BOC Gene

HGNC Family	Fibronectin type III domain containing, Immunoglobulin superfamily domain containing
Name	BOC cell adhesion associated, oncogene regulated
Description	The protein encoded by this gene is a member of the immunoglobulin/fibronectin type III repeat family. It is a component of a cell-surface receptor complex that mediates cell-cell interactions between muscle precursor cells, and promotes myogenic differentiation. Alternative splicing results in multiple transcript variants encoding different isoforms. [provided by RefSeq, Sep 2014]
Summary	{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\n BOC is a transmembrane cell–surface receptor that functions as an essential coreceptor for Hedgehog (HH) ligands in multiple developmental contexts. It acts in concert with CDO and GAS1 to facilitate HH signal reception by localizing within specialized membrane protrusions—such as novel, actin‐based filopodia—that enable long‐range, contact‐mediated ligand transport (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}]}, {"type": "t", "text": "]). In the mammalian neural tube, BOC shows overlapping and indispensable functions with other coreceptors for the specification of neural progenitor fates and for the maintenance of motor neuron progenitors (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "2"}]}, {"type": "t", "text": "]). Beyond early neural patterning, BOC is differentially expressed in distinct neuronal populations; for instance, in the developing cerebral cortex it localizes to presynaptic neurons where its loss reduces the strength of specific synaptic connections, thereby guiding the formation of cortical microcircuits (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "3"}]}, {"type": "t", "text": "]). Biochemical and structural studies further reveal that BOC, similar to CDO, binds HH proteins in a conserved manner to positively regulate HH signaling (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "4"}]}, {"type": "t", "text": "]). In the developing retina, selective expression of BOC in ipsilaterally projecting retinal ganglion cells is required to sustain the expression of transcription factors that counterbalance Shh signaling from nearby contralateral cells, ensuring correct cell fate specification (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "5"}]}, {"type": "t", "text": "]). Moreover, disruptions in the interaction between HH ligands and BOC—such as those caused by mutations in desert hedgehog that alter BOC binding—have been associated with disorders of sex development (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "6"}]}, {"type": "t", "text": "]). BOC expression is also notably enriched in articular cartilage, suggesting a role in modulating HH signaling in differentiated tissues (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "7"}]}, {"type": "t", "text": "]). Finally, genetic studies in mice demonstrate that while loss of BOC alone may not produce overt midline defects, the combined loss of BOC with GAS1 leads to severe craniofacial malformations—including features of holoprosencephaly and disrupted tooth development—thereby underscoring its critical, and sometimes redundant, function in mediating HH signal transduction in craniofacial and neural tissues (["}, {"type": "fg", "children": [{"type": "fg_f", "ref": "8"}]}, {"type": "t", "text": "]).\n "}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Timothy A Sanders, Esther Llagostera, Maria Barna "}, {"type": "b", "children": [{"type": "t", "text": "Specialized filopodia direct long-range transport of SHH during vertebrate tissue patterning."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Nature (2013)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1038/nature12157"}], "href": "https://doi.org/10.1038/nature12157"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "23624372"}], "href": "https://pubmed.ncbi.nlm.nih.gov/23624372"}]}, {"type": "r", "ref": 2, "children": [{"type": "t", "text": "Benjamin L Allen, Jane Y Song, Luisa Izzi, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Overlapping roles and collective requirement for the coreceptors GAS1, CDO, and BOC in SHH pathway function."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Dev Cell (2011)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.devcel.2011.04.018"}], "href": "https://doi.org/10.1016/j.devcel.2011.04.018"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21664576"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21664576"}]}, {"type": "r", "ref": 3, "children": [{"type": "t", "text": "Corey C Harwell, Philip R L Parker, Steven M Gee, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Sonic hedgehog expression in corticofugal projection neurons directs cortical microcircuit formation."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Neuron (2012)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.neuron.2012.02.009"}], "href": "https://doi.org/10.1016/j.neuron.2012.02.009"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "22445340"}], "href": "https://pubmed.ncbi.nlm.nih.gov/22445340"}]}, {"type": "r", "ref": 4, "children": [{"type": "t", "text": "Jennifer M Kavran, Matthew D Ward, Oyindamola O Oladosu, et al. "}, {"type": "b", "children": [{"type": "t", "text": "All mammalian Hedgehog proteins interact with cell adhesion molecule, down-regulated by oncogenes (CDO) and brother of CDO (BOC) in a conserved manner."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "J Biol Chem (2010)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1074/jbc.M110.131680"}], "href": "https://doi.org/10.1074/jbc.M110.131680"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "20519495"}], "href": "https://pubmed.ncbi.nlm.nih.gov/20519495"}]}, {"type": "r", "ref": 5, "children": [{"type": "t", "text": "Luisa Sánchez-Arrones, Francisco Nieto-Lopez, Cristina Sánchez-Camacho, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Shh/Boc signaling is required for sustained generation of ipsilateral projecting ganglion cells in the mouse retina."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "J Neurosci (2013)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1523/JNEUROSCI.2083-12.2013"}], "href": "https://doi.org/10.1523/JNEUROSCI.2083-12.2013"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "23678105"}], "href": "https://pubmed.ncbi.nlm.nih.gov/23678105"}]}, {"type": "r", "ref": 6, "children": [{"type": "t", "text": "Francoise Paris, Delphine Flatters, Sandrine Caburet, et al. "}, {"type": "b", "children": [{"type": "t", "text": "A novel variant of DHH in a familial case of 46,XY disorder of sex development: Insights from molecular dynamics simulations."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Clin Endocrinol (Oxf) (2017)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1111/cen.13420"}], "href": "https://doi.org/10.1111/cen.13420"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "28708305"}], "href": "https://pubmed.ncbi.nlm.nih.gov/28708305"}]}, {"type": "r", "ref": 7, "children": [{"type": "t", "text": "K S Vanderman, M Tremblay, W Zhu, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Brother of CDO (BOC) expression in equine articular cartilage."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Osteoarthritis Cartilage (2011)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1016/j.joca.2011.01.011"}], "href": "https://doi.org/10.1016/j.joca.2011.01.011"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "21262369"}], "href": "https://pubmed.ncbi.nlm.nih.gov/21262369"}]}, {"type": "r", "ref": 8, "children": [{"type": "t", "text": "Maisa Seppala, Guilherme M Xavier, Chen-Ming Fan, et al. "}, {"type": "b", "children": [{"type": "t", "text": "Boc modifies the spectrum of holoprosencephaly in the absence of Gas1 function."}]}, {"type": "t", "text": " "}, {"type": "i", "children": [{"type": "t", "text": "Biol Open (2014)"}]}, {"type": "t", "text": " DOI: "}, {"type": "a", "children": [{"type": "t", "text": "10.1242/bio.20147989"}], "href": "https://doi.org/10.1242/bio.20147989"}, {"type": "t", "text": " PMID: "}, {"type": "a", "children": [{"type": "t", "text": "25063195"}], "href": "https://pubmed.ncbi.nlm.nih.gov/25063195"}]}]}]}
Synonyms	CDON2
Proteins	BOC_HUMAN
NCBI Gene ID	91653
API
Download Associations
Predicted Functions
Co-expressed Genes
Expression in Tissues and Cell Lines

Functional Associations

BOC has 4,791 functional associations with biological entities spanning 9 categories (molecular profile, organism, functional term, phrase or reference, disease, phenotype or trait, chemical, structural feature, cell line, cell type or tissue, gene, protein or microRNA, sequence feature) extracted from 109 datasets.

Click the + buttons to view associations for BOC from the datasets below.

If available, associations are ranked by standardized value

Harmonizome 3.0

BOC Gene

Functional Associations

Please acknowledge the Harmonizome in your publications by citing the following reference(s):

	Dataset	Summary
	Allen Brain Atlas Adult Human Brain Tissue Gene Expression Profiles	tissues with high or low expression of BOC gene relative to other tissues from the Allen Brain Atlas Adult Human Brain Tissue Gene Expression Profiles dataset.
	Allen Brain Atlas Adult Mouse Brain Tissue Gene Expression Profiles	tissues with high or low expression of BOC gene relative to other tissues from the Allen Brain Atlas Adult Mouse Brain Tissue Gene Expression Profiles dataset.
	Allen Brain Atlas Aging Dementia and Traumatic Brain Injury Tissue Sample Gene Expression Profiles	tissue samples with high or low expression of BOC gene relative to other tissue samples from the Allen Brain Atlas Aging Dementia and Traumatic Brain Injury Tissue Sample Gene Expression Profiles dataset.
	Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by Microarray	tissue samples with high or low expression of BOC gene relative to other tissue samples from the Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by Microarray dataset.
	Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by RNA-seq	tissue samples with high or low expression of BOC gene relative to other tissue samples from the Allen Brain Atlas Developing Human Brain Tissue Gene Expression Profiles by RNA-seq dataset.
	Allen Brain Atlas Prenatal Human Brain Tissue Gene Expression Profiles	tissues with high or low expression of BOC gene relative to other tissues from the Allen Brain Atlas Prenatal Human Brain Tissue Gene Expression Profiles dataset.
	BioGPS Human Cell Type and Tissue Gene Expression Profiles	cell types and tissues with high or low expression of BOC gene relative to other cell types and tissues from the BioGPS Human Cell Type and Tissue Gene Expression Profiles dataset.
	BioGPS Mouse Cell Type and Tissue Gene Expression Profiles	cell types and tissues with high or low expression of BOC gene relative to other cell types and tissues from the BioGPS Mouse Cell Type and Tissue Gene Expression Profiles dataset.
	CCLE Cell Line Gene CNV Profiles	cell lines with high or low copy number of BOC gene relative to other cell lines from the CCLE Cell Line Gene CNV Profiles dataset.
	CCLE Cell Line Gene Expression Profiles	cell lines with high or low expression of BOC gene relative to other cell lines from the CCLE Cell Line Gene Expression Profiles dataset.
	CellMarker Gene-Cell Type Associations	cell types associated with BOC gene from the CellMarker Gene-Cell Type Associations dataset.
	ChEA Transcription Factor Binding Site Profiles	transcription factor binding site profiles with transcription factor binding evidence at the promoter of BOC gene from the CHEA Transcription Factor Binding Site Profiles dataset.
	ChEA Transcription Factor Targets	transcription factors binding the promoter of BOC gene in low- or high-throughput transcription factor functional studies from the CHEA Transcription Factor Targets dataset.
	ChEA Transcription Factor Targets 2022	transcription factors binding the promoter of BOC gene in low- or high-throughput transcription factor functional studies from the CHEA Transcription Factor Targets 2022 dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores	cellular components containing BOC protein from the COMPARTMENTS Curated Protein Localization Evidence Scores dataset.
	COMPARTMENTS Curated Protein Localization Evidence Scores 2025	cellular components containing BOC protein from the COMPARTMENTS Curated Protein Localization Evidence Scores 2025 dataset.
	COMPARTMENTS Experimental Protein Localization Evidence Scores	cellular components containing BOC protein in low- or high-throughput protein localization assays from the COMPARTMENTS Experimental Protein Localization Evidence Scores dataset.
	COMPARTMENTS Experimental Protein Localization Evidence Scores 2025	cellular components containing BOC protein in low- or high-throughput protein localization assays from the COMPARTMENTS Experimental Protein Localization Evidence Scores 2025 dataset.
	COMPARTMENTS Text-mining Protein Localization Evidence Scores	cellular components co-occuring with BOC protein in abstracts of biomedical publications from the COMPARTMENTS Text-mining Protein Localization Evidence Scores dataset.
	COMPARTMENTS Text-mining Protein Localization Evidence Scores 2025	cellular components co-occuring with BOC protein in abstracts of biomedical publications from the COMPARTMENTS Text-mining Protein Localization Evidence Scores 2025 dataset.
	COSMIC Cell Line Gene CNV Profiles	cell lines with high or low copy number of BOC gene relative to other cell lines from the COSMIC Cell Line Gene CNV Profiles dataset.
	COSMIC Cell Line Gene Mutation Profiles	cell lines with BOC gene mutations from the COSMIC Cell Line Gene Mutation Profiles dataset.
	CTD Gene-Disease Associations	diseases associated with BOC gene/protein from the curated CTD Gene-Disease Associations dataset.
	DepMap CRISPR Gene Dependency	cell lines with fitness changed by BOC gene knockdown relative to other cell lines from the DepMap CRISPR Gene Dependency dataset.
	DISEASES Experimental Gene-Disease Association Evidence Scores	diseases associated with BOC gene in GWAS datasets from the DISEASES Experimental Gene-Disease Assocation Evidence Scores dataset.
	DISEASES Experimental Gene-Disease Association Evidence Scores 2025	diseases associated with BOC gene in GWAS datasets from the DISEASES Experimental Gene-Disease Assocation Evidence Scores 2025 dataset.
	DISEASES Text-mining Gene-Disease Association Evidence Scores	diseases co-occuring with BOC gene in abstracts of biomedical publications from the DISEASES Text-mining Gene-Disease Assocation Evidence Scores dataset.
	DISEASES Text-mining Gene-Disease Association Evidence Scores 2025	diseases co-occuring with BOC gene in abstracts of biomedical publications from the DISEASES Text-mining Gene-Disease Assocation Evidence Scores 2025 dataset.
	DisGeNET Gene-Disease Associations	diseases associated with BOC gene in GWAS and other genetic association datasets from the DisGeNET Gene-Disease Associations dataset.
	DisGeNET Gene-Phenotype Associations	phenotypes associated with BOC gene in GWAS and other genetic association datasets from the DisGeNET Gene-Phenoptype Associations dataset.
	ENCODE Histone Modification Site Profiles	histone modification site profiles with high histone modification abundance at BOC gene from the ENCODE Histone Modification Site Profiles dataset.
	ENCODE Transcription Factor Binding Site Profiles	transcription factor binding site profiles with transcription factor binding evidence at the promoter of BOC gene from the ENCODE Transcription Factor Binding Site Profiles dataset.
	ENCODE Transcription Factor Targets	transcription factors binding the promoter of BOC gene in ChIP-seq datasets from the ENCODE Transcription Factor Targets dataset.
	ESCAPE Omics Signatures of Genes and Proteins for Stem Cells	PubMedIDs of publications reporting gene signatures containing BOC from the ESCAPE Omics Signatures of Genes and Proteins for Stem Cells dataset.
	GeneRIF Biological Term Annotations	biological terms co-occuring with BOC gene in literature-supported statements describing functions of genes from the GeneRIF Biological Term Annotations dataset.
	GeneSigDB Published Gene Signatures	PubMedIDs of publications reporting gene signatures containing BOC from the GeneSigDB Published Gene Signatures dataset.
	GEO Signatures of Differentially Expressed Genes for Diseases	disease perturbations changing expression of BOC gene from the GEO Signatures of Differentially Expressed Genes for Diseases dataset.
	GEO Signatures of Differentially Expressed Genes for Gene Perturbations	gene perturbations changing expression of BOC gene from the GEO Signatures of Differentially Expressed Genes for Gene Perturbations dataset.
	GEO Signatures of Differentially Expressed Genes for Kinase Perturbations	kinase perturbations changing expression of BOC gene from the GEO Signatures of Differentially Expressed Genes for Kinase Perturbations dataset.
	GEO Signatures of Differentially Expressed Genes for Small Molecules	small molecule perturbations changing expression of BOC gene from the GEO Signatures of Differentially Expressed Genes for Small Molecules dataset.