorigenesis, and its expression correlates with markers of tumor aggressiveness and poor prognosis."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "17"}, {"type": "fg_f", "ref": "15"}]}, {"type": "t", "text": "\n "}]}, {"type": "t", "text": "\n \n "}, {"type": "p", "children": [{"type": "t", "text": "\n In summary, ACKR3 is a multifunctional receptor that integrates chemokine scavenging with active, often β‑arrestin– and G(i/o)–mediated signaling pathways. Through these mechanisms, it plays diverse roles in development, modulation of immune responses, and the progression of cancers, making it an attractive therapeutic target in a range of pathological conditions."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}, {"type": "fg_f", "ref": "10"}, {"type": "fg_f", "ref": "3"}, {"type": "fg_fs", "start_ref": "11", "end_ref": "13"}, {"type": "fg_f", "ref": "6"}, {"type": "fg_f", "ref": "18"}, {"type": "fg_f", "ref": "4"}, {"type": "fg_f", "ref": "7"}, {"type": "fg_f", "ref": "20"}, {"type": "fg_f", "ref": "9"}, {"type": "fg_f", "ref": "14"}, {"type": "fg_f", "ref": "5"}, {"type": "fg_f", "ref": "24"}, {"type": "fg_f", "ref": "8"}, {"type": "fg_f", "ref": "25"}, {"type": "fg_fs", "start_ref": "15", "end_ref": "17"}, {"type": "fg_f", "ref": "21"}, {"type": "fg_f", "ref": "26"}, {"type": "fg_f", "ref": "22"}]}, {"type": "t", "text": "\n "}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Jennifer M Burns, Bretton C Summers, Yu Wang, et al. 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