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Gene Dmel\msl-1

General Information
Symbol	Dmel\msl-1	Species	D. melanogaster
Name	male-specific lethal 1	Annotation symbol	CG10385
Feature type	protein_coding_gene	FlyBase ID	FBgn0005617
Created / Updated	2003-12-01/2003-12-01
Genomic Location
Chromosome (arm)	2L	Recombination map	2-20
Cytogenetic map	36F11-37A1	Sequence location	2L:18,683,720..18,688,796 [-]
Map ( GBrowse )	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene male-specific lethal 1 is referred to in FlyBase by the symbol msl-1 (CG10385, FBgn0005617). It has the cytological map location 36F11-37A1. Its sequence location is 2L:18683720..18688796. Its molecular function is described as: chromatin binding; DNA binding; protein binding. It is involved in the biological processes: dosage compensation; dosage compensation, by hyperactivation of X chromosome; dosage compensation complex assembly during dosage compensation by hyperactivation of X chromosome. 37 alleles are reported. No phenotypic data is available. It has one annotated transcript and one annotated polypeptide.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	kmB Homozygous males die during embryogenesis. msl-1: male-specific lethal Homozygous male embryos hatch but die as much as twelve days later in larval or prepupal stages; females and heterozygous males survive; phenotype slightly more severe in sons of homozygous than of heterozygous mothers. Viability of two-X individuals that develop as phenotypic males (tra2) or intersexes (dsx) is unaffected by msl-1, indicating that the one-X condition is required for msl-1 lethality. No interaction with mle or msl-2 (Belote, 1983, Genetics 105: 881-96). Females heterozygous for Sxlf1 and homozygous for msl-1 show signs of intersexual development [Skripsky and Lucchesi, 1982, Dev. Biol. 94: 153-64 (fig.)]. Pole cells from msl-1 male embryos capable of undergoing normal spermatogenesis when transplanted into wild-type hosts (Bachiller and Sanchez, 1986, Dev. Biol. 118: 379-84). Concluded to be defective in dosage compensation in males based on decreased levels of X-linked-enzyme activities (G6PD, 6GPD, FUM) but not autosomally encoded enzymes (ADH, AO, GPDH, IDH) in homozygous msl-11 and msl-12 male larvae when compared with non-msl-1 controls (Belote and Lucchesi, 1980, Nature 285: 573-75).
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 36F11-37A1   Limits computationally determined from genome sequence between P{EP}CG10413EP2164 and P{lacW}l(2)37Dbk16106&P{lacW}Catsupk05424
Experimentally Determined Cytological Location
Cytogenetic map Notes References 36F7-37A1 (determined by in situ hybridisation)   (Palmer et al., 1993)
Experimentally Determined Recombination Data
Location	2-20 (Pierre, 1972)
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References In direction of increasing cytology: CG31751+ tos- CG31751+ msl-1- (Digilio et al., 1996) Gene Order (overall orientation not stated) References
Gene Model & Products
Please see the GBrowse view of Dmel\msl-1 for information on other features
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) msl-1-RA FBtr0081130  NM_057548   5017   1039
Additional Transcript Data & Comments
Reported size (kB)	4.8, 4.4, 3.6 (northern blot) (Palmer et al., 1993)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein msl-1-PA   FBpp0080674   117.5   1039   5.86     NP_476896   AAF53689
Additional Polypeptide Data & Comments
Reported size (kD)	955 (aa) (Palmer et al., 1993) 170 (kD observed) (Palmer et al., 1994) 1039 (aa) (Kelley et al., 1995)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0005617
Crossreferences
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202011   AA202517   AA264385   AA264645   AA438808   AA696398   AA802707   AA803098   AA949118   AA949166   AA950085   AA950789   AA950803   AC007137   AE003660 AAF53689   AI515054   BG638513   BG639194   BI165044   BI165789   BI212207   BI212848   BI239130   BI355121   BI365580   BI367865   BT023828   CG10385   CL526109   CO188025   CO282844   CO286579   CO288651   CO288874   CO290559   CO290803   CO304429   CO311309   CO321449   CO322839   EC056831   EC063062   EC064401   EC065338   EC068417   EC068823   EC069206   EC070818   EC087498   EC195567   EC196229   EC202471   EC202478   EC207882   EC208277   EC211791   EC219049   EC240504   EC241938   EC242288   EC245486   EC250987   L42514 AAA98918   LD27954   LD46729
UniProtKB/Swiss-Prot	P50535
UniProtKB/TrEMBL
Maps to	FBcl0297581 (FlyBase, 1992-)
Does NOT map to
Identified with	RE18093 (BDGP Project Members, 2000-)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References rescue fragment msl-1[+t15] 2L:18,679,564..18,695,217 linked_to=SalI-SalI_rfrag evidence=experimental (Palmer et al., 1993) rescue fragment msl-1[+tFL] 2L:18,683,246..18,691,225 comment=Approximate location of rescue fragment based on author report that the fragment contains 2.8kb of sequence upstream of the translation start and 2kb of 3'UTR. evidence=experimental (Chang and Kuroda, 1998) rescue fragment msl-1[+tT3UTR] 2L:18,684,871..18,691,225 comment=Approximate location of rescue fragment based on author report that the fragment contains 2.8kb of sequence upstream of the translation start and 99bp of sequence after the first polyA site. evidence=experimental (Chang and Kuroda, 1998) sequence variant msl-1[190-5] 2L:18,685,550..18,686,515 linked_to=BamHI-BamHI_rfrag comment=A 6 kb insertion falls within the BamHI fragment. evidence=experimental (Palmer et al., 1993)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 8014
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  embryonic stage-adult stage   (Palmer et al., 1993)  adult stage | female   (Palmer et al., 1993)  adult stage | male   (Palmer et al., 1993)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  adult stage | male   (Palmer et al., 1994)  larval stage | male   (Palmer et al., 1994)  larval stage | female   (Gorman et al., 1995)  embryonic stage   (Palmer et al., 1994)
Marker for
Subcellular Localization
CV Term	chromosome (Bone et al., 1994)
Microarray Data
	Developmental timecourse, Costello et al., 2008 (Original data from Arbeitman et al., 2002) AA949166 AI515054
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0005617 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0005617 FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for msl-1 Stage(s) 4-6 Stage(s) 7-8 Stage(s) 9-10 Stage(s) 11-12 Stage(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele viable msl-1EY09390 viable | female msl-1Hsp83.PC lethal | male limited msl-1C.hs.T:Zzzz\FLAG msl-1FC.hs.T:Zzzz\FLAG msl-1FN.hs.T:Zzzz\FLAG msl-1Δ84.T:Zzzz\FLAG msl-1Δ84C.T:Zzzz\FLAG lethal | male msl-11 msl-12 lethal | male | recessive msl-1190-5 msl-1Z2681 msl-1Z5150 msl-1Z5823 msl-1kmB msl-1γ216 msl-1γ222 msl-1γ251 msl-1γ269 msl-1γ289 lethal | larval stage | male msl-1γ216 msl-1γ269 semi-lethal | male limited msl-1hs.T:Zzzz\FLAG msl-1Δ84.hs msl-1Δ84C.hs Other Phenotypes Allele wild-type msl-1+t15 msl-1+t5.9 msl-1+tFL msl-1+tT3UTR msl-1Hsp83.PC msl-1MPU Sterility Allele fertile msl-1EY09390 Phenotype manifest in Allele
Classical Alleles ( 16 )
For All Classical Alleles Show
Allele of msl-1 Class Mutagen Stocks Known lesion msl-1190-5 PM hybrid dysgenesis 0 Yes msl-11 amorph ethyl methanesulfonate 0 -- msl-12 ethyl methanesulfonate 0 -- msl-1EY09390 P-element activity 1 -- msl-1L183 0 Yes msl-1L60 γ ray 0 Yes msl-1Z2681 ethyl methanesulfonate 0 -- msl-1Z5150 ethyl methanesulfonate 0 -- msl-1Z5823 ethyl methanesulfonate 0 -- msl-1kmB ethyl methanesulfonate 0 -- msl-1unspecified 0 -- msl-1γ216 γ ray 2 Yes msl-1γ222 γ ray 0 -- msl-1γ251 γ ray 0 -- msl-1γ269 γ ray 0 Yes msl-1γ289 γ ray 0 --
Alleles Carried on Transgenic Constructs ( 21 )
For All Alleles Carried on Transgenic Constructs Show
Allele of msl-1 Class Mutagen Stocks Known lesion msl-1+t15 in vitro construct | genomic fragment 0 Yes msl-1+t5.9 in vitro construct | deletion 0 Yes msl-1+tFL in vitro construct | genomic fragment 0 Yes msl-1+tT3UTR in vitro construct | genomic fragment 0 Yes msl-1618-938.MtnA.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes msl-1940-1039.MtnA.T:Ivir\HA1 in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes msl-1C.hs.T:Zzzz\FLAG in vitro construct | regulatory fusion in vitro construct | deletion 0 Yes msl-1FC.hs.T:Zzzz\FLAG in vitro construct | regulatory fusion in vitro construct | deletion 0 Yes msl-1FN.hs.T:Zzzz\FLAG in vitro construct | regulatory fusion in vitro construct | deletion 0 Yes msl-1GD1242 in vitro construct | RNAi in vitro construct | regulatory fusion 2 Yes msl-1Hsp83.PC in vitro construct | regulatory fusion 0 Yes msl-1MID.hs.T:Zzzz\FLAG in vitro construct | regulatory fusion in vitro construct | deletion 0 Yes msl-1MPU in vitro construct | site directed mutagenesis 0 Yes msl-1MtnA.T:Zzzz\FLAG in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes msl-1dsRNA.cMa in vitro construct | RNAi 0 Yes msl-1hs.PS in vitro construct | regulatory fusion 0 Yes msl-1hs.T:Zzzz\FLAG in vitro construct | coding region fusion 0 Yes msl-1Δ84.T:Zzzz\FLAG in vitro construct | coding region fusion 0 Yes msl-1Δ84.hs in vitro construct | regulatory fusion in vitro construct | deletion 0 Yes msl-1Δ84C.T:Zzzz\FLAG in vitro construct | coding region fusion 0 Yes msl-1Δ84C.hs in vitro construct | coding region fusion in vitro construct | deletion 0 Yes
Aneuploid Aberrations
Useful deficiency	Df(2L)OD15
Disrupted in	Df(2L)BSC112 (Deal and Cook, 2005.6.1) Df(2L)Exel6041 (Ryder, 2004.4.26) Df(2L)OD15 (Palmer et al., 1993) Df(2L)TW137 (Bevan et al., 1993) Df(2L)TW201 (Palmer et al., 1993) Df(2L)TW3 (Thor and Thomas, 1997) Df(2L)TW50 (Bevan et al., 1993) Df(2L)γ244 (Palmer et al., 1993)
Not disrupted in	Df(2L)M36F-S6 (Palmer et al., 1993) Df(2L)OD15 (Thor and Thomas, 1997)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1242} NA characterization construct P{FΔ84.msl-1} NA P{FΔ84C.msl-1} NA P{msl-1+5.9} NA P{msl-1+tFL} NA P{msl-1+tT3UTR} NA P{msl-1MPU} NA P{msl-1} NA heat-shock construct P{C.msl-1} NA P{F.msl-1} NA P{FC.msl-1} NA P{FN.msl-1} NA P{Hsp83-msl-1.C} NA P{MID.msl-1} NA P{hs-msl-1} NA P{Δ84.msl-1} NA P{Δ84C.msl-1} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EPgy2}msl-1EY09390 NA
Related Comments
	Please look at the allele reports for the complete phenotype data Mutants show decreased levels of X-linked-enzyme activities (G6PD, 6GPD, FUM) but not autosomally encoded enzymes (ADH, AO, GPDH, IDH) in homozygous msl-11 and msl-12 male larvae when compared with non-msl controls. (Belote and Lucchesi, 1980) Females heterozygous for Sxlf1 and homozygous for msl-11 show signs of intersexual development. (Skripsky and Lucchesi, 1982) Mutants show no interaction with mle1 or msl-21. (Belote, 1983) Mutants are defective for dosage compensation in males. Homozygous male embryos hatch but die as much as twelve days later in larval or prepupal stages; females and heterozygous males survive; phenotype slightly more severe in sons of homozygous than of heterozygous mothers. Viability of two-X individuals that develop as phenotypic males (tra21) or intersexes (dsx1) is unaffected by msl-11, indicating that the one-X condition is required for msl-11 lethality. (Lindsley and Zimm, 1992) msl-1, like mle and H4Ac16 (an acetylated form of the His4 product), exhibits a wild type male localisation pattern in Sxl- XX nuclei. (Palmer et al., 1994) RNAi screen using dsRNA made from templates generated with primers directed against this gene results in chromosome misalignment on the metaphase spindle when assayed in S2 cells in the presence of Cdc27 dsRNA. This phenotype cannot be observed when the screen is performed without Cdc27 dsRNA. (Goshima et al., 2007)
Gene Ontology: Function, Process & Cellular Component ( 11 )
Molecular Function
CV term Evidence References
DNA binding non-traceable author statement (Wutz, 2003) inferred from direct assay (Gilfillan et al., 2006)
chromatin binding inferred from direct assay (Bone et al., 1994) non-traceable author statement (Palmer, 1993.7.27) non-traceable author statement (Swiss-Prot Project Members, 1996.10.1)
protein binding traceable author statement (Akhtar, 2003) inferred from physical interaction with FLYBASE:tamo; FB:FBgn0041582 (Minakhina et al., 2003)
Biological Process
CV term Evidence References
dosage compensation inferred from mutant phenotype (Bone et al., 1994)
dosage compensation complex assembly during dosage compensation by hyperactivation of X chromosome traceable author statement (Wutz, 2003) traceable author statement (Akhtar, 2003)
dosage compensation, by hyperactivation of X chromosome non-traceable author statement (FlyBase, 1992-) non-traceable author statement (Amrein, 2000) non-traceable author statement (Schutt and Nothiger, 2000) non-traceable author statement (Swiss-Prot Project Members, 1996.10.1)
Cellular Component
CV term Evidence References
X chromosome non-traceable author statement (Amrein, 2000) traceable author statement (Akhtar, 2003)
X chromosome located dosage compensation complex, transcription activating non-traceable author statement (FlyBase, 1992-) traceable author statement (Amrein, 2000) non-traceable author statement (Schutt and Nothiger, 2000)
chromosome non-traceable author statement (Swiss-Prot Project Members, 1996.10.1)
dosage compensation complex inferred from direct assay (Dahlsveen et al., 2006)
nucleus non-traceable author statement (Swiss-Prot Project Members, 1996.10.1)
Sequence Ontology: Class of Gene
	protein_coding_gene   nuclear_gene
Interactions & Pathways
Summary of Genetic Interactions
Interacts with