Gene Symbol: msl-2
Description: male-specific lethal 2
Alias: CG3241, Dmel\CG3241, MSL, MSL-2, MSL2, Msl 1-3, Msl-2, Msl2, km(2)A, kmA, msl, msl2, male-specific lethal 2, CG3241-PA, CG3241-PB, killer of males, killer-of-males-A, male specifc lethal, male specific lethal 2, male specific-lethal 2, male-specific-lethal 2, male-specific-lethal-2, msl-2-PA, msl-2-PB
Species: fruit fly

Top Publications

  1. Gorman M, Franke A, Baker B. Molecular characterization of the male-specific lethal-3 gene and investigations of the regulation of dosage compensation in Drosophila. Development. 1995;121:463-75 pubmed
    ..This hypertranscription requires four autosomal male-specific lethal (msl) genes and is negatively regulated by the Sxl gene in females...
  2. Copps K, Richman R, Lyman L, Chang K, Rampersad Ammons J, Kuroda M. Complex formation by the Drosophila MSL proteins: role of the MSL2 RING finger in protein complex assembly. EMBO J. 1998;17:5409-17 pubmed publisher
    ..We found that the MSL1, MSL2 and MSL3 proteins are associated in immunoprecipitations, chromatographic steps and in the yeast two-hybrid system, ..
  3. 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...
  4. Dahlsveen I, Gilfillan G, Shelest V, Lamm R, Becker P. Targeting determinants of dosage compensation in Drosophila. PLoS Genet. 2006;2:e5 pubmed
    ..Our bioinformatics analysis suggests that DCC binding sites may be composed of variable combinations of degenerate motifs. ..
  5. Maenner S, Müller M, Frohlich J, Langer D, Becker P. ATP-dependent roX RNA remodeling by the helicase maleless enables specific association of MSL proteins. Mol Cell. 2013;51:174-84 pubmed publisher
    ..This RNA remodeling enhanced specific ATP-dependent association of MSL2, the core subunit of the MSL-DCC, providing a link between roX and MSL subunits...
  6. Lyman L, Copps K, Rastelli L, Kelley R, Kuroda M. Drosophila male-specific lethal-2 protein: structure/function analysis and dependence on MSL-1 for chromosome association. Genetics. 1997;147:1743-53 pubmed
    b>MSL-2 is required for the male-specific assembly of a dosage compensation regulatory complex on the X chromosome of Drosophila melanogaster...
  7. Richter L, Bone J, Kuroda M. RNA-dependent association of the Drosophila maleless protein with the male X chromosome. Genes Cells. 1996;1:325-36 pubmed
    ..The potential relationship between interaction with RNA and transcriptional control of the X chromosome suggests that the mechanism of dosage compensation is distinct from classical models for transcriptional activation. ..
  8. Straub T, Becker P. Dosage compensation: the beginning and end of generalization. Nat Rev Genet. 2007;8:47-57 pubmed
  9. Merendino L, Guth S, Bilbao D, Martinez C, Valcarcel J. Inhibition of msl-2 splicing by Sex-lethal reveals interaction between U2AF35 and the 3' splice site AG. Nature. 1999;402:838-41 pubmed
    ..controls dosage compensation in Drosophila by inhibiting the splicing and translation of male-specific-lethal-2 (msl-2) transcripts...

More Information


  1. Meller V, Gordadze P, Park Y, Chu X, Stuckenholz C, Kelley R, et al. Ordered assembly of roX RNAs into MSL complexes on the dosage-compensated X chromosome in Drosophila. Curr Biol. 2000;10:136-43 pubmed transcriptionally upregulated to achieve dosage compensation, in a process that depends on association of the MSL proteins with the X chromosome. A role for non-coding RNAs has been suggested in recent studies...
  2. 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. ..
  3. Franke A, Dernburg A, Bashaw G, Baker B. Evidence that MSL-mediated dosage compensation in Drosophila begins at blastoderm. Development. 1996;122:2751-60 pubmed
    ..This process, dosage compensation, is controlled by a set of male-specific lethal (msl) genes, that appear to act at the level of chromatin structure...
  4. 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
    ..The core DCC subunits MSL1-MSL2 are responsible for this male-specific organization...
  5. Prabhakaran M, Kelley R. A new strategy for isolating genes controlling dosage compensation in Drosophila using a simple epigenetic mosaic eye phenotype. BMC Biol. 2010;8:80 pubmed publisher
    The Drosophila Male Specific Lethal (MSL) complex contains chromatin modifying enzymes and non-coding roX RNA. It paints the male X at hundreds of bands where it acetylates histone H4 at lysine 16...
  6. Fauth T, Müller Planitz F, König C, Straub T, Becker P. The DNA binding CXC domain of MSL2 is required for faithful targeting the Dosage Compensation Complex to the X chromosome. Nucleic Acids Res. 2010;38:3209-21 pubmed publisher
    ..Early genetic studies suggested that MSL1 and MSL2 serve to recognize high-affinity sites (HAS) in vivo, but a direct interaction of these DCC subunits with DNA has ..
  7. Gebauer F, Grskovic M, Hentze M. Drosophila sex-lethal inhibits the stable association of the 40S ribosomal subunit with msl-2 mRNA. Mol Cell. 2003;11:1397-404 pubmed
    The inhibition of male-specific lethal-2 (msl-2) mRNA translation in female flies is essential for X chromosome dosage compensation in Drosophila melanogaster...
  8. Kelley R, Solovyeva I, Lyman L, Richman R, Solovyev V, Kuroda M. Expression of msl-2 causes assembly of dosage compensation regulators on the X chromosomes and female lethality in Drosophila. Cell. 1995;81:867-77 pubmed
    Male-specific lethal-2 (msl-2) is a RING finger protein that is required for X chromosome dosage compensation in Drosophila males...
  9. Kelley R, Kuroda M. The Drosophila roX1 RNA gene can overcome silent chromatin by recruiting the male-specific lethal dosage compensation complex. Genetics. 2003;164:565-74 pubmed
    The Drosophila MSL complex consists of at least six proteins and two noncoding roX RNAs that mediate dosage compensation. It acts to remodel the male's X chromatin by covalently modifying the amino terminal tails of histones...
  10. Kind J, Akhtar A. Cotranscriptional recruitment of the dosage compensation complex to X-linked target genes. Genes Dev. 2007;21:2030-40 pubmed
    ..In Drosophila this is achieved by targeting the dosage compensation complex or the male-specific lethal (MSL) complex to the male X chromosome...
  11. Mar n I, Franke A, Bashaw G, Baker B. The dosage compensation system of Drosophila is co-opted by newly evolved X chromosomes. Nature. 1996;383:160-3 pubmed publisher
    ..melanogaster, dosage compensation is mediated by the binding of the products of the male-specific lethal (msl) genes to the single male X chromosome...
  12. 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 accomplished by hypertranscription of the single male X chromosome mediated by the male-specific lethal (MSL) complex...
  13. 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. ..
  14. Kelley R, Kuroda M. Equality for X chromosomes. Science. 1995;270:1607-10 pubmed
    ..A common theme in the Drosophila melanogaster and Caenorhabditis elegans systems is that a subtle alteration of chromatin structure may impose this modest, but vital adjustment of the X chromosome transcription level. ..
  15. Lim C, Kelley R. Autoregulation of the Drosophila Noncoding roX1 RNA Gene. PLoS Genet. 2012;8:e1002564 pubmed publisher
    ..Ectopic DC can be induced in wild-type (roX1(+) roX2(+)) females if we provide a heterologous source of MSL2. However, in the absence of roX2, we found that roX1 expression failed to come on reliably...
  16. 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. ..
  17. Gorchakov A, Alekseyenko A, Kharchenko P, Park P, Kuroda M. Long-range spreading of dosage compensation in Drosophila captures transcribed autosomal genes inserted on X. Genes Dev. 2009;23:2266-71 pubmed publisher
    Dosage compensation in Drosophila melanogaster males is achieved via targeting of male-specific lethal (MSL) complex to X-linked genes...
  18. Marín I. Evolution of chromatin-remodeling complexes: comparative genomics reveals the ancient origin of "novel" compensasome genes. J Mol Evol. 2003;56:527-39 pubmed
    ..Recent work showed that three of the Drosophila msls (msl-3, mof, and mle) have an ancient origin...
  19. Lundberg L, Kim M, Johansson A, Faucillion M, Josupeit R, Larsson J. Targeting of Painting of fourth to roX1 and roX2 proximal sites suggests evolutionary links between dosage compensation and the regulation of the fourth chromosome in Drosophila melanogaster. G3 (Bethesda). 2013;3:1325-34 pubmed publisher
    ..The male-specific lethal (MSL) complex supports dosage compensation by stimulating gene expression from the male X-chromosome, and the protein ..
  20. Alekseyenko A, Ellison C, Gorchakov A, Zhou Q, Kaiser V, Toda N, et al. Conservation and de novo acquisition of dosage compensation on newly evolved sex chromosomes in Drosophila. Genes Dev. 2013;27:853-8 pubmed publisher
    ..In Drosophila melanogaster, the MSL complex increases male X transcription approximately twofold...
  21. 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. ..
  22. 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
    The male-specific lethal (MSL) complex upregulates the single male X chromosome to achieve dosage compensation in Drosophila melanogaster...
  23. Oh H, Bone J, Kuroda M. Multiple classes of MSL binding sites target dosage compensation to the X chromosome of Drosophila. Curr Biol. 2004;14:481-7 pubmed
    b>MSL complexes bind hundreds of sites along the single male X chromosome to achieve dosage compensation in Drosophila...
  24. Deshpande G, Calhoun G, Schedl P. The N-terminal domain of Sxl protein disrupts Sxl autoregulation in females and promotes female-specific splicing of tra in males. Development. 1999;126:2841-53 pubmed
    ..This novel activity is discussed in light of the blockage model for the tra splicing regulation. ..
  25. Meller V, Wu K, Roman G, Kuroda M, Davis R. roX1 RNA paints the X chromosome of male Drosophila and is regulated by the dosage compensation system. Cell. 1997;88:445-57 pubmed
    ..The novel regulation and subnuclear localization of roX1 RNAs makes them candidates for an RNA component of the dosage compensation machinery. ..
  26. Gebauer F, Corona D, Preiss T, Becker P, Hentze M. Translational control of dosage compensation in Drosophila by Sex-lethal: cooperative silencing via the 5' and 3' UTRs of msl-2 mRNA is independent of the poly(A) tail. EMBO J. 1999;18:6146-54 pubmed
    Translational repression of male-specific-lethal 2 (msl-2) mRNA by Sex-lethal (SXL) controls dosage compensation in Drosophila...
  27. 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
    In Drosophila, dosage compensation is controlled by the male-specific lethal (MSL) complex consisting of at least five proteins and two noncoding RNAs, roX1 and roX2...
  28. 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. ..
  29. Bachtrog D. Positive selection at the binding sites of the male-specific lethal complex involved in dosage compensation in Drosophila. Genetics. 2008;180:1123-9 pubmed publisher
    ..In Drosophila, the male-specific lethal (MSL) complex binds to hundreds of sites along the male X chromosome and mediates twofold hypertranscription of the ..
  30. 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
    ..Genetic and biochemical studies have demonstrated that male-specific lethal (MSL) proteins together with roX RNAs regulate this process...
  31. Yanowitz J, Deshpande G, Calhoun G, Schedl P. An N-terminal truncation uncouples the sex-transforming and dosage compensation functions of sex-lethal. Mol Cell Biol. 1999;19:3018-28 pubmed
    ..This negative autoregulatory activity may be a homeostasis mechanism. ..
  32. Spierer A, Seum C, Delattre M, Spierer P. Loss of the modifiers of variegation Su(var)3-7 or HP1 impacts male X polytene chromosome morphology and dosage compensation. J Cell Sci. 2005;118:5047-57 pubmed
    ..This observation reveals a new and intriguing genetic interaction between epigenetic silencing and compensation of dose. ..
  33. Greenberg A, Yanowitz J, Schedl P. The Drosophila GAGA factor is required for dosage compensation in males and for the formation of the male-specific-lethal complex chromatin entry site at 12DE. Genetics. 2004;166:279-89 pubmed
    ..Here, we report that mutations in Trithorax-like (Trl) lead to the loss of a single chromatin entry site on the X, male lethality, and mislocalization of dosage compensation complexes. ..
  34. 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. ..
  35. 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
  36. Alekseyenko A, Larschan E, Lai W, Park P, Kuroda M. High-resolution ChIP-chip analysis reveals that the Drosophila MSL complex selectively identifies active genes on the male X chromosome. Genes Dev. 2006;20:848-57 pubmed
    X-chromosome dosage compensation in Drosophila requires the male-specific lethal (MSL) complex, which up-regulates gene expression from the single male X chromosome...
  37. Sass G, Pannuti A, Lucchesi J. Male-specific lethal complex of Drosophila targets activated regions of the X chromosome for chromatin remodeling. Proc Natl Acad Sci U S A. 2003;100:8287-91 pubmed
    The male-specific lethal (MSL) complex of Drosophila is responsible for the presence of a monoacetylated isoform of histone H4 (H4Ac16), found exclusively on the X chromosome of males...
  38. Baker B, Gorman M, Mar n I. Dosage compensation in Drosophila. Annu Rev Genet. 1994;28:491-521 pubmed publisher
  39. Lee C, Reichman T, Baik T, Mathews M. MLE functions as a transcriptional regulator of the roX2 gene. J Biol Chem. 2004;279:47740-5 pubmed
    ..In Drosophila, two non-coding RNA, roX1 and roX2, and at least six protein regulators, MSL-1, MSL-2, MSL-3, MLE, MOF, and JIL-1, have been identified as essential for dosage compensation...
  40. 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. ..
  41. Deng X, Rattner B, Souter S, Meller V. The severity of roX1 mutations is predicted by MSL localization on the X chromosome. Mech Dev. 2005;122:1094-105 pubmed
    ..This ability predicts the level of male survival that each allele supports. This points to a peripheral or transient role for roX in the RNA and protein complex that binds to and regulates the X chromosome. ..
  42. McDowell K, Hilfiker A, Lucchesi J. Dosage compensation in Drosophila: the X chromosome binding of MSL-1 and MSL-2 in female embryos is prevented by the early expression of the Sxl gene. Mech Dev. 1996;57:113-9 pubmed
    In wildtype males, binding of the MSL-1 gene product to the X chromosome is first seen at the cellular blastoderm stage (stage 5)...
  43. 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
    In Drosophila, dosage compensation is controlled by the male-specific lethal (MSL) complex consisting of MSL proteins and roX RNAs...
  44. Duncan K, Grskovic M, Strein C, Beckmann K, Niggeweg R, Abaza I, et al. Sex-lethal imparts a sex-specific function to UNR by recruiting it to the msl-2 mRNA 3' UTR: translational repression for dosage compensation. Genes Dev. 2006;20:368-79 pubmed
    b>MSL-2 (male-specific lethal 2) is the limiting component of the Drosophila dosage compensation complex (DCC) that specifically increases transcription from the male X chromosome...
  45. Jin Y, Wang Y, Johansen J, Johansen K. JIL-1, a chromosomal kinase implicated in regulation of chromatin structure, associates with the male specific lethal (MSL) dosage compensation complex. J Cell Biol. 2000;149:1005-10 pubmed
    ..Furthermore, ectopic expression of the MSL complex directed by MSL2 in females causes a concomitant upregulation of JIL-1 to the female X that is abolished in msl mutants unable to ..
  46. 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
    ..To distinguish between the two models, we used RNA interference to deplete Male-Specific Lethal (MSL) complexes from male-like tissue culture cells...
  47. Meiklejohn C, Landeen E, Cook J, Kingan S, Presgraves D. Sex chromosome-specific regulation in the Drosophila male germline but little evidence for chromosomal dosage compensation or meiotic inactivation. PLoS Biol. 2011;9:e1001126 pubmed publisher
    ..These findings help to resolve several previously conflicting reports and have implications for patterns of genome evolution and speciation in Drosophila...
  48. Bashaw G, Baker B. The msl-2 dosage compensation gene of Drosophila encodes a putative DNA-binding protein whose expression is sex specifically regulated by Sex-lethal. Development. 1995;121:3245-58 pubmed
    ..We have cloned the msl-2 gene and shown that MSL-2 protein is co-localized with the other three MSL proteins at hundreds of sites along ..
  49. 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. ..
  50. Grskovic M, Hentze M, Gebauer F. A co-repressor assembly nucleated by Sex-lethal in the 3'UTR mediates translational control of Drosophila msl-2 mRNA. EMBO J. 2003;22:5571-81 pubmed
    Drosophila Sex-lethal (dSXL)-mediated translational repression of male-specific lethal 2 (msl-2) mRNA is essential for X-chromosome dosage compensation...
  51. Amrein H, Axel R. Genes expressed in neurons of adult male Drosophila. Cell. 1997;88:459-69 pubmed
    ..Our data suggest that the multimeric complex of dosage compensation proteins may operate in different ways on different sets of X-linked genes. ..
  52. 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 multisubunit MSL dosage compensation complex binds to hundreds of sites along the Drosophila single male X chromosome, mediating its hypertranscription. The male X chromosome is also coated with noncoding roX RNAs...
  53. Bone J, Lavender J, Richman R, Palmer M, Turner B, Kuroda M. Acetylated histone H4 on the male X chromosome is associated with dosage compensation in Drosophila. Genes Dev. 1994;8:96-104 pubmed
    ..elevated expression requires the function of at least four loci, known collectively as the male-specific lethal (msl) genes...
  54. 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
    In Drosophila, X chromosome dosage compensation requires the male-specific lethal (MSL) complex, which associates with actively transcribed genes on the single male X chromosome to upregulate transcription approximately 2-fold...
  55. Furuhashi H, Nakajima M, Hirose S. DNA supercoiling factor contributes to dosage compensation in Drosophila. Development. 2006;133:4475-83 pubmed
    ..SCF functionally interacts with components of the MSL complex, which are required for dosage compensation via hypertranscription of the male X chromosome...
  56. 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
    ..X chromosome binding also requires a conserved leucine zipper-like motif that binds to MSL2. A glycine-rich motif between the basic and leucine-zipper-like motifs mediates MSL1 self-association in vitro and ..
  57. Rodriguez M, Vermaak D, Bayes J, Malik H. Species-specific positive selection of the male-specific lethal complex that participates in dosage compensation in Drosophila. Proc Natl Acad Sci U S A. 2007;104:15412-7 pubmed
    ..In particular, the MSL1 and MSL2 proteins have undergone dramatic positive selection in D...
  58. Belote J, Lucchesi J. Male-specific lethal mutations of Drosophila melanogaster. Genetics. 1980;96:165-86 pubmed
    ..mle (Fukunaga, Tanaka and Oishi 1975), while the other three mutants identify two new loci: male-specific lethal-1 (msl-1) (two alleles) at map position 2-53.3 and male-specific lethal-2 (msl-2) at 2-9...
  59. Conrad T, Cavalli F, Vaquerizas J, Luscombe N, Akhtar A. Drosophila dosage compensation involves enhanced Pol II recruitment to male X-linked promoters. Science. 2012;337:742-6 pubmed publisher
    Through hyperacetylation of histone H4 lysine 16 (H4K16), the male-specific lethal (MSL) complex in Drosophila approximately doubles transcription from the single male X chromosome in order to match X-linked expression in females and ..
  60. Kelley R, Wang J, Bell L, Kuroda M. Sex lethal controls dosage compensation in Drosophila by a non-splicing mechanism. Nature. 1997;387:195-9 pubmed
    ..Although msl2 RNA is alternatively spliced in males and females, this does not alter its coding potential and splicing is not ..
  61. 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. ..
  62. Cline T, Meyer B. Vive la différence: males vs females in flies vs worms. Annu Rev Genet. 1996;30:637-702 pubmed
    ..Although no overlap has been found among the molecules used by flies and worms to achieve sex determination, striking similarities have been found in the genetic strategies used by these two species to differentiate their sexes. ..
  63. Oliver B, Pauli D. Suppression of distinct ovo phenotypes in the Drosophila female germline by maleless- and Sex-lethal. Dev Genet. 1998;23:335-46 pubmed
    ..Thus, ovo+ is required for at least two distinct functions, one involving mle+, and one mediated by Sxl+ gene products. The existence of ovo+ functions independent of mle+ and Sxl+ is likely. ..
  64. Palmer M, Richman R, Richter L, Kuroda M. Sex-specific regulation of the male-specific lethal-1 dosage compensation gene in Drosophila. Genes Dev. 1994;8:698-706 pubmed
    ..b>msl-1 is one of four genes that are essential for dosage compensation in males, and MSL-1 protein is associated ..
  65. Beckmann K, Grskovic M, Gebauer F, Hentze M. A dual inhibitory mechanism restricts msl-2 mRNA translation for dosage compensation in Drosophila. Cell. 2005;122:529-40 pubmed
    Drosophila MSL-2 is the limiting component of the dosage compensation complex...
  66. Gilfillan G, König C, Dahlsveen I, Prakoura N, Straub T, Lamm R, et al. Cumulative contributions of weak DNA determinants to targeting the Drosophila dosage compensation complex. Nucleic Acids Res. 2007;35:3561-72 pubmed
    ..a 'one-hybrid' assay, we identified minimal DNA elements that direct the interaction of the key DCC subunit, MSL2, in cells. Strikingly, several such novel MSL2 recruitment modules have very different DNA sequences...
  67. Larschan E, Bishop E, Kharchenko P, Core L, Lis J, Park P, et al. X chromosome dosage compensation via enhanced transcriptional elongation in Drosophila. Nature. 2011;471:115-8 pubmed publisher
    ..b>MSL (Male-specific lethal) complex increases transcription on the single X chromosome of Drosophila males to equalize ..
  68. Hilfiker A, Amrein H, Dübendorfer A, Schneiter R, Nothiger R. The gene virilizer is required for female-specific splicing controlled by Sxl, the master gene for sexual development in Drosophila. Development. 1995;121:4017-26 pubmed
    ..This female-specific lethality can be rescued by constitutive expression of Sxl or by mutations in msl (male-specific lethal) genes. Rescued animals develop as strongly masculinized intersexes or pseudomales...
  69. Rogers D, Carr M, Pomiankowski A. Male genes: X-pelled or X-cluded?. Bioessays. 2003;25:739-41 pubmed
    ..A number of evolutionary hypotheses for the expulsion or exclusion of male-biased genes from the X chromosome have been suggested. None is entirely consistent with the available evidence. ..
  70. 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
    ..The MSL2 and MSL1 proteins form a complex and are essential for X chromosome binding...
  71. 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
    The male-specific lethal (MSL) complex, which includes two noncoding RNA on X (roX)1 and roX2 RNAs, induces histone H4-Lys16 acetylation for twofold hypertranscription of the male X chromosome in Drosophila melanogaster...
  72. 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. ..
  73. Vicoso B, Bachtrog D. Reversal of an ancient sex chromosome to an autosome in Drosophila. Nature. 2013;499:332-5 pubmed publisher
  74. Belote J, Lucchesi J. Control of X chromosome transcription by the maleless gene in Drosophila. Nature. 1980;285:573-5 pubmed
    ..Autoradiographic monitoring of RNA synthesis in larval polytene chromosomes of males homozygous for one of these mutations, mlets, reveals a significant reduction in the rate of X chromosome transcription. ..
  75. Corona D, Clapier C, Becker P, Tamkun J. Modulation of ISWI function by site-specific histone acetylation. EMBO Rep. 2002;3:242-7 pubmed
    ..These findings suggest that H4K16 acetylation directly counteracts chromatin compaction mediated by the ISWI ATPase. ..
  76. Bashaw G, Baker B. Dosage compensation and chromatin structure in Drosophila. Curr Opin Genet Dev. 1996;6:496-501 pubmed
    The past year has brought significant advances in our understanding of the male-specific lethal (msl genes and dosage compensation in Drosophila...
  77. Medenbach J, Seiler M, Hentze M. Translational control via protein-regulated upstream open reading frames. Cell. 2011;145:902-13 pubmed publisher
    Analysis of the regulation of msl-2 mRNA by Sex lethal (SXL), which is critical for dosage compensation in Drosophila, has uncovered a mode of translational control based on common 5' untranslated region elements, upstream open reading ..
  78. Fagegaltier D, Baker B. X chromosome sites autonomously recruit the dosage compensation complex in Drosophila males. PLoS Biol. 2004;2:e341 pubmed
    It has been proposed that dosage compensation in Drosophila males occurs by binding of two core proteins, MSL-1 and MSL-2, to a set of 35-40 X chromosome "entry sites" that serve to nucleate mature complexes, termed ..
  79. 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
    ..followed by deep sequencing, we show that two enzymes in the Male-Specific Lethal complex, MLE RNA helicase and MSL2 ubiquitin ligase, bind evolutionarily conserved domains containing tandem stem-loops in roX1 and roX2 RNAs in vivo...
  80. Zhou S, Yang Y, Scott M, Pannuti A, Fehr K, Eisen A, et al. Male-specific lethal 2, a dosage compensation gene of Drosophila, undergoes sex-specific regulation and encodes a protein with a RING finger and a metallothionein-like cysteine cluster. EMBO J. 1995;14:2884-95 pubmed
    ..Three of these genes, maleless (mle), male-specific lethal 1 (msl-1) and male-specific lethal 3 (msl-3), have been cloned and their products have been shown to interact and to bind ..
  81. Gorman M, Kuroda M, Baker B. Regulation of the sex-specific binding of the maleless dosage compensation protein to the male X chromosome in Drosophila. Cell. 1993;72:39-49 pubmed
    ..Additionally, we have found that mle X chromosome binding requires wild-type msl1, msl2, and msl3 functions...
  82. Rastelli L, Richman R, Kuroda M. The dosage compensation regulators MLE, MSL-1 and MSL-2 are interdependent since early embryogenesis in Drosophila. Mech Dev. 1995;53:223-33 pubmed
    We have analyzed the expression pattern and localization of MLE, MLS-1, MSL-2, and histone H4Ac16 during embryogenesis to determine when msl-dependent dosage compensation begins...
  83. Alekseyenko A, Peng S, Larschan E, Gorchakov A, Lee O, Kharchenko P, et al. A sequence motif within chromatin entry sites directs MSL establishment on the Drosophila X chromosome. Cell. 2008;134:599-609 pubmed publisher
    The Drosophila MSL complex associates with active genes specifically on the male X chromosome to acetylate histone H4 at lysine 16 and increase expression approximately 2-fold...
  84. Abaza I, Coll O, Patalano S, Gebauer F. Drosophila UNR is required for translational repression of male-specific lethal 2 mRNA during regulation of X-chromosome dosage compensation. Genes Dev. 2006;20:380-9 pubmed
    The inhibition of male-specific lethal 2 (msl-2) mRNA translation by the RNA-binding protein sex-lethal (SXL) is an essential regulatory step for X-chromosome dosage compensation in Drosophila melanogaster...
  85. 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...