Gene Symbol: rod
Description: rough deal
Alias: CG1569, Dmel\CG1569, EP3408, ROD, Rod, pNB20A, rough deal, CG1569-PA, rod-PA, roughdeal
Species: fruit fly

Top Publications

  1. Williams B, Goldberg M. Determinants of Drosophila zw10 protein localization and function. J Cell Sci. 1994;107 ( Pt 4):785-98 pubmed
    ..c) The localization of zw10 protein is altered by two mitotic mutations, rough deal and abnormal anaphase resolution, that specifically disrupt anaphase...
  2. Basto R, Scaërou F, Mische S, Wojcik E, Lefebvre C, Gomes R, et al. In vivo dynamics of the rough deal checkpoint protein during Drosophila mitosis. Curr Biol. 2004;14:56-61 pubmed
    b>Rough Deal (Rod) and Zw10 are components of a complex required for the metazoan metaphase checkpoint and for recruitment of dynein/dynactin to the kinetochore...
  3. Gatti M, Goldberg M. Mutations affecting cell division in Drosophila. Methods Cell Biol. 1991;35:543-86 pubmed
  4. Basu J, Bousbaa H, Logarinho E, Li Z, Williams B, Lopes C, et al. Mutations in the essential spindle checkpoint gene bub1 cause chromosome missegregation and fail to block apoptosis in Drosophila. J Cell Biol. 1999;146:13-28 pubmed
    ..Bub1's localization to the kinetochore does not depend on the products of the genes zw10, rod, polo, or fizzy, indicating that the kinetochore is constructed from several independent subassemblies.
  5. Scaerou F, Starr D, Piano F, Papoulas O, Karess R, Goldberg M. The ZW10 and Rough Deal checkpoint proteins function together in a large, evolutionarily conserved complex targeted to the kinetochore. J Cell Sci. 2001;114:3103-14 pubmed
    The zeste-white 10 (zw10) and rough deal (rod) genes of Drosophila both encode kinetochore components, and mutations in either gene greatly increase the missegregation of sister chromatids during mitosis...
  6. Basu J, Logarinho E, Herrmann S, Bousbaa H, Li Z, Chan G, et al. Localization of the Drosophila checkpoint control protein Bub3 to the kinetochore requires Bub1 but not Zw10 or Rod. Chromosoma. 1998;107:376-85 pubmed
    ..In contrast, we demonstrate that the kinetochore constituents Zw10 and Rod are not needed for the binding of Bub3 to the kinetochore...
  7. Scaerou F, Aguilera I, Saunders R, Kane N, Blottière L, Karess R. The rough deal protein is a new kinetochore component required for accurate chromosome segregation in Drosophila. J Cell Sci. 1999;112 ( Pt 21):3757-68 pubmed
    Mutations in the rough deal (rod) gene of Drosophila greatly increase the missegregation of sister chromatids during mitosis, suggesting a role for this gene product in spindle or kinetochore function...
  8. Karess R, Glover D. rough deal: a gene required for proper mitotic segregation in Drosophila. J Cell Biol. 1989;109:2951-61 pubmed
    We describe a genetic locus rough deal (rod) in Drosophila melanogaster, identified by mutations that interfere with the faithful transmission of chromosomes to daughter cells during mitosis...
  9. Basto R, Gomes R, Karess R. Rough deal and Zw10 are required for the metaphase checkpoint in Drosophila. Nat Cell Biol. 2000;2:939-43 pubmed
    ..Here we show that the Drosophila kinetochore components Rough deal (Rod) and Zeste-White 10 (Zw10) are required for the proper functioning of the metaphase checkpoint in flies...

More Information


  1. Buffin E, Emre D, Karess R. Flies without a spindle checkpoint. Nat Cell Biol. 2007;9:565-72 pubmed
  2. Blower M, Daigle T, Kaufman T, Karpen G. Drosophila CENP-A mutations cause a BubR1-dependent early mitotic delay without normal localization of kinetochore components. PLoS Genet. 2006;2:e110 pubmed
    ..We propose that SAC components are able to monitor spindle assembly and inhibit cell cycle progression in the absence of sustained kinetochore localization. ..
  3. 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. ..
  4. Siller K, Serr M, Steward R, Hays T, Doe C. Live imaging of Drosophila brain neuroblasts reveals a role for Lis1/dynactin in spindle assembly and mitotic checkpoint control. Mol Biol Cell. 2005;16:5127-40 pubmed
    ..In addition, we develop an improved time-lapse video microscopy technique that allows live imaging of GFP-Lis1, GFP-Rod checkpoint protein, green fluorescent protein (GFP)-labeled chromosomes, or GFP-labeled mitotic spindle dynamics in ..
  5. Williams B, Li Z, Liu S, Williams E, Leung G, Yen T, et al. Zwilch, a new component of the ZW10/ROD complex required for kinetochore functions. Mol Biol Cell. 2003;14:1379-91 pubmed
    The Zeste-White 10 (ZW10) and Rough Deal (ROD) proteins are part of a complex necessary for accurate chromosome segregation. This complex recruits cytoplasmic dynein to the kinetochore and participates in the spindle checkpoint...
  6. Blower M, Karpen G. The role of Drosophila CID in kinetochore formation, cell-cycle progression and heterochromatin interactions. Nat Cell Biol. 2001;3:730-9 pubmed
    ..separate from proteins involved in sister cohesion (MEI-S332), centric condensation (PROD), kinetochore function (ROD, ZW10 and BUB1) and heterochromatin structure (HP1)...
  7. Rahmani Z, Gagou M, Lefebvre C, Emre D, Karess R. Separating the spindle, checkpoint, and timer functions of BubR1. J Cell Biol. 2009;187:597-605 pubmed publisher
    ..Timing is shorter in bubR1-KEN mad2 double mutants, yet in these flies, lacking both critical SAC components, chromosomes still segregate accurately, reconfirming that in Drosophila, reliable mitosis does not need the SAC. ..
  8. Shah J, Cleveland D. Waiting for anaphase: Mad2 and the spindle assembly checkpoint. Cell. 2000;103:997-1000 pubmed
  9. Wojcik E, Basto R, Serr M, Scaerou F, Karess R, Hays T. Kinetochore dynein: its dynamics and role in the transport of the Rough deal checkpoint protein. Nat Cell Biol. 2001;3:1001-7 pubmed
    ..This behaviour is seen for several metaphase checkpoint proteins, including Rough deal (Rod)...
  10. Blower M, Sullivan B, Karpen G. Conserved organization of centromeric chromatin in flies and humans. Dev Cell. 2002;2:319-30 pubmed
    ..We propose that the unique structural arrangement of CID/CENP-A and H3 nucleosomes presents centromeric chromatin to the poleward face of the condensing mitotic chromosome. ..
  11. Dzhindzhev N, Rogers S, Vale R, Ohkura H. Distinct mechanisms govern the localisation of Drosophila CLIP-190 to unattached kinetochores and microtubule plus-ends. J Cell Sci. 2005;118:3781-90 pubmed
    ..These results indicate distinct molecular requirements for CLIP-190 localisation to unattached kinetochores in mitosis and microtubule ends in interphase. ..
  12. Buffin E, Lefebvre C, Huang J, Gagou M, Karess R. Recruitment of Mad2 to the kinetochore requires the Rod/Zw10 complex. Curr Biol. 2005;15:856-61 pubmed
    ..Among these are Rod, Zw10, and Zwilch, components of a 700 Kdal complex (Rod/Zw10) that is required for recruitment of dynein/dynactin ..
  13. Starr D, Williams B, Hays T, Goldberg M. ZW10 helps recruit dynactin and dynein to the kinetochore. J Cell Biol. 1998;142:763-74 pubmed
    ..and (c) the localization of both ZW10 and dynein to the kinetochore is abolished in cells mutant for the gene rough deal. ZW10's role in the recruitment of dynein to the kinetochore is likely to be reasonably direct, because ..
  14. Karess R. Rod-Zw10-Zwilch: a key player in the spindle checkpoint. Trends Cell Biol. 2005;15:386-92 pubmed
    ..But three proteins with no yeast homologs--Rod, Zw10 and Zwilch--also play a crucial, but poorly understood, role in the metazoan spindle checkpoint...
  15. Lince Faria M, Maffini S, Orr B, Ding Y, Cláudia Florindo -, Sunkel C, et al. Spatiotemporal control of mitosis by the conserved spindle matrix protein Megator. J Cell Biol. 2009;184:647-57 pubmed publisher
  16. Schejter E, Wieschaus E. bottleneck acts as a regulator of the microfilament network governing cellularization of the Drosophila embryo. Cell. 1993;75:373-85 pubmed
    ..The expression pattern and mutant phenotype of bnk suggest a direct role for this element in regulation of the dynamic restructuring of the actin-based cytoskeleton of cellularizing Drosophila embryos. ..
  17. Parry D, Hickson G, O Farrell P. Cyclin B destruction triggers changes in kinetochore behavior essential for successful anaphase. Curr Biol. 2003;13:647-53 pubmed
    ..These findings show that, in addition to loss of sister chromosome cohesion, successful anaphase requires a kinetochore behavioral transition triggered by CYC-B destruction. ..
  18. Mottier Pavie V, Cenci G, Vernì F, Gatti M, Bonaccorsi S. Phenotypic analysis of misato function reveals roles of noncentrosomal microtubules in Drosophila spindle formation. J Cell Sci. 2011;124:706-17 pubmed publisher
    ..Collectively, these results indicate that kinetochore-driven MT formation is an essential process for proper spindle assembly in Drosophila somatic cells. ..
  19. Wisotzkey R, Mehra A, Sutherland D, Dobens L, Liu X, Dohrmann C, et al. Medea is a Drosophila Smad4 homolog that is differentially required to potentiate DPP responses. Development. 1998;125:1433-45 pubmed
    ..We propose that this provides a general model for Smad4/MEDEA function in signaling by the TGF-beta family. ..
  20. Parker D, Jemison J, Cadigan K. Pygopus, a nuclear PHD-finger protein required for Wingless signaling in Drosophila. Development. 2002;129:2565-76 pubmed
    ..Our data argue strongly that Pygopus is a new core component of the Wg signaling pathway that acts downstream or at the level of TCF. ..
  21. Reis R, Feijão T, Gouveia S, Pereira A, Matos I, Sampaio P, et al. Dynein and mast/orbit/CLASP have antagonistic roles in regulating kinetochore-microtubule plus-end dynamics. J Cell Sci. 2009;122:2543-53 pubmed publisher
    ..Our results suggest that mast and dynein have antagonistic roles in the local regulation of microtubule plus-end dynamics at kinetochores, which are important for the maintenance of spindle bipolarity and normal spindle length. ..
  22. Torras Llort M, Medina Giró S, Moreno Moreno O, Azorín F. A conserved arginine-rich motif within the hypervariable N-domain of Drosophila centromeric histone H3 (CenH3) mediates BubR1 recruitment. PLoS ONE. 2010;5:e13747 pubmed publisher
  23. Cabernard C, Prehoda K, Doe C. A spindle-independent cleavage furrow positioning pathway. Nature. 2010;467:91-4 pubmed publisher
    ..This spindle-independent cleavage furrow mechanism may be relevant to other highly polarized mitotic cells, such as mammalian neural progenitors. ..
  24. Radford S, Hoang T, GÅ‚uszek A, Ohkura H, McKim K. Lateral and End-On Kinetochore Attachments Are Coordinated to Achieve Bi-orientation in Drosophila Oocytes. PLoS Genet. 2015;11:e1005605 pubmed publisher
    ..Together, these results demonstrate that, in oocytes, kinetochore-dependent and -independent chromosome-microtubule attachments work together to promote the accurate segregation of chromosomes. ..
  25. Savoian M, Goldberg M, Rieder C. The rate of poleward chromosome motion is attenuated in Drosophila zw10 and rod mutants. Nat Cell Biol. 2000;2:948-52 pubmed
    ..Our results show that ZW10 protein, together with Rod, is involved in production and/or regulation of the force responsible for poleward chromosome motion.
  26. Pandey R, Heeger S, Lehner C. Rapid effects of acute anoxia on spindle kinetochore interactions activate the mitotic spindle checkpoint. J Cell Sci. 2007;120:2807-18 pubmed
    ..Although involving adenosine triphosphate (ATP)-consuming kinases, the spindle checkpoint can therefore be activated by spindle damage in response to acute anoxia and protect against aneuploidies. ..
  27. Williams B, Leung G, Maiato H, Wong A, Li Z, Williams E, et al. Mitch a rapidly evolving component of the Ndc80 kinetochore complex required for correct chromosome segregation in Drosophila. J Cell Sci. 2007;120:3522-33 pubmed
    ..Despite the crucial role of Mitch in cell division, the mitch gene has evolved very rapidly among species in the genus Drosophila. ..
  28. Siller K, Doe C. Lis1/dynactin regulates metaphase spindle orientation in Drosophila neuroblasts. Dev Biol. 2008;319:1-9 pubmed publisher
    ..We propose that an early Lis1/dynactin-dependent pathway and a late Lis1/dynactin-independent pathway regulate neuroblast spindle orientation. ..
  29. Schmitt H. Dsl1p/Zw10: common mechanisms behind tethering vesicles and microtubules. Trends Cell Biol. 2010;20:257-68 pubmed publisher
    ..This review brings together evidence to suggest that there could be common mechanisms behind these different activities, giving clues as to how they might have evolved. ..
  30. Hayward D, Metz J, Pellacani C, Wakefield J. Synergy between multiple microtubule-generating pathways confers robustness to centrosome-driven mitotic spindle formation. Dev Cell. 2014;28:81-93 pubmed publisher
    ..It also demonstrates that the contributions of each pathway to spindle formation are integrated, highlighting the remarkable flexibility with which cells can respond to perturbations that limit their capacity to generate microtubules. ..
  31. Emre D, Terracol R, Poncet A, Rahmani Z, Karess R. A mitotic role for Mad1 beyond the spindle checkpoint. J Cell Sci. 2011;124:1664-71 pubmed publisher
    ..This activity does not require the presence of Mad2, but it does depend in some unknown way on key residues in the Mad2-binding domain of Mad1. ..
  32. Glover D. Abbreviated and regulated cell cycles in Drosophila. Curr Opin Cell Biol. 1990;2:258-61 pubmed
  33. Endow S. Microtubule motors in spindle and chromosome motility. Eur J Biochem. 1999;262:12-8 pubmed
    ..The identification of microtubule motors that function in chromosome distribution represents a major advance in understanding the forces that underlie chromosome and spindle movements during cell division. ..
  34. D Avino P, Archambault V, Przewloka M, Zhang W, Laue E, Glover D. Isolation of protein complexes involved in mitosis and cytokinesis from Drosophila cultured cells. Methods Mol Biol. 2009;545:99-112 pubmed publisher
    ..Although this method has proven very successful in isolating mitotic and cytokinetic complexes, it can also be used to characterise protein complexes involved in many other cellular processes. ..
  35. Parker D, Ni Y, Chang J, Li J, Cadigan K. Wingless signaling induces widespread chromatin remodeling of target loci. Mol Cell Biol. 2008;28:1815-28 pubmed
    ..These data suggest that pathway activation induces localized recruitment of TCF/Armadillo/CBP to Wingless response elements, leading to widespread histone acetylation of target loci prior to transcriptional activation. ..
  36. Milan M, Clemente Ruiz M, Dekanty A, Muzzopappa M. Aneuploidy and tumorigenesis in Drosophila. Semin Cell Dev Biol. 2014;28:110-5 pubmed publisher
    ..Here we discuss the role of aneuploidy in tumorigenesis in light of the contribution of Drosophila epithelial cancer models and propose a stress-induced tumor-promoting role of aneuploidy. ..
  37. Défachelles L, Hainline S, Menant A, Lee L, Karess R. A maternal effect rough deal mutation suggests that multiple pathways regulate Drosophila RZZ kinetochore recruitment. J Cell Sci. 2015;128:1204-16 pubmed publisher
    ..Here, we describe rod(Z3), a maternal-effect Drosophila mutation changing a single residue in the Rough Deal (Rod) subunit of RZZ...
  38. Roth M, Roubinet C, Iffländer N, Ferrand A, Cabernard C. Asymmetrically dividing Drosophila neuroblasts utilize two spatially and temporally independent cytokinesis pathways. Nat Commun. 2015;6:6551 pubmed publisher
    ..These results support the model that two spatially and temporally separate pathways control different key aspects during asymmetric cell division, ensuring correct cell fate determinant segregation and neuroblast self-renewal. ..
  39. Griffis E, Stuurman N, Vale R. Spindly, a novel protein essential for silencing the spindle assembly checkpoint, recruits dynein to the kinetochore. J Cell Biol. 2007;177:1005-15 pubmed
    ..eukaryotes, cytoplasmic dynein is involved in silencing the SAC by removing the checkpoint proteins Mad2 and the Rod-Zw10-Zwilch complex (RZZ) from aligned kinetochores (Howell, B.J., B.F. McEwen, J.C. Canman, D.B. Hoffman, E.M...
  40. Dekanty A, Barrio L, Muzzopappa M, Auer H, Milan M. Aneuploidy-induced delaminating cells drive tumorigenesis in Drosophila epithelia. Proc Natl Acad Sci U S A. 2012;109:20549-54 pubmed publisher
  41. Wainman A, Giansanti M, Goldberg M, Gatti M. The Drosophila RZZ complex - roles in membrane trafficking and cytokinesis. J Cell Sci. 2012;125:4014-25 pubmed publisher
    The Zw10 protein, in the context of the conserved Rod-Zwilch-Zw10 (RZZ) complex, is a kinetochore component required for proper activity of the spindle assembly checkpoint in both Drosophila and mammals...
  42. Gull K. From fungi to flies: pattern and form in the cell cycle. Curr Opin Cell Biol. 1991;3:247-52 pubmed
    ..This review shows how, in some lower eukaryotes, recent studies have made important connections between cell cycle control processes and complex events such as organelle positioning and the differentiation of cells and tissues. ..
  43. Cook K, Murphy T, Nguyen T, Karpen G. Identification of trans-acting genes necessary for centromere function in Drosophila melanogaster using centromere-defective minichromosomes. Genetics. 1997;145:737-47 pubmed
    ..The results presented in this study strongly suggest that dominant genetic interactions between mutations and centromere-defective minichromosomes could be used effectively to identify novel genes necessary for centromere function. ..
  44. Philip A. Mitotic sister-chromatid separation: what Drosophila mutants can tell us. Trends Cell Biol. 1998;8:150 pubmed
  45. Wainman A, Creque J, Williams B, Williams E, Bonaccorsi S, Gatti M, et al. Roles of the Drosophila NudE protein in kinetochore function and centrosome migration. J Cell Sci. 2009;122:1747-58 pubmed publisher
    ..However, in the absence of NudE, the 'shedding' of proteins off the kinetochore is abrogated and the SAC cannot be turned off, implying that NudE regulates dynein function at the kinetochore. ..
  46. Godinho S, Tavares A. A role for Drosophila Polo protein in chromosome resolution and segregation during mitosis. Cell Cycle. 2008;7:2529-34 pubmed
    ..In addition, the checkpoint protein ROD fails to stream over the kinetochore microtubules in the lack of Polo activity...
  47. Castellanos E, Dominguez P, Gonzalez C. Centrosome dysfunction in Drosophila neural stem cells causes tumors that are not due to genome instability. Curr Biol. 2008;18:1209-14 pubmed publisher
    ..We propose that such tumors might be caused by impaired asymmetric division of neural stem cells [4]. These results show that centrosome loss, far from being innocuous, is a potentially dangerous condition in flies...
  48. Olszak A, van Essen D, Pereira A, Diehl S, Manke T, Maiato H, et al. Heterochromatin boundaries are hotspots for de novo kinetochore formation. Nat Cell Biol. 2011;13:799-808 pubmed publisher
    ..These data indicate that the characteristics of regions bordering heterochromatin promote de novo kinetochore assembly and thereby contribute to centromere identity. ..
  49. Dobie K, Hari K, Maggert K, Karpen G. Centromere proteins and chromosome inheritance: a complex affair. Curr Opin Genet Dev. 1999;9:206-17 pubmed
    ..A picture is beginning to emerge of the centromere-kinetechore as a complex and dynamic structure with conservation of function at the protein level across diverse species. ..
  50. Défachelles L, Raich N, Terracol R, Baudin X, Williams B, Goldberg M, et al. RZZ and Mad1 dynamics in Drosophila mitosis. Chromosome Res. 2015;23:333-42 pubmed publisher
    ..Among the factors regulating the levels of Mad1 at kinetochores is the Rod, Zw10, and Zwilch (RZZ) complex, which is required for Mad1 recruitment through a mechanism that remains unknown...
  51. Liu J, Lee D, Yu C, Angers S, Harris T. Stepping stone: a cytohesin adaptor for membrane cytoskeleton restraint in the syncytial Drosophila embryo. Mol Biol Cell. 2015;26:711-25 pubmed publisher
    ..Overall we propose that Sstn acts as a cytohesin adaptor that promotes Steppke activity for localized membrane cytoskeleton restraint in the syncytial Drosophila embryo. ..
  52. Peng J, Karpen G. Heterochromatic genome stability requires regulators of histone H3 K9 methylation. PLoS Genet. 2009;5:e1000435 pubmed publisher
    ..Similar effects of lower magnitude were observed in animals that lack the RNA interference pathway component Dcr2. These results suggest that the H3K9 methylation and RNAi pathways ensure heterochromatin stability. ..