Gene Symbol: mof
Description: males absent on the first
Alias: CG3025, Dmel\CG3025, MOF, Mof, dMOF, dmMOF, max, males absent on the first, CG3025-PA, MOF histone acetylase, males absent on first, males-absent on the X, males-absent-on-the-first, mof-PA
Species: fruit fly

Top Publications

  1. Smith E, Pannuti A, Gu W, Steurnagel A, Cook R, Allis C, et al. The drosophila MSL complex acetylates histone H4 at lysine 16, a chromatin modification linked to dosage compensation. Mol Cell Biol. 2000;20:312-8 pubmed
    ..We have partially purified a complex containing MSL1, -2, and -3, MOF, MLE, and roX2 RNA and demonstrated that it exclusively acetylates H4 at lysine 16 on nucleosomal substrates...
  2. Lucchesi J. Dosage compensation in flies and worms: the ups and downs of X-chromosome regulation. Curr Opin Genet Dev. 1998;8:179-84 pubmed
    ..These two different solutions to the common problem of unequal dosage of X-linked genes in different sexes provide invaluable paradigms for the study of gene regulation at the level of chromatin remodeling. ..
  3. Bai X, Alekseyenko A, Kuroda M. Sequence-specific targeting of MSL complex regulates transcription of the roX RNA genes. EMBO J. 2004;23:2853-61 pubmed
    ..Surprisingly, the DHS is not required for initiation of cis spreading of MSL complex, instead local transcription of roX RNAs correlates with extensive spreading. ..
  4. Gu W, Szauter P, Lucchesi J. Targeting of MOF, a putative histone acetyl transferase, to the X chromosome of Drosophila melanogaster. Dev Genet. 1998;22:56-64 pubmed
    ..of at least five genes, maleless (mle), male-specific lethal 1, 2, and 3 (msl-1, msl-2, msl-3) and males absent on the first (mof), are necessary for dosage compensation...
  5. Gupta A, Guerin Peyrou T, Sharma G, Park C, Agarwal M, Ganju R, et al. The mammalian ortholog of Drosophila MOF that acetylates histone H4 lysine 16 is essential for embryogenesis and oncogenesis. Mol Cell Biol. 2008;28:397-409 pubmed
    The mammalian ortholog of the Drosophila MOF (males absent on the first) gene product is a histone H4 lysine 16-specific acetyltransferase...
  6. Kelley R, Meller V, Gordadze P, Roman G, Davis R, Kuroda M. Epigenetic spreading of the Drosophila dosage compensation complex from roX RNA genes into flanking chromatin. Cell. 1999;98:513-22 pubmed
    ..The male X chromosome is also coated with noncoding roX RNAs. When either msl3, mle, or mof is mutant, a partial MSL complex is bound at only approximately 35 unusual sites distributed along the X...
  7. Park S, Kang Y, Sypula J, Choi J, Oh H, Park Y. An evolutionarily conserved domain of roX2 RNA is sufficient for induction of H4-Lys16 acetylation on the Drosophila X chromosome. Genetics. 2007;177:1429-37 pubmed
  8. Furuhashi H, Nakajima M, Hirose S. DNA supercoiling factor contributes to dosage compensation in Drosophila. Development. 2006;133:4475-83 pubmed
    ..This phenotype was dependent on the histone acetyltransferase MOF and was suppressed by simultaneous overexpression of ISWI...
  9. Meller V. Dosage compensation: making 1X equal 2X. Trends Cell Biol. 2000;10:54-9 pubmed
    ..Each organism uses a largely non-overlapping set of molecules to achieve the same outcome: 1X = 2X. ..

More Information


  1. Bell O, Conrad T, Kind J, Wirbelauer C, Akhtar A, Schübeler D. Transcription-coupled methylation of histone H3 at lysine 36 regulates dosage compensation by enhancing recruitment of the MSL complex in Drosophila melanogaster. Mol Cell Biol. 2008;28:3401-9 pubmed publisher
    ..Thus, H3K36me3 marks the body of all active genes yet is utilized in a chromosome-specific manner to enhance histone acetylation at sites of dosage compensation. ..
  2. Stuckenholz C, Kageyama Y, Kuroda M. Guilt by association: non-coding RNAs, chromosome-specific proteins and dosage compensation in Drosophila. Trends Genet. 1999;15:454-8 pubmed
    ..The discovery of non-coding RNAs that 'paint' dosage-compensated X chromosomes in mammals and in Drosophila suggests that RNAs play an intriguing, unexpected role in the regulation of chromatin structure and gene expression. ..
  3. Gelbart M, Larschan E, Peng S, Park P, Kuroda M. Drosophila MSL complex globally acetylates H4K16 on the male X chromosome for dosage compensation. Nat Struct Mol Biol. 2009;16:825-32 pubmed publisher
  4. Bhadra U, Pal Bhadra M, Birchler J. Role of the male specific lethal (msl) genes in modifying the effects of sex chromosomal dosage in Drosophila. Genetics. 1999;152:249-68 pubmed
    ..However, the staining pattern of acetylation and the mof gene product, a putative histone acetylase, in msl mutant males returns to a uniform genome-wide distribution as ..
  5. Straub T, Neumann M, Prestel M, Kremmer E, Kaether C, Haass C, et al. Stable chromosomal association of MSL2 defines a dosage-compensated nuclear compartment. Chromosoma. 2005;114:352-64 pubmed
    ..Our findings have profound implications for the mechanism underlying dosage compensation and furthermore provide a new, conceptual reference of stability in an otherwise highly dynamic nuclear environment...
  6. Bhadra M, Bhadra U, Kundu J, Birchler J. Gene expression analysis of the function of the male-specific lethal complex in Drosophila. Genetics. 2005;169:2061-74 pubmed
    ..In mutant male embryos without either the MSL complex or MOF histone acetylase, dosage compensation is retained but autosomal expression is increased...
  7. Zhang Y, Malone J, Powell S, Periwal V, Spana E, MacAlpine D, et al. Expression in aneuploid Drosophila S2 cells. PLoS Biol. 2010;8:e1000320 pubmed publisher
    ..Our data indicate that expression dosage compensation dampens the effect of altered DNA copy number genome-wide. For the X chromosome, compensation includes fixed and dose-dependent components. ..
  8. Morales V, Straub T, Neumann M, Mengus G, Akhtar A, Becker P. Functional integration of the histone acetyltransferase MOF into the dosage compensation complex. EMBO J. 2004;23:2258-68 pubmed
    ..Crucial for this activation is the acetylation of histone H4 by the histone acetyltransferase (HAT) MOF. In male cells, MOF resides in a complex (dosage compensation complex, DCC) with MSL proteins and noncoding roX RNA...
  9. Stenberg P, Larsson J. Buffering and the evolution of chromosome-wide gene regulation. Chromosoma. 2011;120:213-25 pubmed publisher
    ..In this review, we discuss the origin and function of buffering and compensation using Drosophila as a model. ..
  10. Corona D, Clapier C, Becker P, Tamkun J. Modulation of ISWI function by site-specific histone acetylation. EMBO Rep. 2002;3:242-7 pubmed
    ..Increased expression of MOF, the histone acetyltransferase that acetylates H4K16, strongly enhances phenotypes resulting from the partial loss ..
  11. Nusinow D, Panning B. Recognition and modification of seX chromosomes. Curr Opin Genet Dev. 2005;15:206-13 pubmed
    ..Finally, evidence from mammals suggests that the chromatin modifications that mediate dosage compensation are very dynamic, because they are established, reversed and re-established early in development. ..
  12. Kind J, Akhtar A. Cotranscriptional recruitment of the dosage compensation complex to X-linked target genes. Genes Dev. 2007;21:2030-40 pubmed
    ..In order to study the mechanism of targeting, we have studied two X-chromosomal genes, mof and CG3016, using chromatin immunoprecipitation as well as immuno-FISH analysis on transgenic flies...
  13. Straub T, Becker P. Dosage compensation: the beginning and end of generalization. Nat Rev Genet. 2007;8:47-57 pubmed
  14. Prestel M, Feller C, Straub T, Mitlöhner H, Becker P. The activation potential of MOF is constrained for dosage compensation. Mol Cell. 2010;38:815-26 pubmed publisher
    The H4K16 acetyltransferase MOF plays a crucial role in dosage compensation in Drosophila but has additional, global functions...
  15. Bodai L, Pallos J, Thompson L, Marsh J. Pcaf modulates polyglutamine pathology in a Drosophila model of Huntington's disease. Neurodegener Dis. 2012;9:104-6 pubmed publisher
  16. Akhtar A, Becker P. Activation of transcription through histone H4 acetylation by MOF, an acetyltransferase essential for dosage compensation in Drosophila. Mol Cell. 2000;5:367-75 pubmed
    ..We demonstrate that MOF, a protein required for dosage compensation with significant sequence similarity to the MYST family of ..
  17. Vaquerizas J, Suyama R, Kind J, Miura K, Luscombe N, Akhtar A. Nuclear pore proteins nup153 and megator define transcriptionally active regions in the Drosophila genome. PLoS Genet. 2010;6:e1000846 pubmed publisher
    ..Therefore, we suggest that NAR-binding is used for chromosomal organization that enables gene expression control. ..
  18. Buscaino A, Kocher T, Kind J, Holz H, Taipale M, Wagner K, et al. MOF-regulated acetylation of MSL-3 in the Drosophila dosage compensation complex. Mol Cell. 2003;11:1265-77 pubmed
    ..We show that interaction of MOF with MSL-3 leads to specific acetylation of MSL-3 at a single lysine residue adjacent to one of its chromodomains...
  19. Akhtar A. Dosage compensation: an intertwined world of RNA and chromatin remodelling. Curr Opin Genet Dev. 2003;13:161-9 pubmed
    ..Furthermore, non-coding RNAs appear to play a dual role, being targeting modules as well as encoding for target sites for complex recognition. ..
  20. Regnard C, Straub T, Mitterweger A, Dahlsveen I, Fabian V, Becker P. Global analysis of the relationship between JIL-1 kinase and transcription. PLoS Genet. 2011;7:e1001327 pubmed publisher
    ..We hypothesise that one specific role of JIL-1 may be to reinforce, rather than to establish, the status of active chromatin through the phosphorylation of histone H3 at serine 10. ..
  21. Meller V, Rattner B. The roX genes encode redundant male-specific lethal transcripts required for targeting of the MSL complex. EMBO J. 2002;21:1084-91 pubmed
    ..Therefore, the roX1 and roX2 genes produce redundant, male-specific lethal transcripts required for targeting the MSL complex. ..
  22. Hilfiker A, Hilfiker Kleiner D, Pannuti A, Lucchesi J. mof, a putative acetyl transferase gene related to the Tip60 and MOZ human genes and to the SAS genes of yeast, is required for dosage compensation in Drosophila. EMBO J. 1997;16:2054-60 pubmed
    ..Experimental results and sequence analysis suggest that an additional gene, males-absent on the first (mof), encodes a putative acetyl transferase that plays a direct role in the specific histone acetylation associated ..
  23. Marin I, Siegal M, Baker B. The evolution of dosage-compensation mechanisms. Bioessays. 2000;22:1106-14 pubmed
    ..Some tantalizing similarities in the fly and mammalian mechanisms, however, remain to be explained. ..
  24. Meller V. Initiation of dosage compensation in Drosophila embryos depends on expression of the roX RNAs. Mech Dev. 2003;120:759-67 pubmed
    ..Either roX transcript can support X-localization of the complex, but localization is delayed in roX1 mutants until roX2 expression. These results support a model for the ordered assembly of the complex in embryos. ..
  25. Straub T, Dahlsveen I, Becker P. Dosage compensation in flies: mechanism, models, mystery. FEBS Lett. 2005;579:3258-63 pubmed
    ..We discuss the prevalent models in the context of recent experimental observations. ..
  26. Deng X, Meller V. roX RNAs are required for increased expression of X-linked genes in Drosophila melanogaster males. Genetics. 2006;174:1859-66 pubmed
    ..These results implicate failure to compensate X-linked genes, rather than inappropriate upregulation of autosomal genes at ectopic sites of MSL binding, as the primary cause of male lethality upon loss of roX RNAs. ..
  27. Nielsen P, Nietlispach D, Buscaino A, Warner R, Akhtar A, Murzin A, et al. Structure of the chromo barrel domain from the MOF acetyltransferase. J Biol Chem. 2005;280:32326-31 pubmed
    We report here the structure of the putative chromo domain from MOF, a member of the MYST family of histone acetyltransferases that acetylates histone H4 at Lys-16 and is part of the dosage compensation complex in Drosophila...
  28. Li F, Schiemann A, Scott M. Incorporation of the noncoding roX RNAs alters the chromatin-binding specificity of the Drosophila MSL1/MSL2 complex. Mol Cell Biol. 2008;28:1252-64 pubmed
    ..We suggest that incorporation of the roX RNAs into the MSL complex alters the binding specificity of the chromatin-binding module formed by the amino-terminal domains of MSL1 and MSL2. ..
  29. Schwaiger M, Stadler M, Bell O, Kohler H, Oakeley E, Schübeler D. Chromatin state marks cell-type- and gender-specific replication of the Drosophila genome. Genes Dev. 2009;23:589-601 pubmed publisher
    ..Together, our study reveals novel organizational principles of DNA replication of the Drosophila genome and suggests that H4K16ac is more closely correlated with replication timing than is transcription. ..
  30. Philip P, Pettersson F, Stenberg P. Sequence signatures involved in targeting the Male-Specific Lethal complex to X-chromosomal genes in Drosophila melanogaster. BMC Genomics. 2012;13:97 pubmed publisher
    ..High affinity sites seem to be responsible for initial recruitment of the complex to the X chromosome, but the targeting to and within individual genes is poorly understood...
  31. Ebert A, Schotta G, Lein S, Kubicek S, Krauss V, Jenuwein T, et al. Su(var) genes regulate the balance between euchromatin and heterochromatin in Drosophila. Genes Dev. 2004;18:2973-83 pubmed
  32. Akhtar A, Becker P. The histone H4 acetyltransferase MOF uses a C2HC zinc finger for substrate recognition. EMBO Rep. 2001;2:113-8 pubmed
    Site-specific acetylation of histone H4 by MOF is central to establishing the hyperactive male X chromosome in Drosophila...
  33. Yang X. The diverse superfamily of lysine acetyltransferases and their roles in leukemia and other diseases. Nucleic Acids Res. 2004;32:959-76 pubmed
    ..function as transcriptional co-activators, emerging evidence suggests that MYST proteins, such as Esa1, Sas2, MOF, TIP60, MOZ and MORF, have diverse roles in various nuclear processes...
  34. Sural T, Peng S, Li B, Workman J, Park P, Kuroda M. The MSL3 chromodomain directs a key targeting step for dosage compensation of the Drosophila melanogaster X chromosome. Nat Struct Mol Biol. 2008;15:1318-25 pubmed publisher
    ..Our results support a model in which activating complexes, similarly to their silencing counterparts, use the nucleosomal binding specificity of their respective chromodomains to spread from initiation sites to flanking chromatin. ..
  35. Conrad T, Cavalli F, Holz H, Hallacli E, Kind J, Ilik I, et al. The MOF chromobarrel domain controls genome-wide H4K16 acetylation and spreading of the MSL complex. Dev Cell. 2012;22:610-24 pubmed publisher
    The histone H4 lysine 16 (H4K16)-specific acetyltransferase MOF is part of two distinct complexes involved in X chromosome dosage compensation and autosomal transcription regulation...
  36. Johansson A, Stenberg P, Allgardsson A, Larsson J. POF regulates the expression of genes on the fourth chromosome in Drosophila melanogaster by binding to nascent RNA. Mol Cell Biol. 2012;32:2121-34 pubmed publisher
    ..These results provide useful insights into the means by which genes in heterochromatic regions can overcome the repressive influence of their hostile environment...
  37. Gladstein N, McKeon M, Horabin J. Requirement of male-specific dosage compensation in Drosophila females--implications of early X chromosome gene expression. PLoS Genet. 2010;6:e1001041 pubmed publisher
    ..Our data suggest that before the female state is set, the ground state is male and female X chromosome expression is elevated. Females thus utilize the male dosage compensation process to amplify the signal which determines their fate. ..
  38. Morra R, Smith E, Yokoyama R, Lucchesi J. The MLE subunit of the Drosophila MSL complex uses its ATPase activity for dosage compensation and its helicase activity for targeting. Mol Cell Biol. 2008;28:958-66 pubmed
    ..complex consists of a noncoding RNA and a core of five protein subunits that includes a histone acetyltransferase (MOF) and an ATP-dependent DEXH box RNA/DNA helicase (MLE)...
  39. Bell O, Wirbelauer C, Hild M, Scharf A, Schwaiger M, MacAlpine D, et al. Localized H3K36 methylation states define histone H4K16 acetylation during transcriptional elongation in Drosophila. EMBO J. 2007;26:4974-84 pubmed
    ..Thus di- and trimethylation of H3K36 have opposite effects on H4K16 acetylation, which we propose enable dynamic changes in chromatin compaction during transcript elongation. ..
  40. Ilik I, Quinn J, Georgiev P, Tavares Cadete F, Maticzka D, Toscano S, et al. Tandem stem-loops in roX RNAs act together to mediate X chromosome dosage compensation in Drosophila. Mol Cell. 2013;51:156-73 pubmed publisher
    ..We propose that repetitive structural motifs in lncRNAs could provide plasticity during multiprotein complex assemblies to ensure efficient targeting in cis or in trans along chromosomes...
  41. Weake V, Scott M. The non-dosage compensated Lsp1alpha gene of Drosophila melanogaster escapes acetylation by MOF in larval fat body nuclei, but is flanked by two dosage compensated genes. BMC Mol Biol. 2007;8:35 pubmed
    ..dosage compensation of most X-linked genes is mediated by the male-specific lethal (MSL) complex, which includes MOF. MOF acetylates histone H4 at lysine 16 (H4K16ac)...
  42. Buscaino A, Legube G, Akhtar A. X-chromosome targeting and dosage compensation are mediated by distinct domains in MSL-3. EMBO Rep. 2006;7:531-8 pubmed
    ..Our results demonstrate that MSL-3 localization to the male X chromosome and transcriptional upregulation of X-linked genes are two separable functions of the MSL-3 protein. ..
  43. Kind J, Vaquerizas J, Gebhardt P, Gentzel M, Luscombe N, Bertone P, et al. Genome-wide analysis reveals MOF as a key regulator of dosage compensation and gene expression in Drosophila. Cell. 2008;133:813-28 pubmed publisher
    ..Here we report that the histone H4 lysine 16 (H4K16) specific histone acetyltransferase MOF displays differential binding behavior depending on whether the target gene is located on the X chromosome versus ..
  44. Rattner B, Meller V. Drosophila male-specific lethal 2 protein controls sex-specific expression of the roX genes. Genetics. 2004;166:1825-32 pubmed
    ..We propose that this mechanism maintains a stable MSL/roX ratio that is favorable for localization of the complex to the X chromosome. ..
  45. Grimaud C, Becker P. The dosage compensation complex shapes the conformation of the X chromosome in Drosophila. Genes Dev. 2009;23:2490-5 pubmed publisher
    ..We propose that nuclear organization of HAS is central to the faithful distribution of the DCC along the X chromosome. ..
  46. Larschan E, Alekseyenko A, Gortchakov A, Peng S, Li B, Yang P, et al. MSL complex is attracted to genes marked by H3K36 trimethylation using a sequence-independent mechanism. Mol Cell. 2007;28:121-33 pubmed
    ..Our results support a model in which MSL complex uses high-affinity sites to initially recognize the X chromosome and then associates with many of its targets through sequence-independent features of transcribed genes. ..
  47. Spierer A, Begeot F, Spierer P, Delattre M. SU(VAR)3-7 links heterochromatin and dosage compensation in Drosophila. PLoS Genet. 2008;4:e1000066 pubmed publisher
    ..Taken together, these results bring to light a link between heterochromatin and dosage compensation...
  48. Menon D, Meller V. Imprinting of the Y chromosome influences dosage compensation in roX1 roX2 Drosophila melanogaster. Genetics. 2009;183:811-20 pubmed publisher
    ..We believe that the Y chromosome is likely to act through modulation of a process that is defective in roX1 roX2 mutants: X chromosome recognition or chromatin modification by the MSL complex...
  49. Scott M, Pan L, Cleland S, Knox A, Heinrich J. MSL1 plays a central role in assembly of the MSL complex, essential for dosage compensation in Drosophila. EMBO J. 2000;19:144-55 pubmed
    ..Here we show that one of these proteins, MSL1, interacts with three others, MSL2, MSL3 and MOF. The latter is a putative histone acetyl transferase...
  50. Akhtar A, Zink D, Becker P. Chromodomains are protein-RNA interaction modules. Nature. 2000;407:405-9 pubmed
    ..epigenetic process involving the specific acetylation of histone H4 at lysine 16 by the histone acetyltransferase MOF. Although MOF is expressed in both sexes, it only associates with the X chromosome in males...
  51. Grienenberger A, Miotto B, Sagnier T, Cavalli G, Schramke V, Geli V, et al. The MYST domain acetyltransferase Chameau functions in epigenetic mechanisms of transcriptional repression. Curr Biol. 2002;12:762-6 pubmed
  52. Fagegaltier D, Baker B. X chromosome sites autonomously recruit the dosage compensation complex in Drosophila males. PLoS Biol. 2004;2:e341 pubmed
    ..We propose a new model in which the distribution of compensasomes along the X chromosome is achieved according to the hierarchical affinities of individual binding sites. ..
  53. Roy S, Gilbert M, Hart C. Characterization of BEAF mutations isolated by homologous recombination in Drosophila. Genetics. 2007;176:801-13 pubmed
    ..Combined with the effects on male X polytene chromosomes, we conclude that BEAF function affects chromatin structure or dynamics. ..
  54. Li F, Parry D, Scott M. The amino-terminal region of Drosophila MSL1 contains basic, glycine-rich, and leucine zipper-like motifs that promote X chromosome binding, self-association, and MSL2 binding, respectively. Mol Cell Biol. 2005;25:8913-24 pubmed publisher
    ..We propose that the basic region may mediate DNA binding and that the glycine-rich region may promote the association of MSL complexes to closely adjacent sites on the X chromosome...
  55. Morales V, Regnard C, Izzo A, Vetter I, Becker P. The MRG domain mediates the functional integration of MSL3 into the dosage compensation complex. Mol Cell Biol. 2005;25:5947-54 pubmed
    ..interacts selectively with the X chromosome to restrict the transcription-activating histone H4 acetyltransferase MOF (males-absent-on-the-first) to that chromosome...
  56. Feller C, Prestel M, Hartmann H, Straub T, Söding J, Becker P. The MOF-containing NSL complex associates globally with housekeeping genes, but activates only a defined subset. Nucleic Acids Res. 2012;40:1509-22 pubmed publisher
    The MOF (males absent on the first)-containing NSL (non-specific lethal) complex binds to a subset of active promoters in Drosophila melanogaster and is thought to contribute to proper gene expression...
  57. Gilfillan G, Dahlsveen I, Becker P. Lifting a chromosome: dosage compensation in Drosophila melanogaster. FEBS Lett. 2004;567:8-14 pubmed
    ..Deciphering the principles of X chromosome recognition and the nature of the chromatin configuration, that allows fine-tuning of transcription, remain the most interesting challenges. ..
  58. Mendjan S, Akhtar A. The right dose for every sex. Chromosoma. 2007;116:95-106 pubmed
    ..Taken together, these findings suggest a role of the MSL complex in transcription elongation, RNA processing, and/or nuclear organization. ..
  59. Kageyama Y, Mengus G, Gilfillan G, Kennedy H, Stuckenholz C, Kelley R, et al. Association and spreading of the Drosophila dosage compensation complex from a discrete roX1 chromatin entry site. EMBO J. 2001;20:2236-45 pubmed
    ..When inserted on autosomes, this small roX1 segment is sufficient to produce an ectopic chromatin entry site that can nucleate binding and spreading of the MSL complex hundreds of kilobases into neighboring regions. ..
  60. Hamada F, Park P, Gordadze P, Kuroda M. Global regulation of X chromosomal genes by the MSL complex in Drosophila melanogaster. Genes Dev. 2005;19:2289-94 pubmed
    ..We found that expression of many genes from the X chromosome decreased, while expression from the autosomes was largely unchanged. We conclude that the primary role of the MSL complex is to up-regulate the male X chromosome. ..
  61. Miotto B, Sagnier T, Bérenger H, Bohmann D, Pradel J, Graba Y. Chameau HAT and DRpd3 HDAC function as antagonistic cofactors of JNK/AP-1-dependent transcription during Drosophila metamorphosis. Genes Dev. 2006;20:101-12 pubmed
    ..This study establishes a direct link among JNK signaling, DFos phosphorylation, chromatin modification, and AP-1-dependent transcription and its importance in a developing organism. ..
  62. Bouazoune K, Mitterweger A, Längst G, Imhof A, Akhtar A, Becker P, et al. The dMi-2 chromodomains are DNA binding modules important for ATP-dependent nucleosome mobilization. EMBO J. 2002;21:2430-40 pubmed
    ..Furthermore, we identify DNA binding as a novel chromodomain-associated activity. ..
  63. Gu W, Wei X, Pannuti A, Lucchesi J. Targeting the chromatin-remodeling MSL complex of Drosophila to its sites of action on the X chromosome requires both acetyl transferase and ATPase activities. EMBO J. 2000;19:5202-11 pubmed
    ..the role that the enzymatic activities of two complex components, the histone acetyltransferase activity of MOF and the ATPase activity of MLE, may have in the targeting and association of the complex with the X chromosome...
  64. Franke A, Baker B. Dosage compensation rox!. Curr Opin Cell Biol. 2000;12:351-4 pubmed
  65. Yokoyama R, Pannuti A, Ling H, Smith E, Lucchesi J. A plasmid model system shows that Drosophila dosage compensation depends on the global acetylation of histone H4 at lysine 16 and is not affected by depletion of common transcription elongation chromatin marks. Mol Cell Biol. 2007;27:7865-70 pubmed