Gene Symbol: vtd
Description: verthandi
Alias: 80Fh, CG17436, CG40222, DRAD21, DRad21, DmRAD21, Dmel\CG17436, Drad21, RAD21, Rad21, Rad21/Scc1, Scc1, VTD, dRad21, drad21, l(3)80Fh, l(3)Lh3, lethal 3, rad21, scc1, verthandi, CG17436-PA, CG17436-PB, Rad21, alpha kleisin, alpha-kleisin, cohesin, lethal 3, lethal(3)80Fh, vtd-PA, vtd-PB
Species: fruit fly

Top Publications

  1. Strübbe G, Popp C, Schmidt A, Pauli A, Ringrose L, Beisel C, et al. Polycomb purification by in vivo biotinylation tagging reveals cohesin and Trithorax group proteins as interaction partners. Proc Natl Acad Sci U S A. 2011;108:5572-7 pubmed publisher
    ..Furthermore, our analysis revealed an association of PcG protein complexes with the cohesin complex and showed that Polycomb-dependent silencing of a transgenic reporter depends on cohesin function.
  2. Markov A, Zakharov A, Galkin A, Strunnikov A, Smirnov A. [Localization of cohesin complexes of polytene chromosomes of Drosophila melanogaster located on interbands]. Genetika. 2003;39:1203-11 pubmed
    The distribution of cohesin complex in polytene chromosomes of Drosophila melanogaster was studied. Cohesin is a complicated protein complex which is regulated by the DRAD21 subunit...
  3. Hallson G, Syrzycka M, Beck S, Kennison J, Dorsett D, Page S, et al. The Drosophila cohesin subunit Rad21 is a trithorax group (trxG) protein. Proc Natl Acad Sci U S A. 2008;105:12405-10 pubmed publisher
    ..We show that the Drosophila Rad21 protein is encoded by verthandi (vtd), a member of the trxG gene family that is also involved in regulating the hedgehog (hh) gene...
  4. Schaaf C, Misulovin Z, Gause M, Koenig A, Gohara D, Watson A, et al. Cohesin and polycomb proteins functionally interact to control transcription at silenced and active genes. PLoS Genet. 2013;9:e1003560 pubmed publisher
    b>Cohesin is crucial for proper chromosome segregation but also regulates gene transcription and organism development by poorly understood mechanisms...
  5. Cunningham M, Gause M, Cheng Y, Noyes A, Dorsett D, Kennison J, et al. Wapl antagonizes cohesin binding and promotes Polycomb-group silencing in Drosophila. Development. 2012;139:4172-9 pubmed publisher
    Wapl protein regulates binding of the cohesin complex to chromosomes during interphase and helps remove cohesin from chromosomes at mitosis...
  6. Eichinger C, Kurze A, Oliveira R, Nasmyth K. Disengaging the Smc3/kleisin interface releases cohesin from Drosophila chromosomes during interphase and mitosis. EMBO J. 2013;32:656-65 pubmed publisher
    b>Cohesin's Smc1, Smc3, and kleisin subunits create a tripartite ring within which sister DNAs are entrapped. Evidence suggests that DNA enters through a gate created by transient dissociation of the Smc1/3 interface...
  7. Rollins R, Korom M, Aulner N, Martens A, Dorsett D. Drosophila nipped-B protein supports sister chromatid cohesion and opposes the stromalin/Scc3 cohesion factor to facilitate long-range activation of the cut gene. Mol Cell Biol. 2004;24:3100-11 pubmed
    ..The evolutionarily conserved Cohesin protein complex mediates sister chromatid cohesion, and Scc2 and Mis4 are needed for Cohesin to associate with ..
  8. Gause M, Webber H, Misulovin Z, Haller G, Rollins R, Eissenberg J, et al. Functional links between Drosophila Nipped-B and cohesin in somatic and meiotic cells. Chromosoma. 2008;117:51-66 pubmed
    ..The Nipped-B protein family is required for the cohesin complex that mediates sister chromatid cohesion to bind to chromosomes...
  9. Oliveira R, Coelho P, Sunkel C. The condensin I subunit Barren/CAP-H is essential for the structural integrity of centromeric heterochromatin during mitosis. Mol Cell Biol. 2005;25:8971-84 pubmed
    ..Taken together, our data suggest that the condensin I complex is required not only to promote sister chromatid resolution but also to maintain the structural integrity of centromeric heterochromatin during mitosis. ..

More Information


  1. Schaaf C, Kwak H, Koenig A, Misulovin Z, Gohara D, Watson A, et al. Genome-wide control of RNA polymerase II activity by cohesin. PLoS Genet. 2013;9:e1003382 pubmed publisher
    b>Cohesin is a well-known mediator of sister chromatid cohesion, but it also influences gene expression and development...
  2. Kennison J, Tamkun J. Dosage-dependent modifiers of polycomb and antennapedia mutations in Drosophila. Proc Natl Acad Sci U S A. 1988;85:8136-40 pubmed
    ..e., Sex combs reduced (Scr), Brista (Ba), trithorax (trx), Polycomb (Pc), Polycomblike (Pcl), and Sex comb on midleg (Scm)]. Mutations in several of the additional loci identified here have also been shown to have homoeotic phenotypes. ..
  3. Oliveira R, Hamilton R, Pauli A, Davis I, Nasmyth K. Cohesin cleavage and Cdk inhibition trigger formation of daughter nuclei. Nat Cell Biol. 2010;12:185-92 pubmed publisher
    ..Separase cleaves cohesin, a ring-shaped complex that entraps sister DNAs...
  4. Fay A, Misulovin Z, Li J, Schaaf C, Gause M, Gilmour D, et al. Cohesin selectively binds and regulates genes with paused RNA polymerase. Curr Biol. 2011;21:1624-34 pubmed publisher
    The cohesin complex mediates sister chromatid cohesion and regulates gene transcription...
  5. Coelho P, Queiroz Machado J, Sunkel C. Condensin-dependent localisation of topoisomerase II to an axial chromosomal structure is required for sister chromatid resolution during mitosis. J Cell Sci. 2003;116:4763-76 pubmed
    ..bridges do not result from inappropriate maintenance of sister chromatid cohesion by DRAD21, a subunit of the cohesin complex...
  6. Vass S, Cotterill S, Valdeolmillos A, Barbero J, Lin E, Warren W, et al. Depletion of Drad21/Scc1 in Drosophila cells leads to instability of the cohesin complex and disruption of mitotic progression. Curr Biol. 2003;13:208-18 pubmed
    ..In this study, we examine the consequences of depleting Drad21 and SA, two non-SMC subunits of the cohesin complex, by dsRNA-mediated interference in Drosophila cultured cells...
  7. Warren W, Steffensen S, Lin E, Coelho P, Loupart M, Cobbe N, et al. The Drosophila RAD21 cohesin persists at the centromere region in mitosis. Curr Biol. 2000;10:1463-6 pubmed
    b>Cohesin' is a highly conserved multiprotein complex thought to be the primary effector of sister-chromatid cohesion in all eukaryotes...
  8. Pauli A, van Bemmel J, Oliveira R, Itoh T, Shirahige K, van Steensel B, et al. A direct role for cohesin in gene regulation and ecdysone response in Drosophila salivary glands. Curr Biol. 2010;20:1787-98 pubmed publisher
    ..Developmental abnormalities observed in Cornelia de Lange syndrome have been genetically linked to mutations in the cohesin machinery...
  9. Gause M, Misulovin Z, Bilyeu A, Dorsett D. Dosage-sensitive regulation of cohesin chromosome binding and dynamics by Nipped-B, Pds5, and Wapl. Mol Cell Biol. 2010;30:4940-51 pubmed publisher
    The cohesin protein complex holds sister chromatids together to ensure proper chromosome segregation upon cell division and also regulates gene transcription...
  10. Pauli A, Althoff F, Oliveira R, Heidmann S, Schuldiner O, Lehner C, et al. Cell-type-specific TEV protease cleavage reveals cohesin functions in Drosophila neurons. Dev Cell. 2008;14:239-51 pubmed publisher
    ..At the metaphase to anaphase transition, proteolytic cleavage of the alpha kleisin subunit (Rad21) by separase causes cohesin's dissociation from chromosomes and triggers sister-chromatid ..
  11. Heidmann D, Horn S, Heidmann S, Schleiffer A, Nasmyth K, Lehner C. The Drosophila meiotic kleisin C(2)M functions before the meiotic divisions. Chromosoma. 2004;113:177-87 pubmed
    ..In yeast, the meiosis-specific alpha-kleisin subunit of the cohesin complex, Rec8, is protected from cleavage by separase but only during meiosis I and specifically within the ..
  12. Guo Z, Batiha O, Bourouh M, Fifield E, Swan A. Role of Securin, Separase and Cohesins in female meiosis and polar body formation in Drosophila. J Cell Sci. 2016;129:531-42 pubmed publisher
    ..pathway that culminates in Separase-mediated cleavage of the α-kleisin Rec8, leading to dissolution of cohesin rings...
  13. Schaaf C, Misulovin Z, Gause M, Koenig A, Dorsett D. The Drosophila enhancer of split gene complex: architecture and coordinate regulation by notch, cohesin, and polycomb group proteins. G3 (Bethesda). 2013;3:1785-94 pubmed publisher
    The cohesin protein complex functionally interacts with Polycomb group (PcG) silencing proteins to control expression of several key developmental genes, such as the Drosophila Enhancer of split gene complex [E(spl)-C]...
  14. Carter S, Sjögren C. The SMC complexes, DNA and chromosome topology: right or knot?. Crit Rev Biochem Mol Biol. 2012;47:1-16 pubmed publisher
    ..This review focuses on the details of the relationship between the SMC complexes and topology. It also discusses the possibility that the SMC complexes are united by a capability to sense the geometrical chirality of DNA crossings. ..
  15. Oliveira R, Kotadia S, Tavares A, Mirkovic M, Bowlin K, Eichinger C, et al. Centromere-independent accumulation of cohesin at ectopic heterochromatin sites induces chromosome stretching during anaphase. PLoS Biol. 2014;12:e1001962 pubmed publisher
    ..It contributes to sister chromatid cohesion, a process mediated by the cohesin complex that ensures proper genome segregation during nuclear division...
  16. Sinclair D, Schulze S, Silva E, Fitzpatrick K, Honda B. Essential genes in autosomal heterochromatin of Drosophila melanogaster. Genetica. 2000;109:9-18 pubmed
    ..Our results to date suggest that the three most proximal genes in 3L heterochromatin have key roles in development, and indicate strong effects of combinations of genetic modifiers of PEV on het gene expression. ..
  17. Dimitri P, Corradini N, Rossi F, Vernì F, Cenci G, Belloni G, et al. Vital genes in the heterochromatin of chromosomes 2 and 3 of Drosophila melanogaster. Genetica. 2003;117:209-15 pubmed
    ..Repetitive loci, such as Responder and the ABO elements, which are also located in the heterochromatin of chromosome 2, are not discussed here because they have been reviewed in detail elsewhere. ..
  18. Koryakov D, Zhimulev I, Dimitri P. Cytogenetic analysis of the third chromosome heterochromatin of Drosophila melanogaster. Genetics. 2002;160:509-17 pubmed
    ..Intriguingly, a major part of chromosome 3 heterochromatin was found to be "empty," in that it did not contain either known genes or known satellite DNAs. ..
  19. Orr B, Bousbaa H, Sunkel C. Mad2-independent spindle assembly checkpoint activation and controlled metaphase-anaphase transition in Drosophila S2 cells. Mol Biol Cell. 2007;18:850-63 pubmed
    ..containing a microtubule poison, they arrest with high levels of kinetochore-associated BubR1, properly localized cohesin complex and fail to exit mitosis revealing normal spindle assembly checkpoint activity...
  20. Muraro N, Moffat K. Down-regulation of torp4a, encoding the Drosophila homologue of torsinA, results in increased neuronal degeneration. J Neurobiol. 2006;66:1338-53 pubmed
  21. Schulze S, Sinclair D, FitzPatrick K, Honda B. A genetic and molecular characterization of two proximal heterochromatic genes on chromosome 3 of Drosophila melanogaster. Genetics. 2005;169:2165-77 pubmed
    ..The second gene in this study, Dbp80, is very large and also appears to show a transcriptional dependence upon HP1; however, it does not correspond to any known lethal complementation group and is likely to be a nonessential gene. ..
  22. Nuzhdin S, Reiwitch S. Are the same genes responsible for intra- and interspecific variability for sex comb tooth number in Drosophila?. Heredity (Edinb). 2000;84 ( Pt 1):97-102 pubmed
    ..One intraspecific QTL mapped to the same interval as the QTL for interspecific differences between D. simulans and D. mauritiana. Whether or not these effects result from the same genes requires further examination. ..
  23. Godinho S, Tavares A. A role for Drosophila Polo protein in chromosome resolution and segregation during mitosis. Cell Cycle. 2008;7:2529-34 pubmed
    ..metaphase-anaphase transition that chromosomes lose cohesion completely, by proteolysis of the component of the cohesin complex Scc1 (Sister chromatid cohesion 1)...
  24. Schulze S, McAllister B, Sinclair D, FitzPatrick K, Marchetti M, Pimpinelli S, et al. Heterochromatic genes in Drosophila: a comparative analysis of two genes. Genetics. 2006;173:1433-45 pubmed publisher
    ..This study shows that Dbp80 and RpL15 can function in contrasting chromatin contexts on an evolutionary timescale. The complex history of these genes also provides unique insight into the dynamic nature of genome evolution...
  25. Sollars V, Lu X, Xiao L, Wang X, Garfinkel M, Ruden D. Evidence for an epigenetic mechanism by which Hsp90 acts as a capacitor for morphological evolution. Nat Genet. 2003;33:70-4 pubmed
    ..These findings suggest that Hsp90 acts as a capacitor for morphological evolution through epigenetic and genetic mechanisms. ..
  26. Williams B, Garrett Engele C, Li Z, Williams E, Rosenman E, Goldberg M. Two putative acetyltransferases, san and deco, are required for establishing sister chromatid cohesion in Drosophila. Curr Biol. 2003;13:2025-36 pubmed
    ..Chromatids are linked by the multiprotein cohesin complex, which binds to DNA during G(1) and then establishes cohesion during S phase DNA replication...
  27. Marchant G, Holm D. Genetic Analysis of the Heterochromatin of Chromosome 3 in Drosophila Melanogaster. II. Vital Loci Identified through Ems Mutagenesis. Genetics. 1988;120:519-32 pubmed
    ..This study provides a detailed genetic analysis of chromosome 3 heterochromatin and offers further information on the genetic nature and heterogeneity of Drosophila heterochromatin. ..
  28. Meyer R, Algazeery A, Capri M, Brazier H, Ferry C, Aït Ahmed O. Drosophila Yemanuclein associates with the cohesin and synaptonemal complexes. J Cell Sci. 2014;127:4658-66 pubmed publisher
    ..3 chaperone in complex with Hira. We report here that Yem associates with the synaptonemal complex and the cohesin complex...
  29. Haering C, Nasmyth K. Building and breaking bridges between sister chromatids. Bioessays. 2003;25:1178-91 pubmed
    ..Two multisubunit protein complexes termed 'cohesin' and 'condensin', both composed of SMC (Structural Maintenance of Chromosomes) and kleisin subunits, have emerged ..
  30. Dorsett D. Cohesin, gene expression and development: lessons from Drosophila. Chromosome Res. 2009;17:185-200 pubmed publisher
    The cohesin complex, discovered through its role in sister chromatid cohesion, also plays roles in gene expression and development in organisms from yeast to human...
  31. Benna C, Bonaccorsi S, Wülbeck C, Helfrich Forster C, Gatti M, Kyriacou C, et al. Drosophila timeless2 is required for chromosome stability and circadian photoreception. Curr Biol. 2010;20:346-52 pubmed publisher
    ..tim2's residual role in light entrainment thus provides an evolutionary link that may explain why its derived paralog, tim1, came to play such a major role in both circadian photosensitivity and core clock function. ..
  32. Althoff F, Karess R, Lehner C. Spindle checkpoint-independent inhibition of mitotic chromosome segregation by Drosophila Mps1. Mol Biol Cell. 2012;23:2275-91 pubmed publisher
    ..These findings further expand Mps1's significance for faithful mitotic chromosome segregation and emphasize the importance of its careful regulation...
  33. Mouri K, Horiuchi S, Uemura T. Cohesin controls planar cell polarity by regulating the level of the seven-pass transmembrane cadherin Flamingo. Genes Cells. 2012;17:509-24 pubmed publisher
    ..SMC3 protein is one subunit of the cohesin complex, which regulates sister chromatid cohesion and also plays a role in transcriptional control of gene ..
  34. Kapoun A, Kaufman T. Regulatory regions of the homeotic gene proboscipedia are sensitive to chromosomal pairing. Genetics. 1995;140:643-58 pubmed
    ..Interestingly, the intronic PS regions contain positive regulatory sequences for pb, whereas the upstream PS sites contain pb negative regulatory elements. ..
  35. Kassis J. Unusual properties of regulatory DNA from the Drosophila engrailed gene: three "pairing-sensitive" sites within a 1.6-kb region. Genetics. 1994;136:1025-38 pubmed
    ..I propose that engrailed PS sites normally act to promote interactions between distantly located engrailed regulatory sites and the engrailed promoter. ..
  36. Valdeolmillos A, Rufas J, Suja J, Vass S, Heck M, Martinez A C, et al. Drosophila cohesins DSA1 and Drad21 persist and colocalize along the centromeric heterochromatin during mitosis. Biol Cell. 2004;96:457-62 pubmed
    Sister chromatid cohesion in eukaryotes is maintained mainly by a conserved multiprotein complex termed cohesin. Drad21 and DSA1 are the Drosophila homologues of the yeast Scc1 and Scc3 cohesin subunits, respectively...
  37. Choo K. Domain organization at the centromere and neocentromere. Dev Cell. 2001;1:165-77 pubmed
    ..Studies in different organisms point to a model of conserved pattern of organization for these domains. ..
  38. Warren W, Lin E, Nheu T, Hime G, McKay M. Drad21, a Drosophila rad21 homologue expressed in S-phase cells. Gene. 2000;250:77-84 pubmed
    b>Cohesin is an evolutionarily conserved multiprotein complex required to establish and maintain sister chromatid cohesion...
  39. Coelho P, Queiroz Machado J, Carmo A, Moutinho Pereira S, Maiato H, Sunkel C. Dual role of topoisomerase II in centromere resolution and aurora B activity. PLoS Biol. 2008;6:e207 pubmed publisher
  40. Goshima G, Wollman R, Stuurman N, Scholey J, Vale R. Length control of the metaphase spindle. Curr Biol. 2005;15:1979-88 pubmed
    ..Dis1/XMAP215/TOG]) or depolymerization (Kinesin-8, Kinesin-13), or mediate sister-chromatid cohesion (Rad21) in order to explore how these forces influence spindle length...
  41. Schulze S, Sinclair D, Silva E, Fitzpatrick K, Singh M, Lloyd V, et al. Essential genes in proximal 3L heterochromatin of Drosophila melanogaster. Mol Gen Genet. 2001;264:782-9 pubmed
    ..The third gene to the left of the centromere [lethal 3, l3 or l(3)80Fh] is likely to correspond to verthandi (vtd), a known trx-G gene that plays a role in the regulation of hedgehog (hh) expression and signalling...
  42. Ribeiro A, Silva R, Foyn H, Tiago M, Rathore O, Arnesen T, et al. Naa50/San-dependent N-terminal acetylation of Scc1 is potentially important for sister chromatid cohesion. Sci Rep. 2016;6:39118 pubmed publisher
    ..Our work suggests that Naa50/San is required during tissue proliferation for the correct interaction between the cohesin subunits Scc1 and Smc3...
  43. Liu D, Shaukat Z, Saint R, Gregory S. Chromosomal instability triggers cell death via local signalling through the innate immune receptor Toll. Oncotarget. 2015;6:38552-65 pubmed publisher
    ..This pathway also activates Mmp1, which recruits hemocytes to the CIN tissue to provide local amplification of the immune response that is needed for effective elimination of CIN cells. ..
  44. Kusch T. Brca2-Pds5 complexes mobilize persistent meiotic recombination sites to the nuclear envelope. J Cell Sci. 2015;128:717-27 pubmed publisher
    ..The identification of Brca2-Pds5 complexes as key mediators of this process provides a first mechanistic explanation for the contribution of lamins and cohesins to meiotic recombination. ..
  45. Buchner K, Roth P, Schotta G, Krauss V, Saumweber H, Reuter G, et al. Genetic and molecular complexity of the position effect variegation modifier mod(mdg4) in Drosophila. Genetics. 2000;155:141-57 pubmed
  46. Petruk S, Smith S, Sedkov Y, Mazo A. Association of trxG and PcG proteins with the bxd maintenance element depends on transcriptional activity. Development. 2008;135:2383-90 pubmed publisher
    ..There is, however, no overall synergism or antagonism between and within the trxG and PcG proteins and, instead, only subsets of trxG proteins act synergistically. ..
  47. Oliveira R, Nasmyth K. Cohesin cleavage is insufficient for centriole disengagement in Drosophila. Curr Biol. 2013;23:R601-3 pubmed publisher
  48. Van Bortle K, Nichols M, Li L, Ong C, Takenaka N, Qin Z, et al. Insulator function and topological domain border strength scale with architectural protein occupancy. Genome Biol. 2014;15:R82 pubmed publisher
  49. Kennison J. Transcriptional activation of Drosophila homeotic genes from distant regulatory elements. Trends Genet. 1993;9:75-9 pubmed
    ..Some of the trans-acting proteins may facilitate interactions between cis-regulatory elements and the promoter by bringing together distant chromosomal elements. ..
  50. Fasulo B, Deuring R, Murawska M, Gause M, Dorighi K, Schaaf C, et al. The Drosophila MI-2 chromatin-remodeling factor regulates higher-order chromatin structure and cohesin dynamics in vivo. PLoS Genet. 2012;8:e1002878 pubmed publisher
    ..dMi-2 and the cohesin complex are enriched at sites of active transcription; fluorescence-recovery after photobleaching (FRAP) assays ..
  51. Pimenta Marques A, Tostões R, Marty T, Barbosa V, Lehmann R, Martinho R. Differential requirements of a mitotic acetyltransferase in somatic and germ line cells. Dev Biol. 2008;323:197-206 pubmed publisher
    ..Surprisingly, during oogenesis we cannot detect mitotic defects in germ line cells mutant for san. We hypothesize the female germ line stem cells have differential requirements for mitotic sister chromatid cohesion. ..
  52. Li L, Lyu X, Hou C, Takenaka N, Nguyen H, Ong C, et al. Widespread rearrangement of 3D chromatin organization underlies polycomb-mediated stress-induced silencing. Mol Cell. 2015;58:216-31 pubmed publisher
    ..These results suggest that the TAD organization of metazoan genomes is plastic and can be reconfigured quickly. ..
  53. Mirkovic M, Hutter L, Novák B, Oliveira R. Premature Sister Chromatid Separation Is Poorly Detected by the Spindle Assembly Checkpoint as a Result of System-Level Feedback. Cell Rep. 2015;13:469-478 pubmed publisher
    Sister chromatid cohesion, mediated by the cohesin complex, is essential for faithful mitosis...
  54. Urban E, Nagarkar Jaiswal S, Lehner C, Heidmann S. The cohesin subunit Rad21 is required for synaptonemal complex maintenance, but not sister chromatid cohesion, during Drosophila female meiosis. PLoS Genet. 2014;10:e1004540 pubmed publisher
    ..This association is mediated by the ring-shaped cohesin complex that appears to embrace the sister chromatids...
  55. Lee J, Dej K, Lopez J, Orr Weaver T. Control of centromere localization of the MEI-S332 cohesion protection protein. Curr Biol. 2004;14:1277-83 pubmed
    ..One likely activity of these proteins is to protect the cohesin subunit Rec8 from cleavage at the metaphase I-anaphase I transition...
  56. Swain A, Misulovin Z, Pherson M, Gause M, Mihindukulasuriya K, Rickels R, et al. Drosophila TDP-43 RNA-Binding Protein Facilitates Association of Sister Chromatid Cohesion Proteins with Genes, Enhancers and Polycomb Response Elements. PLoS Genet. 2016;12:e1006331 pubmed publisher
    The cohesin protein complex mediates sister chromatid cohesion and participates in transcriptional control of genes that regulate growth and development...
  57. Manning A, Longworth M, Dyson N. Loss of pRB causes centromere dysfunction and chromosomal instability. Genes Dev. 2010;24:1364-76 pubmed publisher
    ..These observations explain an important consequence of RB1 inactivation, and suggest that subtle defects in centromere function are a frequent source of merotely and CIN in cancer. ..