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Gene 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	7 publicly available
Also Known As	MSL2, MSL
Genomic Location
Chromosome (arm)	2L	Recombination map	2-9.0
Cytogenetic map	23F3-23F3	Sequence location	2L:3,462,213..3,466,111 [-]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE GBrowse	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene male-specific lethal 2 is referred to in FlyBase by the symbol Dmel\msl-2 (CG3241, FBgn0005616). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: chromatin binding; DNA binding. There is experimental evidence that it is involved in the biological process: dosage compensation. 78 alleles are reported. No phenotypic data is available. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Zinc finger, RING-type; Zinc finger, RING-type, conserved site. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of low expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval central nervous system. Comments on Affy2 ProbeSet: ProbeSet 1635840_s_at completely aligns to an exonic region common to each of the 2 FlyBase-annotated transcript isoforms of msl-2. Gene sequence location is 2L:3462213..3466111.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	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).
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2011_10	References Meiklejohn et al., 2011, PLoS Biol. 9(8): e1001126
FB2012_01	References Schiemann et al., 2010, Biochem. Biophys. Res. Commun. 402(4): 699--704 Sequence features BKN30252 HFA00829
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 23F3-23F3   Limits computationally determined from genome sequence between P{lacW}Madk00237 and P{lacW}Pdswk10101
Experimentally Determined Cytological Location
Cytogenetic map Notes References
Experimentally Determined Recombination Data
Location	2-9.0   2-45 (Pierre, 1972)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\msl-2 for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) msl-2-RA FBtr0077571  NM_078743   3847   773 msl-2-RB FBtr0077570  NM_164536   3714   773
Additional Transcript Data & Comments
Reported size (kB)	4.0 (northern blot) (Zhou et al., 1995) 4.0, 3.8 (northern blot) (Kelley et al., 1995) 3.8, 3.7 (northern blot) (Bashaw and Baker, 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein msl-2-PA   FBpp0077260   84.9   773   4.59     NP_523467   AAF51104 msl-2-PB   FBpp0077259   84.9   773   4.59     NP_722907   AAN10368
Additional Polypeptide Data & Comments
Reported size (kDa)	773 (aa); 130 (kD observed); 85 (kD predicted) (Kelley et al., 1995) 773 (aa); 115 (kD observed) (Bashaw and Baker, 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
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Zinc finger, RING-type (IPR001841) Zinc finger, RING-type, conserved site (IPR017907)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202847   AE014134 AAF51104   AE014134 AAN10368   AI403118   AI542398   AW940859   AY128426 AAM75019   BH840712   BI161652   BI231233   BI231878   BI235562   BI374941   BI574729   BI584548   BI624715   CO176918   CO176989   CO190944   CO270366   CO271799   CO275877   CO277040   CO282870   CO283564   CO286849   CO292158   CO302354   CO302415   CO302451   CO305735   CO314759   CO322412   CO330222   CO332557   DN154086   EC054584   EC060423   EC060879   EC062128   EC069416   EC077494   EC078899   EC082705   EC082894   EC083029   EC083126   EC200084   EC203467   EC218396   EC218409   EC221758   EC223020   EC230046   EC234547   EC235001   EC241457   EC245742   EC246990   EC248729   GH748009   GH748106   GH757402   GH771616   GH774425   GH781052   GH783757   GH804981   GH808603   GH810627   GH813991   GH816452   GH820119   GH829373   GH842199   GH854307   GH877989   GH884287   GH893155   GH906165   GH914292   GH915952   GH918419   GH919849   GH929897   L42553 AAA75573   X89241 CAA61529   Z48443 CAA88358
UniProtKB/Swiss-Prot	P50534
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
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     (Kelley et al., 1995, Bashaw and Baker, 1995, Zhou et al., 1995) adult stage | female     (Bashaw and Baker, 1995, Zhou et al., 1995, Kelley et al., 1995)
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage | male   cell   • X chromosome (Franke and Baker, 1999) cell   • nucleus | punctate (Franke and Baker, 1999) Comment:expression is male-specific 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. (Bashaw and Baker, 1995, Kelley et al., 1995) msl-2 protein is detected in extracts from male, but not female, larvae. (Zhou et al., 1995) msl-2 protein is observed in males but not females. (Bashaw and Baker, 1995, Kelley et al., 1995)
Marker for
Subcellular Localization
CV Term	chromosome (Kelley et al., 1995) polytene chromosome (Bashaw and Baker, 1995)
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data for FBgn0005616    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0005616 Very low expression bin max Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of low expression. Peak expression observed within 00-12 hour embryonic stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0005616 expression level Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (189) embryo 02-04hr  (1851) embryo 04-06hr  (1579) embryo 06-08hr  (1539) embryo 08-10hr  (1025) embryo 10-12hr  (755) embryo 12-14hr  (650) embryo 14-16hr  (506) embryo 16-18hr  (428) embryo 18-20hr  (348) embryo 20-22hr  (362) embryo 22-24hr  (377) larva L1  (441) larva L2  (314) larva L3 12hr old  (267) larva L3 puffstage 1-2  (298) larva L3 puffstage 3-6  (556) larva L3 puffstage 7-9  (618) white prepupae new  (670) white prepupae 12hr  (630) white prepupae 24hr  (946) pupae 2d postWPP  (604) pupae 3d postWPP  (410) pupae 4d postWPP  (296) adult male 01day  (567) adult male 05day  (645) adult male 30day  (622) adult female 01day  (208) adult female 05day  (143) adult female 30day  (138) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr  (1851) embryo 04-06hr  (1579) embryo 06-08hr  (1539) embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0005616 expression level Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (189) embryo 02-04hr  (1851) embryo 04-06hr  (1579) embryo 06-08hr  (1539) embryo 08-10hr  (1025) embryo 10-12hr  (755) embryo 12-14hr  (650) embryo 14-16hr  (506) embryo 16-18hr  (428) embryo 18-20hr  (348) embryo 20-22hr  (362) embryo 22-24hr  (377) larva L1  (441) larva L2  (314) larva L3 12hr old  (267) larva L3 puffstage 1-2  (298) larva L3 puffstage 3-6  (556) larva L3 puffstage 7-9  (618) white prepupae new  (670) white prepupae 12hr  (630) white prepupae 24hr  (946) pupae 2d postWPP  (604) pupae 3d postWPP  (410) pupae 4d postWPP  (296) adult male 01day  (567) adult male 05day  (645) adult male 30day  (622) adult female 01day  (208) adult female 05day  143 adult female 30day  138 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr  1851 embryo 04-06hr  1579 embryo 06-08hr  1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    189 embryo 02-04hr    1851 embryo 04-06hr    1579 embryo 06-08hr    1539 embryo 08-10hr    1025 embryo 10-12hr    755 embryo 12-14hr    650 embryo 14-16hr    506 embryo 16-18hr    428 embryo 18-20hr    348 embryo 20-22hr    362 embryo 22-24hr    377 larva L1    441 larva L2    314 larva L3 12hr old    267 larva L3 puffstage 1-2    298 larva L3 puffstage 3-6    556 larva L3 puffstage 7-9    618 white prepupae new    670 white prepupae 12hr    630 white prepupae 24hr    946 pupae 2d postWPP    604 pupae 3d postWPP    410 pupae 4d postWPP    296 adult male 01day    567 adult male 05day    645 adult male 30day    622 adult female 01day    208 adult female 05day    143 adult female 30day    138 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     FlyAtlas Anatomical Expression Data for FBgn0005616    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0005616 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval central nervous system. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (1000 - 25000) Linear, scaled to maximum FBgn0005616 expression level Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0005616 expression level Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    120.9 Larval Midgut    21.8 Larval Hindgut    44.5 Larval Malpighian Tubules    26.9 Larval Fat Body    16.6 Larval Salivary Gland    42.4 Larval Trachea    22.925 Larval Carcass    33.9 Adult Head    34.2 Adult Eye    25.8 Adult Brain    72.7 Adult Thoracic-Abdominal Ganglion    52.2 Adult Crop    28.3 Adult Midgut    16.3 Adult Hindgut    35.9 Adult Malpighian Tubules    40.8 Adult Fat Body    30.4 Adult Salivary Gland    35.6 Adult Heart    16.45 Adult VirginFemale Spermatheca    17 Adult InseminatedFemale Spermatheca    19.5 Adult Ovary    9.8 Adult Testis    5.1 Adult Male Accessory Gland    55.5 Adult Carcass    28.7 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    34.2   NA  Eye    25.8   NA  Brain    72.7   120.9  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    52.2   NA  Crop    28.3   21.8  Midgut    16.3   44.5  Hindgut    35.9   26.9  Malpighian Tubules    40.8   16.6  Fat Body    30.4   42.4  Salivary Gland    35.6   NA  Heart    16.45   22.925  Trachea    NA   NA  VirginFemale Spermatheca    17   NA  InseminatedFemale Spermatheca    19.5   NA  Ovary    9.8   NA  Testis    5.1   NA  Male Accessory Gland    55.5   33.9  Carcass    28.7 modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_11
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0005616 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG3241-RB
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal msl-2γ9 lethal | female msl-2Hsp83.PK lethal | female, with Tn10\tetRHsp26.T:Hsim\VP16 msl-2NOPU.Tn10\tetO lethal | female, with Tn10\tetRYp3.T:Hsim\VP16 msl-2NOPU.Tn10\tetO lethal | female | partially msl-2Hsp83.PK lethal | larval stage | male msl-21 lethal | male msl-21 msl-22 msl-2CA-1 msl-2CA-3 msl-2γ136 lethal | male | recessive msl-2227 msl-2kmA msl-2Z2397 msl-2Z2421 msl-2Z3890 lethal | partially msl-2Hsp83.PK viable, with Scer\GAL4pnr-MD237 msl-2GD14745 viable | female | poor msl-2Hsp83.PK Sterility Allele female sterile msl-2Hsp83.PK female sterile | partially msl-2Hsp83.PK Other Phenotypes Allele developmental rate defective | female msl-2Hsp83.PK visible | female limited msl-2ey.hs wild-type msl-2+t5.0 msl-2+t6 msl-2+t11.0 msl-2+tKa msl-2+tKb msl-2+tKg msl-2MM2SV msl-2NOPU msl-2SM53 msl-2SXB1-2 msl-2SXB1 msl-2SXB2 Phenotype manifest in Allele eye | female msl-2ey.hs
Classical Alleles ( 16 )
For All Classical Alleles Show
Allele of msl-2 Class Mutagen Stocks Known lesion msl-2227 1 Yes msl-2KG06747 P-element activity 1 -- msl-2kmA ethyl methanesulfonate 1 -- msl-2138 0 -- msl-21 ethyl methanesulfonate 0 Yes msl-22 ethyl methanesulfonate 0 -- msl-2g134 0 -- msl-2P17 Delta2-3 0 Yes msl-2P22 Delta2-3 0 Yes msl-2P4 Delta2-3 0 Yes msl-2unspecified 0 -- msl-2Z2397 ethyl methanesulfonate 0 -- msl-2Z2421 ethyl methanesulfonate 0 -- msl-2Z3890 ethyl methanesulfonate 0 -- msl-2γ136 0 Yes msl-2γ9 loss of function allele 0 --
Alleles Carried on Transgenic Constructs ( 62 )
For All Alleles Carried on Transgenic Constructs Show
Allele of msl-2 Class Mutagen Stocks Known lesion msl-2GD14745 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes msl-2GL00309 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes msl-2JF01412 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes msl-2KK110252 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes msl-2+t10 in vitro construct - deletion 0 Yes msl-2+t11.0 in vitro construct - genomic fragment 0 Yes msl-2+t11.5 in vitro construct - deletion 0 Yes msl-2+t33.5 in vitro construct - genomic fragment 0 Yes msl-2+t5.0 in vitro construct - genomic fragment 0 Yes msl-2+t5.1 in vitro construct - deletion 0 Yes msl-2+t6 in vitro construct - genomic fragment in vitro construct 0 Yes msl-2+tKa in vitro construct - genomic fragment 0 Yes msl-2+tKb in vitro construct - genomic fragment 0 Yes msl-2+tKg in vitro construct - genomic fragment 0 Yes msl-2109-6.1 in vitro construct - deletion 0 Yes msl-2110-5.5 in vitro construct - deletion 0 Yes msl-2113-9 in vitro construct - deletion 0 Yes msl-2119-9.5 in vitro construct - deletion 0 Yes msl-2129-7.9 in vitro construct - deletion 0 Yes msl-2CA-1 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2CA-3 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2CC in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2dsRNA.cBa in vitro construct - RNAi 0 Yes msl-2dsRNA.cHa in vitro construct - RNAi 0 Yes msl-2dsRNA.cSa in vitro construct - RNAi 0 Yes msl-2EA in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2EP in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2ey.hs in vitro construct - regulatory fusion 0 Yes msl-2FC.Act5C in vitro construct - regulatory fusion 0 Yes msl-2FC.hs.T:Ecol\lacZ in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes msl-2FCΔ3'.Act5C in vitro construct - deletion 0 Yes msl-2FCΔ3'SX.Act5C in vitro construct - deletion 0 Yes msl-2FCΔspl.Act5C in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔspl.hs.T:Ecol\lacZ in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔSx3.hs.T:Ecol\lacZ in vitro construct - deletion 0 Yes msl-2FCΔSx5.hs.T:Ecol\lacZ in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔSx5ΔSx3.hs.T:Ecol\lacZ in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔSXL5'.Act5C in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔSXL5'Δ3'SX.Act5C in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔuORF.Act5C in vitro construct - site directed mutagenesis 0 Yes msl-2FCΔuORF.hs.T:Ecol\lacZ in vitro construct - site directed mutagenesis 0 Yes msl-2hs.T:Avic\GFP-Venus in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes msl-2Hsp83.PK in vitro construct - regulatory fusion 0 Yes msl-2Hsp83.T:Ecol\dam,T:Hsap\MYC in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes msl-2M63 ethyl methanesulfonate 0 Yes msl-2MC.Act5C in vitro construct - regulatory fusion 0 Yes msl-2MCΔ3'.Act5C in vitro construct - deletion 0 Yes msl-2MM2SV in vitro construct 0 Yes msl-2MtnA.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - regulatory fusion 0 Yes msl-2N.hs.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes msl-2NOPU.Tn10\tetO in vitro construct - regulatory fusion in vitro construct - amino acid replacement 0 Yes msl-2NOPU in vitro construct 0 Yes msl-2SD-KG in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2SK in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes msl-2SM53 in vitro construct 0 Yes msl-2SXB1-2 in vitro construct 0 Yes msl-2SXB1 in vitro construct 0 Yes msl-2SXB2 in vitro construct 0 Yes msl-2tKc in vitro construct - genomic fragment 0 Yes msl-2tKd in vitro construct - genomic fragment 0 Yes msl-2tKe in vitro construct - genomic fragment 0 Yes msl-2tKf in vitro construct - genomic fragment 0 Yes
Aneuploid Aberrations
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)
Duplicated in	Dp(2;3)odd5.1 (Zhou et al., 1995) Ts(YLt;2Lt)G120 (Belote, 1983, Zhou et al., 1995, Bashaw and Baker, 1995) Ts(YLt;2Lt)L126 (Belote, 1983) Ts(YLt;2Lt)P8 (Belote, 1983) Ts(YLt;2Lt)R127 (Belote, 1983) Ts(YSt;2Lt)H116 (Belote, 1983)
Not duplicated in	Ts(Y;2Lt)179 (Bashaw and Baker, 1995) Ts(Y;2Lt)188.9 (Bashaw and Baker, 1995) Ts(Y;2Lt)G146 (Belote, 1983, Bashaw and Baker, 1995) Ts(YSt;2Lt)R136 (Belote, 1983)
Not disrupted in	In(2LR)DTD16LDTD42R (Bashaw and Baker, 1995) Ts(2Lt;3Rt)DTD62+Ts(2Rt;3Lt)T(2;3)DTD30 (Bashaw and Baker, 1995)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data heat-shock construct P{H83M2} NA P{HDM} NA P{hs-msl-2.N.FLAG} NA P{hs-msl-2.Venus} NA P{y+YEM2} NA UAS construct P{GD14745} NA P{KK110252} NA P{tetO.7-msl-2.NOPU} NA P{TRiP.GL00309} NA P{TRiP.JF01412} NA characterization construct P{cosJT3} NA P{K-K11.0} NA P{M2NG11} NA P{MM2SV} NA P{msl-2.CA-1} NA P{msl-2.CA-3} NA P{msl-2.CC} NA P{msl-2.EA} NA P{msl-2.EP} NA P{msl-2.FC} NA P{msl-2.FCΔ3'} NA P{msl-2.FCΔ3'SX} NA P{msl-2.FCΔspl} NA P{msl-2.FCΔSXL5'} NA P{msl-2.FCΔSXL5'Δ3'SX} NA P{msl-2.FCΔuORF} NA P{msl-2.MC} NA P{msl-2.MCΔ3'} NA P{msl-2.SD-KG} NA P{msl-2.SK} NA P{msl-2+tKa} NA P{msl-2+tKb} NA P{msl-2+tKg} NA P{msl-2M63} NA P{msl-2tKc} NA P{msl-2tKd} NA P{msl-2tKe} NA P{msl-2tKf} NA P{NOPU} NA P{pSZ109-6.1} NA P{pSZ110-5.5} NA P{pSZ112-11.5} NA P{pSZ113-9} NA P{pSZ119-9.5} NA P{pSZ122-10} NA P{pSZ129-7.9} NA P{pSZ131-5.1} NA P{SM53} NA P{S-N11.2} NA P{S-S11.5} NA P{SXB1-2} NA P{SXB1} NA P{SXB2} NA P{X-X5.0} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{rib7}msl-2P4 NA P{rib7}msl-2P17 NA P{rib7}msl-2P22 NA P{SUPor-P}msl-2KG06747 NA
Gene Ontology: Function, Process & Cellular Component ( 11 unique terms )
Terms Based on Experimental Evidence ( 5 terms )
Molecular Function
CV term Evidence References
chromatin binding inferred from direct assay (Kelley et al., 1995, Straub et al., 2008) DNA binding inferred from direct assay (Fauth et al., 2010)
Biological Process
CV term Evidence References
dosage compensation inferred from mutant phenotype (Kelley et al., 1995)
Cellular Component
CV term Evidence References
dosage compensation complex inferred from direct assay (Dahlsveen et al., 2006) X chromosome inferred from direct assay (Kotlikova et al., 2006)
Terms Based on Predictions or Assertions ( 9 terms )
Molecular Function
CV term Evidence References
chromatin binding non-traceable author statement (Swiss-Prot Project Members, 1996.10.1) DNA binding traceable author statement (Wutz, 2003) zinc ion binding inferred from electronic annotation with InterPro:IPR001841 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
dosage compensation, by hyperactivation of X chromosome non-traceable author statement (Amrein, 2000, FlyBase, 1992-, Schutt and Nothiger, 2000, Swiss-Prot Project Members, 1996.10.1) dosage compensation complex assembly involved in dosage compensation by hyperactivation of X chromosome traceable author statement (Wutz, 2003)
Cellular Component
CV term Evidence References
chromosome non-traceable author statement (Swiss-Prot Project Members, 1996.10.1) nucleus non-traceable author statement (Swiss-Prot Project Members, 1996.10.1) X chromosome non-traceable author statement (Amrein, 2000) X chromosome located dosage compensation complex, transcription activating non-traceable author statement (FlyBase, 1992-, Schutt and Nothiger, 2000) traceable author statement (Amrein, 2000)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions msl-2 allele Gene References msl-21 Sxl (Skripsky and Lucchesi, 1982) Trl (Greenberg et al., 2004) msl-22 Sxl (Uenoyama et al., 1982) msl-2ey.hs msl-1 (Kelley and Kuroda, 2003) msl-2Hsp83.PK msl-1 (Chang and Kuroda, 1998, Scott et al., 2000, Oh et al., 2003) Sxl (Yanowitz et al., 1999, Cline et al., 1999) Trl (Greenberg et al., 2004) msl-2γ136 msl-1 (Scott et al., 2000) vir (Hilfiker et al., 1995)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 59776 DroID - A comprehensive database of gene and protein interactions. • FBgn0005616 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 33565
Orthologs
Genome-wide drosophilid orthologs	Dana\GF23410 Dere\GG24438 Dgri\GH10188 Dmoj\GI14252 Dper\GL18894 Dpse\GA16882 Dsec\GM11132 Dvir\msl-2 Dwil\GK23860 Dyak\GE14867
Curated drosophilid orthologs	Dana\msl-2   Dfun\msl-2   Dpse\GA16882   Drob\msl-2   Dsim\msl-2   Dvir\msl-2
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AGAP000533-PA Anopheles gambiae (mosquito) BGIBMGA003686-PA Bombyx mori (silk worm) ENSBTAP00000000991 Bos taurus (bovine) ENSCAFP00000010500 Canis familiaris (dog) CPIJ012569 Culex pipens (mosquito) ENSCPOP00000002356 Cavia porcellus (guinea pig) 300921 Daphnia pulex (daphnia) ENSECAP00000015722 Equus caballus (horse) ENSP00000311827 Homo sapiens (human) ISCW015004-PA Ixodes scapularis (deer tick) ENSMUSP00000082270 Mus musculus (house mouse) ENSOANP00000023857 Ornithorynchus anatinus (platypus) ENSORLP00000016604 Oryzias latipes (japanese killifish) ENSPTRP00000026588 Pan troglodytes (chimpanzee) XP_001176841.1 Strongylocentrotus purpuratus (purple sea urchin) XP_001809262.1 Tribolium castaneum (red flour beetle) ENSTNIP00000011553 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000045407 Takifugu rubripes (pufferfish Takifugu) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV11479 LH12855 ISCW015004 FBgn0232457 FBgn0226462 FBgn0225822 FBgn0213748 FBgn0017795 FBgn0187546 FBgn0166077 FBgn0249776 JGI_V11_300921 FBgn0248645 FBgn0076896 FBgn0161088 FBgn0156495 FBgn0153775 FBgn0137005 FBgn0117669 FBgn0116570 FBgn0100404 CPIJ012569 CPIJ010636 BGIBMGA003686 ACYPI49193 GB18291 AGAP000533 AAEL017254 AAEL006714 PB25160 PHUM348150 GLEAN_01753
Stocks & Reagents
Stocks Listed in FlyBase ( 7 )
Bloomington	25158 msl-2kmA/CyO 5871 msl-2227 bw1/CyO 31627 y1 v1; P{TRiP.JF01412}attP2 14313 y1 w67c23; P{SUPor-P}msl-2KG06747 35390 y1 sc* v1; P{TRiP.GL00309}attP2/TM3, Sb1
VDRC	v102386 P{KK110252}VIE-260B v29356 w1118; P{GD14745}v29356
Genomic Clones ( 2 )
	BACR03G07 BACR26A07 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 58 )
	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 GBrowse for alignment of the cDNAs and ESTs to the gene model.
cDNA Clones, Fully Sequenced
BDGP DGC clones	GH22488
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD03176 RE01106 RE20469 RE24832 RE31422 RE62637 RH23787 RH26553 RH64324 SD08701
Other clones	CK01.127D.C10 EC054584 EK050217 EK051617 EK052046 EK064045 EK098632 EK104645 EK119028 EK122730 EK161514 EK163656 EK174937 EK183959 EK184112 EK184161 EK188062 EK199338 EK237729 EK249327 EK294743 EK303903 RP003005242 RP003023967 RP003048075 RP003060721 RP003073425 RP003087887 RP003096985 RP003107598 RP003129572 RP003140595 RP003182671 RP003221006 RP003223827 RP003244481 RP003266960 RP003271561 RP003289577 RP003291428 RP003350412 RP003359978 RP003366310 RP003456209 RP003473947 RP003558031 RP003615759
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0005616 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 33565
Antibody Information
	polyclonal (Kelley et al., 1995, Zhou et al., 1995, Bashaw and Baker, 1995)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
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	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) The binding of mle, msl-1, msl-2 and His4 proteins to the X chromosome are interdependent from early embryogenesis. (Rastelli 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 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) Sxlf1/Sxlfhv1 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 Sxlf1 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)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_078743 NM_164536 RefSeq (Proteins) • NP_523467 NP_722907 DDBJ / EMBL / GenBank • AAA75573 AAF51104 AAM75019 AAN10368 AI403118 AW940859 AY128426 BH840712 CAA61529 CAA88358 L42553 X89241 Z48443 Entrez Gene - A searchable database of RefSeq genes. • 33565 UniProtKB/Swiss-Prot • P50534
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Zinc finger, RING-type (IPR001841) Zinc finger, RING-type, conserved site (IPR017907)
Linkouts
	BioGRID - A database of protein and genetic interactions • 59776 DroID - A comprehensive database of gene and protein interactions. • FBgn0005616 DRSC - Results from RNAi screens. • FBgn0005616 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0005616 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG3241-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0005616 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 33565 InParanoid A subset of ortholog calls from InParanoid. • AGAP000533-PA Anopheles gambiae (mosquito) BGIBMGA003686-PA Bombyx mori (silk worm) ENSBTAP00000000991 Bos taurus (bovine) ENSCAFP00000010500 Canis familiaris (dog) CPIJ012569 Culex pipens (mosquito) ENSCPOP00000002356 Cavia porcellus (guinea pig) 300921 Daphnia pulex (daphnia) ENSECAP00000015722 Equus caballus (horse) ENSP00000311827 Homo sapiens (human) ISCW015004-PA Ixodes scapularis (deer tick) ENSMUSP00000082270 Mus musculus (house mouse) ENSOANP00000023857 Ornithorynchus anatinus (platypus) ENSORLP00000016604 Oryzias latipes (japanese killifish) ENSPTRP00000026588 Pan troglodytes (chimpanzee) XP_001176841.1 Strongylocentrotus purpuratus (purple sea urchin) XP_001809262.1 Tribolium castaneum (red flour beetle) ENSTNIP00000011553 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000045407 Takifugu rubripes (pufferfish Takifugu) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /polycomb/msl2.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 33565 modMine - Data generated by the modENCODE project. • FBgn0005616 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • NV11479 LH12855 ISCW015004 FBgn0232457 FBgn0226462 FBgn0225822 FBgn0213748 FBgn0017795 FBgn0187546 FBgn0166077 FBgn0249776 JGI_V11_300921 FBgn0248645 FBgn0076896 FBgn0161088 FBgn0156495 FBgn0153775 FBgn0137005 FBgn0117669 FBgn0116570 FBgn0100404 CPIJ012569 CPIJ010636 BGIBMGA003686 ACYPI49193 GB18291 AGAP000533 AAEL017254 AAEL006714 PB25160 PHUM348150 GLEAN_01753
Synonyms & Secondary IDs ( 19 )
Reported As
Symbol Synonym	CG3241 (Goshima et al., 2007, Mohammad et al., 2009, Keleman et al., 2009.8.5, Ni et al., 2008, Ni et al., 2011.7.19) km(2)A (Pierre, 1972) kmA (Lindsley and Zimm, 1992) MSL (Pannuti et al., 2005, Minear et al., 2005, Larsson et al., 2003, Morey and Avner, 2004, Park et al., 2004, Rogers et al., 2003, Kelley and Kuroda, 2003, Park and Kuroda, 2001, Kelley and Kuroda, 2001, Kelley and Kuroda, 2000, Amrein and Dunipace, 2000, Meller and Paskus Warden, 2000, Meller and Paskus Warden, 1999, Kuroda, 1998, Hennig, 1999, Kong and Meller, 2008) msl (Johansen et al., 1999, Pal-Bhadra et al., 2000, Amrein, 1999, Bhadra et al., 1999, Baker, 1998, Hodgkin, 1990) Msl 1-3 (Schubeler, 2006) MSL2 (Kotlikova et al., 2006, Nusinow, 2005, Straub et al., 2005, Straub et al., 2005, Perrod and Gasser, 2003, Lee et al., 2004, Morales et al., 2004, Bai et al., 2004, Oh et al., 2004, Gilfillan et al., 2004, Kelley, 2004, Deng and Meller, 2004, Andersen and Panning, 2003, Carrozza et al., 2003, Wutz, 2003, Rattner and Meller, 2003, Deng and Meller, 2002, Wang et al., 2001, Pannuti and Lucchesi, 2000, Kelley and Kuroda, 2000, Smith et al., 2000, Stuckenholz et al., 1999, Copps et al., 1998, Zhang et al., 2006, Dahlsveen et al., 2006, Demakova et al., 2007, Alekseyenko et al., 2006, Gilfillan et al., 2006, Prasanth and Spector, 2007, Furuhashi et al., 2006, Straub et al., 2005, Mendjan and Akhtar, 2007, Bai et al., 2007, Birchler et al., 2008, Kind and Akhtar, 2007, Park et al., 2007, Gilfillan et al., 2007, Larschan et al., 2007, Straub et al., 2008, Worringer et al., 2009, Gorchakov et al., 2009, Grimaud and Becker, 2009, Prestel et al., 2010, Spierer et al., 2008, Fauth et al., 2010, Larschan et al., 2011, Regnard et al., 2011, Sural et al., 2008, Schiemann et al., 2010) msl2 (Hamada et al., 2005, Beckstead, 2005, Meller, 2000, Greenberg et al., 2004, Rattner and Meller, 2004, Meller, 2003, Kelley and Kuroda, 2003, Amrein, 2000, Oh et al., 2003, Rattner and Meller, 2002, Keightley and Eyre-Walker, 2001, Graveley, 2001, Gu et al., 2000, Mahesh and Ranganath, 2000, Kelley et al., 1999, Chang and Kuroda, 1998, Birchler et al., 1998, Ladomery, 1997, Kelley et al., 1997, Cline and Meyer, 1996, Straub and Becker, 2007, Deng et al., 2005, Sun and Birchler, 2007, Demakova et al., 2003, Deng and Meller, 2006, Li et al., 2005, Rodriguez et al., 2007, Morra et al., 2008, Yokoyama et al., 2007, Li et al., 2008, Robida et al., 2007, Levine et al., 2007, Alekseyenko et al., 2008, Gelbart et al., 2009, Banerjee et al., 2003, Zhang et al., 2010, Spierer et al., 2008, Prabhakaran and Kelley, 2010, Schiemann et al., 2010, Meiklejohn et al., 2011) Msl2 (Yang, 2004, Wilkinson and Shyu, 2001, Touchon et al., 2004, Carré et al., 2008) MSL-2 (Bhadra et al., 2000, Franke and Baker, 1999, Meller et al., 1997, Mendjan et al., 2006, Mendjan and Akhtar, 2007, Pal Bhadra et al., 2006, Lalli et al., 2003, Larschan et al., 2007, Corona et al., 2002, Buscaino et al., 2006, Bao et al., 2008) msl-2 (Ray and Fox, 2005, Beckmann et al., 2005, Beckmann et al., 2005, Kozak, 2006, Furuhashi et al., 2006, Lott et al., 2011, Lindsley and Zimm, 1992, Patalano et al., 2009, Pierre et al., 2008, Zhimulev et al., 2003, Forch et al., 2001, Abaza and Gebauer, 2008, Kulkarni-Shukla et al., 2008, Duncan et al., 2009, Barbash, 2010, Graveley et al., 2011, Chatterjee et al., 2011, Gladstein et al., 2010, Medenbach et al., 2011, van Eyk et al., 2011) Msl-2 (Goshima et al., 2007)
Name Synonym	killer of males (Lindsley and Zimm, 1992) killer-of-males-A   male specifc lethal (Rogers et al., 2003) male-specific lethal 2 (Lucchesi, 1995.2.20, Prasanth and Spector, 2007, Prabhakaran and Kelley, 2010, Graveley et al., 2011, Chatterjee et al., 2011) male specific-lethal 2 (Robida et al., 2007) male-specific-lethal-2 (Forch et al., 2001) male-specific-lethal 2 (Abaza and Gebauer, 2008)
Secondary FlyBase IDs
	FBgn0001317 FBgn0002852
References ( 302 )
Generate a list of	All references relating to this gene ( 302 )
List References by type	Research paper  ( 152 ) Supplementary material  ( 4 ) Review  ( 71 ) Erratum  ( 1 ) Personal communication to FlyBase  ( 6 ) Abstract  ( 57 ) FlyBase analysis  ( 1 ) DNA/RNA sequence record  ( 1 ) Book  ( 1 ) Protein sequence record  ( 1 ) Computer file  ( 2 ) Letter  ( 2 ) Automatic genome annotation  ( 1 ) Curated genome annotation  ( 1 )
Recent research papers ( 16 )
	Chatterjee et al., 2011, Development 138(6): 1099--1109 The female-specific Doublesex isoform regulates pleiotropic transcription factors to pattern genital development in Drosophila. [FBrf0213054] Graveley et al., 2011, Nature 471(7339): 473--479 The developmental transcriptome of Drosophila melanogaster. [FBrf0213330] Larschan et al., 2011, Nature 471(7336): 115--118 X chromosome dosage compensation via enhanced transcriptional elongation in Drosophila. [FBrf0213143] Lott et al., 2011, PLoS Biol. 9(2): e1000590 Noncanonical Compensation of Zygotic X Transcription in Early Drosophila melanogaster Development Revealed through Single-Embryo RNA-Seq. [FBrf0213135] Medenbach et al., 2011, Cell 145(6): 902--913 Translational Control via Protein-Regulated Upstream Open Reading Frames. [FBrf0213906] Meiklejohn et al., 2011, PLoS Biol. 9(8): e1001126 Sex Chromosome-Specific Regulation in the Drosophila Male Germline But Little Evidence for Chromosomal Dosage Compensation or Meiotic Inactivation. [FBrf0214756] Regnard et al., 2011, PLoS Genet. 7(3): e1001327 Global Analysis of the Relationship between JIL-1 Kinase and Transcription. [FBrf0213276] van Eyk et al., 2011, Hum. Mol. Genet. 20(14): 2783--2794 Perturbation of the Akt/Gsk3-{beta} signalling pathway is common to Drosophila expressing expanded untranslated CAG, CUG and AUUCU repeat RNAs. [FBrf0213942] Barbash, 2010, Genetics 184(1): 313--316 Genetic testing of the hypothesis that hybrid male lethality results from a failure in dosage compensation. [FBrf0209660] Fauth et al., 2010, Nucleic Acids Res. 38(10): 3209--3221 The DNA binding CXC domain of MSL2 is required for faithful targeting the Dosage Compensation Complex to the X chromosome. [FBrf0210963] Gladstein et al., 2010, PLoS Genet. 6(7): e1001041 Requirement of male-specific dosage compensation in Drosophila females-implications of early X chromosome gene expression. [FBrf0211448] Prabhakaran and Kelley, 2010, BMC Biol. 8: 80 A new strategy for isolating genes controlling dosage compensation in Drosophila using a simple epigenetic mosaic eye phenotype. [FBrf0211122] Prestel et al., 2010, Mol. Cell 38(6): 815--826 The Activation Potential of MOF Is Constrained for Dosage Compensation. [FBrf0211211] Schiemann et al., 2010, BMC Mol. Biol. 11: 80 Sex-biased transcription enhancement by a 5' tethered Gal4-MOF histone acetyltransferase fusion protein in Drosophila. [FBrf0212392] Schiemann et al., 2010, Biochem. Biophys. Res. Commun. 402(4): 699--704 The importance of location and orientation of male specific lethal complex binding sites of differing affinities on reporter gene dosage compensation in Drosophila. [FBrf0212385] Zhang et al., 2010, PLoS Biol. 8(2): e1000320 Expression in aneuploid Drosophila s2 cells. [FBrf0210115] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...