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Gene Dmel\vvl

General Information
Symbol	Dmel\vvl	Species	D. melanogaster
Name	ventral veins lacking	Annotation symbol	CG10037
Feature type	protein_coding_gene	FlyBase ID	FBgn0086680
Gene Model Status	Current	Stock availability	37 publicly available
Also Known As	dfr, Cf1a, ut, Dfr/Vvl, drf
Genomic Location
Chromosome (arm)	3L	Recombination map	3-
Cytogenetic map	65C5-65C5	Sequence location	3L:6,783,252..6,787,637 [+]
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 ventral veins lacking is referred to in FlyBase by the symbol Dmel\vvl (CG10037, FBgn0086680). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: RNA polymerase II regulatory region sequence-specific DNA binding; RNA polymerase II distal enhancer sequence-specific DNA binding; RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription. There is experimental evidence that it is involved in the biological process: motor axon guidance; regulation of dendrite morphogenesis; positive regulation of transcription from RNA polymerase II promoter; brain development; brain segmentation; peripheral nervous system development. 30 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; organ system; adult segment; larval abdominal segment; non-connected developing system; sensory cluster; portion of tissue; adult; peripheral nervous system; region of integument. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Homeobox; Homeobox, conserved site; Homeodomain-like; Homeodomain-related; Lambda repressor-like, DNA-binding; POU; POU-specific; Transcription factor, homeobox/POU. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed within 06-18 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression in all larval and adult organs/tissues ranges from low to undetected. Expression at moderate levels in the following post-embryonic organs or tissues: adult heart, adult fat body, larval trachea, adult carcass. Gene sequence location is 3L:6783252..6787637.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	sep: separated Most of posterior crossvein absent, one-third usually remaining attached to vein L5. RK2A. vvl: ventral veins lacking Prevents differentiation of longitudinal veins 2 and 4 which form on the ventral surface of the wing; in Hw; vvl flies, extra sensillae form in the ventral surface where veins would ordinarily form.
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	Controlled Vocabulary Terms positive regulation of transcription from RNA polymerase II promoter RNA polymerase II regulatory region sequence-specific DNA binding RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription RNA polymerase II distal enhancer sequence-specific DNA binding regulation of dendrite morphogenesis Sequence features BKN22601 HFA11409 Alleles vvl[GAL4]
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 65C5-65C5   Limits computationally determined from genome sequence between P{PZ}l(3)0209402094 and P{lacW}l(3)L4060L4060
Experimentally Determined Cytological Location
Cytogenetic map Notes References 65A2-65E1   (determined by in situ hybridisation)   (Salzberg et al., 1994) 65D-65D   (determined by in situ hybridisation)   (Billin et al., 1991) 65D-65D   (determined by in situ hybridisation)   (Johnson and Hirsh, 1990) 65D-65D   (determined by in situ hybridisation)   (Frigerio et al., 1986)
Experimentally Determined Recombination Data
Location	3-26 (Salzberg et al., 1994) 3-
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\vvl for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	3' UTR contains an ORF that is conserved among close Drosophila species (potential peptide 264 aa in length); possible stop codon suppression or alternative splicing product (see alternative transcript).
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) vvl-RA FBtr0076944  NM_079224   4378   427 vvl-RB FBtr0300260  NM_001169884   2292   763
Additional Transcript Data & Comments
Reported size (kB)	4.5 (northern blot) (Treacy et al., 1991) 3.5, 3.4 (northern blot) (Anderson et al., 1995, Billin 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 vvl-PA   FBpp0076653   45.9   427   8.31     NP_523948   AAF50641 vvl-PB   FBpp0289488   82.7   763   8.22     NP_001163355   ACZ94626
Additional Polypeptide Data & Comments
Reported size (kDa)	549 (aa); 60 (kD) (Treacy et al., 1991) 427 (aa); 46 (kD observed); 46 (kD predicted) (Anderson et al., 1995) 427 (aa) (Billin et al., 1991) 150 (kD) (Johnson and Hirsh, 1990)
Comments	vvl protein is a positive regulator of Ddc gene transcription. Immunoprecipitation and crosslinking studies show that vvl protein associates with acj6 protein in vivo. When bound by acj6 protein as a heterodimer, vvl protein no longer binds DNA. Transfection experiments in CV-1 cells demonstrate that vvl protein activates transcription of a reporter gene driven by the Ddc promoter or a heterologous promoter preceded by the vvl element. This activation does not occur in the presence of acj6 protein. The vvl DNA sequence and predicted amino acid sequence is significantly different than that presented in FBrf 54626. (Treacy et al., 1991) vvl protein encodes a sequence specific DNA-binding protein. The vvl gene product binds to a DNA region required for the expression of Ddc. A high degree of similarity was noted between the vvl protein and Human Brn-1 and Brn-2 POU-domain sequences which extends to the 17aa linker between the homeodomain and the POU domain. (Johnson and Hirsh, 1990)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Homeobox, conserved site (IPR017970) POU-specific (IPR000327) Homeobox (IPR001356) Homeodomain-like (IPR009057) Lambda repressor-like, DNA-binding (IPR010982) Homeodomain-related (IPR012287) POU (IPR013847) Transcription factor, homeobox/POU (IPR016362)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE014296 AAF50641   AE014296 ACZ94626   BI162129   BI162130   BI229419   BI234425   BI237775   BI240977   BI358363   BI362199   BI366413   BI371408   BI372466   BI374683   BI375186   BI483033   BI485408   BI580320   BT003517 AAO39521   BT030760   CO176953   CO178643   CO186454   CO189598   CO189615   CO189852   CO263697   CO263852   CO268493   CO268814   CO269444   CO270394   CO271377   CO273156   CO273695   CO273957   CO276038   CO276474   CO278840   CO286366   CO286930   CO287413   CO293064   CO305171   CO307387   CO309199   CO309842   CO311873   CO316228   CO318628   CO321496   EC060784   EC061190   EC062832   EC063698   EC069178   EC069796   EC071018   EC077802   EC078223   EC082516   EC083420   EC088757   EC196975   EC197077   EC202861   EC204358   EC208681   EC208981   EC209887   EC217481   EC218211   EC220891   EC224918   EC225002   EC230476   EC232720   EC236923   EC238080   EC241259   EC243426   EC243807   GH941510   GH941987   GH945241   GH945384   GH945500   GH945518   GH945997   GH946316   GH946374   GH947281   GH949161   GH949739   GH951957   GH951981   GH952227   GH952658   GH955430   GH955434   GH958111   GH958316   GH959317   GH959962   GH960018   GH960235   GH960684   GH961346   GH961408   GH961471   GH961700   GH962293   GH962444   GH962638   GH962757   GH963528   GH963633   GH963670   GH964083   GH964179   GH964250   GH964650   GH965251   GH966000   GH966095   GH966463   GH966989   GH967309   GH967372   GH967677   GH969947   GH970590   GH970889   GH971524   GH971737   GH973647   GH973686   GH974429   GH974780   GH975521   GH975824   GH975959   GH976189   GH976878   GH977800   GH978858   GH979165   GH979724   GH979971   GH980160   GH980966   GH982942   GH983277   GH984543   M81959 AAA28831   X52252 CAA36496   X58435 CAA41341
UniProtKB/Swiss-Prot	P16241
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References protein binding site FBsf0000160244 3L:6,778,294..6,778,310 bound_moiety=vvl-XP evidence=experimental (Certel et al., 1996, Bergman et al., 2004) protein binding site FBsf0000160245 3L:6,778,066..6,778,086 bound_moiety=vvl-XP evidence=experimental (Certel et al., 1996, Bergman et al., 2004)
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference embryonic stage   ventral midline   (Treacy et al., 1991) tracheal primordium   (Treacy et al., 1991) ventral nerve cord   (Treacy et al., 1991) glial cell   (Treacy et al., 1991) embryonic stage | late   central nervous system | restricted   (Billin et al., 1991) in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   presumptive embryonic/larval peripheral nervous system | restricted   (Billin et al., 1991) embryonic stage | mid   tracheal primordium   (Billin et al., 1991) embryonic stage   ectoderm   (Treacy et al., 1991) mesectoderm   (Treacy et al., 1991) neuron   (Treacy et al., 1991) supraesophageal ganglion   (Treacy et al., 1991) embryonic stage 14   ectoderm of hindgut | anterior   (Hombria et al., 2005) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage 1 -- 14     (Frigerio et al., 1986) Comment:reference states transcript expressed up to 8-12 hr AEL oogenesis     (Frigerio et al., 1986)
Additional Descriptive Data
	vvl transcripts first appear at 5-6hr of development, show a peak of expression between 10hr and 13hr and decline substantially by the end of embryogenesis. (Anderson et al., 1995) vvl transcripts are initially detected in 3-6 hr embryos, progressively increase to maximum levels in late embryos, and decline thereafter. Transcripts are initially detected in the ectoderm of stage 10 embyros at regular, repeating intervals. This correlates with sites of the invaginations that are the origins of the tracheal placodes. Transcripts are also found along the midline of the CNS. vvl transcripts are found in a subset of neurons in the CNS that substantially overlap those that express acj6 transcripts. (Treacy et al., 1991) vvl is expressed in the ectoderm of the anterior hindgut at embryonic stage 14. (Hombria et al., 2005) vvl transcripts are first detected in 3-6 hr embryos, are at a maximum level in 6-12 hr embryos and persist throughout development. They are first detected in the tracheal placodes of germ band extended embryos. They are later observed in cells surrounding the tracheal pits that may be chordotonal organs and in a group of cells on the roof of the stomodeal invagination. A complex pattern of expression is observed in the PNS and CNS. (Billin et al., 1991) vvl transcript is detected in oocytes and early embryos. (Frigerio et al., 1986)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   serotonergic neuron   (Anderson et al., 1995) central nervous system | restricted   (Anderson et al., 1995) RP neuron   (Anderson et al., 1995) embryonic stage   embryonic/larval oenocyte precursor   (Elstob et al., 2001) embryonic stage 10   tracheal primordium   (Anderson et al., 1995, Zelzer and Shilo, 2000) embryonic stage 10 -- 11   procephalic segment | restricted   (Anderson et al., 1995) labrum | restricted   (Anderson et al., 1995) labral segment | restricted   (Anderson et al., 1995) embryonic stage 10 -- 12   ventral midline   (Anderson et al., 1995) embryonic stage 10 -- 17   midline glial cell   (Anderson et al., 1995) embryonic stage 11   embryonic/larval tracheocyte   (Llimargas and Casanova, 1999) embryonic stage 11 -- 17   tracheal pit   (Anderson et al., 1995) embryonic stage 13 -- 15   embryonic/larval hindgut   (Anderson et al., 1995) embryonic stage 13 -- 16   abdominal 1 larval oenocyte group -- abdominal 7 larval oenocyte group   (Anderson et al., 1995) embryonic stage 13 -- 17   epidermis   (Anderson et al., 1995) adult stage   adult antennal lobe projection neuron DA1 lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron DL3 lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron VA7m lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron VA5 lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron DM5 lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron DM2 lPN   (Komiyama and Luo, 2007) adult antennal lobe projection neuron DM1 lPN   (Komiyama and Luo, 2007)
Additional Descriptive Data
	vvl protein expression is first apparent in early stage 10 in two rows of 10-cell patches corresponding to the developing tracheal placodes. vvl expression serves as a marker for the tracheal pits as they undergo extensive morphogenetic movements in stages 10-13. Epidermal vvl expression becomes prominant from stages 13-16/17 and masks the tracheal expression at these stages. During this period, strong staining is seen in the seven laterally symmetrical oenocyte clusters in segments A1-A7. vvl is also expressed in the hindgut in stages 13-15. vvl expression is observed in the developing head region in stage 10/11 embryos in tissues of the procephalon, clypeolabrum, and labium. The most prominant labeling is seen in laterally symmetrical placodes of cells located on both the dorsal and ventral sides of the stomodeal opening. This corresponds to regions containing sensory progenitor cells derived from the clypeolabrum and labrum. Expression is seen in the cells of the mesectoderm in stages 10-12. Mesectoderm expression extends beyond stage 12 in the middle pair of midline glia (MGM). In later stages, vvl is expressed in a uniform repeated pattern of ~25 nuclei per neuromere grouped into 5 distinctive clusters: 1) a midline cluster of four nuclei containing the MGM and a pair of RP motoneurons, 2) two symmetrical mediolateral clusters that contain the paired serotonergic neurons, and 3) two symmetrical lateral clusters. (Anderson et al., 1995)
Marker for
	embryonic/larval tracheocyte
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 FBgn0086680    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0086680 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 high expression to a trough of very low expression. Peak expression observed within 06-18 hour embryonic stages. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0086680 expression level Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 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    36 embryo 02-04hr  (201) embryo 04-06hr  (574) embryo 06-08hr  (3420) embryo 08-10hr  (4563) embryo 10-12hr  (3376) embryo 12-14hr  (3478) embryo 14-16hr  (4469) embryo 16-18hr  (3143) embryo 18-20hr  (2657) embryo 20-22hr  (1896) embryo 22-24hr  (1331) larva L1  (1639) larva L2  (1352) larva L3 12hr old  (871) larva L3 puffstage 1-2  (337) larva L3 puffstage 3-6  (950) larva L3 puffstage 7-9  (844) white prepupae new  (831) white prepupae 12hr  (840) white prepupae 24hr  (1094) pupae 2d postWPP  (722) pupae 3d postWPP  (581) pupae 4d postWPP  (967) adult male 01day  (1431) adult male 05day  (1706) adult male 30day  (1428) adult female 01day  (896) adult female 05day  (237) adult female 30day  (239) Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr  (3420) embryo 08-10hr  (4563) embryo 10-12hr  (3376) embryo 12-14hr  (3478) embryo 14-16hr  (4469) embryo 16-18hr  (3143) embryo 18-20hr  (2657) embryo 20-22hr  (1896) embryo 22-24hr    1331 larva L1  (1639) larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day  1431 adult male 05day  (1706) adult male 30day  1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 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    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0086680 expression level Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr  (201) embryo 04-06hr  (574) embryo 06-08hr  (3420) embryo 08-10hr  (4563) embryo 10-12hr  (3376) embryo 12-14hr  (3478) embryo 14-16hr  (4469) embryo 16-18hr  (3143) embryo 18-20hr  (2657) embryo 20-22hr  (1896) embryo 22-24hr  (1331) larva L1  (1639) larva L2  (1352) larva L3 12hr old  (871) larva L3 puffstage 1-2  (337) larva L3 puffstage 3-6  (950) larva L3 puffstage 7-9  (844) white prepupae new  (831) white prepupae 12hr  (840) white prepupae 24hr  (1094) pupae 2d postWPP  (722) pupae 3d postWPP  (581) pupae 4d postWPP  (967) adult male 01day  (1431) adult male 05day  (1706) adult male 30day  (1428) adult female 01day  (896) adult female 05day  (237) adult female 30day  (239) Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr  (3420) embryo 08-10hr  (4563) embryo 10-12hr  (3376) embryo 12-14hr  (3478) embryo 14-16hr  (4469) embryo 16-18hr  (3143) embryo 18-20hr  (2657) embryo 20-22hr  1896 embryo 22-24hr    1331 larva L1  1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day  1431 adult male 05day  1706 adult male 30day  1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 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    36 embryo 02-04hr    201 embryo 04-06hr    574 embryo 06-08hr    3420 embryo 08-10hr    4563 embryo 10-12hr    3376 embryo 12-14hr    3478 embryo 14-16hr    4469 embryo 16-18hr    3143 embryo 18-20hr    2657 embryo 20-22hr    1896 embryo 22-24hr    1331 larva L1    1639 larva L2    1352 larva L3 12hr old    871 larva L3 puffstage 1-2    337 larva L3 puffstage 3-6    950 larva L3 puffstage 7-9    844 white prepupae new    831 white prepupae 12hr    840 white prepupae 24hr    1094 pupae 2d postWPP    722 pupae 3d postWPP    581 pupae 4d postWPP    967 adult male 01day    1431 adult male 05day    1706 adult male 30day    1428 adult female 01day    896 adult female 05day    237 adult female 30day    239 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 FBgn0086680    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0086680 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression in all larval and adult organs/tissues ranges from low to undetected. Expression at moderate levels in the following post-embryonic organs or tissues: adult heart, adult fat body, larval trachea, 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) Heatmap Tissue   Expression Level Larval Central Nervous System no informative data Larval Midgut     Larval Hindgut no informative data Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea no informative data Larval Carcass no informative data Adult Head no informative data Adult Eye     Adult Brain no informative data Adult Thoracic-Abdominal Ganglion no informative data Adult Crop no informative data Adult Midgut     Adult Hindgut no informative data Adult Malpighian Tubules     Adult Fat Body no informative data Adult Salivary Gland     Adult Heart no informative data Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass no informative data 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_29
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0086680 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • S.C3L000862 FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for vvl Stages(s) 1-3 Stages(s) 4-6 Stages(s) 7-8 Stages(s) 9-10 Stages(s) 11-12 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal vvlM1 vvlM2 vvlM3 lethal | embryonic stage | recessive vvldfr-E82 vvlGA3 vvlH76 vvlH599 vvlM638 vvlZm lethal | recessive vvlH76 vvlH599 vvlM638 semi-lethal | pupal stage, with Scer\GAL4pnr-MD237 vvlGD16468 viable vvlsep viable, with Scer\GAL4pnr-MD237 vvlGD4503 Other Phenotypes Allele increased cell death vvlGA3 vvlH599 neuroanatomy defective vvlGA3 vvlH599 vvlunspecified neuroanatomy defective, with Scer\GAL4ftz.ng, vvldfr-B129 vvldsRNA.Scer\UAS neuroanatomy defective, with Scer\GAL4ftz.ng, vvldsRNA.Scer\UAS vvldfr-B129 neuroanatomy defective, with Scer\GAL4Lim3.B vvlScer\UAS.cCa neuroanatomy defective, with Scer\GAL4Lim3.B, vvldfr-B129 vvldsRNA.Scer\UAS neuroanatomy defective, with Scer\GAL4Lim3.B, vvldsRNA.Scer\UAS vvldfr-B129 neuroanatomy defective | dominant vvldfr-B129 neuroanatomy defective | recessive | somatic clone vvldfr-E82 neurophysiology defective, with Scer\GAL4GH146 vvlScer\UAS.cCa visible, with Scer\GAL4pnr-MD237 vvlGD4503 vvlGD16468 visible, with Scer\GAL4Ser.PGF vvlB157.Scer\UAS vvlScer\UAS.cCa visible (with Df(3L)XBB70) vvlsep visible (with In(3LR)282) vvlunspecified visible | dominant vvldfr-B1.57 visible | recessive | somatic clone vvlH599 visible | somatic clone vvlH599 wild-type vvl+mAa Phenotype manifest in Allele abdominal 1 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 2 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 3 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 4 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 5 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 6 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal 7 lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlH76 vvlM638 abdominal desC neuron vvlH599 abdominal desD neuron vvlH599 abdominal lateral pentascolopidial chordotonal organ lch5 vvlH599 vvlunspecified adult abdominal sensillum (with vvlGA3) vvlM638 adult abdominal sensillum (with vvlH76) vvlM638 adult abdominal sensillum (with vvlM638) vvlGA3 vvlH76 adult thoracic sensillum (with vvlGA3) vvlM638 adult thoracic sensillum (with vvlH76) vvlM638 adult thoracic sensillum (with vvlM638) vvlGA3 vvlH76 antennal glomerulus & neuron, with Scer\GAL4GH146 vvlScer\UAS.cCa antennal glomerulus & neuron | somatic clone vvldfr-B129 antennal lobe glomerulus DM5 | somatic clone vvldfr-B129 antennal sense organ | somatic clone vvlH599 anterior commissure vvlZm commissure vvlGA3 vvlZm vvldfr-E82 vvlhs.PC denticle belt vvlGA3 vvlZm vvldfr-E82 desC neuron vvlunspecified desD neuron vvlunspecified embryonic/larval deuterocerebrum | embryonic stage 14 vvldfr-E82 embryonic/larval dorsal trunk | embryonic stage vvlGA3 vvlZm vvldfr-E82 embryonic/larval protocerebrum | embryonic stage 14 vvldfr-E82 embryonic/larval tracheal system vvlGA3 vvldfr-B1.57 vvldfr-E82 vvlunspecified embryonic/larval tritocerebral commissure | embryonic stage 14 vvldfr-E82 embryonic/larval tritocerebrum | embryonic stage 14 vvldfr-E82 filzkorper vvlGA3 vvlZm vvldfr-E82 glial cell vvldfr-E82 glial cell & embryonic brain | embryonic stage 14 vvldfr-E82 head bristle (with vvlGA3) vvlM638 head bristle (with vvlH76) vvlM638 head bristle (with vvlM638) vvlGA3 vvlH76 head bristle | somatic clone vvlH599 interommatidial bristle | somatic clone vvlH599 intersegmental nerve | embryonic stage 16 vvldfr-B129 intersegmental nerve | embryonic stage 16, with Scer\GAL4ftz.ng, vvldfr-B129 vvldsRNA.Scer\UAS intersegmental nerve | embryonic stage 16, with Scer\GAL4ftz.ng, vvldsRNA.Scer\UAS vvldfr-B129 lateral abdominal cluster vvlGA3 vvlH599 lch5 neuron vvlGA3 vvlH599 lobula | somatic clone vvldfr-E82 longitudinal connective vvlGA3 vvlZm vvldfr-E82 longitudinal connective | embryonic stage 14 vvldfr-E82 macrochaeta | ectopic vvlGA3 vvlZm vvldfr-E82 maxillary palp bristle | somatic clone vvlH599 medulla layer M2 | somatic clone vvldfr-E82 medulla layer M3 | somatic clone vvldfr-E82 medulla layer M8 | somatic clone vvldfr-E82 medulla layer M9 | somatic clone vvldfr-E82 medulla layer M10 | somatic clone vvldfr-E82 mesectoderm vvldfr-B1.57 vvldfr-E82 midline glial cell vvldfr-E82 olfactory sensory organ & neuron & dendrite, with Scer\GAL4GH146 vvlScer\UAS.cCa olfactory sensory organ & neuron & dendrite | somatic clone vvldfr-B129 posterior commissure vvlZm posterior crossvein vvlsep posterior crossvein (with Df(3L)XBB70) vvlsep presumptive embryonic/larval central nervous system vvlGA3 vvlZm vvldfr-E82 presumptive embryonic/larval nervous system vvldfr-B1.57 vvldfr-E82 sensory neuron vvlGA3 socket | somatic clone vvlH599 socket | somatic clone | supernumerary vvlH599 trachea vvlGA3 vvlZm vvldfr-B129 vvldfr-E82 transmedullary neuron Tm27 | somatic clone vvldfr-E82 transverse nerve | embryonic stage 16 vvldfr-B129 transverse nerve | embryonic stage 16, with Scer\GAL4Lim3.B vvlScer\UAS.cCa transverse nerve | embryonic stage 16, with Scer\GAL4Lim3.B, vvldfr-B129 vvldsRNA.Scer\UAS transverse nerve | embryonic stage 16, with Scer\GAL4Lim3.B, vvldsRNA.Scer\UAS vvldfr-B129 trichogen cell, with Scer\GAL4pnr-MD237 vvlGD4503 vvlGD16468 trichogen cell | somatic clone vvlH599 ventral nerve cord | embryonic stage 14 vvldfr-E82 wing vvlGA3 vvlM1 vvlZm vvldfr-E82 wing, with Scer\GAL4Ser.PGF vvlB157.Scer\UAS wing (with Df(3L)XBB70) vvlsep wing & macrochaeta vvlGA3 vvlZm vvldfr-E82 wing | distal, with Scer\GAL4Ser.PGF vvlScer\UAS.cCa wing | posterior compartment vvlGA3 vvlZm vvldfr-E82 wing cell | ectopic, with Scer\GAL4Ser.PGF vvlScer\UAS.cCa wing vein vvlGA3 vvlZm vvldfr-B1.57 vvldfr-E82 wing vein (with Df(3L)XBB70) vvlsep wing vein L2 (with Df(3L)XBB70) vvlsep wing vein L4 vvlsep wing vein L4 (with Df(3L)XBB70) vvlsep wing vein L4 (with In(3LR)282) vvlunspecified
Classical Alleles ( 18 )
For All Classical Alleles Show
Allele of vvl Class Mutagen Stocks Known lesion vvlsep hypomorphic allele - genetic evidence 32 -- vvlM638 loss of function allele ethyl methanesulfonate 1 Yes vvldfr-B1.57 loss of function allele ethyl methanesulfonate 0 -- vvldfr-B108 0 -- vvldfr-B129 ethyl methanesulfonate 0 -- vvldfr-B140 0 -- vvldfr-E82 loss of function allele ethyl methanesulfonate 0 Yes vvldfr-XC185 0 -- vvlE82-2D 0 -- vvlGA3 loss of function allele X ray 0 -- vvlGAL4 hypomorphic allele - molecular evidence ends-out gene targeting 0 Yes vvlH599 loss of function allele ethyl methanesulfonate 0 Yes vvlH76 loss of function allele ethyl methanesulfonate 0 Yes vvlM1 0 -- vvlM2 0 -- vvlM3 0 -- vvlunspecified 0 -- vvlZm loss of function allele spontaneous 0 --
Alleles Carried on Transgenic Constructs ( 12 )
For All Alleles Carried on Transgenic Constructs Show
Allele of vvl Class Mutagen Stocks Known lesion vvlGD16468 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes vvlGD4503 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes vvlJF02126 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes vvl+mAa in vitro construct - regulatory fusion in vitro construct - minigene 0 Yes vvlAct5C.PM in vitro construct - regulatory fusion 0 Yes vvlB157.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 0 Yes vvldsRNA.Scer\UAS in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes vvlhs.PC in vitro construct - regulatory fusion 0 Yes vvlKK102520 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes vvlScer\UAS.cBa in vitro construct - regulatory fusion 0 Yes vvlScer\UAS.cCa in vitro construct - regulatory fusion 0 Yes vvlScer\UAS.cLa in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(3L)Exel6109 (Ryder, 2004.4.26) Df(3L)v65c (Inbal et al., 2003) Df(3L)W5.4 (Anderson et al., 1995) Df(3L)XBB70 (Certel et al., 2000, Anderson et al., 1995, Anderson et al., 1996) Df(3L)XDI98 (Salzberg et al., 1994) Df(3L)ZN47 (Inbal et al., 2003) In(3LR)sep (de Celis et al., 1995)
Not disrupted in	Df(3L)CH12 (Inbal et al., 2003) Df(3L)XAS96 (Anderson et al., 1995)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data reporter construct P{0.8Rdfr-lacZ} No P{4.5Xdfr-lacZ} No P{7.0Rdfr-lacZ} No P{198RVdfr-lacZ} No P{514dfr-lacZ} No P{587dfr-lacZ} No P{650dfr-lacZ} No P{889dfr-lacZ} No P{1047dfr-lacZ} No P{dfr-lacZ} No P{HcXdfr-lacZ} No P{HdHcdfr-lacZ} No P{HdXdfr-lacZ} No P{mutDFRE1-514dfr-lacZ} No P{RHcdfr-lacZ} No P{RHddfr-lacZ} No P{RRVdfr-lacZ} No P{RX-dfr-lacZ} No P{XRdfr-lacZ} No UAS construct P{GD4503} NA P{GD16468} NA P{KK102520} NA P{TRiP.JF02126} NA P{UAS-vvl.B157} NA P{UAS-vvl.B} NA P{UAS-vvl.C} NA P{UAS-vvl.ds} NA P{UAS-vvl.L} NA heat-shock construct P{hsp-dfr} NA characterization construct P{RX-dfr-minigene} NA
Insertions
	Type of insertions Name Expression data
Gene Ontology: Function, Process & Cellular Component ( 15 unique terms )
Terms Based on Experimental Evidence ( 9 terms )
Molecular Function
CV term Evidence References
RNA polymerase II core promoter proximal region sequence-specific DNA binding transcription factor activity involved in positive regulation of transcription inferred from direct assay (Junell et al., 2007) RNA polymerase II distal enhancer sequence-specific DNA binding inferred from direct assay (Miller et al., 2009) RNA polymerase II regulatory region sequence-specific DNA binding inferred from direct assay (Anderson et al., 1996)
Biological Process
CV term Evidence References
brain development inferred from mutant phenotype (Meier et al., 2006) brain segmentation inferred from mutant phenotype (Meier et al., 2006) motor axon guidance inferred from mutant phenotype (Certel and Thor, 2004) peripheral nervous system development inferred from mutant phenotype (Inbal et al., 2003) positive regulation of transcription from RNA polymerase II promoter inferred from direct assay (Miller et al., 2009) inferred from mutant phenotype (Anderson et al., 1996) regulation of dendrite morphogenesis inferred from mutant phenotype (Hasegawa et al., 2011)
Cellular Component ( 0 terms)
Terms Based on Predictions or Assertions ( 7 terms )
Molecular Function
CV term Evidence References
DNA binding non-traceable author statement (Swiss-Prot Project Members, 1990.4.1) sequence-specific DNA binding transcription factor activity traceable author statement (Ghabrial et al., 2003)
Biological Process
CV term Evidence References
dendrite morphogenesis traceable author statement (Grueber and Jan, 2004, Gao and Bogert, 2003) epithelial cell fate determination, open tracheal system traceable author statement (Affolter and Shilo, 2000, Ghabrial et al., 2003) peripheral nervous system development traceable author statement (Salzberg et al., 1994) regulation of transcription, DNA-dependent non-traceable author statement (Swiss-Prot Project Members, 1990.4.1)
Cellular Component
CV term Evidence References
nucleus inferred from sequence or structural similarity with UniProtKB:Q17237 (FlyBase, 1992-) non-traceable author statement (Swiss-Prot Project Members, 1990.4.1)
Sequence Ontology: Class of Gene
	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 vvl allele Gene References vvldfr-B1.57 E(vvl)23 (Certel and Johnson, 1998) Mad (Certel and Johnson, 1998) vvldfr-B108 Mad (Certel and Johnson, 1998) vvldfr-B129 tup (Certel and Thor, 2004) vvlScer\UAS.cCa acj6 (Komiyama et al., 2003) tup (Certel and Thor, 2004) vvlsep dpp (de Celis et al., 1995) Egfr (de Celis et al., 1995) tkv (de Celis et al., 1995) vvlunspecified Egfr (Diaz-Benjumea and Garcia-Bellido, 1990) unspecified btl (Llimargas and Casanova, 1997) N (de Celis et al., 1995) rho (Sturtevant and Bier, 1995, de Celis et al., 1995)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 64202 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0086680 DroID - A comprehensive database of gene and protein interactions. • FBgn0086680 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 38752
Orthologs
Genome-wide drosophilid orthologs
Curated drosophilid orthologs	Dpse\GA10024
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL003560-PA Aedes aegypti (mosquito) AGAP005878-PA Anopheles gambiae (mosquito) XP_393686.2 Apis mellifera (honey bee) XP_001948842.1 Acyrthosiphon pisum (pea aphid) BGIBMGA010868-PA Bombyx mori (silk worm) ENSBTAP00000010443 Bos taurus (bovine) CPIJ002221 Culex pipens (mosquito) ENSDARP00000037034 Danio rerio (zebra fish) ENSECAP00000017373 Equus caballus (horse) ENSGACP00000002556 Gymnopilus aculeatus (mushroom) ENSP00000329170 Homo sapiens (human) ISCW020718-PA Ixodes scapularis (deer tick) ENSMODP00000029696 Monodelphis domestica (gray opossum) ENSMUSP00000063809 Mus musculus (house mouse) ENSPTRP00000032636 Pan troglodytes (chimpanzee) ENSRNOP00000009043 Rattus norvegicus (norway rat) XP_968364.2 Tribolium castaneum (red flour beetle) ENSTNIP00000019394 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000008884 Takifugu rubripes (pufferfish Takifugu) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16730 NV14897 LH23007 ISCW020718 FBgn0200336 FBgn0185665 FBgn0169718 JGI_V11_14402 FBgn0070083 FBgn0135721 FBgn0122836 FBgn0097052 CPIJ002221 CPIJ019261 BGIBMGA010868 ACYPI008866 GB16085 AAEL003560 PHUM605420 GLEAN_14350
Stocks & Reagents
Stocks Listed in FlyBase ( 37 )
Bloomington	26228 y1 v1; P{TRiP.JF02126}attP2 822 In(3LR)sep, vvlsep kniri-1 pp
Kyoto	106087 In(3LR)sep, vvlsep kniri-1 pp, vvlsep/TM6B 101576 AntpD43 / TM3, y+ kniri-1 vvlsep Ubxbx-34e es Ser1 101575 Antp73b-rv21 / TM3, y+ kniri-1 vvlsep Sb1 Ubxbx-34e es 101590 Df(3L)BK9 / TM3, y+ kniri-1 pp vvlsep Sb1 Ubxbx-34e es 101638 flr3 / TM3, y+ kniri-1 pp vvlsep Ubxbx-34e es Ser1 103302 Df(3L)VW3 / TM3, y+ kniri-1 pp vvlsep Sb1 Ubxbx-34e es Ser1 101710 Pc11 / TM3, y+ kniri-1 pp vvlsep Sb1 Ubxbx-34e es Ser1
VDRC	v47182 w1118; P{GD16468}v47182 v47185 w1118; P{GD16468}v47185 v10756 w1118; P{GD4503}v10756/TM3
	More stocks available...
Genomic Clones ( 1 )
	BACR10L09 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 149 )
	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	RE27192
Other clones	DPiM_FI01008 FI01008
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	RE30124 RE34066 RE37946 RE45114 RE46704 RE51947 RE58286 RE59434 RE62339 RE62939 RE65865 RE68727 RE74495
Other clones	10212 11405 11915 14848 15581 18466 28403 59217 72737 83094 97168 103859 109098 144914 149215 158053 165911 172457 CK01.11B.D1 CK01.39D.G12 CK01.55D.F4 CK01.72A.D12 CK01.81A.F5 CK01.107A.E1 CK01.109A.D12 CK01.121C.E9 CK01.124B.D2 CK01.147C.E10 CK01.168B.H1 EK019906 EK044654 EK044724 EK045354 EK051736 EK057246 EK062623 EK064303 EK065808 EK089317 EK091649 EK092819 EK094545 EK098746 EK103229 EK109847 EK111932 EK112726 EK119515 EK120949 EK131843 EK136653 EK185802 EK201619 EK219457 EK227115 EK247602 EK254461 EK260632 EK264042 EK273235 RE01535 RP004004265 RP004037349 RP004043359 RP004097904 RP004097920 RP004140435 RP004146562 RP004150706 RP004151090 RP004183228 RP004191113 RP004197168 RP004211316 RP004211504 RP004215590 RP004220505 RP004221230 RP004225235 RP004230319 RP004231039 RP004231510 RP004232777 RP004235930 RP004244938 RP004246069 RP004247344 RP004253098 RP004254946 RP004260559 RP004267629 RP004277619 RP004307814 RP004325243 RP004331131 RP004335424 RP004340688 RP004356585 RP004363441 RP004376696 RP004377815 RP004408097 RP004410105 RP004420656 RP004425259 RP004434775 RP004464448 RP004468300 RP004469067 RP004473765 RP004479300 RP004484957 RP004486089 RP004494203 RP004500415 RP004507005 RP004507574 RP004507767 RP004512391 RP004512683 RP004513291 RP004521757 RP004522661 RP004525372 RP004527165 RP004534400 RP004534724 RP004535540 RP004536263 RP004542099 RP004547352 RP004549754 RP004550263
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0086680 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 38752 252138
Antibody Information
	polyclonal (Anderson et al., 1995, Hasegawa et al., 2011, Treacy et al., 1991)
Other Information
Discoverer
	Muller.
Etymology
	Transheterozygous vvl mutants show lack of veins with a ventral component. (Diaz-Benjumea and Garcia-Bellido, 1990)
Identification
Relationship to Other Genes
Source for database identity of	Source for identity of: vvl CG10037 (Gabrielian, 1999.11)
Source for database merge of	Source for merge of: vvl prd3 (Gelbart, 2000.9.30) Source for merge of: vvl sep (Diaz-Benjumea and Garcia-Bellido, 1990) Source for merge of: vvl ut (Salzberg and Inbal, 2003)
Additional comments
Other Comments
	The Hrp59 protein binds preferentially to a subset of mRNAs, including vvl mRNA. (Kiesler et al., 2005) vvl may regulate dendritic targeting and coordinate dendritic and axonal connectivity of projection neurons in the olfactory system to ensure the highly stereotypes acquisition and delivery of olfactory information by the central olfactory neurons. (Komiyama et al., 2003) D and vvl act in concert with sim to control midline gene expression in the embryo. (Ma et al., 2000) vvl is required for the maintenance of btl expression during tracheal tree development. vvl is also required for the specific expression in the tracheal cells of tkv and rho. vvl makes the tracheal cells competent to further signalling through activation of its target genes. (Llimargas and Casanova, 1997) Maintenance of high level btl expression requires wild-type vvl function. Characterisation of btl regulatory sequences identifies seven high affinity and one low affinity vvl recognition elements, implying direct transcriptional regulation of btl expression by vvl. (Anderson et al., 1996) Tracheal cell migration defects in vvl mutant embryos are rescued by ubiquitous expression of btl. (Anderson et al., 1996) Two variant POU domain recognition elements, DFRE1 and DFRE2, have been identified within a 514bp vvl autoregulatory enhancer. DFRE1 and DFRE2 show separable tissue-specific functions when independently disrupted or deleted. (Certel et al., 1996) vvl function in CNS development is studied by misexpression of the protein both temporally and spatially. Results suggest that premature vvl expression disrupts initial mesectodermal lineage designation and therefore subsequent midline differentiation. (Certel and Johnson, 1996) acj6 protein does not affect DNA binding by the vvl protein, and no interaction between the acj6 and vvl proteins can be detected. (Turner, 1996) Examination of the mutant CNS and tracheal phenotypes suggests a requirement for vvl protein for the correct differentiation and/or migration of tracheal cells and midline glia in the CNS. (Anderson et al., 1995) The pattern of vvl expression during embryonic development has been analysed in wild-type and mutant embryos. (Billin and Poole, 1995) Examination of CNS formation in vvl loss of function mutants suggest that the vvl gene is required for the correct differentiation of selected neurons and glia in the CNS. Ecol\lacZ reporter gene constructs suggest that vvl is capable of selectively altering mesectodermal cell fates. (Certel and Johnson, 1995) vvl is required for the differentiation of dorsal and ventral veins, and for cell proliferation in internal regions of the wing. (de Celis and Ashburner, 1995) vvl links positional information and cell differentiation during embryonic and imaginal development. During embryogenesis vvl is required for the formation of the tracheal tree and in the patterning of the ventral ectoderm. During imaginal development vvl is required for cell proliferation and the differentiation of the wing veins. vvl expression is dependent on the coordinate activities of dpp, wg and hh. vvl interacts with other genes involved in vein differentiation, including rho, tkv, Egfr, dpp and N. (de Celis et al., 1995) vvl is required in developmental processes as diverse as tracheal tree elongation and ventral ectoderm development during embryonic stages and cell proliferation and vein differentiation in the wing imaginal disc. It is proposed that vvl integrates information from different signaling molecules and in turn regulates the expression of specific cell differentiation genes during tracheal development and vein differentiation. (Llimargas et al., 1995) Mutation in vvl causes abnormal positioning of the chordotonal organs. (Salzberg et al., 1994) vvl is required for PNS development in the embryo. (Salzberg et al., 1994) One of the homeodomain loci identified in a screen for genes encoding DNA binding proteins capable of binding to a consensus Engrailed binding site. (Kalionis and O'Farrell, 1993) An alternative transcript of Ipou, lacking two basic amino acids, is incapable of dimerizing with Cf1a. (Treacy et al., 1992) vvl contains a POU domain, and is expressed at maximum levels during early development. (Billin et al., 1991) Cf1a is a transactivator of the Ddc gene and is coexpressed in subsets of neurons with Ipou during development. Ipou forms a highly stable heterodimeric complex with Cf1a and inhibits its ability to bind DNA and activate transcription of Ddc. (Treacy et al., 1991) Mutations in vvl have no pleiotropic phenotypes in embryonic patterns. (Diaz-Benjumea and Garcia-Bellido, 1990) vvl encodes a sequence specific DNA binding protein. The vvl gene product binds to a DNA region required for the expression of Ddc. (Johnson and Hirsh, 1990) ve, vn, ci, cg, svs, ast(S), H, Vno and vvl belong to the vein phenotypic group (Puro, 1982, Droso. Info. Serv. 58:205--208) within the 'lack-of-vein' mutant class. Loss-of-function alleles at these loci remove stretches of veins in two or more longitudinal veins. Double mutations within members of this group remove all veins, have smaller, slightly lanceolate wings, no sensilla and extra chaetae. Some mutant alleles of vvl prevent ventral vein differentiation. Embryonic lethal alleles also exist. (Diaz-Benjumea et al., 1989) Identified on the basis of similarity to a 600bp prd cDNA fragment which codes for the last 205 amino acids of prd. (Frigerio et al., 1986)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079224 NM_001169884 RefSeq (Proteins) • NP_523948 NP_001163355 DDBJ / EMBL / GenBank • AAA28831 AAF50641 AAO39521 ACZ94626 BI229419 BT003517 CAA36496 CAA41341 M81959 X52252 X58435 Entrez Gene - A searchable database of RefSeq genes. • 38752 UniProtKB/Swiss-Prot • P16241
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Homeobox, conserved site (IPR017970) POU-specific (IPR000327) Homeobox (IPR001356) Homeodomain-like (IPR009057) Lambda repressor-like, DNA-binding (IPR010982) Homeodomain-related (IPR012287) POU (IPR013847) Transcription factor, homeobox/POU (IPR016362)
Linkouts
	BioGRID - A database of protein and genetic interactions • 64202 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0086680 DroID - A comprehensive database of gene and protein interactions. • FBgn0086680 DRSC - Results from RNAi screens. • FBgn0086680 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0086680 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • S.C3L000862 FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0086680 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 38752 252138 InParanoid A subset of ortholog calls from InParanoid. • AAEL003560-PA Aedes aegypti (mosquito) AGAP005878-PA Anopheles gambiae (mosquito) XP_393686.2 Apis mellifera (honey bee) XP_001948842.1 Acyrthosiphon pisum (pea aphid) BGIBMGA010868-PA Bombyx mori (silk worm) ENSBTAP00000010443 Bos taurus (bovine) CPIJ002221 Culex pipens (mosquito) ENSDARP00000037034 Danio rerio (zebra fish) ENSECAP00000017373 Equus caballus (horse) ENSGACP00000002556 Gymnopilus aculeatus (mushroom) ENSP00000329170 Homo sapiens (human) ISCW020718-PA Ixodes scapularis (deer tick) ENSMODP00000029696 Monodelphis domestica (gray opossum) ENSMUSP00000063809 Mus musculus (house mouse) ENSPTRP00000032636 Pan troglodytes (chimpanzee) ENSRNOP00000009043 Rattus norvegicus (norway rat) XP_968364.2 Tribolium castaneum (red flour beetle) ENSTNIP00000019394 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000008884 Takifugu rubripes (pufferfish Takifugu) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /gene/drifter.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 38752 modMine - Data generated by the modENCODE project. • FBgn0086680 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB16730 NV14897 LH23007 ISCW020718 FBgn0200336 FBgn0185665 FBgn0169718 JGI_V11_14402 FBgn0070083 FBgn0135721 FBgn0122836 FBgn0097052 CPIJ002221 CPIJ019261 BGIBMGA010868 ACYPI008866 GB16085 AAEL003560 PHUM605420 GLEAN_14350 REDfly - A database of transcriptional regulatory elements. • FBgn0086680
Synonyms & Secondary IDs ( 33 )
Reported As
Symbol Synonym	Cf1a (Reim et al., 2003, Johnson, 1995.7.19, Nakagoshi et al., 1998, Nakachi et al., 1997, Billin and Poole, 1995, Frasch, 1995, Manning and Krasnow, 1993, Kalionis and O'Farrell, 1993, Billin et al., 1991, Janknecht et al., 1991, Wintermantel et al., 1991, Johnson and Hirsh, 1990, ) Cf1-a (Gehring et al., 1994) Cf1a/drifter (Arendt and Nubler-Jung, 1999) Cfla (Billin, 1991.12.16) CG10037 (Perkins et al., 2009.3.31, Keleman et al., 2009.8.5) dfr (Howard and Davidson, 2004, Grueber and Jan, 2004, Komiyama et al., 2003, Certel et al., 2001, Glazer and Shilo, 2001, Certel et al., 2001, Ma et al., 2000, Certel et al., 2000, Certel et al., 2000, Certel and Johnson, 1999, Koizumi et al., 1997, Certel et al., 1997, Turner, 1996, Certel and Johnson, 1996, Certel et al., 1996, Isaac and Andrew, 1996, Anderson et al., 1996, Anderson et al., 1995, Anderson et al., 1995, Certel and Johnson, 1995, Anderson and Johnson, 1995, Estes et al., 2008, Meier et al., 2006, Sonnenfeld et al., 2010, Morozova et al., 2010) DFR (Bradley and Andrew, 2001, Certel and Johnson, 1999, Certel and Johnson, 1997) Dfr/Vvl (Petit et al., 2002, Junell et al., 2010) dfr/vvl (Zelzer and Shilo, 2000, Junell et al., 2007) dfr-vvl (Affolter and Shilo, 2000) drf (Kambadur et al., 1998, Sotillos et al., 2010, Hasegawa et al., 2011) Drf (Michaud and Tanguay, 2003) prd3   sep (Reichhart and Ferrandon, 1998, ) ut (Inbal et al., 2004, Salzberg and Inbal, 2003, Salzberg et al., 1997, Salzberg et al., 1994, ) Vv1 (Gomes and Schweisguth, 2005) vvl (Diaz-Benjumea and Garcia-Bellido, 1990, Kiesler et al., 2005, Hombria et al., 2005, Wheeler et al., 2006, Jiang and Crews, 2006, Castelli-Gair Hombria, 2006, Arbouzova et al., 2006, Sotillos and De Celis, 2005, Sotillos and de Celis, 2006, Aerts et al., 2007, Meier et al., 2006, Araujo et al., 2007, Sotillos et al., 2008, Weake et al., 2011, Miller et al., 2009, Dworkin et al., 2009, Brodu et al., 2004, Sotillos et al., 2010, Buffin and Gho, 2010, Tolhuis et al., 2011, Letizia et al., 2011, Morozova et al., 2010, Dworkin et al., 2011) VVL (Inbal and Salzberg, 2002, Bradley and Andrew, 2001) vvl/dfr (Junell et al., 2007) vvl/dfr/ut (Matsuda et al., 2006)
Name Synonym	Drifter (Komiyama et al., 2004, Jan and Yan, 2003, Fichelson and Gho, 2004, Gomes and Schweisguth, 2005, Certel et al., 2000, Certel and Johnson, 1999, Michaud and Tanguay, 2003, Hasegawa et al., 2011) drifter (Hallem and Carlson, 2004, Gao and Bogert, 2003, Barolo and Posakony, 2002, Allan et al., 2001, Beitel and Krasnow, 2000, Nikitin and Thomas, 2000, Crews and Fan, 1999, Montell, 1999, Rhee et al., 1998, Ma et al., 1999, Clyne et al., 1999, Zelzer and Shilo, 1998, Crews, 1998, Ma et al., 1998, Odenwald et al., 1997, Klaembt et al., 1996, Odenwald et al., 1996, Kambadur et al., 1995, Grueber and Jan, 2004, Salzberg and Inbal, 2003, Certel and Johnson, 1999, Kambadur et al., 1998, Anderson et al., 1996, Certel and Johnson, 1996, Certel et al., 1996, Anderson et al., 1995, Certel and Johnson, 1995, Anderson and Johnson, 1995, Komiyama and Luo, 2007, Chung et al., 2009, Sonnenfeld et al., 2010) Drifter/Ventral veinless (Petit et al., 2002, Junell et al., 2010) paired-like 3   PRD gene 3 (Frigerio et al., 1986) separated   U-turn (Gomes and Schweisguth, 2005) u-turn (Salzberg et al., 1994, ) ventral veinless (Wheeler et al., 2006, Jiang and Crews, 2006, Diaz-Benjumea and Garcia-Bellido, 1990, Sotillos et al., 2008, Chung et al., 2009, Sotillos et al., 2010, Letizia et al., 2011, Sonnenfeld et al., 2010) Ventral veinless (Sotillos and De Celis, 2005) Ventral Veinless (Morozova et al., 2010) ventral veins lacking (Schwartz and Pirrotta, 2007, Meier et al., 2006, Carpenter et al., 2009, Buffin and Gho, 2010) Ventral veins lacking (Miller et al., 2009)
Secondary FlyBase IDs
	FBgn0000285 FBgn0003146 FBgn0003995 FBgn0010170 FBgn0011727
References ( 191 )
Generate a list of	All references relating to this gene ( 191 )
List References by type	Research paper  ( 102 ) Supplementary material  ( 3 ) Review  ( 29 ) Personal communication to FlyBase  ( 9 ) Abstract  ( 37 ) FlyBase analysis  ( 1 ) DNA/RNA sequence record  ( 2 ) Curated genome annotation  ( 2 ) Protein sequence record  ( 1 ) Book  ( 1 ) Computer file  ( 2 ) Automatic genome annotation  ( 1 )
Recent research papers ( 10 )
	Dworkin et al., 2011, Genetics 187(4): 1171--1184 The effects of weak genetic perturbations on the transcriptome of the wing imaginal disc and its association with wing shape in Drosophila melanogaster. [FBrf0214376] Hasegawa et al., 2011, Development 138(5): 983--993 Concentric zones, cell migration and neuronal circuits in the Drosophila visual center. [FBrf0213020] Letizia et al., 2011, J. Cell Sci. 124(2): 240--251 Regulated Crb accumulation controls apical constriction and invagination in Drosophila tracheal cells. [FBrf0212559] Tolhuis et al., 2011, PLoS Genet. 7(3): e1001343 Interactions among Polycomb Domains Are Guided by Chromosome Architecture. [FBrf0213356] Weake et al., 2011, Genes Dev. 25(14): 1499--1509 Post-transcription initiation function of the ubiquitous SAGA complex in tissue-specific gene activation. [FBrf0214372] Buffin and Gho, 2010, PLoS ONE 5(2): e9285 Laser microdissection of sensory organ precursor cells of Drosophila microchaetes. [FBrf0210039] Junell et al., 2010, Mol. Cell. Biol. 30(14): 3672--3684 The POU transcription factor Drifter/Ventral veinless regulates expression of Drosophila immune defense genes. [FBrf0211117] Morozova et al., 2010, Dev. Genes Evol. 220(7-8): 191--206 The Drosophila jing gene is a downstream target in the Trachealess/Tango tracheal pathway. [FBrf0214147] Sonnenfeld et al., 2010, Dev. Genes Evol. 220(7-8): 207--220 Drosophila Jing is part of the breathless fibroblast growth factor receptor positive feedback loop. [FBrf0214215] Sotillos et al., 2010, Dev. Biol. 340(2): 571--582 An efficient approach to isolate STAT regulated enhancers uncovers STAT92E fundamental role in Drosophila tracheal development. [FBrf0210508] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...