Gene Symbol: Boc
Description: biregional cell adhesion molecule-related/down-regulated by oncogenes (Cdon) binding protein
Alias: 4732455C11, mFLJ00376, brother of CDO, biregional Cdon binding protein
Species: mouse
Products:     Boc

Top Publications

  1. Kang J, Mulieri P, Hu Y, Taliana L, Krauss R. BOC, an Ig superfamily member, associates with CDO to positively regulate myogenic differentiation. EMBO J. 2002;21:114-24 pubmed
    ..Reported here is the identification of BOC, which, with CDO, defines a newly recognized subfamily within the immunoglobulin superfamily...
  2. Okada A, Charron F, Morin S, Shin D, Wong K, Fabre P, et al. Boc is a receptor for sonic hedgehog in the guidance of commissural axons. Nature. 2006;444:369-73 pubmed
    ..Here we show that two Robo-related proteins, Boc and Cdon, bind specifically to Shh and are therefore candidate receptors for the action of Shh as an axon guidance ..
  3. Mulieri P, Kang J, Sassoon D, Krauss R. Expression of the boc gene during murine embryogenesis. Dev Dyn. 2002;223:379-88 pubmed
    b>BOC is a receptor-like protein that, with the related factor CDO, belongs to a newly recognized subfamily within the immunoglobulin superfamily of cell-surface molecules...
  4. Sánchez Arrones L, Nieto Lopez F, Sánchez Camacho C, Carreres M, Herrera E, Okada A, et al. Shh/Boc signaling is required for sustained generation of ipsilateral projecting ganglion cells in the mouse retina. J Neurosci. 2013;33:8596-607 pubmed publisher cRGCs, iRGCs, which are generated in the ventrotemporal crescent (VTC) of the retina, specifically express Boc, a cell adhesion molecule that acts as a high-affinity receptor for Shh...
  5. Allen B, Song J, Izzi L, Althaus I, Kang J, Charron F, et al. Overlapping roles and collective requirement for the coreceptors GAS1, CDO, and BOC in SHH pathway function. Dev Cell. 2011;20:775-87 pubmed publisher
    ..However, recent studies implicate three additional HH-binding, cell-surface proteins, GAS1, CDO, and BOC, as putative coreceptors for HH ligands...
  6. Zhang W, Hong M, Bae G, Kang J, Krauss R. Boc modifies the holoprosencephaly spectrum of Cdo mutant mice. Dis Model Mech. 2011;4:368-80 pubmed publisher
    ..We report here that whereas mice lacking the Cdo paralog Boc do not have HPE, Cdo;Boc double mutants on a largely Cdo-resistant genetic background have lobar HPE with strong ..
  7. Tenzen T, Allen B, Cole F, Kang J, Krauss R, McMahon A. The cell surface membrane proteins Cdo and Boc are components and targets of the Hedgehog signaling pathway and feedback network in mice. Dev Cell. 2006;10:647-56 pubmed
    Cdo and Boc encode cell surface Ig/fibronectin superfamily members linked to muscle differentiation. Data here indicate they are also targets and signaling components of the Sonic hedgehog (Shh) pathway...
  8. Kang J, Feinleib J, Knox S, Ketteringham M, Krauss R. Promyogenic members of the Ig and cadherin families associate to positively regulate differentiation. Proc Natl Acad Sci U S A. 2003;100:3989-94 pubmed
    ..We show here that CDO and BOC, two promyogenic Ig superfamily members that bind to each other in a cis fashion, form complexes with N- and M-..
  9. Kang J, Yi M, Zhang W, Feinleib J, Cole F, Krauss R. Netrins and neogenin promote myotube formation. J Cell Biol. 2004;167:493-504 pubmed
    ..One exception is a cell surface complex containing promyogenic Ig superfamily members (CDO and BOC) and cadherins...

More Information


  1. Zhang W, Kang J, Cole F, Yi M, Krauss R. Cdo functions at multiple points in the Sonic Hedgehog pathway, and Cdo-deficient mice accurately model human holoprosencephaly. Dev Cell. 2006;10:657-65 pubmed
    ..Specific Cdo domains required for its promyogenic effect are dispensable for its Shh signaling role, suggesting that Cdo has multiple, independent functions. ..
  2. Izzi L, Levesque M, Morin S, Laniel D, Wilkes B, Mille F, et al. Boc and Gas1 each form distinct Shh receptor complexes with Ptch1 and are required for Shh-mediated cell proliferation. Dev Cell. 2011;20:788-801 pubmed publisher
    ..Here we show that, unexpectedly, cerebellar granule neuron progenitors (CGNPs) lacking Boc and Cdon, the vertebrate orthologs of Ihog and Boi, still proliferate in response to Hh...
  3. Shin J, Ankamreddy H, Jakka N, Lee S, Kim U, Bok J. Temporal and spatial expression patterns of Hedgehog receptors in the developing inner and middle ear. Int J Dev Biol. 2017;61:557-563 pubmed publisher
    ..arrest-specific 1 (Gas1), cell-adhesion molecules-related/down-regulated by oncogenes (Cdon), and biregional Cdon binding protein (Boc). However, little is known about the role of these Hh co-receptors in inner ear development...
  4. Peng J, Fabre P, Dolique T, Swikert S, Kermasson L, Shimogori T, et al. Sonic Hedgehog Is a Remotely Produced Cue that Controls Axon Guidance Trans-axonally at a Midline Choice Point. Neuron. 2018;97:326-340.e4 pubmed publisher
    ..In vitro, contralateral RGC axons, which secrete Shh, repel ipsilateral RGCs in a Boc- and Smo-dependent manner...
  5. Xavier G, Seppala M, Papageorgiou S, Fan C, Cobourne M. Genetic interactions between the hedgehog co-receptors Gas1 and Boc regulate cell proliferation during murine palatogenesis. Oncotarget. 2016;7:79233-79246 pubmed publisher
    ..Gas1 and Boc are membrane co-receptors for Shh, which demonstrate overlapping domains of expression in the early face...
  6. Yung A, Nishitani A, Goodrich L. Phenotypic analysis of mice completely lacking netrin 1. Development. 2015;142:3686-91 pubmed publisher
  7. Lee H, Jo S, Romer A, Lim H, Kim M, Koo S, et al. Overweight in mice and enhanced adipogenesis in vitro are associated with lack of the hedgehog coreceptor boc. Diabetes. 2015;64:2092-103 pubmed publisher
    ..We report that mice lacking the HH coreceptor BOC displayed age-related overweight and excess WAT...
  8. Camp D, Haitian He B, Li S, Althaus I, Holtz A, Allen B, et al. Ihog and Boi elicit Hh signaling via Ptc but do not aid Ptc in sequestering the Hh ligand. Development. 2014;141:3879-88 pubmed publisher
  9. Kavran J, Ward M, Oladosu O, Mulepati S, Leahy D. All mammalian Hedgehog proteins interact with cell adhesion molecule, down-regulated by oncogenes (CDO) and brother of CDO (BOC) in a conserved manner. J Biol Chem. 2010;285:24584-90 pubmed publisher
    ..Cell adhesion molecule-related, down-regulated by oncogenes (CDO) and Brother of CDO (BOC) are the closest mammalian relatives of Drosophila Ihog, and CDO binds Sonic Hh with micromolar affinity ..
  10. Fabre P, Shimogori T, Charron F. Segregation of ipsilateral retinal ganglion cell axons at the optic chiasm requires the Shh receptor Boc. J Neurosci. 2010;30:266-75 pubmed publisher
    ..Here we show that the Sonic Hedgehog (Shh) receptor Boc is enriched in ipsilateral RGCs of the developing retina...
  11. Seppala M, Xavier G, Fan C, Cobourne M. Boc modifies the spectrum of holoprosencephaly in the absence of Gas1 function. Biol Open. 2014;3:728-40 pubmed publisher
    ..Here, we have elucidated the contribution of Gas1 and an additional hedgehog co-receptor, Boc during early development of the craniofacial midline, by generating single and compound mutant mice...
  12. Vuong T, Leem Y, Kim B, Cho H, Lee S, Bae G, et al. A Sonic hedgehog coreceptor, BOC regulates neuronal differentiation and neurite outgrowth via interaction with ABL and JNK activation. Cell Signal. 2017;30:30-40 pubmed publisher
    ..An immunoglobulin superfamily member, BOC functions as Sonic hedgehog (Shh) coreceptor in canonical and noncanonical Shh signaling in neuronal development ..
  13. Wu Y, Kim J, Zhou S, Smas C. Differential screening identifies transcripts with depot-dependent expression in white adipose tissues. BMC Genomics. 2008;9:397 pubmed publisher
    ..5-fold or greater enrichment in EP vs. SC adipocytes. Boc, a component of the hedgehog signaling pathway demonstrated highest enrichment (approximately 12-fold) in EP ..
  14. Kang J, Bae G, Yi M, Yang Y, Oh J, Takaesu G, et al. A Cdo-Bnip-2-Cdc42 signaling pathway regulates p38alpha/beta MAPK activity and myogenic differentiation. J Cell Biol. 2008;182:497-507 pubmed publisher
    ..Furthermore, interaction with multiple scaffold-type proteins is a distinctive mode of cell surface receptor signaling and provides one mechanism for specificity of p38alpha/beta activation during cell differentiation. ..
  15. Harwell C, Parker P, Gee S, Okada A, McConnell S, Kreitzer A, et al. Sonic hedgehog expression in corticofugal projection neurons directs cortical microcircuit formation. Neuron. 2012;73:1116-26 pubmed publisher
    ..Shh is expressed in layer V corticofugal projection neurons and the Shh receptor, Brother of CDO (Boc), is expressed in local and callosal projection neurons of layer II/III that synapse onto the subcortical ..
  16. Mille F, Tamayo Orrego L, Lévesque M, Remke M, Korshunov A, Cardin J, et al. The Shh receptor Boc promotes progression of early medulloblastoma to advanced tumors. Dev Cell. 2014;31:34-47 pubmed publisher
    ..Although the Shh-binding protein Boc associates with the Shh receptor Ptch1 to mediate Shh signaling, whether Boc plays a role in medulloblastoma is ..
  17. Jin S, Martinelli D, Zheng X, Tessier Lavigne M, Fan C. Gas1 is a receptor for sonic hedgehog to repel enteric axons. Proc Natl Acad Sci U S A. 2015;112:E73-80 pubmed publisher
    ..Together, our data uncover a previously unsuspected mechanism underlying development of centrifugal tubular organization and identify a previously unidentified effector of Shh in axon guidance. ..
  18. Chambers D, Wilson L, Alfonsi F, Hunter E, Saxena U, Blanc E, et al. Rhombomere-specific analysis reveals the repertoire of genetic cues expressed across the developing hindbrain. Neural Dev. 2009;4:6 pubmed publisher
    ..These findings give the first genome-wide insight into the complexity of gene expression during patterning of the developing hindbrain. ..
  19. Sánchez Camacho C, Bovolenta P. Autonomous and non-autonomous Shh signalling mediate the in vivo growth and guidance of mouse retinal ganglion cell axons. Development. 2008;135:3531-41 pubmed publisher
  20. Tucker E, Segall S, Gopalakrishna D, Wu Y, Vernon M, Polleux F, et al. Molecular specification and patterning of progenitor cells in the lateral and medial ganglionic eminences. J Neurosci. 2008;28:9504-18 pubmed publisher
    ..restricted in either the LGE [biregional cell adhesion molecule-related/downregulated by oncogenes binding protein (Boc), Frizzled homolog 8 (Fzd8), Ankrd43 (ankyrin repeat domain-containing protein 43), and Ikzf1 (Ikaros family zinc ..
  21. Ikeda Y, Zabbarova I, Schaefer C, Bushnell D, de Groat W, Kanai A, et al. Fgfr2 is integral for bladder mesenchyme patterning and function. Am J Physiol Renal Physiol. 2017;312:F607-F618 pubmed publisher
    ..Moreover, E16.5Fgfr2BM-/- bladders exhibit higher levels of Cdo and Boc, hedgehog coreceptors that enhance sensitivity to Shh, than controls...