{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nMacrophage receptor with collagenous structure (MARCO) is a class A scavenger receptor prominently expressed by macrophages that is endowed with a specialized extracellular architecture. Its short cytoplasmic and transmembrane regions are juxtaposed to a large extracellular portion comprising a spacer, extended collagenous domain, and a C‐terminal scavenger receptor cysteine‐rich (SRCR) domain. Functional studies have demonstrated that this SRCR domain, particularly an arginine‐rich RXR motif, is critical for high‐affinity binding to a broad range of unopsonized ligands such as environmental particles, bacteria, lipopolysaccharide, and even endogenous proteins like the uteroglobin-related protein UGRP1 in the lung. Structural analysis of soluble MARCO has revealed it adopts a triple‐helical collagenous conformation that facilitates multimerization and ligand clustering. Moreover, evolutionary analyses have identified human‐specific residues under positive selection that augment its binding and signaling capacity."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "5"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond structural attributes, MARCO plays a central role in innate immune defense through the recognition, binding, and phagocytic clearance of diverse pathogens. In human alveolar and decidual macrophages, MARCO mediates the uptake of unopsonized bacteria—including Escherichia coli, Staphylococcus aureus, and pathogens such as Clostridium sordellii—and contributes to host resistance during microbial challenge. Its engagement can also trigger downstream pro‐inflammatory signaling cascades that modulate cytokine production and antimicrobial peptide expression, as observed during central nervous system infections. Moreover, MARCO influences postinfluenza recovery by enhancing phagocytosis and bacterial clearance, participates in the uptake of fungal pathogens like Cryptococcus neoformans, and facilitates endocytosis of nano‐sized exosomes. Together, these findings underscore MARCO’s multifaceted role as a pattern recognition receptor that orchestrates both uptake and immune activation. In some contexts, however, pathogens such as herpes simplex virus type 1 may hijack MARCO to promote adhesion and infection, highlighting a dualistic role in host–pathogen interactions."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "6", "end_ref": "16"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nMARCO’s influence extends into the tumor microenvironment and genetically determined host responses. In non‐small cell lung cancer (NSCLC) and other neoplastic settings, MARCO expression by tumor‐associated macrophages correlates with immunosuppressive phenotypes—promoting regulatory T‐cell expansion, impairing cytotoxic lymphocyte functions, and thereby contributing to tumor progression. In addition, studies in hepatocellular carcinoma and gliomas suggest that altered MARCO expression may modulate antitumor immunity as well as tumor cell uptake by macrophages. A growing body of genetic association studies has linked single‐nucleotide polymorphisms in MARCO with susceptibility to infections such as tuberculosis, respiratory syncytial virus, and chronic obstructive pulmonary disease, as well as with cardiovascular and neoplastic disorders like carotid atherosclerosis and hepatocellular carcinoma. In the context of cholera vaccination, MARCO’s ability to bind lipopolysaccharide underpins host immune responses to microbial antigens. Collectively, these genetic and functional studies emphasize MARCO’s dual role as an essential mediator of pathogen clearance and as a modulatory factor in inflammatory and neoplastic disease processes."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "17", "end_ref": "30"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Mohamed S Arredouani, Aiyappa Palecanda, Henry Koziel, et al. 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