FlyBase Gene Report: Dmel\inv

General Information

Symbol

Dmel\inv

Species

D. melanogaster

Name

invected

Annotation Symbol

CG17835

Feature Type

protein_coding_gene

FlyBase ID

FBgn0001269

Gene Model Status

Current

Stock Availability

21 publicly available

Gene Summary

invected (inv) encodes a homeodomain transcription factor that is closely related to, co-expressed with, and functionally redundant with the product of en. Although dispensable in the laboratory, the product of inv is a transcriptional repressor important for segmentation in the embryo and formation of the posterior compartment in imaginal discs. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Gene Group
UniProtKB
Red Book
Interactive Fly

Also Known As

in, engrailed-related

Key Links

Genomic Location

Cytogenetic map

47F15-47F17

Sequence location

2R:11,474,465..11,509,695 [+]

Recombination map

2-64

RefSeq locus

NT_033778 REGION:11474465..11509695

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

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

Function

Gene Ontology (GO) Annotations (11 terms)

Molecular Function (3 terms)

Terms Based on Experimental Evidence (0 terms)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

enables DNA binding

inferred from electronic annotation with InterPro:IPR000047, InterPro:IPR000747, InterPro:IPR001356, InterPro:IPR020479

(InterPro Project Members, 2004-)

enables DNA-binding transcription factor activity, RNA polymerase II-specific

inferred from electronic annotation with InterPro:IPR017970

(InterPro Project Members, 2004-)

inferred from biological aspect of ancestor with PANTHER:PTN000678675

(GO Reference Genome Project, 2011-)

enables RNA polymerase II cis-regulatory region sequence-specific DNA binding

inferred from biological aspect of ancestor with PANTHER:PTN000678675

(GO Reference Genome Project, 2011-)

Biological Process (7 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

involved_in imaginal disc-derived wing vein specification

inferred from mutant phenotype

(DeVido et al., 2008)

involved_in neuroblast fate determination

inferred from genetic interaction with FLYBASE:en; FB:FBgn0000577,FLYBASE:gsb; FB:FBgn0001148

(Bhat and Schedl, 1997)

inferred from genetic interaction with FLYBASE:en; FB:FBgn0000577,FLYBASE:gsb; FB:FBgn0001148,FLYBASE:ptc; FB:FBgn0003892

(Bhat and Schedl, 1997)

Terms Based on Predictions or Assertions (5 terms)

CV Term

Evidence

References

involved_in compartment pattern specification

traceable author statement

(Milan and Cohen, 2000)

involved_in neuron differentiation

inferred from biological aspect of ancestor with PANTHER:PTN000678675

(GO Reference Genome Project, 2011-)

involved_in regulation of DNA-templated transcription

inferred from electronic annotation with InterPro:IPR017970

(InterPro Project Members, 2004-)

involved_in regulation of transcription by RNA polymerase II

inferred from biological aspect of ancestor with PANTHER:PTN000678674

(GO Reference Genome Project, 2011-)

involved_in wing disc anterior/posterior pattern formation

traceable author statement

(Milan and Cohen, 2000)

Cellular Component (1 term)

Terms Based on Experimental Evidence (0 terms)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

is_active_in nucleus

inferred from biological aspect of ancestor with PANTHER:PTN000678674

(GO Reference Genome Project, 2011-)

located_in nucleus

inferred from electronic annotation with InterPro:IPR000747

(InterPro Project Members, 2004-)

Gene Group (FlyBase)

NK-LIKE HOMEOBOX TRANSCRIPTION FACTORS

Protein Family (UniProt)

Belongs to the engrailed homeobox family. (P05527)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

Gene Group (FlyBase)

NK-LIKE HOMEOBOX TRANSCRIPTION FACTORS -: NK-like (NKL) homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. NKL transcription factors are homeobox genes closely related to Hox-like genes, a number of which are found in the NK cluster. Many of the NKL members contain an Engrailed Homology 1 (EH1) motif. (Adapted from FBrf0232555 and PMID:22094586).

Protein Function (UniProtKB)

Engrailed (en) and invected (inv) are functionally redundant transcription factors in neuronal precursor cell NB5-3 specification. Inv is unable to substitute for en in other regulatory processes such as maintaining gsb expression in the neuroectoderm after stage 10 of embryogenesis. Maintenance of gsb expression in row 5 of the neuroectoderm involves an as yet unidentified short range signaling molecule.

(UniProt, P05527)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

inv: invected

Although a developmental role for inv is indicated by hybridization, experiments with embryonic and larval cells using cDNA probes, the function in development of the protein encoded by inv is not known (Coleman et al., 1987). No mutant alleles were reported.

Summary (Interactive Fly)

transcription factor - homeodomain - engrailed class - cooperates with engrailed in establishing posterior compartment identity

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\inv for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)

Isoform displayed:

Pfam protein domains

InterPro name

classification

start

end

Protein Domains (via SMART)

Isoform displayed:

SMART protein domains

InterPro name

classification

start

end

Structure

Protein 3D structure (Predicted by AlphaFold) (AlphaFold entry P05527)

If you don't see a structure in the viewer, refresh your browser.

Model Confidence:

Very high (pLDDT > 90)
Confident (90 > pLDDT > 70)
Low (70 > pLDDT > 50)
Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Experimentally Determined Structures

Crossreferences

Comments on Gene Model

Gene model reviewed during 5.49

Gene model reviewed during 5.56

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

inv-RB

FBtr0088069

NM_078975

2261

576

inv-RD

FBtr0088068

NM_165839

3404

576

inv-RE

FBtr0345673

NM_165838

4776

576

Additional Transcript Data and Comments

Reported size (kB)

2.7 (northern blot)

(Coleman et al., 1987)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

inv-PB

FBpp0087175

60.9

576

6.25

FBpp0087174

60.9

576

6.25

FBpp0311724

60.9

576

6.25

P05527

NP_725056

AAM68707

Polypeptides with Identical Sequences

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

576 aa isoforms: inv-PB, inv-PD, inv-PE

Additional Polypeptide Data and Comments

Reported size (kDa)

576 (aa); 60 (kD predicted)

(Coleman et al., 1987)

Comments

External Data

Crossreferences

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

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

P05527

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\inv using the Feature Mapper tool.

External Data

Crossreferences

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

inv_1
inv_2

Linkouts

Expression Data

Testis-specificity index

The testis specificity index was calculated from modENCODE tissue expression data by Vedelek et al., 2018 to indicate the degree of testis enrichment compared to other tissues. Scores range from -2.52 (underrepresented) to 5.2 (very high testis bias).

0.77

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

dorsal ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

ventral ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

embryonic stage 7 -- 8

dorsal ectoderm

(Fisher et al., 2012)

organism | segmentally repeated

(Fisher et al., 2012)

ventral ectoderm

(Fisher et al., 2012)

embryonic stage 9 -- 10

ventral ectoderm

(Fisher et al., 2012)

embryonic stage 11 -- 12

larval foregut atrium | precursor

Comment: reported as atrium primordium

(Fisher et al., 2012)

clypeo-labral primordium

(Fisher et al., 2012)

ventral epidermis primordium

(Fisher et al., 2012)

embryonic stage 13 -- 16

larval foregut atrium

(Fisher et al., 2012)

embryonic epipharynx

(Fisher et al., 2012)

embryonic head epidermis

(Fisher et al., 2012)

embryonic hindgut

(Fisher et al., 2012)

embryonic ventral epidermis

(Fisher et al., 2012)

ventral midline of embryo

(Fisher et al., 2012)

radioisotope in situ

Stage

Tissue/Position (including subcellular localization)

Reference

extended germ band stage

organism | segmentally repeated

(Coleman et al., 1987)

embryonic stage 13

inclusive hindgut primordium

(Coleman et al., 1987)

organism | posterior | segmentally repeated

(Coleman et al., 1987)

third instar larval stage

wing disc posterior compartment

(Coleman et al., 1987)

Additional Descriptive Data

inv transcripts first appear in embryogensis midway through germ-band elongation, at low levels. At the completion of germ band elongation, inv transcript is expressed in a repeated pattern of 2 to 4 segments. Subsequently, inv is expressed in a pattern coincident with that of its neighboring gene, en, in the posterior compartment of each segment. Both en and inv are also expressed in the hindgut, the labral segment, and ganglia. In third instar larvae, en and inv are expressed in the posterior compartment of wing imaginal discs.

(Coleman et al., 1987)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 8 -- 11

organism | segment boundary

(Wech et al., 1999)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

Expression Deduced from Reporters

Reporter: P{en-lacZ(wXba)}inv^Xba21

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage

(Sun et al., 1995)

third instar larval stage

wing disc | restricted

(Sun et al., 1995)

leg disc | restricted

(Sun et al., 1995)

antennal disc | restricted

(Sun et al., 1995)

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\inv in JBrowse

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\inv

View exonic expression by tissue for Dmel\inv

Bulk Downloads

RNA-Seq RPKM values for all genes

Reference

See Gelbart and Emmert, 2013 for analysis details and data files for all genes.

Developmental Proteome: Life Cycle

Developmental Proteome: Embryogenesis

External Data and Images

Linkouts

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0001269

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

36239/tissue=All

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

FBgn0001269

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0001269

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

FBgn0001269

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

CG17835

Flygut - An atlas of the Drosophila adult midgut

FBgn0001269

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\inv

Stages(s) 1-3
Stages(s) 4-6
Stages(s) 7-8
Stages(s) 9-10
Stages(s) 11-12
Stages(s) 13-16

FlyBase Wiki

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 50 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 12 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of inv

Transgenic constructs containing regulatory region of inv

Aberrations (Deficiencies and Duplications) ( 20 )

Inferred from experimentation ( 20 )

Gene disrupted in

Df(2R)Δ110

(De et al., 2016)

Df(2R)en-11

(Sanicola et al., 1995)

Df(2R)en^C

(Saenz-Robles et al., 1995)

Df(2R)en^D

(Saenz-Robles et al., 1995)

Df(2R)en^E

(Zhai et al., 2009, Lawrence et al., 1999, Blair and Ralston, 1997, Chen and Baker, 1997, Whiteley and Kassis, 1997, Azpiazu et al., 1996, Gustavson et al., 1996, Lawrence et al., 1996, Martin-Bermudo and Brown, 1996, Matsuzaki and Saigo, 1996, Strutt and Mlodzik, 1996, Tabata et al., 1995, Zecca et al., 1995, Cadigan et al., 1994, Hidalgo, 1994)

Df(2R)en-Dgr;110

(De et al., 2019)

Df(2R)en-SFX31

(Joly et al., 2007, Whiteley and Kassis, 1997, Serrano et al., 1995)

In(2R)vg^W

(Simmonds et al., 1997, Williams et al., 1990)

In(2R)vg^WR1

(Williams et al., 1990)

In(2R)vg^WR3

(Williams et al., 1990)

In(2R)vg^WR4

(Williams et al., 1990)

In(2R)vg^WRK1

(Simmonds et al., 1997)

In(2R)vg^WRK2

(Simmonds et al., 1997)

In(2R)vg^WRK3

(Simmonds et al., 1997)

In(2R)vg^WRK4

(Simmonds et al., 1997)

In(2R)vg^WRK5

(Simmonds et al., 1997)

In(2R)vg^WRK6

(Simmonds et al., 1997)

T(2;?)inv^l77

(Gustavson et al., 1996)

Gene not disrupted in

Df(2R)en^F

(Gustavson et al., 1996)

Gene partially disrupted in

Df(2R)en^Δ530

(DeVido et al., 2008)

Inferred from location ( 10 )

Variants

Variant Molecular Consequences

Alleles Representing Disease-Implicated Variants

Phenotypes

For more details about a specific phenotype click on the relevant allele symbol.

Lethality

Allele

partially lethal - majority die

inv^Δ45

partially lethal - majority die (with Df(2R)en^E)

inv^Δ45

viable

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

inv^GD16474

viable, with Scer\GAL4^tin.CΔ4

inv^GD16474

Sterility

Allele

fertile

Other Phenotypes

Allele

abnormal immune response | adult stage

inv^GE10665

majority die during pharate adult stage

inv^Δ45

majority die during pharate adult stage (with Df(2R)en^E)

inv^Δ45

visible

visible (with Df(2R)en^Δ530)

inv^17en26

visible (with Df(2R)en-SFX31)

inv^17en26

visible | dominant

visible | dominant | homeotic | heat sensitive

inv^W

Phenotype manifest in

Allele

anterior crossvein

inv³⁰

anterior-posterior compartment boundary of the wing blade

cuticle

inv^12A-11

embryonic hindgut

inv³⁰

larval longitudinal connective

inv^E

presumptive embryonic/larval muscle system

inv^E

segment

inv^12A-11

symmetrical commissure | embryonic stage

inv^E

ventral denticle belt

inv^hs.PG

wing

wing (with Df(2R)en^E)

inv^Δ45

wing | anterior

inv^W

wing vein

wing vein (with Df(2R)en^Δ530)

inv^17en26

wing vein (with Df(2R)en-SFX31)

inv^17en26

wing vein | ectopic

wing vein | ectopic (with Df(2R)en^E)

inv^Δ45

wing vein L1

inv^W

wing vein L2

inv^W

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (127)

Hsap\EN1

9 of 14

Yes

Hsap\EN2

8 of 14

Yes

Hsap\CPHXL

2 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Hsap\DUX3

1 of 14

Yes

Hsap\DUX4

1 of 14

Hsap\DUX4L4

1 of 14

Yes

Hsap\DUX4L7

1 of 14

Yes

Hsap\DUX4L9

1 of 14

Yes

Hsap\DUX5

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

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1 of 14

Hsap\LOC112268164

1 of 14

Yes

1 of 14

1 of 14

1 of 14

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1 of 14

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1 of 14

Yes

1 of 14

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Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (60)

Rnor\En1

8 of 14

Yes

Rnor\En2

8 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Rnor\Hoxa5

1 of 14

Rnor\Hoxa11

1 of 14

Rnor\Hoxa13

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Mus musculus (laboratory mouse) (78)

Mmus\En1

8 of 14

Yes

Mmus\En2

7 of 14

Mmus\Cphx1

2 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

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1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

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1 of 14

1 of 14

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1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Mmus\Rhox4a

1 of 14

Yes

Mmus\Rhox4c

1 of 14

Yes

Mmus\Rhox4f

1 of 14

Yes

Mmus\Rhox4g

1 of 14

Yes

Mmus\Rhox6

1 of 14

Yes

Mmus\Rhox7a

1 of 14

Yes

Mmus\Rhox8

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (107)

Xtro\en2

7 of 13

Yes

Xtro\en1

5 of 13

Yes

1 of 13

1 of 13

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1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Xtro\homeobox100496651-provisional

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Xtro\hoxc13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

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1 of 13

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1 of 13

Yes

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1 of 13

Danio rerio (Zebrafish) (106)

Drer\en1a

8 of 14

Yes

Drer\en1b

8 of 14

Yes

7 of 14

7 of 14

1 of 14

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1 of 14

Yes

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Yes

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Caenorhabditis elegans (Nematode, roundworm) (36)

Cele\ceh-16

7 of 14

Yes

1 of 14

1 of 14

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1 of 14

1 of 14

Anopheles gambiae (African malaria mosquito) (56)

Agam\AgaP_AGAP008025

6 of 12

Yes

Agam\AgaP_AGAP008024

5 of 12

Yes

4 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Arabidopsis thaliana (thale-cress) (12)

Atha\ATML1

1 of 13

Yes

Atha\HB-7

1 of 13

Yes

Atha\HB51

1 of 13

Yes

Atha\HDG2

1 of 13

Yes

Atha\HDG3

1 of 13

Yes

Atha\HDG4

1 of 13

Yes

Atha\HDG8

1 of 13

Yes

Atha\HDG9

1 of 13

Yes

Atha\HDG10

1 of 13

Yes

Atha\HDG11

1 of 13

Yes

Atha\HDG12

1 of 13

Yes

Atha\PDF2

1 of 13

Yes

Saccharomyces cerevisiae (Brewer's yeast) (3)

Scer\RSC58

1 of 13

Yes

Scer\YHP1

1 of 13

Yes

Scer\YOX1

1 of 13

Yes

Schizosaccharomyces pombe (Fission yeast) (2)

Spom\phx1

1 of 12

Yes

Spom\rsc58

1 of 12

Yes

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:inv. Refer to their site for version information.

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (79)

8 of 13

bap

3 of 13

bsh

3 of 13

btn

3 of 13

cad

3 of 13

CG18599

3 of 13

Dbx

3 of 13

Dfd

3 of 13

ems

3 of 13

eve

3 of 13

exex

3 of 13

HGTX

3 of 13

HHEX

3 of 13

Hmx

3 of 13

lab

3 of 13

lms

3 of 13

NK7.1

3 of 13

unpg

3 of 13

zen2

3 of 13

zen

3 of 13

abd-A

2 of 13

Antp

2 of 13

Awh

2 of 13

bcd

2 of 13

B-H1

2 of 13

B-H2

2 of 13

C15

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

Dll

2 of 13

Drgx

2 of 13

ftz

2 of 13

Gsc

2 of 13

H2.0

2 of 13

hbn

2 of 13

ind

2 of 13

lbe

2 of 13

Lim1

2 of 13

Lmx1a

2 of 13

Scr

2 of 13

slou

2 of 13

tin

2 of 13

Ubx

2 of 13

unc-4

2 of 13

vnd

2 of 13

2 of 13

2 of 13

1 of 13

1 of 13

1 of 13

gsb

1 of 13

gsb-n

1 of 13

lbl

1 of 13

Lim3

1 of 13

OdsH

1 of 13

otp

1 of 13

PHDP

1 of 13

Pph13

1 of 13

prd

1 of 13

Ptx1

1 of 13

repo

1 of 13

scro

1 of 13

toy

1 of 13

tup

1 of 13

zfh1

1 of 13

zfh2

1 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 0 )

Allele

Disease

Evidence

References

Potential Models Based on Orthology ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 0 )

Allele

Disease

Interaction

References

Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)

Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.

Homo sapiens (Human)

Gene name

Score

OMIM

OMIM Phenotype

DO term

Complementation?

Transgene?

EN1; engrailed homeobox 1

9 of 14

131290

ENDOVE SYNDROME, LIMB-BRAIN TYPE; ENDOVESLB

EN2; engrailed homeobox 2

8 of 14

131310

Functional Complementation Data

Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.

Interactions

Summary of Physical Interactions

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

inv - CG12112

anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based

(Rhee et al., 2014)

inv - CG31955

two hybrid array

(Shokri et al., 2019)

inv - Rel

two hybrid array

(Shokri et al., 2019)

inv - drm

two hybrid array

(Shokri et al., 2019)

Summary of Genetic Interactions

Interaction Browsers

View in FlyBase Interactions Browser View in MIST tool (external link)

Please look at the allele data for full details of the genetic interactions

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

inv

suppressible

ph-d

(Simmonds and Bell, 1998)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

enhanceable

inv

(De et al., 2016, Whiteley and Kassis, 1997)

External Data

Linkouts

BioGRID - A database of protein and genetic interactions.

62027

DroID - A comprehensive database of gene and protein interactions.

FBgn0001269

MIST (genetic) - An integrated Molecular Interaction Database

36239

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

36239

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P05527

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

2-64

Cytogenetic map

47F15-47F17

Sequence location

2R:11,474,465..11,509,695 [+]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

47F15-47F17

Limits computationally determined from genome sequence between P{lacW}Tapδ^k17005&P{lacW}k05103 and P{lacW}l(2)k13403^k13403&P{lacW}Egm^k14708

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

Experimentally Determined Recombination Data

Location

2-64

(FlyBase, 2016)

2-64.5

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (21)

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

Bloomington

7088

inv³⁰/CyO

26891

w^*; P{EP}inv^GE10665

40519

w¹¹¹⁸; P{GMR88C02-GAL4}attP2

46851

w¹¹¹⁸; P{GMR88B12-GAL4}attP2

93587

y¹ w^*; PBac{inv-GFP.FPTB}VK00031

52163

y¹ w^*; Mi{MIC}inv^MI09433

14433

y¹ w^67c23; P{SUPor-P}inv^KG04405/SM6a

FlyORF

F004775

M{UAS-inv.ORF-VN}ZH-86Fb

F000258

M{UAS-inv.ORF.3xHA.GW}ZH-86Fb

VDRC

v101934

P{KK110281}VIE-260B

v49951

w¹¹¹⁸; P{GD16474}v49951

v342973

P{hsFLP}1, y1 w1118; P{HD_CFD02561}attP40/CyO-GFP

More stocks available...

Genomic Clones (27)

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

cDNA Clones (8)

Please Note This section lists cDNAs and ESTs that fall within the genomic extent of the gene model, which may include cDNAs and ESTs of genes within introns, or of overlapping genes. Please see JBrowse for alignment of the cDNAs and ESTs to the gene model.

cDNA clones, fully sequenced

BDGP DGC clones

LD31887

Other clones

DPiM_LD31887

Drosophila Genomics Resource Center cDNA clones

For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

FBgn0001269

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0001269

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Cheng et al., 2014)

Commercially Available Antibodies

Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research

4D9_anti-engrailed/invected

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

DNA-protein interactions: genome-wide binding profile assayed for inv protein in 0-12 hr embryos; see mE1_TFBS_inv collection report.

(Negre et al., 2011)

Either en or inv is required to uncouple the ptc-gsb regulatory circuit in row 5 neuroblasts during neurogenesis.

(Bhat and Schedl, 1997)

Pc, Psc and polyhomeotic proteins are associated with identical regulatory elements of en in tissue culture cells and differentially distributed on regulatory sequences of inv.

(Strutt and Paro, 1997)

inv gene can play a role in proper development.

(Whiteley and Kassis, 1997)

The varied phenotypes of en alleles can be explained by the differential effects these mutants have on the combination of en and inv activities.

(Gustavson et al., 1996)

en and inv share a regulatory region and encode redundant functions.

(Gustavson et al., 1996)

inv may be involved in determining posterior identity in the developing wing, alongside en.

(Guillen et al., 1995)

inv can perform a subset of functions attributed to en including the ability to confer posterior compartment identity, but does not appear to organize the anterior-posterior compartment boundary.

(Simmonds and Bell, 1995)

The establishment of the anterior-posterior organiser and control of compartment identity are genetically distinguishable and inv may perform a discrete subset of functions previously ascribed to en.

(Simmonds et al., 1995)

Expression pattern of Ecol\lacZ from silencer detector insertions P{lacW}65 and P{en-lacZ(wXba)}inv^Xba21 reflects that of inv.

(Sun et al., 1995)

inv is not essential for embryonic development, but contributes to an activity that is redundant with en.

(Tabata et al., 1995)

inv product differs from en in that it cannot block dpp transcription or activate expression of hh.

(Zecca et al., 1995)

inv has a region of extensive homology (that includes the homeobox) with en.

(Coleman et al., 1987)

Relationship to Other Genes

Source for database merge of

Additional comments

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (13)

Reported As

Symbol Synonym

CG17835

(Ibrahim et al., 2013, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Poole, 2004.9.20)

Engrailed/Invected

(Sedaghat et al., 2002)

INV

(Blatti et al., 2015)

Inv

(Klaus et al., 2023, Webber et al., 2013, Blagburn, 2008, Lebreton et al., 2008, Noyes et al., 2008, Peterson-Nedry et al., 2008)

(Poole et al., 1985)

(Herz et al., 2014, Gunawardena and Rykowski, 2000)

inv

(Berardi et al., 2025, Brown et al., 2024, Li et al., 2024, Cheng et al., 2023, Molina-Gil et al., 2023, Özsoy et al., 2021, Pang et al., 2021, Bridi et al., 2020, De et al., 2020, Iijima et al., 2020, Mahmud et al., 2020, Miscopein Saler et al., 2020, Bageritz et al., 2019, Bailetti et al., 2019, Bridi et al., 2019, De et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Eagen et al., 2017, Karaiskos et al., 2017, Kline et al., 2017, Liu et al., 2017, Transgenic RNAi Project members, 2017-, Bürglin and Affolter, 2016, Gene Disruption Project members, 2016-, Pearson et al., 2016, Peng et al., 2016, Ray et al., 2016, Baëza et al., 2015, Gurdziel et al., 2015, Kavi et al., 2015, Schertel et al., 2015, Cheng et al., 2014, Ghavi-Helm et al., 2014, Jiang and Singh, 2014, Toshima et al., 2014, Ibrahim et al., 2013, Saunders et al., 2013, Langlais et al., 2012, Carreira et al., 2011, Layalle et al., 2011, Nègre et al., 2011, Pérez et al., 2011, Biehs et al., 2010, Cunningham et al., 2010, Kannan et al., 2010, Schwartz et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Yassin et al., 2010, Carreira et al., 2009, Kwon et al., 2009, Levis, 2009.3.18, Mulinari and Häcker, 2009, Seugnet et al., 2009, Song et al., 2009, Zhai et al., 2009, Casso et al., 2008, Colomb et al., 2008, DeVido et al., 2008, Durant and Kassis, 2008, Jin et al., 2008, Beltran et al., 2007, Joly et al., 2007, Comet et al., 2006, Negre et al., 2006, Peel et al., 2006, Schwartz et al., 2006, Copley, 2005, Brodsky et al., 2004)

Name Synonyms

Invected

(Wang and Dahmann, 2020, Dahmann et al., 2011, Landsberg et al., 2009, Mirth et al., 2009)

engrailed-invected

engrailed-related

(Poole et al., 1985)

transcript group IV

Secondary FlyBase IDs

Datasets (1)

Study focus (1)

Experimental Role

Project

Project Type

Title

transgene_used

modENCODE_regulation_TFs

genome binding

Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 73 )

Sequence Crossreferences

NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.

36239

GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.

GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.

RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.

UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.

P05527

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

P05527

Other crossreferences

AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.

P05527

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0001269

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

36239/tissue=All

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

FBgn0001269

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

FBgn0001269

FlyMine - An integrated database for Drosophila genomics

FBgn0001269

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

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

dme:Dmel_CG17835

MARRVEL_MODEL - MARRVEL (model organism gene)

36239

Linkouts

BioGRID - A database of protein and genetic interactions.

62027

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

FBgn0001269

DroID - A comprehensive database of gene and protein interactions.

FBgn0001269

DRSC - Results frm RNAi screens

FBgn0001269

Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research

4D9_anti-engrailed/invected

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

inv_1
inv_2

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0001269

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0001269

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

CG17835

Flygut - An atlas of the Drosophila adult midgut

FBgn0001269

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

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

36239

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

36239

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

P05527

References (281)

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

0.77

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