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Gene Dmel\fwd

General Information
Symbol	Dmel\fwd	Species	D. melanogaster
Name	four wheel drive	Annotation symbol	CG7004
Feature type	protein_coding_gene	FlyBase ID	FBgn0004373
Gene Model Status	Current	Stock availability	12 publicly available
Genomic Location
Chromosome (arm)	3L	Recombination map	3-
Cytogenetic map	61C1-61C1	Sequence location	3L:306,207..319,880 [-]
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 four wheel drive is referred to in FlyBase by the symbol Dmel\fwd (CG7004, FBgn0004373). It is a protein_coding_gene from Drosophila melanogaster. Based on sequence similarity, it is predicted to have molecular function: 1-phosphatidylinositol 4-kinase activity. There is experimental evidence that it is involved in the biological process: male meiosis cytokinesis; contractile ring contraction involved in cell cycle cytokinesis; male germline ring canal formation; determination of adult lifespan; meiotic spindle stabilization; cytokinesis after meiosis II; spermatocyte division; spermatogenesis; cytokinesis after meiosis I. 24 alleles are reported. The phenotypes of these alleles are annotated with: secondary spermatocyte; Nebenkern; Nebenkern derivative; spermatid; spermatid axoneme; spermatozoon. It has 3 annotated transcripts and 3 annotated polypeptides. Protein features are: Phosphatidylinositol 3-/4-kinase, catalytic; Phosphatidylinositol 3/4-kinase, conserved site; Phosphatidylinositol Kinase; Protein kinase-like 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 at stages throughout embryogenesis, during late larval stages, at stages throughout the pupal period, in stages of adults of both sexes. Summary of FlyAtlas Anatomical Expression Data: Nearly all larval and adult tissues/organs expressed at moderate levels. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval/adult salivary gland, larval trachea, adult female reproductive system, adult male reproductive system, adult carcass. Comments on Affy2 ProbeSet: ProbeSet 1634509_s_at completely aligns to an exonic region common to each of the 3 FlyBase-annotated transcript isoforms of fwd. Gene sequence location is 3L:306207..319880.
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	fwd: four wheel drive (M. Fuller) Recessive male sterile. Failure of cytokinesis after meiosis I and meiosis II in males, usually resulting in onion stage early spermatids with four equal sized nuclei associated with a single, large mitochondrial derivative. Cells in meiosis II contain two spindles. Females fertile.
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
FB2012_01	Insertions Mi{MIC}fwd[MI04139] Alleles fwd[MI04139] Sequence features BKN50050 BKN22161 BKN50849 HFA08580 References Bellen, 2011.11.10, set958.xml
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 61C1-61C1   Limits computationally determined from genome sequence between P{PZ}trh10512 and P{EP}EP3104EP3104
Experimentally Determined Cytological Location
Cytogenetic map Notes References 61D-61D   (determined by in situ hybridisation)   (Berg and Spradling, 1991) 61B1-61C1   Location based on deficiency mapping.   (Giansanti et al., 2004)
Experimentally Determined Recombination Data
Location	3- (Giansanti et al., 2004)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\fwd 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) fwd-RA FBtr0072516  NM_080083   6357   1674 fwd-RB FBtr0072515  NM_167826   5601   1674 fwd-RC FBtr0072517  NM_167827   4434   1338
Additional Transcript Data & Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein fwd-PA   FBpp0072416   182.0   1674   7.00     NP_524822   AAF47375 fwd-PB   FBpp0072415   182.0   1674   7.00     NP_728518   AAN11437 fwd-PC   FBpp0072417   147.2   1338   6.93     NP_728519   AAN11438
Additional Polypeptide Data & Comments
Reported size (kDa)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Phosphatidylinositol 3/4-kinase, conserved site (IPR018936) Phosphatidylinositol 3-/4-kinase, catalytic (IPR000403) Protein kinase-like domain (IPR011009) Phosphatidylinositol Kinase (IPR015433)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA246711   AA263708   AA264278   AA567194   AA696613   AA941468   AA949868   AE014296 AAF47375   AE014296 AAN11437   AE014296 AAN11438   AF242375 AAK27793   AI114377   AI257794   AI293824   AI295515   AI297362   AI518802   AI520161   AW944068   AX094135   AY052049 AAK93473   BF488884   BF497062   BG632603   BG637836   BG637878   BG638448   BG638875   BI353325   BI362230   BI364104   BI364186   CL527805   CO180861   CO187781   CO196066   CO265607   CO267766   CO273392   CO273775   CO274815   CO278159   CO290876   CO295560   CO296251   CO297873   CO304936   CO305571   CO309192   CO317295   CO318931   CO320817   CO322601   CO326388   CO330257   CO332623   CO333994   CO339716   CO339771   CZ486416   EC057195   EC060633   EC061370   EC065779   EC066853   EC067446   EC070419   EC073982   EC078110   EC086955   EC087299   EC090799   EC090992   EC199907   EC204556   EC216121   EC218001   EC226301   EC232467   EC243893   EC244515   EC246494   EC249674   EC254051   EC258842   EC259026   EV591340
UniProtKB/Swiss-Prot
UniProtKB/TrEMBL	Q9BKJ2 Q9W0R8
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
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
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 FBgn0004373    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0004373 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 at stages throughout embryogenesis, during late larval stages, at stages throughout the pupal period, in stages of adults of both sexes. [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 FBgn0004373 expression level Developmental Stage   Expression Level embryo 00-02hr    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr  (3440) embryo 02-04hr  (3366) embryo 04-06hr  (3770) embryo 06-08hr  (3360) embryo 08-10hr  (3317) embryo 10-12hr  (1875) embryo 12-14hr  (2339) embryo 14-16hr  (2393) embryo 16-18hr  (2527) embryo 18-20hr  (1847) embryo 20-22hr  (2688) embryo 22-24hr  (2759) larva L1  (1078) larva L2  (806) larva L3 12hr old  (879) larva L3 puffstage 1-2  (611) larva L3 puffstage 3-6  (1352) larva L3 puffstage 7-9  (1816) white prepupae new  (1819) white prepupae 12hr  (2546) white prepupae 24hr  (2291) pupae 2d postWPP  (2306) pupae 3d postWPP  (1673) pupae 4d postWPP  (1102) adult male 01day  (1480) adult male 05day  (1631) adult male 30day  (1448) adult female 01day  (1596) adult female 05day  (2599) adult female 30day  (2512) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3440) embryo 02-04hr  (3366) embryo 04-06hr  (3770) embryo 06-08hr  (3360) embryo 08-10hr  (3317) embryo 10-12hr  (1875) embryo 12-14hr  (2339) embryo 14-16hr  (2393) embryo 16-18hr  (2527) embryo 18-20hr  (1847) embryo 20-22hr  (2688) embryo 22-24hr  (2759) larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9  (1816) white prepupae new  (1819) white prepupae 12hr  (2546) white prepupae 24hr  (2291) pupae 2d postWPP  (2306) pupae 3d postWPP  (1673) pupae 4d postWPP    1102 adult male 01day  1480 adult male 05day  (1631) adult male 30day  1448 adult female 01day  (1596) adult female 05day  (2599) adult female 30day  (2512) Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 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    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0004373 expression level Developmental Stage   Expression Level embryo 00-02hr    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 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  (3440) embryo 02-04hr  (3366) embryo 04-06hr  (3770) embryo 06-08hr  (3360) embryo 08-10hr  (3317) embryo 10-12hr  (1875) embryo 12-14hr  (2339) embryo 14-16hr  (2393) embryo 16-18hr  (2527) embryo 18-20hr  (1847) embryo 20-22hr  (2688) embryo 22-24hr  (2759) larva L1  (1078) larva L2  (806) larva L3 12hr old  (879) larva L3 puffstage 1-2  (611) larva L3 puffstage 3-6  (1352) larva L3 puffstage 7-9  (1816) white prepupae new  (1819) white prepupae 12hr  (2546) white prepupae 24hr  (2291) pupae 2d postWPP  (2306) pupae 3d postWPP  (1673) pupae 4d postWPP  (1102) adult male 01day  (1480) adult male 05day  (1631) adult male 30day  (1448) adult female 01day  (1596) adult female 05day  (2599) adult female 30day  (2512) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr  (3440) embryo 02-04hr  (3366) embryo 04-06hr  (3770) embryo 06-08hr  (3360) embryo 08-10hr  (3317) embryo 10-12hr  1875 embryo 12-14hr  2339 embryo 14-16hr  2393 embryo 16-18hr  (2527) embryo 18-20hr  1847 embryo 20-22hr  (2688) embryo 22-24hr  (2759) larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9  1816 white prepupae new  1819 white prepupae 12hr  (2546) white prepupae 24hr  2291 pupae 2d postWPP  2306 pupae 3d postWPP  1673 pupae 4d postWPP    1102 adult male 01day  1480 adult male 05day  1631 adult male 30day  1448 adult female 01day  1596 adult female 05day  (2599) adult female 30day  (2512) Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 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    3440 embryo 02-04hr    3366 embryo 04-06hr    3770 embryo 06-08hr    3360 embryo 08-10hr    3317 embryo 10-12hr    1875 embryo 12-14hr    2339 embryo 14-16hr    2393 embryo 16-18hr    2527 embryo 18-20hr    1847 embryo 20-22hr    2688 embryo 22-24hr    2759 larva L1    1078 larva L2    806 larva L3 12hr old    879 larva L3 puffstage 1-2    611 larva L3 puffstage 3-6    1352 larva L3 puffstage 7-9    1816 white prepupae new    1819 white prepupae 12hr    2546 white prepupae 24hr    2291 pupae 2d postWPP    2306 pupae 3d postWPP    1673 pupae 4d postWPP    1102 adult male 01day    1480 adult male 05day    1631 adult male 30day    1448 adult female 01day    1596 adult female 05day    2599 adult female 30day    2512 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 FBgn0004373    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0004373 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Nearly all larval and adult tissues/organs expressed at moderate levels. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval/adult central nervous system, adult crop, larval/adult midgut, larval/adult hindgut, larval/adult Malpighian tubules, adult heart, larval/adult fat body, larval/adult salivary gland, larval trachea, adult female reproductive system, adult male reproductive system, adult carcass. [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 FBgn0004373 expression level Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to maximum FBgn0004373 expression level Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    208.625 Larval Midgut    103.5 Larval Hindgut    190.8 Larval Malpighian Tubules    148 Larval Fat Body    180.9 Larval Salivary Gland    138.3 Larval Trachea    116.925 Larval Carcass    98.275 Adult Head    129.3 Adult Eye    249.4 Adult Brain    301.9 Adult Thoracic-Abdominal Ganglion    232.1 Adult Crop    309.7 Adult Midgut    139.4 Adult Hindgut    297.2 Adult Malpighian Tubules    115.9 Adult Fat Body    187.4 Adult Salivary Gland    159.3 Adult Heart    129.95 Adult VirginFemale Spermatheca    75.3 Adult InseminatedFemale Spermatheca    101.3 Adult Ovary    241.2 Adult Testis    420.4 Adult Male Accessory Gland    183.4 Adult Carcass    104.1 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    129.3   NA  Eye    249.4   NA  Brain    301.9   208.625  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    232.1   NA  Crop    309.7   103.5  Midgut    139.4   190.8  Hindgut    297.2   148  Malpighian Tubules    115.9   180.9  Fat Body    187.4   138.3  Salivary Gland    159.3   NA  Heart    129.95   116.925  Trachea    NA   NA  VirginFemale Spermatheca    75.3   NA  InseminatedFemale Spermatheca    101.3   NA  Ovary    241.2   NA  Testis    420.4   NA  Male Accessory Gland    183.4   98.275  Carcass    104.1 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_01
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004373 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG7004-RB
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | pupal stage, with Scer\GAL4pnr-MD237 fwdGD12100 viable fwdEY05397 viable (with Df(3L)7C) fwd2 fwd3 fwd4 fwdneo1 Sterility Allele female fertile fwdunspecified fertile fwdEY05397 male sterile fwdneo1 male sterile (with Df(3L)7C) fwd3 Other Phenotypes Allele cytokinesis defective fwdneo1 cytokinesis defective (with Df(3L)7C) fwd3 cytokinesis defective | male (with Df(3L)emc-E12) fwd2 cytokinesis defective | male (with fwd3) fwdZ0453 cytokinesis defective | male (with fwdZ0453) fwd3 cytokinesis defective | male | recessive fwdZ0453 long lived, with Tn10\tetRAct5C.T:Hsim\VP16 fwdPdL8S25 meiotic cell cycle defective (with Df(3L)7C) fwd3 meiotic cell cycle defective | male (with Df(3L)7C) fwd2 fwd3 fwd4 fwd5 fwdneo1 meiotic cell cycle defective | male (with Df(3L)emc-E12) fwd2 meiotic cell cycle defective | male | recessive fwdZ0453 mitotic cell cycle defective | male | recessive fwdZ0453 Phenotype manifest in Allele contractile ring & meiotic telophase I | male (with Df(3L)7C) fwdZ0453 contractile ring & meiotic telophase II | male (with Df(3L)7C) fwdZ0453 Nebenkern fwdZ0453 Nebenkern (with Df(3L)emc-E12) fwd2 Nebenkern derivative fwdunspecified secondary spermatocyte (with Df(3L)emc-E12) fwd2 spermatid fwd3 fwdunspecified spermatid (with Df(3L)7C) fwd3 spermatid (with Df(3L)emc-E12) fwd2 spermatid (with fwd3) fwdZ0453 spermatid (with fwdZ0453) fwd3 spermatid & nucleus (with Df(3L)7C) fwd2 fwd3 fwd4 fwd5 fwdneo1 spermatid & nucleus (with Df(3L)emc-E12) fwd2 spermatid & nucleus | supernumerary fwdZ0453 spermatid axoneme fwdunspecified spermatozoon fwdunspecified spindle & meiotic telophase I | male (with Df(3L)7C) fwdZ0453 spindle & meiotic telophase II | male (with Df(3L)7C) fwdZ0453
Classical Alleles ( 16 )
For All Classical Alleles Show
Allele of fwd Class Mutagen Stocks Known lesion fwdB347 1 -- fwdd09794 1 -- fwde03508 1 -- fwdEY05397 P-element activity 1 -- fwdf03822 piggyBac transposase 1 -- fwdGS7301 1 Yes fwdneo1 P-element activity 1 Yes fwdZ0453 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes fwd2 ethyl methanesulfonate 0 Yes fwd3 ethyl methanesulfonate 0 Yes fwd4 ethyl methanesulfonate 0 Yes fwd5 ethyl methanesulfonate 0 Yes fwdCB-5432-3 0 -- fwdMI04139 0 -- fwdPdL8S25 P-element activity 0 Yes fwdunspecified 0 --
Alleles Carried on Transgenic Constructs ( 8 )
For All Alleles Carried on Transgenic Constructs Show
Allele of fwd Class Mutagen Stocks Known lesion fwdGD12100 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes fwdGL00145 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes fwdJF01701 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes fwdJF03329 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes fwddsRNA.cSa in vitro construct - RNAi 0 Yes fwdKD.T:Avic\GFP in vitro construct - amino acid replacement in vitro construct - coding region fusion 0 Yes fwdKK102095 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes fwdT:Avic\GFP in vitro construct - coding region fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(3L)2D (Brill et al., 2000) Df(3L)7C (Brill et al., 2000, Brill, 2000.8.14, Giansanti et al., 2004) Df(3L)17E (Brill et al., 2000, Brill, 2000.8.14) Df(3L)B71 (Brill et al., 2000, Brill, 2000.8.14) Df(3L)emc-E12 (Brill et al., 2000) Df(3L)Exel6084 (Ryder, 2004.4.26)
Not disrupted in	Df(3L)2D (Brill, 2000.8.14) Df(3L)Ar14-8 (Brill et al., 2000)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD12100} NA P{KK102095} NA P{TRiP.GL00145} NA P{TRiP.JF01701} NA P{TRiP.JF03329} NA characterization construct P{Fwd-GFP.KD} NA P{fwd-GFP} NA
Insertions
	Type of insertions Name Expression data insertion of mobile activating element P{EPgy2}fwdEY05397 NA P{GSV2}fwdGS7301 NA P{PdL}fwdPdL8S25 NA P{XP}fwdd09794 NA miscellaneous insertions Mi{MIC}fwdMI04139 NA P{RS3}fwdCB-5432-3 NA PBac{5HPw+}fwdB347 NA PBac{RB}fwde03508 NA PBac{WH}fwdf03822 NA
Gene Ontology: Function, Process & Cellular Component ( 13 unique terms )
Terms Based on Experimental Evidence ( 9 terms )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
contractile ring contraction involved in cell cycle cytokinesis inferred from mutant phenotype (Giansanti et al., 2004) cytokinesis after meiosis I inferred from mutant phenotype (Giansanti et al., 2004) cytokinesis after meiosis II inferred from mutant phenotype (Giansanti et al., 2004) determination of adult lifespan inferred from mutant phenotype (Landis et al., 2003) male germline ring canal formation inferred from mutant phenotype (Brill et al., 2000) male meiosis cytokinesis inferred from mutant phenotype (Brill et al., 2000, Giansanti et al., 2007, Polevoy et al., 2009) meiotic spindle stabilization inferred from mutant phenotype (Giansanti et al., 2004) spermatocyte division inferred from mutant phenotype (Giansanti et al., 2007) spermatogenesis inferred from mutant phenotype (Polevoy et al., 2009)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 4 terms )
Molecular Function
CV term Evidence References
1-phosphatidylinositol 4-kinase activity inferred from sequence or structural similarity with EMBL:U81802 (FlyBase, 1992-) non-traceable author statement (Morrison et al., 2000, Brill, 2000.3.7)
Biological Process
CV term Evidence References
phosphatidylinositol-mediated signaling inferred from electronic annotation with InterPro:IPR015433 (FlyBase Curators et al., 2004-) phosphatidylinositol phosphorylation inferred from electronic annotation with InterPro:IPR015433 (FlyBase Curators et al., 2004-) phosphorylation non-traceable author statement (Morrison et al., 2000)
Cellular Component ( 0 terms)
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 fwd allele Gene References unspecified pbl (Gregory et al., 2007)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 69631 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0004373 DroID - A comprehensive database of gene and protein interactions. • FBgn0004373 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 45374
Orthologs
Genome-wide drosophilid orthologs	Dana\GF25052 Dere\GG14670 Dgri\GH15747 Dmoj\GI24147 Dper\GL16101 Dpse\GA20022 Dsec\GM14285 Dsim\GD25715 Dvir\GJ23452 Dwil\GK17653 Dyak\GE21031
Curated drosophilid orthologs	Dmir\CG7004 (Thornton, 2005.12.20) Dpse\GA20022
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL009294-PA Aedes aegypti (mosquito) XP_391922.1 Apis mellifera (honey bee) 87115 Branchiostoma floridae (lancelet) BGIBMGA007985-PA Bombyx mori (silk worm) CPIJ019027 Culex pipens (mosquito) 156264 Capitella spl (polychaete worm) DDB0191346 Dictyostelium discoideum (amoeba) ENSDARP00000058665 Danio rerio (zebra fish) ENSORLP00000025819 Oryzias latipes (japanese killifish) PHUM009336-PA Pediculus humanus (human body louse) XP_972821.1 Tribolium castaneum (red flour beetle) ENSTRUP00000017982 Takifugu rubripes (pufferfish Takifugu) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16834 LH23293 ISCW021014 FBgn0238311 FBgn0219652 FBgn0210551 FBgn0209894 FBgn0196999 FBgn0169210 JGI_V11_324178 FBgn0080018 FBgn0153705 FBgn0146870 FBgn0140016 FBgn0123218 FBgn0106926 FBgn0102044 CPIJ008596 CPIJ019027 BGIBMGA007985 BGIBMGA007986 ACYPI33367 ACYPI001833 GB19870 AGAP007512 AAEL009294 PHUM409140 PHUM445530 GLEAN_11124
Stocks & Reagents
Stocks Listed in FlyBase ( 12 )
Bloomington	29396 y1 v1; P{TRiP.JF03329}attP2 31187 y1 v1; P{TRiP.JF01701}attP2 16654 y1 w67c23; P{EPgy2}fwdEY05397 35257 y1 sc* v1; P{TRiP.GL00145}attP2 16204 y1 w1118; PBac{5HPw+}fwdB347/TM3, Sb1 Ser1 10069 P{hsneo}AS69, l(3)AS691, fwdneo1 mwh1 red1 e1/TM3, Ser1
Harvard	- P{XP}fwdd09794 - PBac{RB}fwde03508 - PBac{WH}fwdf03822
Kyoto	201157 y1 w67c23; P{GSV2}fwdGS7301/TM3, Sb1 Ser1
VDRC	v27786 w1118; P{GD12100}v27786 v27785 w1118; P{GD12100}v27785/TM3
Genomic Clones ( 2 )
	BACR11J11 BACR13F06 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 74 )
	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	LP07057
Other clones	DPiM_LP07057
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT11124 AT24552 GM01278 GM08014 GM23144 LD07966 LD29916 LD40223 LP06268 LP09174 LP11648 RE46741 RE49150 RE49250
Other clones	10759 11600 13575 18393 32717 53470 54854 62807 77496 82286 124294 146093 157426 163487 172785 175594 183045 191973 214277.27 214379.14 214437.26 282633 CK01.10C.H2 CK01.39D.G5 CK01.97B.C6 CK01.111B.A8 Dmel_neb_012926_0340_1776 EC13043 EK000425 EK033321 EK075433 EK110818 EK112137 EK115540 EK129025 EK143211 EK151808 EK170816 EK188460 EK210711 EK213228 EK246751 EK248957 EK260621 EK267630 EK275245 EK288325 EK294827 EK304104 EK308546 EP03828 EP09361 EP22355 GH11214 LD05780 LD06959 LD25627 LD38593
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0004373 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 45374
Antibody Information
Other Information
Discoverer
	Wolf, 1988.
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: fwd anon-WO0118547.259 (FlyBase, 1996-) Source for merge of: fwd CG7004 (Brill, 2000.8.14)
Additional comments	Source for merge of fwd anon-WO0118547.259 was sequence comparison (date:051113). (FlyBase, 1996-)
Other Comments
	dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology. (Kiger et al., 2003) Area matching Drosophila EST AI297362. (Davies, 2001.3.15) fwd is required for intercellular bridge formation during cytokinesis in male meiosis. (Brill et al., 2000) Mutations of fwd affect the morphology and behaviour of the mitotic spindles of embryonic cleavage divisions to produce multipolar spindles in male meiosis and generate abnormal mitotic figures in larval neuroblasts. (Wilson and Fuller, 1991)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_080083 NM_167826 NM_167827 RefSeq (Proteins) • NP_524822 NP_728518 NP_728519 DDBJ / EMBL / GenBank • AA263708 AAF47375 AAK27793 AAK93473 AAN11437 AAN11438 AF242375 AI114377 AI293824 AI297362 AW944068 AX094135 AY052049 BG632603 BG637878 CL527805 CZ486416 Entrez Gene - A searchable database of RefSeq genes. • 45374 UniProtKB/TrEMBL • Q9BKJ2 Q9W0R8
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Phosphatidylinositol 3/4-kinase, conserved site (IPR018936) Phosphatidylinositol 3-/4-kinase, catalytic (IPR000403) Protein kinase-like domain (IPR011009) Phosphatidylinositol Kinase (IPR015433)
Linkouts
	BioGRID - A database of protein and genetic interactions • 69631 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0004373 DroID - A comprehensive database of gene and protein interactions. • FBgn0004373 DRSC - Results from RNAi screens. • FBgn0004373 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004373 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG7004-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0004373 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 45374 InParanoid A subset of ortholog calls from InParanoid. • AAEL009294-PA Aedes aegypti (mosquito) XP_391922.1 Apis mellifera (honey bee) 87115 Branchiostoma floridae (lancelet) BGIBMGA007985-PA Bombyx mori (silk worm) CPIJ019027 Culex pipens (mosquito) 156264 Capitella spl (polychaete worm) DDB0191346 Dictyostelium discoideum (amoeba) ENSDARP00000058665 Danio rerio (zebra fish) ENSORLP00000025819 Oryzias latipes (japanese killifish) PHUM009336-PA Pediculus humanus (human body louse) XP_972821.1 Tribolium castaneum (red flour beetle) ENSTRUP00000017982 Takifugu rubripes (pufferfish Takifugu) InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 45374 modMine - Data generated by the modENCODE project. • FBgn0004373 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16834 LH23293 ISCW021014 FBgn0238311 FBgn0219652 FBgn0210551 FBgn0209894 FBgn0196999 FBgn0169210 JGI_V11_324178 FBgn0080018 FBgn0153705 FBgn0146870 FBgn0140016 FBgn0123218 FBgn0106926 FBgn0102044 CPIJ008596 CPIJ019027 BGIBMGA007985 BGIBMGA007986 ACYPI33367 ACYPI001833 GB19870 AGAP007512 AAEL009294 PHUM409140 PHUM445530 GLEAN_11124
Synonyms & Secondary IDs ( 6 )
Reported As
Symbol Synonym	anon-WO0118547.259   CG7004 (Brill, 2001.3.29, Morrison et al., 2000, Quinones-Coello, 2007, Perkins et al., 2009, Ni et al., 2008, Keleman et al., 2009.8.5, Ni et al., 2011.7.19) fwd (Polevoy et al., 2007, Gregory et al., 2007, Quinones-Coello, 2007, Bakal et al., 2008, Giansanti et al., 2007, Muraro and Moffat, 2006, Leandro et al., 2007, Giansanti et al., 2006, Anderson et al., 2009, Mattila et al., 2008, Dason et al., 2009, Polevoy et al., 2009, Robinett et al., 2009, Davidson et al., 2009, Goldbach et al., 2010, Yavari et al., 2010) PI4K (Morrison et al., 2000)
Name Synonym	four wheel drive (Giansanti et al., 2007, Giansanti et al., 2006, Anderson et al., 2009, Robinett et al., 2009, Yavari et al., 2010) Four wheel drive (Polevoy et al., 2009)
Secondary FlyBase IDs
	FBgn0035129 FBgn0062139
References ( 66 )
Generate a list of	All references relating to this gene ( 66 )
List References by type	Research paper  ( 21 ) Supplementary material  ( 5 ) Review  ( 2 ) Personal communication to FlyBase  ( 11 ) Abstract  ( 14 ) FlyBase analysis  ( 3 ) Patent  ( 1 ) DNA/RNA sequence record  ( 3 ) Computer file  ( 3 ) Automatic genome annotation  ( 1 ) Book  ( 1 ) Curated genome annotation  ( 1 )
Recent research papers ( 2 )
	Goldbach et al., 2010, Mol. Biol. Cell 21(9): 1482--1493 Stabilization of the actomyosin ring enables spermatocyte cytokinesis in Drosophila. [FBrf0210683] Yavari et al., 2010, Dev. Cell 19(1): 54--65 Role of Lipid Metabolism in Smoothened Derepression in Hedgehog Signaling. [FBrf0211286] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...