FlyBase Gene Report: Dmel\msl-2

General Information

Symbol

Dmel\msl-2

Species

D. melanogaster

Name

male-specific lethal 2

Annotation Symbol

CG3241

Feature Type

protein_coding_gene

FlyBase ID

FBgn0005616

Gene Model Status

Current

Stock Availability

10 publicly available

Enzyme Name (EC)

Gene Summary

male-specific lethal 2 (msl-2) encodes the key male-specific subunit of the male-specific-lethal dosage compensation complex. It induces or stabilizes all other components. Homozygous mutant males die, while msl-2 ectopic expression kills females. [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

MSL2, MSL, kmA

Key Links

Genomic Location

Cytogenetic map

23F3-23F3

Sequence location

2L:3,462,218..3,466,112 [-]

Recombination map

2-10

RefSeq locus

NT_033779 REGION:3462218..3466112

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 (19 terms)

Molecular Function (9 terms)

Terms Based on Experimental Evidence (9 terms)

CV Term

Evidence

References

enables chromatin binding

inferred from direct assay

(Hallacli et al., 2012, Straub et al., 2008, Kelley et al., 1995)

enables chromatin-protein adaptor activity

inferred from direct assay

(Tikhonova et al., 2024)

inferred from direct assay

(Valsecchi et al., 2021, Villa et al., 2016, Alekseyenko et al., 2008)

enables DNA binding

inferred from direct assay

(Villa et al., 2016, Zheng et al., 2014, Alekseyenko et al., 2008)

inferred from mutant phenotype

(Tikhonova et al., 2019, Soruco et al., 2013)

inferred from direct assay

(Fauth et al., 2010)

enables histone H2B ubiquitin ligase activity

inferred from direct assay

(Villa et al., 2016)

enables lncRNA binding

inferred from direct assay

(Valsecchi et al., 2021, Ilik et al., 2013, Li et al., 2008)

enables molecular condensate scaffold activity

inferred from direct assay

(Valsecchi et al., 2021)

enables protein-RNA adaptor activity

inferred from direct assay

(Valsecchi et al., 2021)

enables sequence-specific double-stranded DNA binding

inferred from direct assay

(Tikhonova et al., 2024)

enables ubiquitin protein ligase activity

inferred from direct assay

(Schunter et al., 2017, Hallacli et al., 2012)

inferred from direct assay

(Villa et al., 2012)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

enables ubiquitin protein ligase activity

inferred from biological aspect of ancestor with PANTHER:PTN000411633

(GO Reference Genome Project, 2011-)

inferred from electronic annotation with InterPro:IPR032043, InterPro:IPR037922

(InterPro Project Members, 2004-)

Biological Process (4 terms)

Terms Based on Experimental Evidence (4 terms)

CV Term

Evidence

References

involved_in dosage compensation by hyperactivation of X chromosome

inferred from direct assay

(Valsecchi et al., 2021, Villa et al., 2016, Ilik et al., 2013, Alekseyenko et al., 2008)

inferred from mutant phenotype

(Tikhonova et al., 2019, Kelley et al., 1995)

involved_in protein localization to chromatin

inferred from direct assay

(Valsecchi et al., 2021, Villa et al., 2016, Alekseyenko et al., 2008)

involved_in protein polyubiquitination

inferred from direct assay

(Schunter et al., 2017)

involved_in protein ubiquitination

inferred from direct assay

(Hallacli et al., 2012)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

involved_in dosage compensation by hyperactivation of X chromosome

non-traceable author statement

(Schutt and Nothiger, 2000)

involved_in protein ubiquitination

inferred from biological aspect of ancestor with PANTHER:PTN000411633

(GO Reference Genome Project, 2011-)

Cellular Component (6 terms)

Terms Based on Experimental Evidence (6 terms)

CV Term

Evidence

References

located_in chromosome

inferred from mutant phenotype

(Tikhonova et al., 2019)

part_of dosage compensation complex

inferred from direct assay

(Dahlsveen et al., 2006)

part_of MSL complex

inferred from mutant phenotype

(Tikhonova et al., 2019, Soruco et al., 2013)

inferred from direct assay

(Meller et al., 2000)

inferred from direct assay

(Tikhonova et al., 2024, Babosha et al., 2023, Hallacli et al., 2012)

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

(Villa et al., 2012, Alekseyenko et al., 2008, Akhtar et al., 2000, Meller et al., 2000)

is_active_in X chromosome

inferred from direct assay

(Valsecchi et al., 2021, Villa et al., 2016, Zheng et al., 2014, Ilik et al., 2013, Villa et al., 2012, Alekseyenko et al., 2008)

located_in X chromosome

inferred from direct assay

(Kotlikova et al., 2006)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

part_of MSL complex

inferred from biological aspect of ancestor with PANTHER:PTN000411633

(GO Reference Genome Project, 2011-)

inferred from electronic annotation with InterPro:IPR037922

(InterPro Project Members, 2004-)

part_of X chromosome located dosage compensation complex, transcription activating

non-traceable author statement

(Schutt and Nothiger, 2000)

Gene Group (FlyBase)

UNCLASSIFIED RING DOMAIN UBIQUITIN LIGASES

MALE SPECIFIC LETHAL COMPLEX

Protein Family (UniProt)

Belongs to the MSL2 family. (P50534)

Protein Signatures (InterPro)

Catalytic Activity (EC/Rhea)

ubiquitin protein ligase activity

Summaries

Gene Snapshot

Gene Group (FlyBase)

UNCLASSIFIED RING DOMAIN UBIQUITIN LIGASES -: This group comprises RING domain ubiquitin ligases that do not classify under other groups in 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)

Limiting 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:18724933, PubMed:23084835, PubMed:23870142, PubMed:27580037, PubMed:31320325, PubMed:33208948, PubMed:7588059, PubMed:7781064, PubMed:7796814). 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). Msl-2 is only produced in males, constituting the limiting component of the MSL complex (PubMed:7781064). Within the MSL complex, msl-2 mediates the selective binding to the X chromosome and recruitment of the MSL complex via two different mechanisms (PubMed:18724933, PubMed:20139418, PubMed:25452275, PubMed:27580037, PubMed:31320325, PubMed:33208948, PubMed:37602401). Recognizes DNA motifs that are enriched on X chromosome, named PionX sites, which are characterized by sequence features and distinct DNA conformation (base roll) (PubMed:18724933, PubMed:25452275, PubMed:27580037, PubMed:31320325, PubMed:37602401). Specific recognition of the X chromosome is also mediated by the formation of a gel-like state: msl-2 undergoes liquid-liquid phase separation upon binding to roX1 and roX2 non-coding RNAs, leading to nucleate the MSL complex on the X chromosome (PubMed:33208948). Msl-2 is also required for translation and/or stability of msl-1 in males (PubMed:7588059, PubMed:7781064, PubMed:7796814). Also acts as an E3 ubiquitin ligase: in complex with msl-1, mediates ubiquitination of histone H2B at 'Lys-34' (H2BK34Ub) (PubMed:21726816). Also catalyzes ubiquitination of msl-1, msl-3 and mof components of the MSL complex (PubMed:23084834, PubMed:28510597).

(UniProt, P50534)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

kmA: killer of males

Homozygous males die during embryogenesis.

msl-2

Homozygous male embryos hatch but die as much as fourteen days later in larval or prepupal stages; females and heterozygous males survive; no maternal effect. 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-1 (Belote, 1983, Genetics 105: 881-96). Few homozygous msl-22 gynandromorphs survive; XO patches small, with small bristles, and mostly confined to abdomen (Uenoyama, Uchida, Fukunaga, and Oishi, 1982, Genetics 102: 223-31). Pole cells from msl-2 male embryos capable of undergoing normal spermatogenesis when transplanted into wild-type hosts (Bachiller and Sanchez, 1986, Dev. Biol. 118: 379-84). Females heterozygous for Sxlf1 and homozygous (and to a lesser extent heterozygous) for msl-2 show signs of intersexual development; effects greater when mother homozygous for msl-2 [Uenoyama, Fukunaga, and Oishi, 1982, Genetics 102: 233-43 (fig.); Skripsky and Lucchesi, 1982, Dev. Biol. 94: 153-64 (fig.)]. 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-21 male larvae when compared with non-msl controls (Belote and Lucchesi, 1980, Nature 285: 573-75).

Summary (Interactive Fly)

chromatin component - ring finger - metallothionein motif - the key male-specific subunit of the Male-Specific-Lethal dosage compensation complex that induces or stabilizes all other components

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\msl-2 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 P50534)

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

PDB - An information portal to biological macromolecular structures

Comments on Gene Model

Gene model reviewed during 5.52

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

msl-2-RA

FBtr0077571

NM_078743

3843

773

msl-2-RB

FBtr0077570

NM_164536

3710

773

Additional Transcript Data and Comments

Reported size (kB)

3.8, 3.7 (northern blot)

(Bashaw and Baker, 1995)

4.0, 3.8 (northern blot)

(Kelley et al., 1995)

4.0 (northern blot)

(Zhou et al., 1995)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

msl-2-PA

FBpp0077260

84.9

773

4.59

FBpp0077259

84.9

773

4.59

P50534

NP_722907

AAN10368

Polypeptides with Identical Sequences

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

773 aa isoforms: msl-2-PA, msl-2-PB

Additional Polypeptide Data and Comments

Reported size (kDa)

773 (aa); 115 (kD observed)

(Bashaw and Baker, 1995)

773 (aa); 130 (kD observed); 85 (kD predicted)

(Kelley et al., 1995)

769 (aa); 135, 105 (kD observed); 84 (kD predicted)

(Zhou et al., 1995)

Comments

The binding of msl-2 protein to polytene chromosomes requires the other three MSL proteins (mle, msl-1, and msl-3 proteins).

(Bashaw and Baker, 1995)

The predicted msl-2 amino acid sequence is 769-773aa in length depending on strain variations.

(Kelley et al., 1995)

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:18086881, PubMed:21551218, PubMed:23084835, PubMed:23870142, PubMed:7588059, PubMed:7781064, PubMed:7796814). When not associated with chromatin, the MSL complex associates with msl-2 mRNAs, possibly to regulate the amount of available MSL complex (PubMed:21551218). Interacts with Clamp; promoting cooperative binding to DNA PionX sites and recruitment of the MSL complex to chromatin (PubMed:31320325, PubMed:35648444, PubMed:37602401).

(UniProt, P50534)

Post Translational Modification

Autoubiquitinated.

(UniProt, P50534)

Domain

The C-terminal disordered region (CTD) undergoes liquid-liquid phase separation (LLPS) upon roX (roX1 and roX2) non-coding RNAs to form a stably condensed state, leading to nucleate the MSL complex on the X chromosome (PubMed:33208948). The C-terminal disordered region (CTD) forms phase-separated droplets with either roX2 or, less efficiently, with roX1 non-coding RNAs (PubMed:18086881, PubMed:33208948).

The MSL2-type CXC domain and the Pro/Bas region recognize and bind specific DNA motifs on the X-chromosome.

(UniProt, P50534)

Crossreferences

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

PDB - An information portal to biological macromolecular structures

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-2 using the Feature Mapper tool.

External Data

Crossreferences

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

msl-2_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).

-0.50

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

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 9 -- 10

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 11 -- 12

organism | ubiquitous

(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

larval stage | male

(Kelley et al., 1995, Zhou et al., 1995)

larval stage | female

(Kelley et al., 1995, Zhou et al., 1995)

adult stage | male

(Bashaw and Baker, 1995, Kelley et al., 1995, Zhou et al., 1995)

adult stage | female

(Bashaw and Baker, 1995, Kelley et al., 1995, Zhou et al., 1995)

Additional Descriptive Data

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)

adult stage | male

adult posterior midgut

(Hudry et al., 2016)

intestinal stem cell of posterior adult midgut epithelium

(Hudry et al., 2016)

enteroblast of posterior adult midgut epithelium

(Hudry et al., 2016)

enteroendocrine cell of posterior adult midgut epithelium

(Hudry et al., 2016)

enterocyte of posterior adult midgut epithelium

(Hudry et al., 2016)

western blot

Stage

Tissue/Position (including subcellular localization)

Reference

larval stage | male

(Bashaw and Baker, 1995, Zhou et al., 1995)

adult stage | male

(Kelley et al., 1995)

Additional Descriptive Data

msl-2 protein was shown to bind to sites along the male X chromosome. The binding pattern is identical to that of msl-1 protein.

(Hudry et al., 2016, Kelley et al., 1995)

Observed in males but not females.

(Bashaw and Baker, 1995)

msl-2 protein associates with numerous sites along the length of the male X chromosome. The pattern of binding is indistinguishable from that of mle and msl-1 proteins.

(Kelley et al., 1995, Zhou et al., 1995)

msl-2 protein binds to hundreds of sites along the length of the male X chromosome where it colocalizes with msl-1 protein. In addition, msl-2 protein is associated with 20-30 sites along the autosomes.

(Bashaw and Baker, 1995, Zhou et al., 1995)

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in chromosome

inferred from mutant phenotype

(Tikhonova et al., 2019)

part_of dosage compensation complex

inferred from direct assay

(Dahlsveen et al., 2006)

part_of MSL complex

inferred from mutant phenotype

(Tikhonova et al., 2019, Soruco et al., 2013)

inferred from direct assay

(Meller et al., 2000)

inferred from direct assay

(Tikhonova et al., 2024, Babosha et al., 2023, Hallacli et al., 2012)

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

(Villa et al., 2012, Alekseyenko et al., 2008, Akhtar et al., 2000, Meller et al., 2000)

is_active_in X chromosome

inferred from direct assay

(Valsecchi et al., 2021, Villa et al., 2016, Zheng et al., 2014, Ilik et al., 2013, Villa et al., 2012, Alekseyenko et al., 2008)

located_in X chromosome

inferred from direct assay

(Kotlikova et al., 2006)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\msl-2 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-2

View exonic expression by tissue for Dmel\msl-2

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

FBgn0005616

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

33565/tissue=All

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

FBgn0005616

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0005616

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

FBgn0005616

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

CG3241

Flygut - An atlas of the Drosophila adult midgut

FBgn0005616

Images

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 21 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 78 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of msl-2

Transgenic constructs containing regulatory region of msl-2

Aberrations (Deficiencies and Duplications) ( 17 )

Inferred from experimentation ( 17 )

Gene disrupted in

Df(2L)BSC31

(Deal and Cook, 2003.4.2)

Df(2L)Exel7016

(Ryder, 2004.4.26)

Ts(2Lt;3Rt)188.16+Ts(2Rt;3Lt)DTD62

(Bashaw and Baker, 1995)

Ts(2Lt;3Rt)HVD7+Ts(2Rt;3Lt)DTD62

(Bashaw and Baker, 1995)

Ts(2Rt;3Lt)DTD62+Ts(2Lt;3Rt)HVD7

(Myers et al., 1995)

Gene duplicated in

Dp(2;3)odd^5.1

(Zhou et al., 1995)

Ts(YLt;2Lt)G120

(Bashaw and Baker, 1995, Zhou et al., 1995, Belote, 1983)

(Belote, 1983)

(Belote, 1983)

(Belote, 1983)

(Belote, 1983)

Gene not duplicated in

Ts(Y;2Lt)179

(Bashaw and Baker, 1995)

Ts(Y;2Lt)188.9

(Bashaw and Baker, 1995)

Ts(Y;2Lt)G146

(Bashaw and Baker, 1995, Belote, 1983)

Ts(YSt;2Lt)R136

(Belote, 1983)

Gene not disrupted in

In(2LR)DTD16^LDTD42^R

(Bashaw and Baker, 1995)

Ts(2Lt;3Rt)DTD62+Ts(2Rt;3Lt)T(2;3)DTD30

(Bashaw and Baker, 1995)

Inferred from location ( 5 )

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

msl-2^γ9

lethal | female

msl-2^Hsp83.PK

lethal | female, with Tn10\tetR^tTA.Hsp26

msl-2^NOPU.tetO

lethal | female, with Tn10\tetR^tTA.Yp3

msl-2^NOPU.tetO

lethal | female | heat sensitive

msl-2^Hsp83.PK

lethal | male

lethal | male | recessive

lethal - all die before end of larval stage | male

msl-2¹

partially lethal

msl-2^Hsp83.PK

partially lethal | female

msl-2^Hsp83.PK

partially lethal | male, with msl-2²²⁷

partially lethal | male, with msl-2^{R543A.Ubi.Tag:FLAG}

msl-2²²⁷

partially lethal | male, with msl-2^{Δ13d.Ubi.Tag:FLAG}

msl-2²²⁷

partially lethal | male, with msl-2^{Δ36.Ubi.Tag:FLAG}

msl-2²²⁷

partially lethal - majority die | female

msl-2^Ubi.Tag:FLAG

partially lethal - majority die | male, with msl-2²²⁷

msl-2^{R543A.Δ13d.Ubi.Tag:FLAG}

partially lethal - majority die | male, with msl-2^{R543A.Δ13d.Ubi.Tag:FLAG}

msl-2²²⁷

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

msl-2^GD14745

viable, with Scer\GAL4^pnr-MD237

msl-2^GD14745

viable, with Scer\GAL4^tin.CΔ4

msl-2^GD14745

viable | female

viable | male, with msl-2²²⁷

msl-2^{ΔCXC.Ubi.Tag:FLAG}

viable | male, with msl-2^{ΔCXC.Ubi.Tag:FLAG}

msl-2²²⁷

Sterility

Allele

female semi-sterile

msl-2^Hsp83.PK

female sterile

msl-2^Hsp83.PK

Other Phenotypes

Allele

abnormal developmental rate | female

msl-2^Hsp83.PK

abnormal flight, with Scer\GAL4^Mef2.PR

msl-2^GD14745

abnormal pain response | adult stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

msl-2^GD14745

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

msl-2^RNAi.UAS.cUa

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

msl-2^RNAi.UAS.cUa

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

msl-2^KK110252

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

msl-2^GD14745

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

msl-2^RNAi.UAS.cUa

short lived | male, with msl-2²²⁷

msl-2^{R543A.Δ13d.Ubi.Tag:FLAG}

short lived | male, with msl-2^{R543A.Δ13d.Ubi.Tag:FLAG}

msl-2²²⁷

some die during larval stage | male

msl-2¹

visible | female limited

msl-2^ey.hs

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

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

msl-2^RNAi.UAS.cUa

wild-type

Phenotype manifest in

Allele

eye | female

msl-2^ey.hs

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

msl-2^KK110252

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

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

msl-2^RNAi.UAS.cUa

wing disc posterior compartment, with Scer\GAL4^e22c

msl-2^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\MSL2

9 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\Msl2

9 of 14

Yes

Mus musculus (laboratory mouse) (1)

Mmus\Msl2

9 of 14

Yes

Xenopus tropicalis (Western clawed frog) (1)

Xtro\msl2

3 of 13

Yes

Danio rerio (Zebrafish) (2)

Drer\msl2b

10 of 14

Yes

Drer\msl2a

9 of 14

Yes

Caenorhabditis elegans (Nematode, roundworm) (0)

Anopheles gambiae (African malaria mosquito) (1)

Agam\AgaP_AGAP000533

7 of 12

Yes

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-2. 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) (0)

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?

MSL2; MSL complex subunit 2

9 of 14

614802

KARAYOL-BORROTO-HAGHSHENAS NEURODEVELOPMENTAL SYNDROME; KBHS

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

RNA-protein

Physical Interaction

Assay

References

msl-2 - Hrb27C

anti bait coimmunoprecipitation, quantitative reverse transcription pcr, pull down, Identification by mass spectrometry, western blot, affinity chromatography technology

(Szostak et al., 2018)

msl-2 - Unr

isothermal titration calorimetry, predetermined participant, nucleic acid uv cross-linking assay, autoradiography, electrophoretic mobility shift assay, affinity chromatography technology, western blot, static light scattering, x-ray crystallography, pull down, anti bait coimmunoprecipitation, quantitative reverse transcription pcr, Identification by mass spectrometry

(Szostak et al., 2018, Hennig et al., 2014, Graindorge et al., 2013, Abaza et al., 2006, Duncan et al., 2006)

msl-2 - eIF3d1

pull down, autoradiography, Identification by mass spectrometry

(Szostak et al., 2018)

msl-2 - how

pull down, western blot, peptide massfingerprinting

(Graindorge et al., 2013)

msl-2 - lncRNA:roX1

nucleic acid uv cross-linking assay, anti bait coimmunoprecipitation, full identification by DNA sequencing, clip, quantitative reverse transcription pcr, anti tag coimmunoprecipitation

(Valsecchi et al., 2018, Ilik et al., 2013, Li et al., 2008)

msl-2 - lncRNA:roX2

electrophoretic mobility shift assay, autoradiography, clip, quantitative reverse transcription pcr, pull down, anti tag western blot, nucleic acid uv cross-linking assay, anti bait coimmunoprecipitation, full identification by DNA sequencing, western blot, cross-linking study, Identification by mass spectrometry

(Li et al., 2025, Müller et al., 2020, Valsecchi et al., 2018, Ilik et al., 2013, Maenner et al., 2013)

msl-2 - loqs

pull down, peptide massfingerprinting

(Graindorge et al., 2013)

msl-2 - me31B

pull down, peptide massfingerprinting

(Graindorge et al., 2013)

msl-2 - ssx

electrophoretic mobility shift assay, autoradiography, anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Moschall et al., 2018)

msl-2 - tral

pull down, peptide massfingerprinting

(Graindorge et al., 2013)

protein-protein

Physical Interaction

Assay

References

msl-2

enzymatic study, western blot, anti tag western blot

(Hallacli et al., 2012, Villa et al., 2012)

msl-2 - JIL-1

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

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

msl-2 - Rpn10

enzymatic study, anti tag western blot

(Hallacli et al., 2012)

msl-2 - mle

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

(Lindehell et al., 2015, Maenner et al., 2013, Li et al., 2008, Copps et al., 1998)

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-2

suppressible

AGO2

(Menon and Meller, 2012)

msl-2

suppressible

lncRNA:roX1

(Lim and Kelley, 2012, Meller and Rattner, 2002)

msl-2|Polr2A

suppressible

lncRNA:roX1

(Meller and Rattner, 2002)

msl-2|msl-1

suppressible

lncRNA:roX1

(Deng et al., 2005)

msl-2

enhanceable

msl-1

(Kelley and Kuroda, 2003)

msl-2

suppressible

Polr2A

(Meller and Rattner, 2002)

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-2

suppressible

Spt5

(Prabhakaran and Kelley, 2012)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

Bub3

enhanceable

msl-2

(Clemente-Ruiz et al., 2016)

enhanceable

(Oh et al., 2003)

suppressible

(Oh et al., 2003)

suppressible

(Scott et al., 2000)

msl-1

enhanceable

msl-2

(Scott et al., 2000)

ocm

suppressible

msl-2

(Lim and Kelley, 2013)

Polr2A|lncRNA:roX1

suppressible

msl-2|msl-1

(Deng et al., 2005)

sc|sisA

enhanceable

msl-2

(Gladstein et al., 2010)

Sxl

enhanceable

msl-2

(Cline et al., 1999)

Sxl

suppressible

msl-2

(Cline et al., 1999, Yanowitz et al., 1999)

Sxl|Mettl3

suppressible

msl-2

(Haussmann et al., 2016)

Trl

suppressible

msl-2

(Greenberg et al., 2004)

Trl

enhanceable

msl-2

(Greenberg et al., 2004)

Trl

enhanceable

msl-1|msl-2|mle

(Greenberg et al., 2004)

vir

suppressible

msl-2

(Hilfiker et al., 1995)

External Data

Subunit Structure (UniProtKB)

(UniProt, P50534 )

Linkouts

BioGRID - A database of protein and genetic interactions.

59776

DroID - A comprehensive database of gene and protein interactions.

FBgn0005616

MIST (genetic) - An integrated Molecular Interaction Database

33565

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

33565

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

2-10

Cytogenetic map

23F3-23F3

Sequence location

2L:3,462,218..3,466,112 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

23F3-23F3

Limits computationally determined from genome sequence between P{lacW}Mad^k00237 and P{lacW}Pdsw^k10101

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

Experimentally Determined Recombination Data

Location

2-10

(FlyBase, 2016)

2-8.3

(Comeron et al., 2012)

2-45

(Pierre, 1972)

2-9.0

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (10)

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

Bloomington

25158

msl-2^kmA/CyO

5871

msl-2²²⁷ bw¹/CyO

31627

y¹ v¹; P{TRiP.JF01412}attP2

67411

w^*; sna^Sco/CyO; M{UAS-msl-2.HA}ZH-68E/TM6B, Tb¹

67410

w^*; msl-2^kmA/CyO; M{UAS-msl-2.HA}ZH-68E/TM6B, Tb¹

NIG-Fly

M2L-0128

y¹ w¹¹¹⁸; msl-2^SK1 P{neoFRT}40A/CyO-Tb

M2L-0129

y¹ w¹¹¹⁸; msl-2^SK2 P{neoFRT}40A/CyO-Tb

2LG-0103

y² VhaAC39-1^cho-1 v¹; P{U6-msl-2.NIG.sgRNA}attP40/CyO

VDRC

v102386

P{KK110252}VIE-260B

v29356

w¹¹¹⁸; P{GD14745}v29356

Genomic Clones (6)

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

cDNA Clones (61)

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

GH22488

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.

FBgn0005616

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0005616

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Tikhonova et al., 2019, Szostak et al., 2018, Smith et al., 2000, Bashaw and Baker, 1995, Kelley et al., 1995, Zhou et al., 1995)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

S2R+-msl2-KO

Other Stable Cell Lines

Other Comments

The MSL complex does not mediate dosage compensation directly, but rather its activity overrides the high level of histone acetylation and counteracts the potential overexpression of X-linked genes to achieve the proper twofold up-regulation in males.

(Sun et al., 2013)

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)

Sxl protein requires the obligatory cofactor Unr protein for 3'-UTR mediated translational repression of msl-2 in females. Sxl protein recruits Unr protein to the 3' UTR of msl-2 mRNA specifically in female cells.

(Duncan et al., 2006)

msl-2 protein controls male-specific transcription of roX1 and does not require any other MSL protein for this role.

(Rattner and Meller, 2004)

The MSL complex (including msl-1, msl-2 and msl-3) targets activated regions of te chromosome for chromatin remodelling.

(Sass et al., 2003)

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)

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)

msl-2 mRNA translation is inhibited by the Sxl gene product. Sxl binding sites in both the 5' and 3' UTRs of msl-2 RNA cooperate in this translational repression.

(Gebauer 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)

Sxl acts synergistically through sequences in both the 5' and 3' untranslated regions of msl-2 to mediate repression of msl-2 protein expression.

(Bashaw and Baker, 1997)

Association of Sxl protein with multiple sites in the 5' and 3' untranslated regions of msl-2 transcript represses its translation in females.

(Kelley et al., 1997)

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)

Mutation analysis of the RING finger domain and second cysteine-rich motif confirms that the RING finger is essential for msl-2 function, while suggesting a less stringent requirement for an intact second motif.

(Lyman et al., 1997)

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.

(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)

The expression of msl-2 is sex-specifically regulated by Sxl.

(Bashaw and Baker, 1995)

The msl-2 gene is needed for dosage compensation. The msl-2 protein binds specifically to the X chromosome, only in males.

(Kelley et al., 1995)

msl-2 is the key male-limited regulator of dosage compensation.

(Kelley et al., 1995)

msl-2 transcripts contain multiple Sxl binding sites and the protein is male specific. Sxl may negatively regulate translation of msl-2 in females.

(Kelley 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)

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 be the target of Sxl sex specific regulation and may play a role in male specific binding of mle, msl-3 and msl-1 to the X chromosome.

(Zhou et al., 1995)

Association of H4Ac16 protein with the male X chromosome requires wild type function of msl-1, msl-2, mle and msl-3.

(Bone et al., 1994)

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

(Hilfiker et al., 1994)

msl-2 negatively regulates the stability of msl-1 protein in males.

(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)

Sxl^f1/Sxl^fhv1 females that are also homozygous for mutants at msl-1, msl-2 or msl-3 develop as intersexes of the mosaic type.

(Gorman et al., 1993)

Mutants are defective for dosage compensation in males. Homozygous male embryos hatch but die as much as fourteen 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-2¹ 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-1¹.

(Belote, 1983)

Females heterozygous for Sxl^f1 and homozygous for msl-2¹ show signs of intersexual development. Effect is greater when the mother is homozygous for msl-2¹.

(Skripsky and Lucchesi, 1982, Uenoyama et al., 1982)

Few homozygous msl-2¹ gynandromorphs survive; X0 patches small, with small bristles and mostly confined to abdomen.

(Uenoyama et al., 1982)

msl-2 is involved in dosage compensation in males.

(Belote and Lucchesi, 1980)

Mutants show decreased levels of X-linked-enzyme activities (G6PD, 6GPD, FUM) but not autosomally encoded enzymes (ADH, AO, GPDH, IDH) in homozygous msl-2¹ male larvae when compared with non-msl controls.

(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 (22)

Reported As

Symbol Synonym

CG3241

(National Institute of Genetics Fly Stocks, 2022-, Ni et al., 2011.7.19, Keleman et al., 2009.8.5, Mohammad et al., 2009, Ni et al., 2008, Goshima et al., 2007)

MSL

(Lladser et al., 2017, Kong and Meller, 2008, Minear et al., 2005, Pannuti et al., 2005, Morey and Avner, 2004, Park et al., 2004, Kelley and Kuroda, 2003, Larsson et al., 2003, Rogers et al., 2003, Kelley and Kuroda, 2001, Park and Kuroda, 2001, Amrein and Dunipace, 2000, Kelley and Kuroda, 2000, Meller and Paskus Warden, 2000, Hennig, 1999, Meller and Paskus Warden, 1999, Kuroda, 1998)

MSL-2

(Vensko and Stone, 2015, Alekseyenko et al., 2014, Lee et al., 2014, Vicoso and Bachtrog, 2013, Georgiev et al., 2011, Bao et al., 2008, Larschan et al., 2007, Mendjan and Akhtar, 2007, Buscaino et al., 2006, Mendjan et al., 2006, Pal Bhadra et al., 2006, Lalli et al., 2003, Marin, 2003, Corona et al., 2002, Bhadra et al., 2000, Franke and Baker, 1999, Meller et al., 1997)

MSL2

(Choudhury et al., 2024, Kiss et al., 2024, Tikhonova et al., 2024, Wang et al., 2024, Eggers et al., 2023, Zhang et al., 2023, Tikhonova et al., 2022, Tikhonova et al., 2022, Babosha et al., 2021, Birchler and Veitia, 2021, Dai et al., 2021, Makki and Meller, 2021, Valsecchi et al., 2021, Villa et al., 2021, Zhang et al., 2021, Samata et al., 2020, Albig et al., 2019, Bhardwaj et al., 2019, Lv et al., 2019, Pal et al., 2019, Prayitno et al., 2019, Tikhonova et al., 2019, Lucchesi, 2018, Valsecchi et al., 2018, Ilik et al., 2017, Joshi and Meller, 2017, Schauer et al., 2017, Schunter et al., 2017, Chlamydas et al., 2016, Cugusi et al., 2016, Zee et al., 2016, Cugusi et al., 2015, Ellison and Bachtrog, 2015, Keller and Akhtar, 2015, Li et al., 2015, Lindehell et al., 2015, Lucchesi and Kuroda, 2015, Ramírez et al., 2015, Chen et al., 2014, Comoglio and Paro, 2014, Dias et al., 2014, Ferrari et al., 2014, Figueiredo et al., 2014, McElroy et al., 2014, Alekseyenko et al., 2013, Ferrari et al., 2013, Ilik et al., 2013, Maenner et al., 2013, Sun et al., 2013, Thomae et al., 2013, Wang et al., 2013, Alekseyenko et al., 2012, Conrad et al., 2012, Dunlap et al., 2012, Maenner et al., 2012, Prabhakaran and Kelley, 2012, Villa et al., 2012, Zheng et al., 2012, Larschan et al., 2011, Regnard et al., 2011, Stenberg and Larsson, 2011, Straub and Becker, 2011, Deng and Disteche, 2010, Fauth et al., 2010, Prestel et al., 2010, Schiemann et al., 2010, Schiemann et al., 2010, Gorchakov et al., 2009, Grimaud and Becker, 2009, Worringer et al., 2009, Bachtrog, 2008, Birchler et al., 2008, Spierer et al., 2008, Straub et al., 2008, Sural et al., 2008, Bai et al., 2007, Demakova et al., 2007, Gilfillan et al., 2007, Kind and Akhtar, 2007, Larschan et al., 2007, Mendjan and Akhtar, 2007, Park et al., 2007, Prasanth and Spector, 2007, Alekseyenko et al., 2006, Dahlsveen et al., 2006, Furuhashi et al., 2006, Gilfillan et al., 2006, Kotlikova et al., 2006, Zhang et al., 2006, Nusinow and Panning, 2005, Spierer et al., 2005, Straub et al., 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, Perrod and Gasser, 2003, Rattner and Meller, 2003, Wutz, 2003, Deng and Meller, 2002, Wang et al., 2001, Kelley and Kuroda, 2000, Pannuti and Lucchesi, 2000, Smith et al., 2000, Stuckenholz et al., 1999, Copps et al., 1998)

Msl 1-3

(Schubeler, 2006)

Msl-2

(Joy et al., 2024, Goshima et al., 2007)

Msl2

(Surkova et al., 2021, Hallacli et al., 2012, Venables et al., 2012, Kadlec et al., 2011, Carré et al., 2008, Touchon et al., 2004, Yang, 2004, Wilkinson and Shyu, 2001)

km(2)A

(Pierre, 1972)

kmA

(Lindsley and Zimm, 1992)

msl

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

msl-2

(Kalita and Keller Valsecchi, 2025, Toups and Vicoso, 2025, Fan et al., 2023, Jalloh et al., 2023, Pérez-Mojica et al., 2023, Shingleton and Vea, 2023, National Institute of Genetics Fly Stocks, 2022-, Seong and Kang, 2022, Higareda Alvear et al., 2021, Hopkins and Kopp, 2021, Keller Valsecchi et al., 2021, Lakhotia, 2021, Ota et al., 2021, Witt et al., 2021, Leitner and Ben-Shahar, 2020, Leatham-Jensen et al., 2019, Cheetham and Brand, 2018, Moschall et al., 2018, Sandler et al., 2018, Szostak et al., 2018, Valsecchi et al., 2018, Mohr et al., 2017, Moschall et al., 2017, Cheng et al., 2016, Haussmann et al., 2016, Fear et al., 2015, Paris et al., 2015, Gonzalez et al., 2014, Lee et al., 2014, Stern et al., 2014, Evans and Cline, 2013, Hennig et al., 2013, Ruiz et al., 2013, Wang et al., 2013, Chintapalli et al., 2012, Tarone et al., 2012, Chatterjee et al., 2011, Du et al., 2011, Graham et al., 2011, Graveley et al., 2011, Hartmann et al., 2011, Lott et al., 2011, Lu and Wu, 2011, McNeil et al., 2011, Medenbach et al., 2011, Morra et al., 2011, Yao and Fox, 2011, Barbash, 2010, Gladstein et al., 2010, Duncan et al., 2009, Patalano et al., 2009, Sun and Birchler, 2009, Abaza and Gebauer, 2008, Kulkarni-Shukla et al., 2008, Pierre et al., 2008, Furuhashi et al., 2006, Kozak, 2006, Beckmann et al., 2005, Beckmann et al., 2005, Ray and Fox, 2005, Zhimulev et al., 2003, Forch et al., 2001, Kelley et al., 1995, Lindsley and Zimm, 1992)

msl2

(Xie et al., 2025, Becker, 2024, Makki and Meller, 2024, McCarthy et al., 2022, Demakova et al., 2020, Ota et al., 2017, Urban et al., 2017, Birchler, 2016, Erickson, 2016, Haussmann et al., 2016, Kuroda et al., 2016, Lucchesi and Kuroda, 2015, Hennig et al., 2014, Graindorge et al., 2013, Lim and Kelley, 2013, Philip and Stenberg, 2013, Bateman et al., 2012, Larschan et al., 2012, Lim and Kelley, 2012, Menon and Meller, 2012, Mihailovich et al., 2012, Venables et al., 2012, Birchler et al., 2011, Laverty et al., 2011, Meiklejohn et al., 2011, Morra et al., 2011, Stenberg and Larsson, 2011, Prabhakaran and Kelley, 2010, Schiemann et al., 2010, Zhang et al., 2010, Deng et al., 2009, Gelbart et al., 2009, Sun and Birchler, 2009, Alekseyenko et al., 2008, Li et al., 2008, Morra et al., 2008, Spierer et al., 2008, Levine et al., 2007, Robida et al., 2007, Rodriguez et al., 2007, Straub and Becker, 2007, Sun and Birchler, 2007, Yokoyama et al., 2007, Deng and Meller, 2006, Beckstead et al., 2005, Deng et al., 2005, Hamada et al., 2005, Li et al., 2005, Greenberg et al., 2004, Rattner and Meller, 2004, Scott et al., 2004, Banerjee et al., 2003, Demakova et al., 2003, Kelley and Kuroda, 2003, Meller, 2003, Oh et al., 2003, Rattner and Meller, 2002, Graveley, 2001, Keightley and Eyre-Walker, 2001, Amrein, 2000, Gu et al., 2000, Mahesh and Ranganath, 2000, Meller, 2000, Kelley et al., 1999, Birchler et al., 1998, Chang and Kuroda, 1998, Kelley et al., 1997, Ladomery, 1997, Cline and Meyer, 1996)

Name Synonyms

Male-Specific Lethal 2

(Vensko and Stone, 2015)

killer of males

(Lindsley and Zimm, 1992)

killer-of-males-A

male specifc lethal

(Rogers et al., 2003)

male specific lethal 2

(Sun and Birchler, 2009)

male specific-lethal 2

(Robida et al., 2007)

male-specific lethal 2

(Haussmann et al., 2016, Comoglio and Paro, 2014, Chatterjee et al., 2011, Georgiev et al., 2011, Graveley et al., 2011, Prabhakaran and Kelley, 2010, Prasanth and Spector, 2007, Lucchesi, 1995.2.20)

male-specific-lethal 2

(Abaza and Gebauer, 2008)

male-specific-lethal-2

(Evans and Cline, 2013, Forch et al., 2001)

males specific lethal 2

(Straub and Becker, 2011)

Secondary FlyBase IDs

FBgn0001317
FBgn0002852

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 53 )

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.

33565

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.

P50534

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

P50534

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

A4V038

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.

P50534

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0005616

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

33565/tissue=All

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

FBgn0005616

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

FBgn0005616

FlyMine - An integrated database for Drosophila genomics

FBgn0005616

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

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

dme:Dmel_CG3241

MARRVEL_MODEL - MARRVEL (model organism gene)

33565

PDB - An information portal to biological macromolecular structures

Linkouts

BioGRID - A database of protein and genetic interactions.

59776

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

FBgn0005616

DroID - A comprehensive database of gene and protein interactions.

FBgn0005616

DRSC - Results frm RNAi screens

FBgn0005616

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

msl-2_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0005616

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0005616

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

CG3241

Flygut - An atlas of the Drosophila adult midgut

FBgn0005616

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

TC001753

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

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

33565

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

33565

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

HATs acetylate histones

References (489)

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

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