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Gene Dmel\mle

General Information
Symbol	Dmel\mle	Species	D. melanogaster
Name	maleless	Annotation symbol	CG11680
Feature type	protein_coding_gene	FlyBase ID	FBgn0002774
Gene Model Status	Current	Stock availability	9 publicly available
Also Known As	nap, napts
Genomic Location
Chromosome (arm)	2R	Recombination map	2-55.2
Cytogenetic map	42A6-42A6	Sequence location	2R:1,862,243..1,868,277 [-]
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 maleless is referred to in FlyBase by the symbol Dmel\mle (CG11680, FBgn0002774). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: chromatin binding. There is experimental evidence that it is involved in the biological process: male courtship behavior, veined wing generated song production; determination of adult lifespan; dosage compensation; axon extension. 51 alleles are reported. The phenotypes of these alleles are annotated with: giant fiber neuron; NMJ bouton; polytene chromosome; embryonic/larval neuromuscular junction; wing; adult brain; spermatozoon. It has 3 annotated transcripts and 3 annotated polypeptides. Protein features are: DEAD-like helicase; DNA/RNA helicase, ATP-dependent, DEAH-box type, conserved site; DNA/RNA helicase, DEAD/DEAH box type, N-terminal; Domain of unknown function DUF1605; Double-stranded RNA-binding; Double-stranded RNA-binding-like; Helicase, C-terminal; Helicase-associated domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of moderate expression. Peak expression observed within 00-18 hour embryonic stages, at stages throughout the pupal period. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval/adult central nervous system, adult fat body, adult spermathecae. Comments on Affy2 ProbeSet: ProbeSet 1629507_a_at completely aligns to an exonic region common to each of the 3 FlyBase-annotated transcript isoforms of mle. Gene sequence location is 2R:1862243..1868277.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	mle: maleless Homozygous males die, but homozygous females survive. Males produced by homozygous females die during the third larval instar, whereas those produced by heterozygous females are late pupal lethals. Females transformed into phenotypic males (tra) or intersexes (dsx) unaffected by mle, i.e. mle acts only upon single-X-bearing flies. No interaction with msl-1 or msl-2 (Belote, 1983, Genetics 96: 165-86). mle4 males surviving at 18 are sterile, small, and slow developing. Concluded to be defective in dosage compensation in males based on decreased levels of X-linked-enzyme activities (G6PD, 6GPD, FUM, β-HAD) but not autosomally encoded enzymes (ADH, AO, GPDH, IDH) in homozygous mle4 male larvae and escaping adults, e.g. β-HAD. The incorporation of labeled uridine by the polytene X chromosome relative to that of 2R is lower than normal in mle4 males (Belote and Lucchesi, 1980, Nature 285: 573-75); steady-rate level of Sgs4 mRNA incompletely compensated in mle4 and mle4/mle6 male larvae (Breen and Lucchesi, 1986, Genetics 112: 483-91). Polytene X chromosome of mle males appears narrower and more densely stained than that of control males. Few homozygous mle gynandromorphs survive; XO patches small, with small bristles, and mostly confined to abdomen (Uenoyama, Uchida, Fukunaga, and Oishi, 1982, Genetics 102: 223-31). mle pole cell transplanted into wild-type hosts incapable of undergoing normal spermatogenesis (Bachiller and Sanchez, 1986, Dev. Biol. 118: 379-84). Homozygous (and to a lesser extent heterozygous) mle females that are heterozygous for SxlF1 (Uenoyama, Fukunaga, and Oishi, 1982, Genetics 102: 233-43; Skripsky and Lucchesi, 1982, Dev. Biol. 94: 153-64) or are the surviving progeny raised at 17 of homozygous da mothers (Cline, 1982, Genetics 100: 641-63) develop as intersexes. napts: no action potential (J.C. Hall; M. Kernan) Larvae or adults become rapidly paralyzed when exposed to 37 and rapidly recover on return to lower temperatures. Rearing stocks chronically at room temperature or above causes napts to "adapt" such that higher temperatures (> 40) are required for paralysis (Kyriacou and Hall, 1985). Experiments involving one-time rearing at low temperature caused napts to paralyze at relatively low temperatures (Nelson and Wyman, 1990). Axonal conduction (but not synaptic transmission) fails in larvae at high temperatures (Wu et al., 1978; Wu and Ganetzky, 1980), but action potentials in the giant fiber (GF) pathway of adults are not blocked at temperatures up to 43 (Elkins and Ganetzky, 1990; Nelson and Wyman, 1990), though the latency from brain stimulation to response of thoracic muscles are aberrantly long, even at low temperatures (Nelson and Wyman, 1990), and this long-latency disappears as the temperature is raised to 35 (Elkins and Ganetzky, 1990). "Following frequency" of napts thoracic muscle responses (re. GF pathway stimulation) reduced at elevated temperatures, an effect which can be reversed by injection of 4-amino-pyridine (Nelson and Wyman, 1990). At permissive temperatures, refractory period for elicitation of a series of action potentials is abnormally long (Ganetzky and Wu, 1980); at these low temperatures, napts suppresses effects of "hyperexcitability" mutations such as Sh, bas, bss, eas, Hk, kdn, and tko (Ganetzky and Wu, 1982, Genetics 100: 597-614). napts is unconditionally lethal (Ganetzky and Wu, 1980) in a double mutant with parats1 (death occurring during 1st larval instar) and the viability of other para; napts combinations is poor (Ganetzky, 1984); two doses of para+ (in males) suppresses high-temperature paralysis of napts (Stern et al., 1990). In mosaic experiments, cuticular clones of parats1 in a napts background (after low-temperature development) have non-functioning sensory cells, probably due to lack of nerve conduction which, however, did not cause any anatomical abnormalities involving the central projections of these sensory neurons (Burg and Wu, 1986). Another developmental study, examining larval nerve terminal innervating body-wall muscles (Budnik et al., 1990), showed slight reduction in the extent of branching caused by napts at permissive temperature; the increase in branching (and higher than normal number of varicosities on motor-neurites) induced by an eag Sh double mutant was suppressed by napts (re. low-temperature rearing). napts in combination with tipE leads to poor viability at permissive temperature for both mutants (Ganetzky, 1986, J. Neurogenet. 3: 19-31; Jackson, Wilson, and Hall, 1986, J. Neurogenet. 3: 1-17). napts is, at permissive temperature, hypersensitive to blocking effects of tetrodotoxin (TTX) on action potentials (Ganetzky and Wu, 1980); brain membrane extracts of napts, assayed at low or high temperatures, have subnormal levels of tetrodotoxin (Kauvar, 1982) or saxitoxin (Jackson et al., 1984) binding activity; the latter study reports that there are no qualitative alterations of this binding activity (kd is normal). Cultured neurons from napts larvae are 4 to 5-fold more resistant than wild-type cells to killing effects of veratridine, irrespective of temperature (22 vs 35) (Suzuki and Wu 1984, J. Neurogenet. 1: 225-38), but TTX has no effects on these mutant cells, whose general growth characteristics are also normal (Wu, Suzuki and Poo, 1983, J. Neurosci. 3: 1888-99). The mutation does not seem to modify the expression of sodium currents in embryonic neurons (O'Dowd and Aldrich, 1988, J. Neurosci. 8: 3633-43). Exposure of napts males to high temperature causes arrest of oscillator underlying rhythmic component of courtship song (Kyriacou and Hall, 1985); in experiments on conditioned courtship, napts males learn normally but have shortened memory spans, and napts suppresses Sh-induced decrements in courtship learning (Cowan and Siegel, 1984, J. Neurogenet. 1: 333-44; 1986, J. Neurogenet. 3: 187-201).
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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FB2012_01	References Schiemann et al., 2010, Biochem. Biophys. Res. Commun. 402(4): 699--704 Sequence features BKN28045 HFA04921
FB2011_10
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 42A6-42A6   Limits computationally determined from genome sequence between P{PZ}l(2)0985109851&P{lacW}Src42Ak10108 and P{lacW}l(2)k09848k09848&P{EP}EP407
Experimentally Determined Cytological Location
Cytogenetic map Notes References 42A-42A   (determined by in situ hybridisation)   (Kuroda et al., 1991)
Experimentally Determined Recombination Data
Location	2-55.2   2-56.8 (Cicchetti and Loverre, 1989)
Left of (cM)
Right of (cM)	ap lt (Cicchetti and Loverre, 1989)
Notes	Maps just distal to ap based on deficiency mapping.
Gene Model & Products
Please see the GBrowse view of Dmel\mle 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) mle-RA FBtr0086031  NM_057293   4412   1293 mle-RB FBtr0086030  NM_057294   4528   226 mle-RC FBtr0100576  NM_165451   3901   936
Additional Transcript Data & Comments
Reported size (kB)	4.0 (northern blot) (Kuroda et al., 1991)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein mle-PA   FBpp0085367   143.7   1293   7.23     NP_476641   AAF57297 mle-PB   FBpp0085366   25.1   226   7.34     NP_476642   AAM68336 mle-PC   FBpp0100031   104.1   936   7.15     NP_724440   AAM68335
Additional Polypeptide Data & Comments
Reported size (kDa)	1293, 226 (aa); 140 (kD observed); 144 (kD predicted) (Kuroda et al., 1991)
Comments	The truncated form of mle protein is predicted by one of the cDNAs. It has 198aa in common with the larger form of mle and has 28 distinct amino acids at its carboxy terminus. (Kuroda et al., 1991)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Double-stranded RNA-binding (IPR001159) Helicase, C-terminal (IPR001650) DNA/RNA helicase, ATP-dependent, DEAH-box type, conserved site (IPR002464) Helicase-associated domain (IPR007502) Domain of unknown function DUF1605 (IPR011709) Double-stranded RNA-binding-like (IPR014720) DEAD-like helicase (IPR014001) DNA/RNA helicase, DEAD/DEAH box type, N-terminal (IPR011545)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA697368   AE013599 AAF57297   AE013599 AAM68335   AE013599 AAM68336   AC009847   AI512625   AI514561   AI531582   BI214603   BI215346   BI353811   BI611458   BI636383   BT003785   BT010267 AAQ23585   CK135045   CO188484   CO283139   CO303878   EC057790   EC065119   EC199655   EC236880   EV587140   EV588655   EV589146   M74121 AAC41573
UniProtKB/Swiss-Prot	P24785
UniProtKB/TrEMBL	A1Z6I8
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 embryonic stage -- adult stage     (Kuroda et al., 1991)
Additional Descriptive Data
	mle transcripts are not sex-specific. (Kuroda et al., 1991) mle transcripts are detected throughout development and are found in males and females. (Kuroda et al., 1991)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
western blot Stage Tissue/Position (including subcellular localization) Reference adult stage | female     (Kuroda et al., 1991) adult stage | male     (Kuroda et al., 1991)
Additional Descriptive Data
	Strong anti-mle protein staining is observed at hundreds of sites along the length of the male X chromosome. Weak staining of the autosomes and the female X is also observed. (Kuroda et al., 1991) mle protein binds to the same sites on the male X chromosome as msl-1 protein. The pattern of binding of H4Ac16 along the X chromosome is largely coincident with that of mle and msl-1. (Bone et al., 1994)
Marker for
Subcellular Localization
CV Term	polytene chromosome (Kuroda et al., 1991) chromosome (Bone et al., 1994)
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 FBgn0002774    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0002774 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 moderate expression. Peak expression observed within 00-18 hour embryonic stages, at stages throughout the pupal period. [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 FBgn0002774 expression level Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 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  (886) embryo 02-04hr  (1701) embryo 04-06hr  (3062) embryo 06-08hr  (2577) embryo 08-10hr  (2088) embryo 10-12hr  (1695) embryo 12-14hr  (1637) embryo 14-16hr  (1198) embryo 16-18hr  (1023) embryo 18-20hr  (819) embryo 20-22hr  (679) embryo 22-24hr  (797) larva L1  (839) larva L2  (683) larva L3 12hr old  (495) larva L3 puffstage 1-2  (595) larva L3 puffstage 3-6  (968) larva L3 puffstage 7-9  (1284) white prepupae new  (1300) white prepupae 12hr  (1509) white prepupae 24hr  (1846) pupae 2d postWPP  (1588) pupae 3d postWPP  (897) pupae 4d postWPP  (525) adult male 01day  (775) adult male 05day  (919) adult male 30day  (910) adult female 01day  (818) adult female 05day  (981) adult female 30day  (904) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr  (1701) embryo 04-06hr  (3062) embryo 06-08hr  (2577) embryo 08-10hr  (2088) embryo 10-12hr  (1695) embryo 12-14hr  (1637) embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr  1509 white prepupae 24hr  (1846) pupae 2d postWPP  (1588) pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 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    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0002774 expression level Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (886) embryo 02-04hr  (1701) embryo 04-06hr  (3062) embryo 06-08hr  (2577) embryo 08-10hr  (2088) embryo 10-12hr  (1695) embryo 12-14hr  (1637) embryo 14-16hr  (1198) embryo 16-18hr  (1023) embryo 18-20hr  (819) embryo 20-22hr  (679) embryo 22-24hr  (797) larva L1  (839) larva L2  (683) larva L3 12hr old  (495) larva L3 puffstage 1-2  (595) larva L3 puffstage 3-6  (968) larva L3 puffstage 7-9  (1284) white prepupae new  (1300) white prepupae 12hr  (1509) white prepupae 24hr  (1846) pupae 2d postWPP  (1588) pupae 3d postWPP  (897) pupae 4d postWPP  (525) adult male 01day  (775) adult male 05day  (919) adult male 30day  (910) adult female 01day  (818) adult female 05day  (981) adult female 30day  (904) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr  1701 embryo 04-06hr  (3062) embryo 06-08hr  (2577) embryo 08-10hr  2088 embryo 10-12hr  1695 embryo 12-14hr  1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr  1509 white prepupae 24hr  1846 pupae 2d postWPP  1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 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    886 embryo 02-04hr    1701 embryo 04-06hr    3062 embryo 06-08hr    2577 embryo 08-10hr    2088 embryo 10-12hr    1695 embryo 12-14hr    1637 embryo 14-16hr    1198 embryo 16-18hr    1023 embryo 18-20hr    819 embryo 20-22hr    679 embryo 22-24hr    797 larva L1    839 larva L2    683 larva L3 12hr old    495 larva L3 puffstage 1-2    595 larva L3 puffstage 3-6    968 larva L3 puffstage 7-9    1284 white prepupae new    1300 white prepupae 12hr    1509 white prepupae 24hr    1846 pupae 2d postWPP    1588 pupae 3d postWPP    897 pupae 4d postWPP    525 adult male 01day    775 adult male 05day    919 adult male 30day    910 adult female 01day    818 adult female 05day    981 adult female 30day    904 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 FBgn0002774    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0002774 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/adult central nervous system, adult fat body, adult spermathecae. [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 FBgn0002774 expression level Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0002774 expression level Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    232.6 Larval Midgut    41.9 Larval Hindgut    41.7 Larval Malpighian Tubules    68 Larval Fat Body    30.5 Larval Salivary Gland    73.2 Larval Trachea    34.425 Larval Carcass    46.225 Adult Head    94.3 Adult Eye    59 Adult Brain    284.1 Adult Thoracic-Abdominal Ganglion    243.2 Adult Crop    47.9 Adult Midgut    31.6 Adult Hindgut    33.3 Adult Malpighian Tubules    51.5 Adult Fat Body    120.8 Adult Salivary Gland    55.4 Adult Heart    40.475 Adult VirginFemale Spermatheca    155.8 Adult InseminatedFemale Spermatheca    148 Adult Ovary    59.2 Adult Testis    22.1 Adult Male Accessory Gland    60.3 Adult Carcass    31.6 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    94.3   NA  Eye    59   NA  Brain    284.1   232.6  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    243.2   NA  Crop    47.9   41.9  Midgut    31.6   41.7  Hindgut    33.3   68  Malpighian Tubules    51.5   30.5  Fat Body    120.8   73.2  Salivary Gland    55.4   NA  Heart    40.475   34.425  Trachea    NA   NA  VirginFemale Spermatheca    155.8   NA  InseminatedFemale Spermatheca    148   NA  Ovary    59.2   NA  Testis    22.1   NA  Male Accessory Gland    60.3   46.225  Carcass    31.6 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_30
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0002774 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG11680-RB
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal mle9 lethal | larval stage | male mle1 mle8 mle15 lethal | larval stage | male | recessive mle1 lethal | male mle1 mle2 mle5 mle9 mle10 mle11 mle12 mle13 mle14 mle15 lethal | male | heat sensitive mle1 mle4 mle6 lethal | male | partially mleGET lethal | male | pupal stage mle1 mle15 lethal | male | recessive mle1 mle1a mle1b mle3 mle4 mle10 mle11 mle12 mle13 mle14 mle15 mleI926Z mleZ1536 mleZ3543 mleZ5317 lethal | male limited | recessive mleCo lethal | third instar larval stage mle1 semi-lethal | pupal stage, with Scer\GAL4pnr-MD237 mleGD1488 semi-viable | male mleGNT.T:Ivir\HA1 viable mlee04702 viable | male | poor mleDQIH.T:Ivir\HA1 Sterility Allele fertile mlee04702 male fertile | poor mleDQIH.T:Ivir\HA1 male sterile mle1 mle4 mle6 Other Phenotypes Allele behavior defective mlenap-ts1 behavior defective | recessive | heat sensitive mlenap-rv1 mlenap-ts1 mlenap-ts2 mlenap-ts3 mlenap-ts4 mlenap-ts5 mlenap-ts6 mlenap-ts7 chemical resistant mlenap-ts1 chemical sensitive mlenap-ts1 chemical sensitive | heat sensitive mlenap-ts1 chemical sensitive | partially | recessive mlenap-ts1 locomotor behavior defective | third instar larval stage mlenap-ts1 neuroanatomy defective | heat sensitive mlenap-ts1 neuroanatomy defective | third instar larval stage mlenap-ts1 neurophysiology defective mlenap-ts1 neurophysiology defective | heat sensitive mlenap-ts1 neurophysiology defective | recessive | heat sensitive mlenap-ts1 paralytic mlenap-ts1 paralytic | heat sensitive mlenap-ts1 paralytic | recessive | heat sensitive mlenap-ts1 short lived mlenap-ts1 short lived | recessive | heat sensitive mlenap-ts1 song defective | recessive mlenap-ts1 wild-type mle+20.T:Ivir\HA1 mle+t10.5 mleAAA.T:Ivir\HA1 mleATG.T:Ivir\HA1 mleDQIH.T:Ivir\HA1 mleGNT.T:Ivir\HA1 mlehs.PR mlehs.T:Ivir\HA1 mleSTOP.T:Ivir\HA1 mlets.T:Ivir\HA1 Phenotype manifest in Allele abdomen & macrochaeta mle1 adult brain mlenap-ts1 axon & motor neuron | conditional ts mlenap-ts1 chromosome mle1 embryonic/larval heart | heat sensitive mlenap-ts1 embryonic/larval neuromuscular junction mlenap-ts1 giant fiber neuron mlenap-ts1 neuromuscular junction & synapse | conditional ts mlenap-ts1 NMJ bouton mlenap-ts1 polytene chromosome mle8 spermatozoon mle4 spermatozoon | heat sensitive mle1 mle6 synapse | ectopic | heat sensitive mlenap-ts1 wing mle1
Classical Alleles ( 34 )
For All Classical Alleles Show
Allele of mle Class Mutagen Stocks Known lesion mle1 loss of function allele spontaneous natural population 2 Yes mle9 loss of function allele gamma ray 2 Yes mlee04702 piggyBac transposase 2 -- mleMB11906 1 -- mle10 gamma ray 0 Yes mle11 gamma ray 0 Yes mle12 gamma ray 0 -- mle13 gamma ray 0 Yes mle14 gamma ray 0 Yes mle15 ethyl methanesulfonate 0 Yes mle1a spontaneous 0 -- mle1b spontaneous 0 -- mle2 spontaneous 0 Yes mle3 ethyl methanesulfonate 0 -- mle4 ethyl methanesulfonate 0 Yes mle5 spontaneous 0 Yes mle6 ethyl methanesulfonate 0 -- mle7 gamma ray ethyl methanesulfonate 0 -- mle8 gamma ray ethyl methanesulfonate 0 Yes mleck 0 -- mleCo natural population 0 -- mleI926Z ethyl methanesulfonate 0 Yes mlenap-rv1 gain of function allele gamma ray 0 -- mlenap-ts1 gain of function allele ethyl methanesulfonate 0 Yes mlenap-ts2 gain of function allele PM hybrid dysgenesis 0 Yes mlenap-ts3 gain of function allele ethyl methanesulfonate gamma ray 0 Yes mlenap-ts4 gain of function allele ethyl methanesulfonate 0 Yes mlenap-ts5 gain of function allele ethyl methanesulfonate 0 Yes mlenap-ts6 gain of function allele ethyl methanesulfonate 0 -- mlenap-ts7 gain of function allele gamma ray 0 Yes mleunspecified 0 -- mleZ1536 ethyl methanesulfonate 0 -- mleZ3543 ethyl methanesulfonate 0 -- mleZ5317 ethyl methanesulfonate 0 --
Alleles Carried on Transgenic Constructs ( 17 )
For All Alleles Carried on Transgenic Constructs Show
Allele of mle Class Mutagen Stocks Known lesion mleGD1488 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes mleHMS00182 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes mle+20.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes mle+t10.5 in vitro construct - genomic fragment 0 Yes mleAAA.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mleATG.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes mleDQIH.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mledsRNA.N in vitro construct - RNAi 0 Yes mleGET in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mleGNT.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mlehs.PR in vitro construct - regulatory fusion 0 Yes mlehs.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes mleMtnA.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes mleRK.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mleSTOP.T:Ivir\HA1 in vitro construct - coding region fusion 0 Yes mlets.T:Ivir\HA1 in vitro construct - coding region fusion in vitro construct - site directed mutagenesis in vitro construct - amino acid replacement 0 Yes mleΔgly in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)Exel7085 (Ryder, 2004.4.26) Df(2R)G-2 (Dowse et al., 1995) Df(2R)nap2 (Ringo et al., 1991) Df(2R)nap3 (Ringo et al., 1991) In(2R)bwVDe2LCyR (Cicchetti and Loverre, 1989, Kuroda et al., 1991)
Not disrupted in	Df(2L)C (Cicchetti and Loverre, 1989) Df(2L)C' (Cicchetti and Loverre, 1989) Df(2L)D (Cicchetti and Loverre, 1989) Df(2L)D' (Cicchetti and Loverre, 1989) Df(2L)TW65 (Cicchetti and Loverre, 1989) Df(2L)TW161 (Cicchetti and Loverre, 1989) Df(2R)42 (Cicchetti and Loverre, 1989) Df(2R)M41A4 (Cicchetti and Loverre, 1989) Df(2R)M41A8 (Cicchetti and Loverre, 1989) Df(2R)M41A10 (Cicchetti and Loverre, 1989) Df(2R)rl-D (Cicchetti and Loverre, 1989) Df(2R)stmA-12 (Chandrashekaran and Sarla, 1993)
Duplicated in	Dp(1;4)r+l (Reenan and Rogina, 2008)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1488} NA P{TRiP.HMS00182} NA heat-shock construct P{hsHA-COOH} NA P{hsMLE25} NA characterization construct P{HA+20} NA P{HA-AAA} NA P{HA-ATG} NA P{HA-DQIH} NA P{HA-GNT} NA P{HA-RK} NA P{HA-STOP} NA P{HA-ts} NA P{mle.Δgly} NA P{mle+T6.1} NA P{mle-GET} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{}mlenap-ts2 NA PBac{RB}mlee04702 NA insertion of enhancer trap binary system Mi{ET1}mleMB11906 No
Gene Ontology: Function, Process & Cellular Component ( 17 unique terms )
Terms Based on Experimental Evidence ( 7 terms )
Molecular Function
CV term Evidence References
chromatin binding inferred from direct assay (Bone et al., 1994)
Biological Process
CV term Evidence References
axon extension inferred from mutant phenotype (Zhong and Wu, 2004) determination of adult lifespan inferred from mutant phenotype (Fergestad et al., 2006) dosage compensation inferred from mutant phenotype (Bone et al., 1994) male courtship behavior, veined wing generated song production inferred from genetic interaction with cac (Peixoto and Hall, 1998)
Cellular Component
CV term Evidence References
chromatin inferred from direct assay (Bone et al., 1994) X chromosome inferred from direct assay (Kotlikova et al., 2006, Smith et al., 2008)
Terms Based on Predictions or Assertions ( 11 terms )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR001650, InterPro:IPR002464, InterPro:IPR011545 (FlyBase Curators et al., 2004-) ATP-dependent helicase activity inferred from electronic annotation with InterPro:IPR002464, InterPro:IPR011545 (FlyBase Curators et al., 2004-) traceable author statement (Bone et al., 1994) double-stranded RNA binding inferred from electronic annotation with InterPro:IPR001159 (FlyBase Curators et al., 2004-) inferred from sequence or structural similarity (Lasko, 2000) helicase activity inferred from sequence or structural similarity (Lasko, 2000) RNA helicase activity non-traceable author statement (Wutz, 2003)
Biological Process
CV term Evidence References
dosage compensation non-traceable author statement (Swiss-Prot Project Members, 1992.3.1) traceable author statement (Williamson and Lehmann, 1996) dosage compensation, by hyperactivation of X chromosome non-traceable author statement (FlyBase, 1992-, Williamson and Lehmann, 1996, Schutt and Nothiger, 2000) 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, 1992.3.1) nucleus non-traceable author statement (Swiss-Prot Project Members, 1992.3.1) 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 mle allele Gene References mle1 ovo (Oliver et al., 1993, Oliver and Pauli, 1998) Sxl (Skripsky and Lucchesi, 1982, Uenoyama et al., 1982) mle4 Sxl (Skripsky and Lucchesi, 1982) mle5 ovo (Oliver et al., 1993, Oliver and Pauli, 1998) mle9 ovo (Oliver and Pauli, 1998) vir (Hilfiker et al., 1995) mle14 ovo (Oliver and Pauli, 1998) mlenap-ts1 cac (Peixoto and Hall, 1998) eag (Wang et al., 2000) Khc (Hurd et al., 1996) para (Ganetzky, 1984, Dowse et al., 1995) unspecified bas (Ganetzky and Wu, 1982) bss (Ganetzky and Wu, 1982) da (Cline, 1986) eag (Budnik et al., 1990) eas (Ganetzky and Wu, 1982) Hk (Ganetzky and Wu, 1982) kdn (Ganetzky and Wu, 1982) para (Wu and Ganetzky, 1980) Sh (Ganetzky and Wu, 1982, Budnik et al., 1990, Wang et al., 2000) shi (Johnson et al., 2001) tipE (Jackson et al., 1986, Ganetzky, 1986) tko (Ganetzky and Wu, 1982)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 61429 DroID - A comprehensive database of gene and protein interactions. • FBgn0002774 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35523
Orthologs
Genome-wide drosophilid orthologs	Dana\GF13828 Dere\GG10841 Dgri\GH20409 Dmoj\GI19596 Dper\GL17788 Dpse\GA11141 Dsec\GM16493 Dsim\GD10344 Dvir\GJ18398 Dwil\GK21754 Dyak\GE19435
Curated drosophilid orthologs	Dmir\mle   Dpse\GA11141   Dpse\mle (Bachtrog, 2003)
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL004859-PA Aedes aegypti (mosquito) AGAP006599-PA Anopheles gambiae (mosquito) XP_396525.2 Apis mellifera (honey bee) XP_001943592.1 Acyrthosiphon pisum (pea aphid) 124637 Branchiostoma floridae (lancelet) BGIBMGA010840-PA Bombyx mori (silk worm) ENSBTAP00000026409 Bos taurus (bovine) CN00840 Caenorhabditis brenneri (small nematode) CBP00773 Caenorhabditis briggsae (nematode) CE39177 Caenorhabditis elegans (worm) ENSCAFP00000019476 Canis familiaris (dog) ENSCINP00000015974 Ciona intestinalis (sea squirt) JA01841 Caenorhabditis japonica (nematode) cn13189 Cryptococcus neoformans (yeast-like fungus) RP26542 Caenorhabditis remanei (nematode) ENSCSAVP00000002531 Ciona savignyi (sedentary tunicate) 220258 Capitella spl (polychaete worm) DDB0233740 Dictyostelium discoideum (amoeba) 329366 Daphnia pulex (daphnia) 64508 Hypsipyla robusta (cedar shoot borer) ENSP00000356520 Homo sapiens (human) ISCW009103-PA Ixodes scapularis (deer tick) 116243 Lottia gigantea (owl limpet) ENSMODP00000008627 Monodelphis domestica (gray opossum) ENSMMUP00000007450 Macaca mulatta (rhesus monkey) ENSMUSP00000038135 Mus musculus (house mouse) 121518 Nematostella vectensis (starlet anemone) XP_001604965.1 Nasonia vitripennis (wasp) ENSOANP00000001567 Ornithorynchus anatinus (platypus) ENSORLP00000009349 Oryzias latipes (japanese killifish) PHUM005676-PA Pediculus humanus (human body louse) ENSPPYP00000000490 Pongo pygmaeus (orangutan) ENSPTRP00000002937 Pan troglodytes (chimpanzee) ENSRNOP00000003707 Rattus norvegicus (norway rat) Smp_035030 Schistosoma mansoni (trematode parasite) 20896 Trichoplax adhaerens (metazoa) XP_973454.1 Tribolium castaneum (red flour beetle) ENSTNIP00000007003 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000004908 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000027390 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16128 NV14917 LH19851 ISCW009103 FBgn0236778 FBgn0223739 FBgn0205557 FBgn0182116 FBgn0171408 JGI_V11_329366 FBgn0071196 FBgn0155391 FBgn0142333 FBgn0127872 FBgn0103146 FBgn0090855 CPIJ013504 BGIBMGA010840 ACYPI003650 ACYPI003167 ACYPI000545 GB14139 AGAP006599 AAEL004859 PHUM607970 GLEAN_03184
Stocks & Reagents
Stocks Listed in FlyBase ( 9 )
Bloomington	18284 w1118; PBac{RB}mlee04702 29260 w1118; Mi{ET1}mleMB11906 5873 mle9 cn1 bw1/CyO 4235 y1 w1 Nspl-1/Dp(1;Y)BS; mle1/SM1 34864 y1 sc* v1; P{TRiP.HMS00182}attP2/TM3, Sb1
Harvard	- PBac{RB}mlee04702
Kyoto	108473 mle9 cn1 bw1/CyO 107647 y1 w1 Nspl-1/Dp(1;Y)BS; mle1/SM1
VDRC	v19691 w1118; P{GD1488}v19691
Genomic Clones ( 2 )
	BACR03L10 BACR39E09 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 19 )
	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	LD44547 RE21725
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	GM25238 HL02337 LD41112 RE20562 SD02678 SD18044
Other clones	45992 103333 105735 CK01.50C.F10 Dmel_neb_013623_0395_0563 Dmel_neb_013843_0081_2688 Dmel_neb_014912_0131_1727 EK040455 EK162334 EK243041 RH17678
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0002774 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35523
Antibody Information
	polyclonal (Kuroda et al., 1991, Bone et al., 1994)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: mle CG11680 (Li, 1999.11)
Source for database merge of
Additional comments	The paralysis of 'nap' is not complemented by mle alleles. (Lindsley and Zimm, 1992) "mle" alleles fail to complement the paralysis caused by "nap" alleles, indicating that "nap" and "mle" are allelic. (Kernan et al., 1991) Molecular analysis demonstrated that "mle" mutants are allelic to "nap" mutants. (Kuroda et al., 1991)
Other Comments
	The mle protein interacts with the most prominent transcriptionally active regions of chromosomes independently of other MSL proteins. (Kotlikova et al., 2006) More than 80% of para transcripts in a mlenap-ts1 background are aberrant, owing to internal deletions that include the edited para exon. (Reenan et al., 2000) The mlenap-ts1 mutation results in the occurrence of a "splicing catastrophe" of the para transcript in the region of the RNA editing site. (Reenan et al., 2000) 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) Used as a 'bait' in the yeast two-hybrid system to screen for interactors from an imaginal disc cDNA library: Dbp80 is identified. (Eisen et al., 1998) 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. mle is necessary but only as a structural component for the recruitment of mof to the X chromosome. (Gu et al., 1998) In the germline mle is not involved in chromosomal dosage compensation but may be involved in post-transcriptional gene regulation. Loss of mle has no detectable effect on expression or localisation of acetylated His4. (Rastelli and Kuroda, 1998) mle has NTPase and both RNA and DNA helicase activities. (Lee et al., 1997) The NTPase/helicase activities of mle are essential for dosage compensation. (Lee et al., 1997) Loss of mle NTPase and helicase activities results in male lethality without elimination of the localisation of the mle-msl-1 complex to the male X chromosome. (Lee 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) The roX1 gene is positively regulated by genes of the dosage compensation system such as mle. (Meller et al., 1997) X chromosome proteins associated with dosage compensation in melanogaster are sufficiently conserved to allow significant antibody cross-reaction to D.simulans, D.virilis, D.americana.americana and D.pseudoobscura.pseudoobscura chromosomes. Cross reaction is also observed in the X chromosome and the X2 chromosome (2 copies in females and 1 in males) of D.miranda. These results provide evidence that the male-specific lethal proteins can be acquired on previously unrelated chromosome arms during evolution. (Bone and Kuroda, 1996) Male-specific lethal (MSL) proteins accumulate in a subregion of male nuclei (the X chromosome) beginning at late blastoderm stage. X chromosomal binding of the MSLs is observed throughout embryonic and larval development in both diploid and polytene tissues. His4 colocalises with the MSLs in embryos. Binding of the MSLs is interdependent in diploid cells and is prevented in female embryonic cells by Sxl. (Franke et al., 1996) Mutations in Khc enhance the para and mel and suppress the Sh and eag mutant phenotypes. (Hurd et al., 1996) Association of mle with the polytene X chromosome is RNase sensitive and mutations in the ATPase motifs affect mle function. The carboxyl terminus of mle may have a potential role in general affinity to RNA. (Richter 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) Heartbeat of mlenap-ts1 individuals is seriously impaired, becoming arrhythmic at elevated temperatures. (Dowse et al., 1995) Immunostaining of embryonic and larval stages demonstrates that His4, msl-1 and msl-3 are associated with the male X chromosome as early as gastrulation, while mle binding is not detected until the late embryonic/late larval stages. (Franke and Baker, 1995) The msl-3, mle and msl-1 gene products may associate with one another in a male-specific heteromeric complex on the X chromosome to achieve its hyperactivation. (Gorman et al., 1995) The expression pattern of mle suggests mle has a maternal component that appears at the beginning of embryogenesis and localises to the male X chromosome at stage 8. (Rastelli and Kuroda, 1995) Elements needed for dosage compensation are localised to the X chromosome only after blastoderm and msl-dependent dosage compensation is not necessary during the first part of embryogenesis. This suggest the existance of an additional msl-independent dosage compensation mechanism; dosage compensation of run expression at blastoderm is not dependent on male specific lethal genes. (Rastelli and Kuroda, 1995) The binding of mle, msl-1, msl-2 and His4 proteins to the X chromosome are interdependent from early embryogenesis. (Rastelli et al., 1995) The msl-2 primary transcript may play a role in male specific binding of mle, msl-3 and msl-1 to the X chromosome. (Zhou et al., 1995) The gene products of mle, msl-1 bind to the male X chromosome in an identical pattern, and the binding sites of H4Ac16 acetylated form of the His4 product are largely coincident with the mle/msl-1 binding sites. 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) Antisera to mle protein label the euchromatic X chromosome through mitosis, but neither the X heterochromatin nor autosomes. (Lavender et al., 1994) msl-1, like mle and H4Ac16 (an acetylated form of the His4 product), exhibits a wild type male localisation pattern in Sxl- XX nuclei. (Palmer et al., 1994) In females, the Sxl product functions to prevent mle from binding to the two X chromosomes. The X chromosome binding of mle requires wild type msl-1, msl-2 and msl-3 functions. (Gorman et al., 1993) mle was identified by two very different mutant phenotypes, male-lethality ('mle') and rapid paralysis of larvae or adults when exposed to 37oC ('napts'). (Lindsley and Zimm, 1992) The 'mle' group phenotype: Mutants are defective for dosage compensation in males. Homozygous males die, but homozygous females survive. Males produced by homozygous females die during the third larval instar, whereas those produced by heterozygous females are late pupal lethals. Females transformed into phenotypic males (tra1) or intersexes (dsx1) unaffected by mle1, i.e. mle acts only upon single-X-bearing flies. mle4 males surviving at 18oC are sterile, small and slow developing. Polytene X chromosome of mle1 males appears narrower and more densely stained than that of control males. The 'nap' group phenotype: Larvae or adults become rapidly paralyzed when exposed to 37oC and rapidly recover on return to lower temperatures. (Lindsley and Zimm, 1992) mle plays a direct role in dosage compensation of the X chromosome. (Kuroda et al., 1991) Slight reduction in the extent of branching caused by mlenap-ts1 at permissive temperature; the increase in branching (and higher than normal number of varicosities on motor-neurites) induced by an eag Sh double mutant was suppressed by mlenap-ts1 (re. low-temperature rearing). (Budnik et al., 1990) Action potentials in the giant fiber (GF) pathway of adults are not blocked at temperatures up to 43oC. The long-latency phenotype disappears as the temperature is raised to 35oC. (Elkins and Ganetzky, 1990) Experiments involving one-time rearing at low temperature caused mlenap-ts1 to paralyze at relatively low temperatures. Action potentials in the giant fiber (GF) pathway of adults are not blocked at temperatures up to 43oC, though the latency from brain stimulation to response of thoracic muscles are aberrantly long, even at low temperatures. 'Following frequency' of mlenap-ts1 thoracic muscle responses (re. GF pathway stimulation) is reduced at elevated temperatures, an effect which can be reversed by injection of 4-amino-pyridine. (Nelson and Wyman, 1990) Two doses of para+ (in males) suppresses high-temperature paralysis of mlenap-ts1. (Stern et al., 1990) Mutation does not seem to modify the expression of sodium currents in embryonic neurons. (O'Dowd and Aldrich, 1988) mle1 pole cell transplanted into wild-type hosts are incapable of undergoing normal spermatogenesis. (Bachiller and Sanchez, 1986) Steady-rate level of Sgs4 mRNA incompletely compensated in mle4 and mle4/mle6 male larvae. (Breen and Lucchesi, 1986) In mosaic experiments, cuticular clones of parats1 in a mlenap-ts1 background (after low-temperature development) have non-functioning sensory cells, probably due to lack of nerve conduction which, however, did not cause any anatomical abnormalities involving the central projections of these sensory neurons. (Burg and Wu, 1986) mlenap-ts1 in combination with tipE1 leads to poor viability at permissive temperature. (Jackson et al., 1986, Ganetzky, 1986) Rearing stocks chronically at room temperature or above causes mlenap-ts1 to 'adapt' such that higher temperatures (> 40oC) are required for paralysis. Exposure of mlenap-ts1 males to high temperature causes arrest of oscillator underlying rhythmic component of courtship song. (Kyriacou and Hall, 1985) In experiments on conditioned courtship, mlenap-ts1 males learn normally but have shortened memory spans and mlenap-ts1 suppresses Sh-induced decrements in courtship learning. (Cowan and Siegel, 1984, Cowan and Siegel, 1986) mle and para are involved in the function of sodium channels. (Ganetzky, 1984) Genetic, electrophysiological, behavioral and pharmacological studies of mlenap and para mutants suggest that they effect sodium channels. (Ganetzky, 1984) Brain membrane extracts of mlenap-ts1, assayed at low or high temperatures, have subnormal levels of saxitoxin, though there are no qualitative alterations of this binding activity (kD is normal). (Jackson et al., 1984) Cultured neurons from mlenap-ts1 larvae are 4 to 5-fold more resistant than wild-type cells to killing effects of veratridine, irrespective of temperature (22oC vs 35oC). (Suzuki and Wu, 1984) Mutants show no interaction with msl-11 or msl-21. (Belote, 1983) Cultured neurons from mlenap-ts1 larvae are not affected by TTX, their general growth characteristics are normal. (Wu et al., 1983) Brain membrane extracts of mlenap-ts1, assayed at low or high temperatures, have subnormal levels of tetrodotoxin. (Kauvar, 1982) Few homozygous mle1 gynandromorphs survive; X0 patches small, with small bristles, and mostly confined to abdomen. (Uenoyama et al., 1982) mle 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 mle4 male larvae when compared with non-msl controls. (Belote and Lucchesi, 1980) The incorporation of labeled uridine by the polytene X chromosome relative to that of 2R is lower than normal in mle4 males. (Belote and Lucchesi, 1980) Axonal conduction (but not synaptic transmission) fails in mutant larvae at high temperatures. At permissive temperatures, refractory period for elicitation of a series of action potentials is abnormally long and mutants are hypersensitive to blocking effects of tetrodotoxin (TTX) on action potentials. mlenap-ts1 is unconditionally lethal in a double mutant with parats1 (death occurring during 1st larval instar). (Wu and Ganetzky, 1980) Axonal conduction (but not synaptic transmission) fails in larvae at high temperatures. (Wu et al., 1978)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057293 NM_057294 NM_165451 RefSeq (Proteins) • NP_476641 NP_476642 NP_724440 DDBJ / EMBL / GenBank • AAC41573 AAF57297 AAF57297.1 AAM68335 AAM68336 AAM68336.2 AAQ23585 AC009847 AI512625 AI531582 BI215346 BT010267 M74121 Entrez Gene - A searchable database of RefSeq genes. • 35523 UniProtKB/Swiss-Prot • P24785 UniProtKB/TrEMBL • A1Z6I8
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Double-stranded RNA-binding (IPR001159) Helicase, C-terminal (IPR001650) DNA/RNA helicase, ATP-dependent, DEAH-box type, conserved site (IPR002464) Helicase-associated domain (IPR007502) Domain of unknown function DUF1605 (IPR011709) Double-stranded RNA-binding-like (IPR014720) DEAD-like helicase (IPR014001) DNA/RNA helicase, DEAD/DEAH box type, N-terminal (IPR011545)
Linkouts
	BioGRID - A database of protein and genetic interactions • 61429 DroID - A comprehensive database of gene and protein interactions. • FBgn0002774 DRSC - Results from RNAi screens. • FBgn0002774 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0002774 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG11680-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0002774 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 35523 InParanoid A subset of ortholog calls from InParanoid. • AAEL004859-PA Aedes aegypti (mosquito) AGAP006599-PA Anopheles gambiae (mosquito) XP_396525.2 Apis mellifera (honey bee) XP_001943592.1 Acyrthosiphon pisum (pea aphid) 124637 Branchiostoma floridae (lancelet) BGIBMGA010840-PA Bombyx mori (silk worm) ENSBTAP00000026409 Bos taurus (bovine) CN00840 Caenorhabditis brenneri (small nematode) CBP00773 Caenorhabditis briggsae (nematode) CE39177 Caenorhabditis elegans (worm) ENSCAFP00000019476 Canis familiaris (dog) ENSCINP00000015974 Ciona intestinalis (sea squirt) JA01841 Caenorhabditis japonica (nematode) cn13189 Cryptococcus neoformans (yeast-like fungus) RP26542 Caenorhabditis remanei (nematode) ENSCSAVP00000002531 Ciona savignyi (sedentary tunicate) 220258 Capitella spl (polychaete worm) DDB0233740 Dictyostelium discoideum (amoeba) 329366 Daphnia pulex (daphnia) 64508 Hypsipyla robusta (cedar shoot borer) ENSP00000356520 Homo sapiens (human) ISCW009103-PA Ixodes scapularis (deer tick) 116243 Lottia gigantea (owl limpet) ENSMODP00000008627 Monodelphis domestica (gray opossum) ENSMMUP00000007450 Macaca mulatta (rhesus monkey) ENSMUSP00000038135 Mus musculus (house mouse) 121518 Nematostella vectensis (starlet anemone) XP_001604965.1 Nasonia vitripennis (wasp) ENSOANP00000001567 Ornithorynchus anatinus (platypus) ENSORLP00000009349 Oryzias latipes (japanese killifish) PHUM005676-PA Pediculus humanus (human body louse) ENSPPYP00000000490 Pongo pygmaeus (orangutan) ENSPTRP00000002937 Pan troglodytes (chimpanzee) ENSRNOP00000003707 Rattus norvegicus (norway rat) Smp_035030 Schistosoma mansoni (trematode parasite) 20896 Trichoplax adhaerens (metazoa) XP_973454.1 Tribolium castaneum (red flour beetle) ENSTNIP00000007003 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000004908 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000027390 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /polycomb/malels1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 35523 modMine - Data generated by the modENCODE project. • FBgn0002774 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16128 NV14917 LH19851 ISCW009103 FBgn0236778 FBgn0223739 FBgn0205557 FBgn0182116 FBgn0171408 JGI_V11_329366 FBgn0071196 FBgn0155391 FBgn0142333 FBgn0127872 FBgn0103146 FBgn0090855 CPIJ013504 BGIBMGA010840 ACYPI003650 ACYPI003167 ACYPI000545 GB14139 AGAP006599 AAEL004859 PHUM607970 GLEAN_03184
Synonyms & Secondary IDs ( 16 )
Reported As
Symbol Synonym	CG11680 (Fulton et al., 2001, Ni et al., 2011.7.19) mak   MLE (Nusinow, 2005, Kelley, 2004, Rattner and Meller, 2001, Kelley and Kuroda, 2000, Copps et al., 1998, Panning and Jaenisch, 1998, Franke and Baker, 1995, Straub and Becker, 2007, Dahlsveen et al., 2006, Mendjan et al., 2006, Morra et al., 2007, Gilfillan et al., 2006, Lucchesi et al., 2007, Prasanth and Spector, 2007, Furuhashi et al., 2006, Deng and Meller, 2006, Mendjan and Akhtar, 2007, Park et al., 2007, Smith et al., 2008, Patalano et al., 2009, Izzo et al., 2008, Straub et al., 2008, Li et al., 2005, Buscaino et al., 2006, Roy et al., 2011, Schiemann et al., 2010) mle (Deng and Meller, 2006, Ray and Fox, 2005, Schubeler, 2006, Song and Tanouye, 2007, Feng et al., 2004, Kuebler and Tanouye, 2000, Kind and Akhtar, 2007, Rodriguez et al., 2007, Morra et al., 2008, Worringer and Panning, 2007, Li et al., 2008, Levine et al., 2007, Doheny et al., 2008, Spierer et al., 2005, Pierre et al., 2008, Sandstrom, 2008, Worringer et al., 2009, Zhimulev et al., 2003, Carré et al., 2008, Reenan and Rogina, 2008, Kuebler et al., 2001, Lee and Wu, 2006, Spierer et al., 2008, Barbash, 2010, Corona et al., 2002, Prabhakaran and Kelley, 2010, Di Stefano et al., 2011, Gladstein et al., 2010, Gleason, 2005) mll   mls (Lukacsovich et al., 1999) nap (Johnson et al., 2001, Hebbar and Fernandes, 2001, Greenspan and Ferveur, 2000, Wang et al., 2000, Lee and Wu, 2000, Yamamoto and Nakano, 1999, Tolar and Pallanck, 1998, Peixoto and Hall, 1998, Budnik, 1996, Dowse et al., 1995, Morgan, 1996, Leibovitch et al., 1995, Jarecki and Keshishian, 1995, Pauron and Castella, 1995, Leibovitch et al., 1993, Hall et al., 1982, Burg et al., 1993, Palka, 1993, Engel and Wu, 1992, Budnik et al., 1990, Elkins and Ganetzky, 1990, Nelson and Wyman, 1990, Stern et al., 1990, Sakai et al., 1989, Bloomquist et al., 1989, Jackson et al., 1984, Cowan and Siegel, 1984, Suzuki and Wu, 1984, Ganetzky, 1984, Ganetzky and Wu, 1982, Wang et al., 2002, Hebbar and Fernandes, 2004, Zhong and Wu, 2004, Lnenicka et al., 2003, Hebbar et al., 2006, Reenan and Rogina, 2008, Lee and Wu, 2010) napts (Mehren et al., 2004, Mir and Krishnan, 1995, Lindsley and Zimm, 1992, )
Name Synonym	male-killer   maleless (Prasanth and Spector, 2007, Izzo et al., 2008, Worringer et al., 2009, Kuebler et al., 2001, Prabhakaran and Kelley, 2010, Lee and Wu, 2010, Gleason, 2005) Maleless (Cho et al., 2005, Deng and Meller, 2006, Patalano et al., 2009, Gerbasi et al., 2010) maleless-no-action potential (Kuebler and Tanouye, 2000) male-lethal   male lethal (Lee and Wu, 2006) no action potential (Johnson et al., 1998, ) no-action-potential (Wang et al., 2002, Reenan and Rogina, 2008)
Secondary FlyBase IDs
References ( 309 )
Generate a list of	All references relating to this gene ( 309 )
List References by type	Research paper  ( 178 ) Supplementary material  ( 3 ) Review  ( 63 ) Personal communication to FlyBase  ( 6 ) Abstract  ( 49 ) FlyBase analysis  ( 1 ) Letter  ( 2 ) Book  ( 1 ) Curated genome annotation  ( 1 ) Protein sequence record  ( 1 ) Computer file  ( 2 ) Automatic genome annotation  ( 1 )
Recent research papers ( 8 )
	Di Stefano et al., 2011, Genes Dev. 25(1): 17--28 Functional antagonism between histone H3K4 demethylases in vivo. [FBrf0212709] Roy et al., 2011, Mol. Genet. Genomics 285(2): 113--123 Lack of the Drosophila BEAF insulator proteins alters regulation of genes in the Antennapedia complex. [FBrf0212912] Barbash, 2010, Genetics 184(1): 313--316 Genetic testing of the hypothesis that hybrid male lethality results from a failure in dosage compensation. [FBrf0209660] Gerbasi et al., 2010, Mol. Cell. Proteomics 9(9): 1866--1872 Proteomics Identification of Drosophila Small Interfering RNA-associated Factors. [FBrf0211707] 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] Lee and Wu, 2010, J. Neurosci. 30(47): 15821--15833 Orchestration of stepwise synaptic growth by k+ and Ca2+ channels in Drosophila. [FBrf0212405] 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] 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] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...