FlyBase Gene Report: 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

Gene Model Status

Current

Stock Availability

15 publicly available

Gene Summary

male-specific lethal 1 (msl-1) encodes a protein that is thought to form a scaffold to organize the full male-specific-lethal dosage compensation complex, which increases male X chromosome transcription approximately two-fold. msl-1 homozygous mutant males die as larvae, while females are viable. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Gene Group
UniProtKB
Red Book
Interactive Fly

Also Known As

MSL1, MSL

Key Links

Genomic Location

Cytogenetic map

36F11-37A1

Sequence location

2L:18,683,720..18,688,800 [-]

Recombination map

2-54

RefSeq locus

NT_033779 REGION:18683720..18688800

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

Gene Ontology (GO) Annotations (15 terms)

Molecular Function (5 terms)

Terms Based on Experimental Evidence (4 terms)

CV Term

Evidence

References

contributes_to chromatin binding

inferred from direct assay

(Hallacli et al., 2012)

enables chromatin binding

inferred from direct assay

(Straub et al., 2008, Bone et al., 1994)

enables DNA binding

inferred from direct assay

(Gilfillan et al., 2006)

inferred from mutant phenotype

(Tikhonova et al., 2019)

enables protein binding

inferred from physical interaction with FLYBASE:tamo; FB:FBgn0041582

(Minakhina et al., 2003)

enables transcription cis-regulatory region binding

inferred from direct assay

(Hallacli et al., 2012)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

contributes_to chromatin binding

inferred from biological aspect of ancestor with PANTHER:PTN000493939

(GO Reference Genome Project, 2011-)

enables protein-macromolecule adaptor activity

inferred from biological aspect of ancestor with PANTHER:PTN000493939

(GO Reference Genome Project, 2011-)

inferred from sequence or structural similarity with UniProtKB:Q9HCI7

(Gene Ontology Curators, 2002-)

Biological Process (3 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

involved_in chromatin organization

inferred from mutant phenotype

(Deng et al., 2009)

involved_in dosage compensation by hyperactivation of X chromosome

inferred from mutant phenotype

(Bone et al., 1994)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

involved_in chromatin remodeling

inferred from biological aspect of ancestor with PANTHER:PTN000493939

(GO Reference Genome Project, 2011-)

involved_in dosage compensation by hyperactivation of X chromosome

non-traceable author statement

(Schutt and Nothiger, 2000)

Cellular Component (7 terms)

Terms Based on Experimental Evidence (6 terms)

CV Term

Evidence

References

located_in chromosome

inferred from mutant phenotype

(Tikhonova et al., 2019)

inferred from direct assay

(Bone et al., 1994)

part_of dosage compensation complex

inferred from direct assay

(Dahlsveen et al., 2006)

part_of MSL complex

inferred from direct assay

(Tikhonova et al., 2024, Babosha et al., 2023, Hallacli et al., 2012, Franke and Baker, 1999)

inferred from mutant phenotype

(Tikhonova et al., 2019)

inferred from direct assay

(Meller et al., 2000)

located_in nuclear chromosome

inferred from direct assay

(Figueiredo et al., 2014)

part_of X chromosome located dosage compensation complex, transcription activating

inferred from direct assay

(Kind et al., 2008, Akhtar et al., 2000, Meller et al., 2000)

is_active_in X chromosome

inferred from direct assay

(Kind et al., 2008, Akhtar et al., 2000)

Terms Based on Predictions or Assertions (4 terms)

CV Term

Evidence

References

located_in chromosome

inferred from experiment

(Kind et al., 2008, Mendjan et al., 2006, Akhtar et al., 2000)

part_of MSL complex

inferred from electronic annotation with InterPro:IPR026711

(InterPro Project Members, 2004-)

inferred from biological aspect of ancestor with PANTHER:PTN000493939

(GO Reference Genome Project, 2011-)

located_in nucleus

inferred from experiment

(Mendjan et al., 2006)

part_of X chromosome located dosage compensation complex, transcription activating

non-traceable author statement

(Schutt and Nothiger, 2000)

Gene Group (FlyBase)

MALE SPECIFIC LETHAL COMPLEX

Protein Family (UniProt)

Belongs to the msl-1 family. (P50535)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

Gene Group (FlyBase)

MALE SPECIFIC LETHAL COMPLEX -: The Male Specific Lethal (MSL) complex is a chromatin modifying complex composed of five protein subunits and two non-coding RNAs. MSL is involved in X chromosome dosage compensation in males. (Adapted from FBrf0228243).

Protein Function (UniProtKB)

Component of the male-specific lethal (MSL) histone acetyltransferase complex, a multiprotein complex essential for elevating transcription of the single X chromosome in the male (X chromosome dosage compensation) (PubMed:21726816, PubMed:7781064). The MSL complex specifically associates with the single X chromosome in males and mediates formation of H4K16ac, promoting a two-fold activation of X chromosome (PubMed:16543150, PubMed:18510926). In complex with msl-2, promotes ubiquitination of histone H2B (PubMed:21726816). In addition to its role in dosage compensation in males, regulates the activity of gene promoters: acts together with Cdk7 to promote phosphorylation of 'Ser-5' of the C-terminal heptapeptide repeat domain (CTD) of the largest RNA polymerase II subunit Polr2A (PubMed:27183194).

(UniProt, P50535)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

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).

Summary (Interactive Fly)

chromatin component - dosage compensation - basic motif, leucine zipper-like motif, glycine-rich motif protein that directly binds DNA - Msl-1 is thought to form a scaffold to organize the full Male-Specific-Lethal dosage compensation complex, which increases male X chromosome transcription approximately two-fold - functional interplay between MSL1 and CDK7 controls RNA polymerase II Ser5 phosphorylation

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\msl-1 for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)

Isoform displayed:

Pfam protein domains

InterPro name

classification

start

end

Protein Domains (via SMART)

Isoform displayed:

SMART protein domains

InterPro name

classification

start

end

Structure

Protein 3D structure (Predicted by AlphaFold) (AlphaFold entry P50535)

If you don't see a structure in the viewer, refresh your browser.

Model Confidence:

Very high (pLDDT > 90)
Confident (90 > pLDDT > 70)
Low (70 > pLDDT > 50)
Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures

Crossreferences

Comments on Gene Model

Gene model reviewed during 5.51

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

msl-1-RA

FBtr0081130

NM_057548

5021

1039

msl-1-RB

FBtr0343118

NM_001299125

3757

1039

Additional Transcript Data and 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 (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

msl-1-PA

FBpp0080674

117.5

1039

5.86

FBpp0309817

117.5

1039

5.86

P50535

NP_001286054

AHN54568

Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

1039 aa isoforms: msl-1-PA, msl-1-PB

Additional Polypeptide Data and Comments

Reported size (kDa)

1039 (aa)

(Kelley et al., 1995)

170 (kD observed)

(Palmer et al., 1994)

955 (aa)

(Palmer et al., 1993)

Comments

External Data

Subunit Structure (UniProtKB)

Component of the male-specific lethal (MSL) histone acetyltransferase complex, composed of mof, mle, msl-1, msl-2 and msl-3 proteins, as well as roX1 and roX2 non-coding RNAs (PubMed:10679323, PubMed:11014199, PubMed:16543150, PubMed:18510926). Interacts (via PEHE domain) with mof (via HAT domain) and msl-3 (via MRG domain); both interactions are direct. Interacts with tamo via the nuclear localization signal (PubMed:12653959). Component of a maternal MSL subcomplex composed of mof, msl-1 and msl-3 (PubMed:32502394).

(UniProt, P50535)

Post Translational Modification

Phosphorylation at Ser-18, Thr743, Thr-747 and Thr-751 is required to promote phosphorylation of 'Ser-5' of the C-terminal heptapeptide repeat domain (CTD) of the largest RNA polymerase II subunit Polr2A (PubMed:27183194). Phosphorylated by Cdk7 in vitro (PubMed:27183194). In contrast, phosphorylation at Ser-18, Thr743, Thr-747 and Thr-751 does not affect its role in dosage compensation in males (PubMed:34426916).

Ubiquitinated by msl-2.

(UniProt, P50535)

Crossreferences

InterPro - A database of protein families, domains and functional sites

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\msl-1 using the Feature Mapper tool.

External Data

Crossreferences

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

msl-1_1

Linkouts

Expression Data

Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

-0.48

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

organism

Comment: rapidly degraded

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | faint | ubiquitous

(Fisher et al., 2012)

embryonic stage 9 -- 10

organism | faint | ubiquitous

(Fisher et al., 2012)

trunk mesoderm

(Fisher et al., 2012)

embryonic stage 11 -- 12

central brain primordium

(Fisher et al., 2012)

organism | faint | ubiquitous

(Fisher et al., 2012)

posterior midgut primordium

(Fisher et al., 2012)

embryonic stage 13 -- 16

embryonic brain

(Fisher et al., 2012)

larval ventral nerve cord

(Fisher et al., 2012)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage -- adult stage

(Palmer et al., 1993)

adult stage | female

(Palmer et al., 1993)

adult stage | male

(Palmer et al., 1993)

Additional Descriptive Data

msl-1 transcripts are detected throughout development and are present in adult males and females.

(Palmer et al., 1993)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage | male

cell

• X chromosome

(Franke and Baker, 1999)

cell

• nucleus | punctate

Comment: expression is male-specific

(Franke and Baker, 1999)

western blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage

(Palmer et al., 1994)

larval stage | female

(Gorman et al., 1995)

larval stage | male

(Gorman et al., 1995, Palmer et al., 1994)

adult stage | male

(Palmer et al., 1994)

Additional Descriptive Data

msl-1 protein is present in male and female larvae but is much less abundant in females.

(Gorman et al., 1995, Palmer et al., 1993)

msl-1 protein binds to the same sites on the male X chromosome as mle protein. The pattern of binding of H4Ac16 along the X chromosome is largely coincident with that of mle and msl-1.

(Bone et al., 1994, Palmer et al., 1994)

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in chromosome

inferred from mutant phenotype

(Tikhonova et al., 2019)

inferred from direct assay

(Bone et al., 1994)

part_of dosage compensation complex

inferred from direct assay

(Dahlsveen et al., 2006)

part_of MSL complex

inferred from direct assay

(Tikhonova et al., 2024, Babosha et al., 2023, Hallacli et al., 2012, Franke and Baker, 1999)

inferred from mutant phenotype

(Tikhonova et al., 2019)

inferred from direct assay

(Meller et al., 2000)

located_in nuclear chromosome

inferred from direct assay

(Figueiredo et al., 2014)

part_of X chromosome located dosage compensation complex, transcription activating

inferred from direct assay

(Kind et al., 2008, Akhtar et al., 2000, Meller et al., 2000)

is_active_in X chromosome

inferred from direct assay

(Kind et al., 2008, Akhtar et al., 2000)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\msl-1 in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\msl-1

View exonic expression by tissue for Dmel\msl-1

Bulk Downloads

RNA-Seq RPKM values for all genes

Reference

See Gelbart and Emmert, 2013 for analysis details and data files for all genes.

Developmental Proteome: Life Cycle

Developmental Proteome: Embryogenesis

External Data and Images

Linkouts

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0005617

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

35121/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0005617

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0005617

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0005617

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG10385

Flygut - An atlas of the Drosophila adult midgut

FBgn0005617

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\msl-1

Stages(s) 4-6
Stages(s) 7-8
Stages(s) 9-10
Stages(s) 11-12
Stages(s) 13-16

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 26 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 55 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of msl-1

Transgenic constructs containing regulatory region of msl-1

Aberrations (Deficiencies and Duplications) ( 10 )

Inferred from experimentation ( 10 )

Gene disrupted in

Df(2L)BSC112

(Deal and Cook, 2005.6.1)

Df(2L)ED1203

(Kahsai and Cook, 2018)

Df(2L)Exel6041

(Ryder, 2004.4.26)

Df(2L)γ244

(Palmer et al., 1993)

Df(2L)OD15

(Palmer et al., 1993)

Df(2L)TW3

(Thor and Thomas, 1997)

Df(2L)TW50

(Bevan et al., 1993)

Df(2L)TW137

(Bevan et al., 1993)

Df(2L)TW201

(Palmer et al., 1993)

Gene not disrupted in

Df(2L)M36F-S6

(Palmer et al., 1993)

Df(2L)OD15

(Thor and Thomas, 1997)

Inferred from location ( 9 )

Variants

Variant Molecular Consequences

Alleles Representing Disease-Implicated Variants

Phenotypes

For more details about a specific phenotype click on the relevant allele symbol.

Lethality

Allele

lethal, with Scer\GAL4^Mef2.PR

msl-1^GD1242

lethal | female | heat sensitive, with Scer\GAL4^αTub84B.PL

lethal | male

lethal | male, with Scer\GAL4^αTub84B.PL

lethal | male (with Df(2L)ED1203)

msl-1^kmB

lethal | male (with msl-1^AFG)

lethal | male (with msl-1^F979Z)

lethal | male (with msl-1^K1009X)

lethal | male (with msl-1^L60)

lethal | male (with msl-1^P864L)

msl-1^L60

lethal | male (with msl-1^S943F)

msl-1^L60

lethal | male | heat sensitive, with Scer\GAL4^αTub84B.PL

msl-1^{mut4.UAS.Tag:FLAG}

lethal | male | recessive

lethal | male limited

lethal | male limited (with msl-1^L60)

msl-1^γ269

lethal | male limited (with msl-1^γ269)

msl-1^L60

lethal | male limited | heat sensitive, with Scer\GAL4^αTub84B.PL

msl-1^{mut1.UAS.Tag:FLAG}

lethal - all die before end of larval stage | male

partially lethal | male | recessive

partially lethal - majority die | male | recessive

partially lethal - majority die | male limited

viable

viable, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

msl-1^GD1242

viable, with Scer\GAL4^pnr-MD237

msl-1^GD1242

viable, with Scer\GAL4^tin.CΔ4

msl-1^GD1242

viable, with Scer\GAL4^αTub84B.PL

msl-1^UAS.Tag:FLAG

viable | female

msl-1^Hsp83.PC

viable | male

msl-1^N.hs.Tag:HA

Sterility

Allele

fertile

msl-1^EY09390

Other Phenotypes

Allele

abnormal size | adult stage | male limited, with Scer\GAL4^ey.PH

abnormal size | adult stage | male limited, with Scer\GAL4^nub-AC-62

msl-1^RNAi.UAS.cUa

increased cell death | larval stage | male limited, with Scer\GAL4^e22c

msl-1^RNAi.UAS.cUa

short lived | male (with msl-1^P864L)

msl-1^S943F

short lived | male (with msl-1^S943F)

msl-1^P864L

some die during larval stage | male

visible, with Scer\GAL4^pnr-MD237

msl-1^GD1242

visible | adult stage | male limited, with Scer\GAL4^ey.PH

visible | male limited, with Scer\GAL4^ey.PH

msl-1^RNAi.UAS.cUa

visible | male limited, with Scer\GAL4^nub-AC-62

msl-1^RNAi.UAS.cUa

wild-type

Phenotype manifest in

Allele

eye | male limited, with Scer\GAL4^ey.PH

mesonotum, with Scer\GAL4^pnr-MD237

msl-1^GD1242

trichogen cell, with Scer\GAL4^pnr-MD237

msl-1^GD1242

wing | male limited, with Scer\GAL4^nub-AC-62

msl-1^RNAi.UAS.cUa

wing disc posterior compartment, with Scer\GAL4^e22c

msl-1^RNAi.UAS.cUa

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (1)

Hsap\MSL1

6 of 14

Yes

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (1)

Rnor\Msl1

6 of 14

Yes

Mus musculus (laboratory mouse) (1)

Mmus\Msl1

6 of 14

Yes

Xenopus tropicalis (Western clawed frog) (0)

Danio rerio (Zebrafish) (2)

Drer\msl1a

5 of 14

Yes

Drer\msl1b

4 of 14

Yes

Caenorhabditis elegans (Nematode, roundworm) (0)

Anopheles gambiae (African malaria mosquito) (0)

Arabidopsis thaliana (thale-cress) (0)

Saccharomyces cerevisiae (Brewer's yeast) (0)

Schizosaccharomyces pombe (Fission yeast) (0)

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:msl-1. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (1)

CG12316

2 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 0 )

Allele

Disease

Evidence

References

Potential Models Based on Orthology ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 0 )

Allele

Disease

Interaction

References

Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)

Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.

Homo sapiens (Human)

Gene name

Score

OMIM

OMIM Phenotype

DO term

Complementation?

Transgene?

MSL1; MSL complex subunit 1

6 of 14

614801

Functional Complementation Data

Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.

Interactions

Summary of Physical Interactions

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

msl-1

anti tag coimmunoprecipitation, anti tag western blot, western blot

(Hallacli et al., 2012, Li et al., 2005)

msl-1 - Cdk7

anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation

(Chlamydas et al., 2016)

msl-1 - JIL-1

anti bait coimmunoprecipitation, western blot, anti tag western blot, anti tag coimmunoprecipitation, pull down, proximity ligation assay, fluorescence microscopy

(Albig et al., 2019, Lindehell et al., 2015, Jin et al., 2000)

msl-1 - Jasper

anti bait coimmunoprecipitation, western blot

(Albig et al., 2019)

msl-1 - MBD-R2

anti bait coimmunoprecipitation, western blot

(Raja et al., 2010)

msl-1 - Mat1

anti bait coimmunoprecipitation, western blot

(Chlamydas et al., 2016)

msl-1 - Spt5

pull down, anti tag western blot

(Prabhakaran and Kelley, 2012)

msl-1 - Top2

proximity ligation assay, fluorescence microscopy

(Lindehell et al., 2015)

msl-1 - mle

anti tag coimmunoprecipitation, western blot, proximity ligation assay, fluorescence microscopy, anti bait coimmunoprecipitation

(Lindehell et al., 2015, Cugusi et al., 2013, Ilik et al., 2013, Kadlec et al., 2011, Morra et al., 2008, Smith et al., 2000, Copps et al., 1998)

msl-1 - stumps

two hybrid

(Battersby et al., 2003)

msl-1 - tamo

pull down, autoradiography

(Minakhina et al., 2003)

RNA-protein

Physical Interaction

Assay

References

msl-1 - lncRNA:roX2

anti bait coimmunoprecipitation, primer specific pcr, nucleic acid uv cross-linking assay, western blot, anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Babosha et al., 2024, Müller et al., 2020, Hallacli et al., 2012, Meller et al., 2000, Smith et al., 2000)

Summary of Genetic Interactions

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please look at the allele data for full details of the genetic interactions

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

msl-1

enhanceable

lic

(Clemente-Ruiz et al., 2016)

msl-2|msl-1

suppressible

lncRNA:roX1

(Deng et al., 2005)

msl-1

suppressible

msl-2

(Scott et al., 2000)

msl-1

enhanceable

msl-2

(Scott et al., 2000)

lncRNA:roX1|msl-1

suppressible

ocm

(Lim and Kelley, 2013)

msl-1

suppressible

ocm

(Lim and Kelley, 2013)

msl-1|msl-2

enhanceable

Polr2A|lncRNA:roX1

(Deng et al., 2005)

msl-2|msl-1

suppressible

Polr2A|lncRNA:roX1

(Deng et al., 2005)

msl-1

suppressible

puc

(Clemente-Ruiz et al., 2016)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

Bub3

enhanceable

msl-1

(Clemente-Ruiz et al., 2016)

enhanceable

(Oh et al., 2003)

suppressible

(Oh et al., 2003)

enhanceable

(Prabhakaran and Kelley, 2012)

msl-2

enhanceable

msl-1

(Kelley and Kuroda, 2003)

ocm

suppressible

msl-1

(Lim and Kelley, 2013)

Polr2A|lncRNA:roX1

suppressible

msl-2|msl-1

(Deng et al., 2005)

rod

enhanceable

msl-1

(Clemente-Ruiz et al., 2016)

sc|sisA

enhanceable

msl-1

(Gladstein et al., 2010)

Sxl

suppressible

msl-1

(Clemente-Ruiz et al., 2016)

Trl

enhanceable

msl-1|msl-2|mle

(Greenberg et al., 2004)

Trl

enhanceable

msl-1

(Greenberg et al., 2004)

vir

suppressible

msl-1

(Hilfiker et al., 1995)

External Data

Subunit Structure (UniProtKB)

(UniProt, P50535 )

Linkouts

BioGRID - A database of protein and genetic interactions.

61115

DroID - A comprehensive database of gene and protein interactions.

FBgn0005617

MIST (genetic) - An integrated Molecular Interaction Database

35121

MIST (protein-protein) - An integrated Molecular Interaction Database

35121

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

HATs acetylate histones

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

2-54

Cytogenetic map

36F11-37A1

Sequence location

2L:18,683,720..18,688,800 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

36F11-37A1

Limits computationally determined from genome sequence between P{EP}CG10413^EP2164 and P{lacW}l(2)37Db^k16106&P{lacW}Catsup^k05424

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

36F7-37A1

(determined by in situ hybridisation)

(Palmer et al., 1993)

Experimentally Determined Recombination Data

Location

2-54

(FlyBase, 2016)

2-53.4

(Comeron et al., 2012)

2-20

(Pierre, 1972)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (15)

Aberrations including deletions of this gene Aberrations including duplications of this gene

Bloomington

25157

msl-1^kmB/CyO

31626

y¹ v¹; P{TRiP.JF01411}attP2

5870

msl-1^γ216 cn¹ bw¹/CyO

17564

y¹ w^67c23; P{EPgy2}msl-1^EY09390

35345

y¹ sc^* v¹ sev²¹; P{TRiP.GL00256}attP2

39012

y¹ sc^* v¹ sev²¹; P{TRiP.HMS01930}attP40

Kyoto

108472

msl-1^γ216 cn¹ bw¹/CyO

NIG-Fly

10385R-3

P{NIG.10385R}3

M2L-1803

y¹ w¹¹¹⁸; msl-1^SK2 P{neoFRT}40A/CyO-Tb

M2L-1804

y¹ w¹¹¹⁸; msl-1^SK7 P{neoFRT}40A/CyO-Tb

VDRC

v9239

w¹¹¹⁸; P{GD1242}v9239

v9240

w¹¹¹⁸; P{GD1242}v9240

More stocks available...

Genomic Clones (24)

Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

cDNA Clones (65)

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequenced

BDGP DGC clones

RE18093

Other clones

Drosophila Genomics Resource Center cDNA clones

For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

FBgn0005617

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0005617

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Tikhonova et al., 2019, Smith et al., 2000, Gorman et al., 1995, Bone et al., 1994, Hilfiker et al., 1994, Palmer et al., 1994, Palmer et al., 1993)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

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)

The msl-1 protein shows a clear bias for binding the exons of genes, rather than binding intergenic regions of the chromosome.

(Gilfillan et al., 2006)

msl-1 has a suggested central role in assembly of the MSL complex.

(Scott et al., 2000)

Gene products of the male specific lethal (msl) group of genes preferentially associate with the male X chromosome and may have a role in dosage compensation. This may be achieved by regulating an inverse dosage effect, which would be maintained on the male X and nullified on the autosomes.

(Bhadra et al., 1999)

Gene products of the male specific lethal (msl) group of genes including msl-1, msl-2, msl-3, mle, and mof are associated with all female chromosomes at a low level but are sequestered to the X chromosome in males. There is evidence for the presence of nucleation sites for association of msl proteins with the X chromosome rather than individual gene binding sites.

(Bhadra et al., 1999)

mof colocalises with the MSL complex on the X chromosome: a sequence of binding events results in the formation of the MSL complex on the X chromosome in males and in the targeting of mof to its presumed site of action.

(Gu et al., 1998)

msl-2 and msl-1 colocalise to a reproducible subset of their wild-type X chromosome sites in the absence of either mle or msl-3.

(Lyman et al., 1997)

X chromosome proteins associated with dosage compensation in melanogaster are sufficiently conserved to allow significant antibody cross-reaction to D.simulans, D.virilis, D.americana.americana and D.pseudoobscura.pseudoobscura chromosomes. Cross reaction is also observed in the X chromosome and the X2 chromosome (2 copies in females and 1 in males) of D.miranda. These results provide evidence that the male-specific lethal proteins can be acquired on previously unrelated chromosome arms during evolution.

(Bone and Kuroda, 1996)

Male-specific lethal (MSL) proteins accumulate in a subregion of male nuclei (the X chromosome) beginning at late blastoderm stage. X chromosomal binding of the MSLs is observed throughout embryonic and larval development in both diploid and polytene tissues. His4 colocalises with the MSLs in embryos. Binding of the MSLs is interdependent in diploid cells and is prevented in female embryonic cells by Sxl.

(Franke et al., 1996)

Sex- and chromosome-specific binding of the male-specific lethal (msl) proteins occurs in Drosophilid species spanning 4 genera. msl binding correlates with the evolution of the sex chromosomes.

(Marin et al., 1996)

msl-1 requires msl-2 in order to become associated with the X chromosome. msl-1 binding is prevented in females by the Sxl products derived from the activation of the early Sxl promoter. Association of msl-1 with the X chromosome is stable.

(McDowell et al., 1996)

The products of msl-1, msl-2, mle and msl-3 loci specifically associate with hundreds of sites along the X chromosome in males, but not in females. The binding of each of the four proteins requires the functional products from the other three. 2X3A individuals are mosaic for both Sxl expression and msl-1, msl-2, mle and msl-3 binding to the X chromosome, with a perfect inverse correlation at the cellular level between Sxl expression and msl-1, msl-2, mle and msl-3 X chromosome binding.

(Baker, 1995)

Immunostaining of embryonic and larval stages demonstrates that His4, msl-1 and msl-3 are associated with the male X chromosome as early as gastrulation, while mle binding is not detected until the late embryonic/late larval stages.

(Franke and Baker, 1995)

The msl-3, mle and msl-1 gene products may associate with one another in a male-specific heteromeric complex on the X chromosome to achieve its hyperactivation.

(Gorman et al., 1995)

msl-2 gene product specifically interacts with the male X chromosome, as do mle, msl-1 and msl-3. msl-2 colocalises with msl-1 and antibodies directed against either msl-2 or msl-1 co-immunoprecipitate both proteins from male nuclear extracts.

(Lyman et al., 1995)

The expression pattern of msl-1 suggests msl-1 has a maternal component that appears at the beginning of embryogenesis and localises to the male X chromosome at blastoderm.

(Rastelli and Kuroda, 1995)

Elements needed for dosage compensation are localised to the X chromosome only after blastoderm and msl-dependent dosage compensation is not necessary during the first part of embryogenesis. This suggest the existance of an additional msl-independent dosage compensation mechanism; dosage compensation of run expression at blastoderm is not dependent on male specific lethal genes.

(Rastelli and Kuroda, 1995)

The binding of mle, msl-1, msl-2 and His4 proteins to the X chromosome are interdependent from early embryogenesis.

(Rastelli et al., 1995)

The msl-2 primary transcript may play a role in male specific binding of mle, msl-3 and msl-1 to the X chromosome.

(Zhou et al., 1995)

The gene products of mle and msl-1 bind to the male X chromosome in an identical pattern. The binding sites of H4Ac16 acetylated form of the His4 product are largely coincident with the mle/msl-1 binding sites. This localisation of H4Ac16 protein is dependent on the dosage compensation regulatory pathway.

(Bone et al., 1994)

Sxl is necessary to prevent the association of msl-1 with the X chromosome in females.

(Hilfiker et al., 1994)

The four msl gene products interact to form a multiprotein complex.

(Hilfiker et al., 1994)

Antisera to msl-1 protein label the euchromatic X chromosome through mitosis, but neither the X heterochromatin nor autosomes.

(Lavender et al., 1994)

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)

Sxl negatively regulates the level of msl-1 protein. msl-1 is transcribed in somatic tissues of females and negatively regulated at the post-transcriptional level by msl-2.

(Palmer et al., 1994)

The X chromosome binding of mle requires wild type msl-1, msl-2 and msl-3 functions.

(Gorman et al., 1993)

The msl-1 protein is associated with hundreds of sites along the X chromosomes in males, but not females, consistent with its proposed role in increasing the level of X linked transcription in male nuclei.

(Palmer et al., 1993)

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)

Pole cells from msl-1¹ male embryos are capable of undergoing normal spermatogenesis when transplanted into wild-type hosts.

(Bachiller and Sanchez, 1986)

Mutants show no interaction with mle¹ or msl-2¹.

(Belote, 1983)

Females heterozygous for Sxl^f1 and homozygous for msl-1¹ show signs of intersexual development.

(Skripsky and Lucchesi, 1982)

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)

msl-1 is involved in dosage compensation in males.

(Belote and Lucchesi, 1980)

Relationship to Other Genes

Source for database merge of

Additional comments

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (20)

Reported As

Symbol Synonym

CG10385

(National Institute of Genetics Fly Stocks, 2022-, Hosono et al., 2015, Ni et al., 2011.7.19, Ni et al., 2010.12.1, Ni et al., 2008, Goshima et al., 2007, Battersby et al., 2003)

MSL1

(Kalita and Keller Valsecchi, 2025, Babosha et al., 2024, Choudhury et al., 2024, Kiss et al., 2024, Tikhonova et al., 2024, Tikhonova et al., 2024, Tikhonova et al., 2022, Dai et al., 2021, Makki and Meller, 2021, Villa et al., 2021, Samata et al., 2020, Albig et al., 2019, Albig et al., 2019, Prayitno et al., 2019, Tikhonova et al., 2019, Harumoto and Lemaitre, 2018, Schunter et al., 2017, Chlamydas et al., 2016, Cugusi et al., 2016, Zee et al., 2016, Cugusi et al., 2015, Keller and Akhtar, 2015, Lindehell et al., 2015, Lucchesi and Kuroda, 2015, Apte et al., 2014, Chen et al., 2014, Comoglio and Paro, 2014, Dias et al., 2014, Ferrari et al., 2014, Figueiredo et al., 2014, McElroy et al., 2014, Ferrari et al., 2013, Ilik et al., 2013, Jennings, 2013, Rohrbaugh et al., 2013, Alekseyenko et al., 2012, Conrad et al., 2012, Dunlap et al., 2012, Hohl et al., 2012, Maenner et al., 2012, Menon and Meller, 2012, Philip et al., 2012, Villa et al., 2012, Morra et al., 2011, Regnard et al., 2011, Stenberg and Larsson, 2011, Straub and Becker, 2011, Wu et al., 2011, Fauth et al., 2010, Prestel et al., 2010, Raja et al., 2010, Schiemann et al., 2010, Vaquerizas et al., 2010, Deng and Meller, 2009, Grimaud and Becker, 2009, Bachtrog, 2008, Bell et al., 2008, Izzo et al., 2008, Kelley et al., 2008, Kind et al., 2008, Matyunina et al., 2008, Semeshin et al., 2008, Straub et al., 2008, Sural et al., 2008, Bai et al., 2007, Gilfillan et al., 2007, Kind and Akhtar, 2007, Mendjan and Akhtar, 2007, Park et al., 2007, Prasanth and Spector, 2007, Straub and Becker, 2007, Weake and Scott, 2007, Dahlsveen et al., 2006, Furuhashi et al., 2006, Gilfillan et al., 2006, Hamada et al., 2005, Morales et al., 2005, Nusinow and Panning, 2005, Straub et al., 2005, Straub et al., 2005, Bai et al., 2004, Deng and Meller, 2004, Gilfillan et al., 2004, Kelley, 2004, Lee et al., 2004, Morales et al., 2004, Oh et al., 2004, Andersen and Panning, 2003, Carrozza et al., 2003, Dahlsveen et al., 2003, Park et al., 2003, Perrod and Gasser, 2003, Stuckenholz et al., 2003, Wutz, 2003, Deng and Meller, 2002, Smith et al., 2001, Wang et al., 2001, Gu et al., 2000, Kelley and Kuroda, 2000, Pannuti and Lucchesi, 2000, Scott et al., 2000, Scott et al., 2000, Smith et al., 2000, Stuckenholz et al., 1999, Copps et al., 1998)

Msl 1-3

(Schubeler, 2006)

Msl-1

(Goshima et al., 2007, Minakhina et al., 2003)

Msl1

(Demakova et al., 2020, Hallacli et al., 2012, Kadlec et al., 2011, Carré et al., 2008, Yang, 2004, Minakhina et al., 2003, Meller et al., 2000)

i94

(Battersby et al., 2003)

km(2)B

(Pierre, 1972)

kmB

mls-1

(Loidl, 1994)

msl

(Kotlikova et al., 2006, Moore et al., 2002, Pal-Bhadra et al., 2000, Amrein, 1999, Bhadra et al., 1999, Johansen et al., 1999, Baker, 1998, Hodgkin, 1990)

msl1

(Xie et al., 2025, McCarthy et al., 2022, Babosha et al., 2021, Benhra et al., 2018, Birchler, 2016, Erickson, 2016, Kuroda et al., 2016, Koya and Meller, 2015, Lucchesi and Kuroda, 2015, Ramírez et al., 2015, Lim and Kelley, 2013, Philip and Stenberg, 2013, Bateman et al., 2012, Hallacli et al., 2012, Larschan et al., 2012, Prabhakaran and Kelley, 2012, Birchler et al., 2011, Laverty et al., 2011, Stenberg and Larsson, 2011, Prabhakaran and Kelley, 2010, Schiemann et al., 2010, Schiemann et al., 2010, Deng et al., 2009, Menon and Meller, 2009, Li et al., 2008, Morra et al., 2008, Spierer et al., 2008, Levine et al., 2007, Rodriguez et al., 2007, Worringer and Panning, 2007, Deng and Meller, 2006, Beckstead et al., 2005, Deng et al., 2005, Li et al., 2005, Greenberg et al., 2004, Rattner and Meller, 2004, Demakova et al., 2003, Kelley and Kuroda, 2003, Meller, 2003, Oh et al., 2003, Park et al., 2002, Amrein, 2000, Jin et al., 2000, Mahesh and Ranganath, 2000, Meller, 2000, Kelley et al., 1999, Birchler et al., 1998, Chang and Kuroda, 1998, Cline and Meyer, 1996)

Name Synonyms

Male-Specific Lethal 1

(Vensko and Stone, 2015)

killer-of-males-B

male specifc lethal

(Rogers et al., 2003)

male specific lethal 1

(Raja et al., 2010)

male-specific lethal 1

(Comoglio and Paro, 2014, Georgiev et al., 2011, Prabhakaran and Kelley, 2010, Prasanth and Spector, 2007, Gilfillan et al., 2006)

males specific lethal 1

(Straub and Becker, 2011)

Secondary FlyBase IDs

FBgn0001318
FBgn0002851

Datasets (1)

Study focus (1)

Experimental Role

Project

Project Type

Title

bait_protein

modENCODE_chromatin_proteins_cells

genome binding

Genome-wide localization of chromosomal proteins in cell lines by ChIP-chip and ChIP-Seq.

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 39 )

Sequence Crossreferences

NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.

35121

GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.

GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.

RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.

UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.

P50535

UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information

P50535

UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

X2J6S0

Other crossreferences

AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.

P50535

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0005617

DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species

35121/tissue=All

EMBL-EBI Single Cell Expression Atlas - Single cell expression across species

FBgn0005617

FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data

FBgn0005617

FlyMine - An integrated database for Drosophila genomics

FBgn0005617

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

dme:Dmel_CG10385

MARRVEL_MODEL - MARRVEL (model organism gene)

35121

Linkouts

BioGRID - A database of protein and genetic interactions.

61115

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0005617

DroID - A comprehensive database of gene and protein interactions.

FBgn0005617

DRSC - Results frm RNAi screens

FBgn0005617

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

msl-1_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0005617

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0005617

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG10385

Flygut - An atlas of the Drosophila adult midgut

FBgn0005617

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC014045

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

35121

MIST (protein-protein) - An integrated Molecular Interaction Database

35121

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

HATs acetylate histones

References (373)

The following external sites may use different assemblies or annotations than FlyBase.

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