Gene Symbol: cav
Description: caravaggio
Alias: 1G5, Anon1G5, CG6219, Caravaggio, Dmel\CG6219, HOAP, Hoap, anon fe 1G5, anon-EST:fe1G5, anon1G5, hoap, p55, caravaggio, CG6219-PA, CG6219-PB, CG6219-PD, CG6219-PE, HP1 and Orc-associated protein, HP1-ORC associated protein, HP1/ORC-associated protein, anon-fast-evolving-1G5, anonymous fast evolving gene IG5, cav-PA, cav-PB, cav-PD, cav-PE, drosophila anonymous fast evolving 1G5
Species: fruit fly

Top Publications

  1. Bi X, Wei S, Rong Y. Telomere protection without a telomerase; the role of ATM and Mre11 in Drosophila telomere maintenance. Curr Biol. 2004;14:1348-53 pubmed
    ..Since Drosophila telomeres are not added by a telomerase, our findings support an additional role for both ATM and Mre11 in telomere maintenance that is independent of telomerase regulation. ..
  2. Raffa G, Ciapponi L, Cenci G, Gatti M. Terminin: a protein complex that mediates epigenetic maintenance of Drosophila telomeres. Nucleus. 2011;2:383-91 pubmed publisher
    ..Recent work has shown that Drosophila telomeres are capped by a complex, we call terminin, which includes HOAP, HipHop, Moi and Ver; these are fast-evolving proteins that prevent telomere fusion, directly interact with each ..
  3. Dubruille R, Orsi G, Delabaere L, Cortier E, Couble P, Marais G, et al. Specialization of a Drosophila capping protein essential for the protection of sperm telomeres. Curr Biol. 2010;20:2090-9 pubmed publisher
    ..somatic cells, assembly of the protecting capping complex at telomeres notably involves the recruitment of HOAP, HP1, and their recently identified partner, HipHop...
  4. Raffa G, Raimondo D, Sorino C, Cugusi S, Cenci G, Cacchione S, et al. Verrocchio, a Drosophila OB fold-containing protein, is a component of the terminin telomere-capping complex. Genes Dev. 2010;24:1596-601 pubmed publisher
    ..Drosophila telomeres are protected by terminin, a complex that includes the HOAP (Heterochromatin Protein 1/origin recognition complex-associated protein) and Moi (Modigliani) proteins and shares ..
  5. Oikemus S, Queiroz Machado J, Lai K, McGinnis N, Sunkel C, Brodsky M. Epigenetic telomere protection by Drosophila DNA damage response pathways. PLoS Genet. 2006;2:e71 pubmed
    ..atm/tefu, mre11, or rad50 disrupt telomere protection and localization of the telomere-associated proteins HP1 and HOAP, suggesting that recognition of chromosome ends contributes to telomere protection...
  6. Raffa G, Siriaco G, Cugusi S, Ciapponi L, Cenci G, Wojcik E, et al. The Drosophila modigliani (moi) gene encodes a HOAP-interacting protein required for telomere protection. Proc Natl Acad Sci U S A. 2009;106:2271-6 pubmed publisher
    Several proteins have been identified that protect Drosophila telomeres from fusion events. They include UbcD1, HP1, HOAP, the components of the Mre11-Rad50-Nbs (MRN) complex, the ATM kinase, and the putative transcription factor Woc...
  7. Li Y, Kirschmann D, Wallrath L. Does heterochromatin protein 1 always follow code?. Proc Natl Acad Sci U S A. 2002;99 Suppl 4:16462-9 pubmed
    ..Such sequences are likely to include HP1 "target genes" whose discovery will aid in our understanding of HP1 lethality in Drosophila and metastasis of breast cancer cells. ..
  8. Cenci G, Ciapponi L, Gatti M. The mechanism of telomere protection: a comparison between Drosophila and humans. Chromosoma. 2005;114:135-45 pubmed
    ..These proteins include the telomere capping factors HP1/ORC-associated protein (HOAP) and heterochromatin protein 1 (HP1), the Rad50 and Mre11 DNA repair proteins that are required for HOAP and HP1 ..
  9. Ciapponi L, Cenci G, Gatti M. The Drosophila Nbs protein functions in multiple pathways for the maintenance of genome stability. Genetics. 2006;173:1447-54 pubmed
    ..In contrast, analysis of chromosome breakage indicated that the three components of the MRN complex function in a single pathway for the repair of the DNA damage leading to chromosome aberrations. ..

More Information


  1. Badugu R, Shareef M, Kellum R. Novel Drosophila heterochromatin protein 1 (HP1)/origin recognition complex-associated protein (HOAP) repeat motif in HP1/HOAP interactions and chromocenter associations. J Biol Chem. 2003;278:34491-8 pubmed
    ..The Drosophila HOAP protein is a DNA-binding protein that was identified as a component of a multiprotein complex of HP1 containing ..
  2. Biessmann H, Mason J. Telomerase-independent mechanisms of telomere elongation. Cell Mol Life Sci. 2003;60:2325-33 pubmed
    ..We will summarize current knowledge on these telomerase-independent elongation mechanisms in yeast and mammalian cells and will discuss in more detail the telomere elongation mechanism by retrotransposons in Drosophila melanogaster. ..
  3. Bi X, Srikanta D, Fanti L, Pimpinelli S, Badugu R, Kellum R, et al. Drosophila ATM and ATR checkpoint kinases control partially redundant pathways for telomere maintenance. Proc Natl Acad Sci U S A. 2005;102:15167-72 pubmed
  4. Shareef M, King C, Damaj M, Badagu R, Huang D, Kellum R. Drosophila heterochromatin protein 1 (HP1)/origin recognition complex (ORC) protein is associated with HP1 and ORC and functions in heterochromatin-induced silencing. Mol Biol Cell. 2001;12:1671-85 pubmed publisher
    ..Moreover, the gene encoding HP1/ORC-associated protein was found to display reciprocal dose-dependent variegation modifier phenotypes, similar to those for mutants in HP1 and the ORC 2 subunit...
  5. Rong Y. Loss of the histone variant H2A.Z restores capping to checkpoint-defective telomeres in Drosophila. Genetics. 2008;180:1869-75 pubmed publisher
    ..Loss of H2A.Z partially restores the loading of the HOAP capping protein, possibly accounting for the partial restoration in capping. I propose that, in the absence of H2A...
  6. Shareef M, Badugu R, Kellum R. HP1/ORC complex and heterochromatin assembly. Genetica. 2003;117:127-34 pubmed
    ..A complex of HP1 associated with the origin recognition complex (ORC) and an HP1/ORC-associated protein (HOAP) was purified from the maternally loaded cytoplasm of early Drosophila embryo...
  7. Badugu R, Yoo Y, Singh P, Kellum R. Mutations in the heterochromatin protein 1 (HP1) hinge domain affect HP1 protein interactions and chromosomal distribution. Chromosoma. 2005;113:370-84 pubmed
    ..with the DNA-binding activities of the origin recognition complex (ORC) and an HMG-like HP1/ORC-Associated Protein (HOAP)...
  8. Rashkova S, Karam S, Kellum R, PARDUE M. Gag proteins of the two Drosophila telomeric retrotransposons are targeted to chromosome ends. J Cell Biol. 2002;159:397-402 pubmed
    ..They also imply a symbiotic relationship between the two elements, with HeT-A Gag directing the telomere-specific targeting of the elements, whereas TART provides reverse transcriptase for transposition. ..
  9. Gao G, Walser J, Beaucher M, Morciano P, Wesolowska N, Chen J, et al. HipHop interacts with HOAP and HP1 to protect Drosophila telomeres in a sequence-independent manner. EMBO J. 2010;29:819-29 pubmed publisher
    ..sequence-independent capping mechanism, we isolated proteins that interact with the HP1/ORC-associated protein (HOAP) capping protein, and identified HipHop as a subunit of the complex...
  10. Musarò M, Ciapponi L, Fasulo B, Gatti M, Cenci G. Unprotected Drosophila melanogaster telomeres activate the spindle assembly checkpoint. Nat Genet. 2008;40:362-6 pubmed publisher
    ..Previous work has shown that the Drosophila HOAP protein, encoded by the caravaggio (cav) gene, is required to prevent telomeric fusions...
  11. Cenci G, Siriaco G, Raffa G, Kellum R, Gatti M. The Drosophila HOAP protein is required for telomere capping. Nat Cell Biol. 2003;5:82-4 pubmed
    ..Here, we show that caravaggio, a mutation in the HOAP-encoding gene, causes extensive telomere-telomere fusions in larval brain cells, ..
  12. Oikemus S, McGinnis N, Queiroz Machado J, Tukachinsky H, Takada S, Sunkel C, et al. Drosophila atm/telomere fusion is required for telomeric localization of HP1 and telomere position effect. Genes Dev. 2004;18:1850-61 pubmed
    ..We propose that recognition of chromosome ends by ATM prevents telomere fusion and apoptosis by recruiting chromatin-modifying complexes to telomeres. ..
  13. Ciapponi L, Cenci G, Ducau J, Flores C, Johnson Schlitz D, Gorski M, et al. The Drosophila Mre11/Rad50 complex is required to prevent both telomeric fusion and chromosome breakage. Curr Biol. 2004;14:1360-6 pubmed
    ..roles of rad50 and mre11 in telomere protection, mutant chromosome preparations were immunostained for both HP1 and HOAP, two proteins that protect Drosophila telomeres from fusion...
  14. Raffa G, Cenci G, Siriaco G, Goldberg M, Gatti M. The putative Drosophila transcription factor woc is required to prevent telomeric fusions. Mol Cell. 2005;20:821-31 pubmed
    ..and rad50, four genes that prevent telomeric fusions; Su(var)205 and cav encode HP1 and HP1/ORC Associated Protein (HOAP), respectively...
  15. Gao G, Cheng Y, Wesolowska N, Rong Y. Paternal imprint essential for the inheritance of telomere identity in Drosophila. Proc Natl Acad Sci U S A. 2011;108:4932-7 pubmed publisher
  16. Rong Y. Telomere capping in Drosophila: dealing with chromosome ends that most resemble DNA breaks. Chromosoma. 2008;117:235-42 pubmed publisher
    ..I review recent progress in Drosophila telomere studies, and challenge the notion that Drosophila may not be a relevant model for the study of telomere maintenance. ..
  17. Messina G, Damia E, Fanti L, Atterrato M, Celauro E, Mariotti F, et al. Yeti, an essential Drosophila melanogaster gene, encodes a protein required for chromatin organization. J Cell Sci. 2014;127:2577-88 pubmed publisher
    ..V at chromatin sites. Overall, our findings suggest an unanticipated role of YETI protein in chromatin organization and provide, for the first time, mechanistic clues on how BCNT proteins control development in multicellular organisms. ..
  18. Wesolowska N, Amariei F, Rong Y. Clustering and protein dynamics of Drosophila melanogaster telomeres. Genetics. 2013;195:381-91 pubmed publisher
    ..We followed the EGFP-labeled telomeric protein HOAP in vivo and found that the 16 telomeres yield four to six foci per nucleus, indicative of clustering...
  19. Li H, Rodriguez J, Yoo Y, Shareef M, Badugu R, Horabin J, et al. Cooperative and antagonistic contributions of two heterochromatin proteins to transcriptional regulation of the Drosophila sex determination decision. PLoS Genet. 2011;7:e1002122 pubmed publisher
    ..Here we report a similar phenotype for the telomeric partner of HP1, HOAP, and roles for both proteins in regulating the Drosophila sex determination pathway...
  20. Zhang Y, Zhang L, Tang X, Bhardwaj S, Ji J, Rong Y. MTV, an ssDNA Protecting Complex Essential for Transposon-Based Telomere Maintenance in Drosophila. PLoS Genet. 2016;12:e1006435 pubmed publisher
    ..Drosophila uses a transposon-based mechanism for end protection. We showed that the HOAP-HipHop complex occupies the duplex region. Whether an ssDNA-binding complex exists is not known...
  21. Cipressa F, Morciano P, Bosso G, Mannini L, Galati A, Raffa G, et al. A role for Separase in telomere protection. Nat Commun. 2016;7:10405 pubmed publisher
    ..Finally, we find that siRNA-induced depletion of ESPL1, the Sse human orthologue, causes telomere dysfunction and HP1 level reduction in primary fibroblasts, highlighting a conserved role of Separase in telomere protection. ..
  22. Dubruille R, Marais G, Loppin B. Repeated evolution of testis-specific new genes: the case of telomere-capping genes in Drosophila. Int J Evol Biol. 2012;2012:708980 pubmed publisher
    ..Finally, we discuss the plausibility of different evolutionary scenarios that could explain the diversification of this gene family. ..
  23. PARDUE M, DeBaryshe P. Retrotransposons provide an evolutionarily robust non-telomerase mechanism to maintain telomeres. Annu Rev Genet. 2003;37:485-511 pubmed
    ..These telomeres are now known to predate the separation of extant Drosophila species, allowing ample time for elements and hosts to coevolve interesting mechanisms. ..
  24. Dubruille R, Loppin B. Protection of Drosophila chromosome ends through minimal telomere capping. J Cell Sci. 2015;128:1969-81 pubmed publisher
    ..Here, we investigate the role of the large capping domain by manipulating HOAP (encoded by caravaggio) capping-protein expression in the male germ cells, where telomere protection can be challenged without ..
  25. Schmid K, Nigro L, Aquadro C, Tautz D. Large number of replacement polymorphisms in rapidly evolving genes of Drosophila. Implications for genome-wide surveys of DNA polymorphism. Genetics. 1999;153:1717-29 pubmed
  26. Cipressa F, Romano S, Centonze S, zur Lage P, Vernì F, Dimitri P, et al. Effete, a Drosophila chromatin-associated ubiquitin-conjugating enzyme that affects telomeric and heterochromatic position effect variegation. Genetics. 2013;195:147-58 pubmed publisher
    ..Eff is enriched at several euchromatic bands and interbands, the TAS regions, and the chromocenter. Our results suggest that Eff associates with different types of chromatin affecting their abilities to regulate gene expression. ..
  27. Purdy A, Su T. Telomeres: not all breaks are equal. Curr Biol. 2004;14:R613-4 pubmed
    ..How do these proteins help chromosomal termini escape fusion through 'repair' while promoting repair of induced DNA breaks? ..
  28. Vedelek B, Blastyák A, Boros I. Cross-Species Interaction between Rapidly Evolving Telomere-Specific Drosophila Proteins. PLoS ONE. 2015;10:e0142771 pubmed publisher
    ..The five proteins supposed to form the terminin complex are HP1-ORC associated protein, HP1-HOAP interacting protein, Verrocchio, Drosophila Telomere Loss/Modigliani and Heterochromatic Protein 1...
  29. Frydrychova R, Mason J, Archer T. HP1 is distributed within distinct chromatin domains at Drosophila telomeres. Genetics. 2008;180:121-31 pubmed publisher
    ..Heterochromatin protein 1 (HP1) and HP1-ORC-associated protein (HOAP) are major protein components of the telomere cap in Drosophila and are required for telomere stability...
  30. Zaessinger S, Busseau I, Simonelig M. Oskar allows nanos mRNA translation in Drosophila embryos by preventing its deadenylation by Smaug/CCR4. Development. 2006;133:4573-83 pubmed
    ..This study provides insights into molecular mechanisms of regulated deadenylation by specific proteins and demonstrates its importance in development. ..
  31. Hediger F, Gasser S. Heterochromatin protein 1: don't judge the book by its cover!. Curr Opin Genet Dev. 2006;16:143-50 pubmed
    ..Not only do HP1 isoforms have specific binding sites in both heterochromatic and euchromatic domains but they might also participate in the repression and activation of transcription in both compartments. ..
  32. Cenci G, Ciapponi L, Marzullo M, Raffa G, Morciano P, Raimondo D, et al. The Analysis of Pendolino (peo) Mutants Reveals Differences in the Fusigenic Potential among Drosophila Telomeres. PLoS Genet. 2015;11:e1005260 pubmed publisher
    ..Fly telomeres are protected by the terminin complex (HOAP-HipHop-Moi-Ver) that localizes and functions exclusively at telomeres and by non-terminin proteins that do not ..
  33. Raffa G, Cenci G, Ciapponi L, Gatti M. Organization and Evolution of Drosophila Terminin: Similarities and Differences between Drosophila and Human Telomeres. Front Oncol. 2013;3:112 pubmed publisher
    ..has shown that Drosophila telomeres are capped by the terminin complex, which includes the fast-evolving proteins HOAP, HipHop, Moi, and Ver...
  34. Singh A, Lakhotia S. The hnRNP A1 homolog Hrb87F/Hrp36 is important for telomere maintenance in Drosophila melanogaster. Chromosoma. 2016;125:373-88 pubmed publisher
    ..mutant-derived expanded telomeres carry Hrb87F on cap and HTT arrays while Hrb87F-null telomeres have HP1 and HOAP on caps and expanded HTT arrays...
  35. Fay J, Wyckoff G, Wu C. Testing the neutral theory of molecular evolution with genomic data from Drosophila. Nature. 2002;415:1024-6 pubmed
    ..A higher A/S ratio of divergence than of polymorphism is also observed in other species, which suggests a rate of adaptive evolution that is far higher than permitted by the neutral theory of molecular evolution. ..
  36. Shaffer C, Cenci G, Thompson B, Stephens G, Slawson E, Adu Wusu K, et al. The large isoform of Drosophila melanogaster heterochromatin protein 2 plays a critical role in gene silencing and chromosome structure. Genetics. 2006;174:1189-204 pubmed
    ..These results indicate a role for HP2 in the structural organization of chromosomes and in heterochromatin-induced gene silencing and show that the larger isoform plays a critical role in these processes. ..
  37. Wallace J, Orr Weaver T. Replication of heterochromatin: insights into mechanisms of epigenetic inheritance. Chromosoma. 2005;114:389-402 pubmed
    ..Potential mechanistic links between these factors are discussed. In some cells, replication of the heterochromatin is blocked, and in Drosophila this inhibition is mediated by a chromatin binding protein SuUR. ..
  38. Temme C, Zaessinger S, Meyer S, Simonelig M, Wahle E. A complex containing the CCR4 and CAF1 proteins is involved in mRNA deadenylation in Drosophila. EMBO J. 2004;23:2862-71 pubmed
    ..The data show that the CCR4-NOT complex is conserved in Drosophila melanogaster and plays a role in general and regulated mRNA deadenylation. ..
  39. Morciano P, Zhang Y, Cenci G, Rong Y. A hypomorphic mutation reveals a stringent requirement for the ATM checkpoint protein in telomere protection during early cell division in Drosophila. G3 (Bethesda). 2013;3:1043-8 pubmed publisher
    ..The amenability of Drosophila embryos to molecular and biochemical investigations ensures that this newly identified mutation will facilitate future studies of ATM in telomere maintenance. ..
  40. Joppich C, Scholz S, Korge G, Schwendemann A. Umbrea, a chromo shadow domain protein in Drosophila melanogaster heterochromatin, interacts with Hip, HP1 and HOAP. Chromosome Res. 2009;17:19-36 pubmed publisher
    ..shown that HP1 interacts with the telomere capping factor HP1/ORC (origin recognition complex)-associated protein (HOAP)...
  41. Heun P, Erhardt S, Blower M, Weiss S, Skora A, Karpen G. Mislocalization of the Drosophila centromere-specific histone CID promotes formation of functional ectopic kinetochores. Dev Cell. 2006;10:303-15 pubmed
    ..Thus, CENP-A mislocalization is one possible mechanism for genome instability during cancer progression, as well as centromere plasticity during evolution. ..
  42. Zhang L, Beaucher M, Cheng Y, Rong Y. Coordination of transposon expression with DNA replication in the targeting of telomeric retrotransposons in Drosophila. EMBO J. 2014;33:1148-58 pubmed publisher
    ..Our study also supports a co-adaptive relationship between the Drosophila host and HeT-A mobile elements. ..
  43. Kurzhals R, Titen S, Xie H, Golic K. Chk2 and p53 are haploinsufficient with dependent and independent functions to eliminate cells after telomere loss. PLoS Genet. 2011;7:e1002103 pubmed publisher
    ..In spite of these mechanisms to eliminate cells that have lost a telomere, we find that such cells can make a substantial contribution to differentiated adult tissues. ..
  44. Burgio G, Cipressa F, Ingrassia A, Cenci G, Corona D. The histone deacetylase Rpd3 regulates the heterochromatin structure of Drosophila telomeres. J Cell Sci. 2011;124:2041-8 pubmed publisher
    ..Our work suggests that Drosophila telomere structure is epigenetically regulated by the histone deacetylase Rpd3. ..
  45. Shaffer C, Stephens G, Thompson B, Funches L, Bernat J, Craig C, et al. Heterochromatin protein 2 (HP2), a partner of HP1 in Drosophila heterochromatin. Proc Natl Acad Sci U S A. 2002;99:14332-7 pubmed
    ..Mutations recovered in HP2 act as dominant suppressors of position effect variegation, confirming a role in heterochromatin spreading and gene silencing. ..
  46. López Panadès E, Casacuberta E. NAP-1, Nucleosome assembly protein 1, a histone chaperone involved in Drosophila telomeres. Insect Biochem Mol Biol. 2016;70:111-5 pubmed publisher
    ..All these results facilitate the understanding of the transposon telomere maintenance mechanism, as well as the integration of telomere biology with the rest of the cell metabolism. ..
  47. Komonyi O, Schauer T, Papai G, Deak P, Boros I. A product of the bicistronic Drosophila melanogaster gene CG31241, which also encodes a trimethylguanosine synthase, plays a role in telomere protection. J Cell Sci. 2009;122:769-74 pubmed publisher
    ..Thus, uncovering the role of DTL will help to dissect the evolutionary conserved pathway(s) controlling ATM-ATR-related telomere protection. ..