FlyBase Gene Report: Dmel\ara

General Information

Symbol

Dmel\ara

Species

D. melanogaster

Name

araucan

Annotation Symbol

CG10571

Feature Type

protein_coding_gene

FlyBase ID

FBgn0015904

Gene Model Status

Current

Stock Availability

51 publicly available

Gene Summary

araucan (ara) encodes a homeodomain-containing protein of the TALE subfamily and one of the three components of the Iroquois gene complex. ara product acts as a transcriptional regulator to control territorial and cell fate specification, cell sorting behaviour and pattern formation. It also plays a transcription independent cell-autonomous role in cell proliferation control. [Date last reviewed: 2018-09-20] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Gene Group
UniProtKB
Interactive Fly

Also Known As

iro, iroquois, Iro-C, IroC

Key Links

Genomic Location

Cytogenetic map

69C8-69C10

Sequence location

3L:12,580,672..12,596,018 [+]

Recombination map

3-38

RefSeq locus

NT_037436 REGION:12580672..12596018

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 (19 terms)

Molecular Function (3 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

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

inferred from direct assay

(Gomez-Skarmeta et al., 1996)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

enables DNA binding

inferred from electronic annotation with InterPro:IPR001356, InterPro:IPR003893, InterPro:IPR008422

(InterPro Project Members, 2004-)

inferred from electronic annotation with InterPro:IPR001356, InterPro:IPR008422

(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:PTN002257217

(GO Reference Genome Project, 2011-)

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

inferred from biological aspect of ancestor with PANTHER:PTN002257217

(GO Reference Genome Project, 2011-)

Biological Process (14 terms)

Terms Based on Experimental Evidence (8 terms)

CV Term

Evidence

References

involved_in compound eye morphogenesis

inferred from mutant phenotype

(Tomlinson, 2003)

involved_in dendrite morphogenesis

inferred from mutant phenotype

(Iyer et al., 2013)

involved_in imaginal disc-derived wing morphogenesis

inferred from mutant phenotype

(Gomez-Skarmeta et al., 1996)

involved_in imaginal disc-derived wing vein specification

inferred from genetic interaction with FLYBASE:rho; FB:FBgn0004635

(Gomez-Skarmeta et al., 1996)

inferred from genetic interaction with FLYBASE:caup; FB:FBgn0015919

(Gomez-Skarmeta et al., 1996)

involved_in muscle cell fate commitment

inferred from genetic interaction with FLYBASE:caup; FB:FBgn0015919

(Carrasco-Rando et al., 2011)

involved_in negative regulation of growth

inferred from mutant phenotype

(Barrios et al., 2015)

involved_in notum cell fate specification

inferred from mutant phenotype

(Wang et al., 2000)

involved_in positive regulation of transcription by RNA polymerase II

inferred from mutant phenotype

(Gomez-Skarmeta et al., 1996)

Terms Based on Predictions or Assertions (6 terms)

CV Term

Evidence

References

involved_in cell development

inferred from biological aspect of ancestor with PANTHER:PTN002257217

(GO Reference Genome Project, 2011-)

involved_in equator specification

traceable author statement

(McNeill, 2000)

involved_in neuron differentiation

inferred from biological aspect of ancestor with PANTHER:PTN000129978

(GO Reference Genome Project, 2011-)

involved_in positive regulation of DNA-templated transcription

non-traceable author statement

(Calleja et al., 2002)

involved_in regulation of DNA-templated transcription

inferred from electronic annotation with InterPro:IPR003893, InterPro:IPR008422, InterPro:IPR017970

(InterPro Project Members, 2004-)

inferred from electronic annotation with InterPro:IPR008422, InterPro:IPR017970

(InterPro Project Members, 2004-)

involved_in regulation of transcription by RNA polymerase II

inferred from biological aspect of ancestor with PANTHER:PTN002257217

(GO Reference Genome Project, 2011-)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

located_in cytoplasm

inferred from high throughput direct assay

(Sarov et al., 2016)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

is_active_in nucleus

inferred from biological aspect of ancestor with PANTHER:PTN002257217

(GO Reference Genome Project, 2011-)

located_in nucleus

inferred from sequence or structural similarity with UniProtKB:P40424

(Gomez-Skarmeta et al., 1996)

inferred by curator from GO:0000981

(Gomez-Skarmeta et al., 1996)

inferred from electronic annotation with InterPro:IPR003893

(InterPro Project Members, 2004-)

Gene Group (FlyBase)

TALE HOMEOBOX TRANSCRIPTION FACTORS

Protein Family (UniProt)

Belongs to the TALE/IRO homeobox family. (Q24248)

Protein Signatures (InterPro)

Summaries

Gene Snapshot

Gene Group (FlyBase)

TALE HOMEOBOX TRANSCRIPTION FACTORS -: TALE homeobox transcription factors are sequence-specific DNA binding proteins that regulate transcription. These proteins are characterized by an atypical homeodomain that possesses a three amino acid insertion in the first loop. (Adapted from FBrf0232555 and PMID:18797923).

Protein Function (UniProtKB)

Controls proneural and vein forming genes. Positive transcriptional controller of AC-SC (achaete-scute). May act as an activator that interacts with the transcriptional complex assembled on the AC and SC promoters and participates in transcription initiation.

(UniProt, Q24248)

Summary (Interactive Fly)

transcription factor in the Iroquois Complex- homeodomain Pbx class - a prepattern gene that determines the size, shape and number of sensory organ mother cells in imaginal discs - required in the dorsal mesoderm to ensure normal heart development

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\ara 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 Q24248)

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 5.45

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

ara-RA

FBtr0075908

NM_079321

2942

717

ara-RB

FBtr0300551

NM_001169961

2918

709

Additional Transcript Data and Comments

Reported size (kB)

3.0, 1.8, 0.6 (northern blot)

(Gomez-Skarmeta et al., 1996)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

ara-PA

FBpp0075640

75.4

717

6.39

FBpp0289778

74.7

709

6.39

E1JHZ0

NP_001163432

ACZ94703

Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments

Reported size (kDa)

716 (aa)

(Gomez-Skarmeta et al., 1996)

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

Mapped Features

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

External Data

Crossreferences

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

ara_1

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).

-0.71

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage

embryonic/larval proventriculus

(Hernández de Madrid and Casanova, 2018, Gomez-Skarmeta et al., 1996)

procephalic segment

(Gomez-Skarmeta et al., 1996)

embryonic stage 4 -- 6

dorsal ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

organism | gap expression pattern

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

trunk mesoderm 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 | gap expression pattern

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

trunk mesoderm

(Fisher et al., 2012)

embryonic stage 9 -- 10

dorsal ectoderm

(Fisher et al., 2012)

organism | gap expression pattern

(Fisher et al., 2012)

organism | faint | ubiquitous

(Fisher et al., 2012)

embryonic stage 10

cardiogenic mesoderm

(Mirzoyan and Pandur, 2013)

visceral trunk mesoderm

(Mirzoyan and Pandur, 2013)

embryonic stage 11 -- 12

dorsal epidermis primordium

(Fisher et al., 2012)

organism | faint | ubiquitous

(Fisher et al., 2012)

proventriculus primordium

(Fisher et al., 2012)

tracheal primordium

(Fisher et al., 2012)

ventral epidermis primordium

(Fisher et al., 2012)

embryonic stage 11 -- 17

embryonic epidermis | lateral

(Gomez-Skarmeta et al., 1996)

larval stage

larval ventral nerve cord

(Gomez-Skarmeta et al., 1996)

imaginal disc

(Gomez-Skarmeta et al., 1996)

second instar larval stage -- third instar larval stage

haltere disc

(Gomez-Skarmeta et al., 1996)

third instar larval stage

presumptive wing vein L1

(Gomez-Skarmeta et al., 1996)

presumptive wing vein L3

(Gomez-Skarmeta et al., 1996)

presumptive wing vein L5

(Gomez-Skarmeta et al., 1996)

presumptive tegula

(Gomez-Skarmeta et al., 1996)

alar lobe | precursor

(Gomez-Skarmeta et al., 1996)

mesothoracic pleurum | precursor

(Gomez-Skarmeta et al., 1996)

northern blot

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage -- adult stage

organism

(Gomez-Skarmeta et al., 1996)

Additional Descriptive Data

At stage 10 of embryogenesis, ara mRNA transcripts are detected in a similar pattern albeit at a lower level compared to the mesodermal expression of caup mRNA transcripts.

(Mirzoyan and Pandur, 2013)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 10

ectoderm | dorsal

(Mirzoyan and Pandur, 2013)

mesoderm | dorsal

(Mirzoyan and Pandur, 2013)

embryonic stage 11

trunk mesoderm

(Carrasco-Rando et al., 2011)

visceral trunk mesoderm

(Carrasco-Rando et al., 2011)

ectoderm | dorso-lateral

(Carrasco-Rando et al., 2011)

embryonic stage 12

somatic trunk mesoderm

(Carrasco-Rando et al., 2011)

muscle founder cell

(Carrasco-Rando et al., 2011)

abdominal dorsal transverse muscle 1 founder cell

(Carrasco-Rando et al., 2011)

embryonic stage 13 -- 15

heart primordium | restricted

(Mirzoyan and Pandur, 2013)

embryonic stage 15

muscle cell of lateral transverse muscle 1

(Carrasco-Rando et al., 2011)

muscle cell of lateral transverse muscle 2

(Carrasco-Rando et al., 2011)

muscle cell of lateral transverse muscle 3

(Carrasco-Rando et al., 2011)

muscle cell of lateral transverse muscle 4

(Carrasco-Rando et al., 2011)

dorsal transverse muscle cell

(Carrasco-Rando et al., 2011)

segment border muscle cell

(Carrasco-Rando et al., 2011)

Additional Descriptive Data

Mirzoyan and Pandur (FBrf0222881) use an antibody which detects both ara and caup proteins, which have similar expression patterns. During stage 13, single ara/caup-expressing cells form alongside the row of Mef2-positive myocardial cells, and are between the eve-positive pericardial cells by stage 15.

(Mirzoyan and Pandur, 2013)

Using an anti-caup antibody that recognizes both ara and caup, expression is detected in groups of cells in the presumptive visceral trunk mesoderm at early embryonic stage 11. By mid stage 11, they are expressed at the same dorsoventral level in the visceral mesoderm and the in the dorso-lateral ectoderm. Expression declines in the visceral mesoderm at late stage 11 and then is detected in the somatic mesoderm in promuscular clusters from which a subset of muscle progenitors expressing ara and caup will segregate in stage 12. Expression is maintained in sibling muscle founder myoblasts derived from araFBtr0091350:pb-XRcaup-expressing progenitors and in the muscles they give rise to, namely LT1-4, dorsal transverse 1, and segment border muscle. ara and caup muscle expression was compared that of several muscle identity genes.

(Carrasco-Rando et al., 2011)

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in cytoplasm

inferred from high throughput direct assay

(Sarov et al., 2016)

Expression Deduced from Reporters

Reporter: P{ara-GAL4.H}

Stage

Tissue/Position (including subcellular localization)

Reference

first instar larval stage

wing disc | proximal | restricted

(Paul et al., 2013)

Reporter: P{GawB}ara^rF209

Stage

Tissue/Position (including subcellular localization)

Reference

pupal stage P6

photoreceptor neuron | subset

(Mazzoni et al., 2008)

adult stage

photoreceptor cell R7

(Mazzoni et al., 2008)

photoreceptor cell R8

(Mazzoni et al., 2008)

Reporter: P{PZ}ara^rF209

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 13

ectoderm | lateral | restricted of embryonic thoracic segment

(Calleja et al., 2000)

ectoderm | lateral | restricted of embryonic abdominal segment | subset

Comment: not expressed in A8 or A10-11.

(Calleja et al., 2000)

second instar larval stage

wing disc | restricted

(Gomez-Skarmeta et al., 1996)

third instar larval stage

wing disc | restricted

(Ikmi et al., 2008, Gomez-Skarmeta et al., 1996)

mesonotum | restricted

(Ikmi et al., 2008)

adult stage

retina | dorsal

(Wernet et al., 2003)

adult abdominal segment | lateral

(Calleja et al., 2000)

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\ara in JBrowse

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

View exonic expression by developmental stage for Dmel\ara

View exonic expression by tissue for Dmel\ara

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

FBgn0015904

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

39439/tissue=All

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

FBgn0015904

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0015904

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

FBgn0015904

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

CG10571

Flygut - An atlas of the Drosophila adult midgut

FBgn0015904

Images

FlyExpress - Embryonic expression images (BDGP data)

Show all FlyExpress processed images for Dmel\ara

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 ( 25 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 14 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of ara

Transgenic constructs containing regulatory region of ara

Aberrations (Deficiencies and Duplications) ( 11 )

Inferred from experimentation ( 11 )

Gene disrupted in

Df(3L)iro-2

(Gomez-Skarmeta et al., 1996)

Df(3L)iro^EGP6

(Carrasco-Rando et al., 2011)

Df(3L)iro-DFM1

(Kehl et al., 1998, Gomez-Skarmeta et al., 1996)

Df(3L)iro-DFM3

(Gomez-Skarmeta et al., 1996)

In(3L)iro-DFM2

(Gomez-Skarmeta et al., 1996)

Tp(3;3)iro-1

(Gomez-Skarmeta et al., 1996)

Gene not disrupted in

Df(3L)iro^EGP5

(Carrasco-Rando et al., 2011)

Df(3L)iro^EGPΔ1

(Carrasco-Rando et al., 2011)

Df(3L)sowah^EGP1

(Maeso et al., 2012)

Gene partially disrupted in

Df(3L)iro^EGP7

(Barrios et al., 2015, Andreu et al., 2012)

Df(3L)sowah^EGP3

(Maeso et al., 2012)

Inferred from location ( 5 )

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

lethal, with Scer\GAL4^insc-Mz1407

ara^GD17184

lethal | recessive

lethal - all die before end of larval stage, with Scer\GAL4^Dll-md23

ara^UAS.cGa

partially lethal - majority die, with Scer\GAL4^pnr-MD237

ara^GD17184

partially lethal - majority die | recessive

ara^B6.8

viable

ara¹

viable, with Scer\GAL4^Mef2.PR

ara^GD17184

viable (with ara^B6.8)

ara^rF209

viable (with ara^rF209)

ara^B6.8

Sterility

Allele

female sterile | recessive

ara¹

male sterile | recessive

ara¹

Other Phenotypes

Allele

abnormal neuroanatomy, with Scer\GAL4²²¹

abnormal neuroanatomy, with Scer\GAL4^ppk.1.9

abnormal neuroanatomy, with Scer\GAL4^ppk.PG

ara^UAS.cGa

abnormal planar polarity, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

decreased occurrence of cell division | larval stage, with Scer\GAL4^nub.PU

ara^UAS.cGa

some die during larval stage, with Scer\GAL4^Dll-md23

ara^UAS.cGa

some die during pupal stage, with Scer\GAL4^pnr-MD237

ara^GD17184

visible

ara¹

visible, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

visible, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

visible, with Scer\GAL4^Bx-MS1096

visible, with Scer\GAL4^Bx-MS1096-KE

ara^FAR

visible, with Scer\GAL4^C-765

ara^UAS.cGa

visible, with Scer\GAL4^dpp.blk1

ara^UAS.cGa

visible, with Scer\GAL4^ey.PH

visible, with Scer\GAL4^GMR.long

ara^UAS.cGa

visible, with Scer\GAL4^GMR.PF

ara^FAR

visible, with Scer\GAL4^GMR.PU

ara^FAR

visible, with Scer\GAL4^kni.L2

ara^UAS.cGa

visible, with Scer\GAL4^MD-638

ara^UAS.cGa

visible, with Scer\GAL4^sca-C253

ara^UAS.cGa

visible (with ara¹)

ara^rF209

visible (with ara^B6.8)

ara^rF209

visible (with ara^rF209)

visible | cold sensitive, with Scer\GAL4^Bx-MS1096

ara^{UAS.ORF.GW.Tag:HA}

visible | recessive

ara^B6.8

Phenotype manifest in

Allele

abdominal anterior ventral multidendritic neuron vdaa, with Scer\GAL4^ppk.1.9

abdominal dorsal multidendritic neuron ddaC | third instar larval stage, with Scer\GAL4^ppk.PG

ara^UAS.cGa

abdominal dorsal multidendritic neuron ddaD | third instar larval stage, with Scer\GAL4²²¹

abdominal dorsal multidendritic neuron ddaE | third instar larval stage, with Scer\GAL4²²¹

adult cuticle, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

alar lobe

antenna, with Scer\GAL4^dpp.blk1

ara^UAS.cGa

anterior postalar bristle, with Scer\GAL4^sca-C253

ara^UAS.cGa

anterior supraalar bristle, with Scer\GAL4^C-765

ara^UAS.cGa

arista, with Scer\GAL4^dpp.blk1

ara^UAS.cGa

axillary sclerite, with Scer\GAL4^ap-md544

ara^UAS.cGa

campaniform sensillum of anterior crossvein

ara^DFM3

campaniform sensillum of dorsal radius

ara^DFM3

dendritic tree | third instar larval stage, with Scer\GAL4²²¹

dendritic tree | third instar larval stage, with Scer\GAL4^ppk.PG

ara^UAS.cGa

dorsal margin photoreceptor cell | ectopic, with Scer\GAL4^GMR.PF

ara^UAS.cGa

eye, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

eye, with Scer\GAL4^dpp.blk1

ara^UAS.cGa

eye, with Scer\GAL4^ey.PH

eye, with Scer\GAL4^GMR.long

ara^UAS.cGa

eye, with Scer\GAL4^GMR.PF

eye, with Scer\GAL4^GMR.PU

ara^FAR

eye | dorsal, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

eye | ectopic, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

eye | ventral, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

eye disc, with Scer\GAL4^ey.PH

ara^UAS.cGa

eye disc | dorsal, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

eye disc | somatic clone, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

eye disc | ventral, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

larval brain | larval stage, with Scer\GAL4^insc-Mz1407

ara^GD17184

larval dorsal multidendritic neuron ddaD, with Scer\GAL4²²¹

macrochaeta

ara¹

macrochaeta, with Scer\GAL4^C-765

ara^UAS.cGa

macrochaeta, with Scer\GAL4^sca-C253

ara^UAS.cGa

macrochaeta (with ara¹)

macrochaeta (with ara^DFM1)

ara¹

macrochaeta (with ara^DFM2)

ara¹

macrochaeta (with ara^rF209)

ara¹

macrochaeta & adult thorax

ara^B6.8

macrochaeta & scutum

ara¹

macrochaeta | ectopic, with Scer\GAL4^C-765

ara^UAS.cGa

macrochaeta | ectopic, with Scer\GAL4^sca-C253

ara^UAS.cGa

maxillary palp, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

mesonotum | ectopic, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

mesonotum | ectopic, with Scer\GAL4^MD-638

ara^UAS.cGa

mesothoracic segment

ara¹

microchaeta

ara¹

notopleural suture

ara¹

ommatidium, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

photoreceptor neuron, with Scer\GAL4^unspecified

ara^UAS.cGa

posterior scutellar bristle, with Scer\GAL4^C-765

ara^UAS.cGa

posterior scutellar bristle, with Scer\GAL4^sca-C253

ara^UAS.cGa

posterior supraalar bristle, with Scer\GAL4^C-765

ara^UAS.cGa

rostral membrane, with Scer\GAL4^Act5C.PI

ara^UAS.cGa

scutellum

ara¹

scutum & microchaeta | medial, with Scer\GAL4^sca-537.4

ara^UAS.cGa

sense organ & wing | anterior, with Scer\GAL4^C-765

ara^UAS.cGa

twin sensillum of margin 1

ara^DFM3

twin sensillum of margin 2

ara^DFM3

vibrissal bristle, with Scer\GAL4^dpp.blk1

ara^UAS.cGa

vibrissal bristle, with Scer\GAL4^ey.PH

ara^UAS.cGa

wing

wing, with Scer\GAL4^ap-md544

ara^UAS.cGa

wing, with Scer\GAL4^bi-omb-Gal4

ara^UAS.cGa

wing, with Scer\GAL4^Bx-MS1096

wing, with Scer\GAL4^Bx-MS1096-KE

ara^FAR

wing, with Scer\GAL4^C-765

ara^UAS.cGa

wing, with Scer\GAL4^MD-638

ara^UAS.cGa

wing (with ara^B6.8)

ara^rF209

wing (with ara^rF209)

ara^B6.8

wing | cold sensitive, with Scer\GAL4^Bx-MS1096

ara^{UAS.ORF.GW.Tag:HA}

wing disc | larval stage, with Scer\GAL4^nub.PU

ara^UAS.cGa

wing hinge, with Scer\GAL4^ap-md544

ara^UAS.cGa

wing vein, with Scer\GAL4^Bx-MS1096

ara^FAR

wing vein, with Scer\GAL4^C-765

ara^UAS.cGa

wing vein L1

ara^DFM3

wing vein L2, with Scer\GAL4^kni.L2

ara^UAS.cGa

wing vein L3

ara^DFM3

wing vein L5

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) (27)

Hsap\IRX4

9 of 14

Yes

Hsap\IRX6

9 of 14

Yes

Hsap\IRX1

6 of 14

Yes

Hsap\IRX3

5 of 14

Yes

Hsap\IRX2

4 of 14

Yes

Hsap\IRX5

4 of 14

Yes

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 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

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (18)

Rnor\Irx4

9 of 14

Yes

Rnor\Irx6

9 of 14

Yes

Rnor\Irx1

6 of 14

Yes

Rnor\Irx5

4 of 14

Rnor\Irx2

3 of 14

Rnor\Irx3

2 of 14

Yes

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

Mus musculus (laboratory mouse) (20)

Mmus\Irx4

8 of 14

Yes

Mmus\Irx6

8 of 14

Yes

Mmus\Irx1

6 of 14

Yes

Mmus\Irx2

5 of 14

Yes

Mmus\Irx3

5 of 14

Yes

Mmus\Irx5

5 of 14

Yes

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (15)

Xtro\irx4

7 of 13

Yes

Xtro\irx1

6 of 13

Yes

Xtro\irx3

6 of 13

Yes

Xtro\irx5

3 of 13

Xtro\irx2

2 of 13

Xtro\arl15

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

Xtro\pknox2

1 of 13

Xtro\ventx2.1

1 of 13

Danio rerio (Zebrafish) (26)

Drer\irx4b

9 of 14

Yes

Drer\irx4a

8 of 14

Yes

Drer\irx3b

7 of 14

Yes

Drer\irx6a

7 of 14

Drer\irx1a

6 of 14

Yes

Drer\irx1b

6 of 14

Yes

Drer\irx3a

6 of 14

Yes

Drer\irx7

5 of 14

Yes

Drer\irx5a

4 of 14

Yes

Drer\irx2a

3 of 14

Yes

3 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 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

Caenorhabditis elegans (Nematode, roundworm) (2)

Cele\irx-1

6 of 14

Yes

Cele\unc-62

1 of 14

Anopheles gambiae (African malaria mosquito) (9)

Agam\AgaP_AGAP005010

11 of 12

Yes

6 of 12

2 of 12

2 of 12

2 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Arabidopsis thaliana (thale-cress) (21)

Atha\KNAT2

3 of 13

Yes

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 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

Saccharomyces cerevisiae (Brewer's yeast) (5)

Scer\TOS8

2 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

Schizosaccharomyces pombe (Fission yeast) (0)

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:ara. 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) (8)

caup

11 of 13

mirr

10 of 13

CG11617

3 of 13

hth

2 of 13

achi

1 of 13

Antp

1 of 13

PHDP

1 of 13

vis

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?

IRX4; iroquois homeobox 4

9 of 14

606199

IRX6; iroquois homeobox 6

9 of 14

606196

IRX1; iroquois homeobox 1

6 of 14

606197

IRX3; iroquois homeobox 3

5 of 14

612985

IRX2; iroquois homeobox 2

4 of 14

606198

IRX5; iroquois homeobox 5

4 of 14

606195

HAMAMY SYNDROME; HMMS

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

ara - B-H1

two hybrid array

(Shokri et al., 2019)

ara - CG8478

two hybrid array

(Shokri et al., 2019)

ara - Irbp18

two hybrid array

(Shokri et al., 2019)

ara - mirr

electrophoretic mobility shift assay, autoradiography

(Bilioni et al., 2005)

ara - pdm2

two hybrid array

(Shokri et al., 2019)

ara - toy

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

ara

suppressible

CycE

(Barrios et al., 2015)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

Ada3

suppressible

ara

(Grau et al., 2008)

Delta

enhanceable

ara

(Barrios et al., 2015)

enhanceable

ara

(Singh et al., 2005)

External Data

Linkouts

BioGRID - A database of protein and genetic interactions.

64788

DroID - A comprehensive database of gene and protein interactions.

FBgn0015904

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

39439

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

Class of Gene

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

3-38

Cytogenetic map

69C8-69C10

Sequence location

3L:12,580,672..12,596,018 [+]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

69C8-69C10

Limits computationally determined from genome sequence between P{PZ}l(3)05088⁰⁵⁰⁸⁸ and P{PZ}l(3)06924⁰⁶⁹²⁴

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

69D-69D

(determined by in situ hybridisation)

(Gomez-Skarmeta et al., 1996, Leyns et al., 1996)

Distance between ara and mirr is less than 50kb.

(Kehl et al., 1998)

Experimentally Determined Recombination Data

Location

3-38

(FlyBase, 2016)

3-32.6

(Comeron et al., 2012)

Left of (cM)

mirr

(Netter et al., 1998)

Right of (cM)

Notes

Can be 1.5% recombination between ara and mirr, with none between ara and caup.

(Netter et al., 1998)

Stocks and Reagents

Stocks (51)

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

Bloomington

99028

w^*; P{UAS-ara.M}3

FlyORF

F003436

M{UAS-ara.ORF-CC}ZH-21F

F004806

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

Korea

12677

P{VT029749-GAL4}attP2

12678

P{VT029753-GAL4}attP2

13280

P{VT029750-GAL4}attP2

13281

P{VT029752-GAL4}attP2

13475

P{VT029741-GAL4}attP2

15156

P{VT029754-GAL4}attP2

10571R-2

10571R-3

v101903

More stocks available...

Genomic Clones (33)

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

cDNA Clones (24)

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

Other clones

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.

FBgn0015904

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0015904

Antibody Information

Laboratory Generated Antibodies

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

The acquisition of lateral transverse muscle fate requires the repression mediated by ara/caup of other muscle identity genes (such as slou and vg).

(Carrasco-Rando et al., 2011)

RNAi generated by PCR using primers directed to this gene causes a cell growth and viability phenotype when assayed in Kc167 and S2R⁺ cells.

(Boutros et al., 2004)

Iroquois complex genes (ara, caup and mirr) are required for the specification of dorsal head structures.

(Cavodeassi et al., 2000)

The localised expression of the three genes of the iroquois complex - IRO-C (ara, caup and mirr) specifies the identity of dorsal cells in the eye. Juxtaposition of IRO-C-expressing and non-expressing cells forms a straight border that promotes growth and serves as a pattern-organising centre in the eye disc.

(Cavodeassi et al., 1999)

The genes of the Iroquois complex (ara, caup and mirr) are necessary for notum specification. The Iro-C genes establish a signalling system that appears to organize development in the notum and dorsal wing hinge (tegula and sclerite) territories.

(Diez del Corral et al., 1999)

When the Iroquois complex is absent, notum cells are transformed into hinge (tegula and sclerite) cells.

(Diez del Corral et al., 1999)

A member of the IRO-C (Iroquois complex).

(Netter et al., 1998)

The genes of the IRO-C are coordinately regulated and implicated in similar developmental processes. They are regulated in the eye by the Pc-group and trx-group genes, but not by classical modifiers of position effect variegation. Ventral silencing of the whole IRO-C in the eye occurs at the level of chromatin structure in a manner similar to that of the homeotic gene complexes, perhaps by local compaction of the region into a heterochromatin-like structure involving Pc-group products.

(Netter et al., 1998)

The ara gene maps to the Iroquois complex and contributes to the prepattern that regulates expression of ac-sc complex genes and rho. ara is a positive, direct regulator of the ac-sc genes.

(Gomez-Skarmeta et al., 1996)

Gene titration method has been used to search for genes involved in the determination of sense organs. A mutation of ara and caup (formerly iro) demonstrates an interaction with Df(4)M101-62f, a chromosome known to alter the development of the PNS.

(Dambly-Chaudiere and Leyns, 1992)

In mutants for the Iroquois gene complex, two lateral bands of the notum are completely devoid of sense organs.

(Leyns et al., 1991)

Relationship to Other Genes

Source for database merge of

Additional comments

Nomenclature History

Source for database identify of

Source for identity of: ara CG10571

(Bayraktaroglu, 1999.11)

Nomenclature comments

Etymology

The "ara" gene is named after the Araucanian American-Indian tribe.

(Gomez-Skarmeta et al., 1996)

The 'Iroquois' gene complex is named after the native American Iroquois tribe, also called Mohawks, which shaved all but a medial stripe of hairs on the head.

(Leyns et al., 1996)

"Iroquois" is the name given to the gene complex by analogy with the American tribe that have a haircut ascribed to the Mohawk nation.

(Dambly-Chaudiere and Leyns, 1992)

Synonyms and Secondary IDs (18)

Reported As

Symbol Synonym

Ara

(Poovathumkadavil and Jagla, 2020, Sarov et al., 2016, Barrios and Campuzano, 2015, Kumar et al., 2015, Sorge et al., 2012, Luque and Milan, 2007)

CG10571

(Fukuoh et al., 2014, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009.3.31)

IRO-C

(Villa-Cuesta and Modolell, 2004, Wernet and Desplan, 2004, Cavodeassi et al., 1999, Cavodeassi et al., 1999)

Iro

(Benítez et al., 2009, Sato and Tomlinson, 2007, Callejo et al., 2006, Hooper, 2003, Tomlinson, 2003)

Iro-C

(Chen and Desplan, 2020, Singari et al., 2014, Curtis et al., 2013, Cruz et al., 2009, Navascues and Modolell, 2007, Villa-Cuesta et al., 2007, Villa-Cuesta and Modolell, 2005, Singh and Choi, 2004, Baena-Lopez et al., 2003, D'Costa and Moses, 2003, Arnosti, 2002, Cavodeassi et al., 2002, Cavodeassi et al., 2000, de Celis and Barrio, 1999, Irvine, 1999)

IroC

(Miller et al., 2021, Viets et al., 2016, Wernet et al., 2015, Thanawala et al., 2013)

ara

(Walsh et al., 2025, Collins et al., 2024, Hursh, 2023.6.13, Corrales et al., 2022, Deshpande et al., 2022, Junion and Jagla, 2022, Saha et al., 2022, Hagen et al., 2021, Keder et al., 2020, Seong et al., 2020, Seong et al., 2019, Shokri et al., 2019, Ahmed-de-Prado et al., 2018, Bischof et al., 2018, Gene Disruption Project members, 2018-, Karaiskos et al., 2017, Transgenic RNAi Project members, 2017-, Bürglin and Affolter, 2016, Sarov et al., 2016, Wang et al., 2016, Barrios et al., 2015, Javeed et al., 2015, Schertel et al., 2015, Dobi et al., 2014, Curtis et al., 2013, Hattori et al., 2013, Iyer et al., 2013, Paul et al., 2013, Andreu et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Maeso et al., 2012, Zhou et al., 2012, Carrasco-Rando et al., 2011, Chatterjee et al., 2011, de Navas et al., 2011, King et al., 2011, Neumüller et al., 2011, Kazemian et al., 2010, Oros et al., 2010, Raisin et al., 2010, Grieder et al., 2009, Kerner et al., 2009, Benchabane et al., 2008, Grau et al., 2008, Mazzoni et al., 2008, Bejarano et al., 2007, Beltran et al., 2007, de Navascués and Modolell, 2007, González-Pérez et al., 2007, IKMI and COEN, 2007, Kankel et al., 2007, Letizia et al., 2007, Mukherjee and Bürglin, 2007, Villa-Cuesta et al., 2007, Choksi et al., 2006, Sato et al., 2006, Simpson et al., 2006, Sotillos and de Celis, 2006, Bilioni et al., 2005, Stanyon et al., 2004)

iro-C

(Paul et al., 2013, Rudel and Sommer, 2003, Calleja et al., 2002, Skaer et al., 2002, Klein, 2001, de Celis and Barrio, 2000, Ghysen and Dambly-Chaudiere, 2000)

Name Synonyms

Aracucan

(Ishihara and Shibata, 2008)

Araucan

(Terriente-Félix et al., 2011, Bejarano et al., 2007, Luque and Milan, 2007, Minervini et al., 2007)

Auracan

(Sarov et al., 2016)

Iroquis complex

(Brennan and Gardner, 2002)

Iroquois

(Sato and Tomlinson, 2007, Bilioni et al., 2004, de Celis and Sotillos, 2001, Zaffran-Maurel et al., 1999, Bellefroid et al., 1998)

iroquois gene complex

(de Celis, 2003)

iroquois-complex

(Pichaud and Casares, 2000)

Secondary FlyBase IDs

FBgn0014195

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 45 )

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.

39439

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.

Q24248

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

Q24248

UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

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.

Q24248

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0015904

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

39439/tissue=All

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

FBgn0015904

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

FBgn0015904

FlyMine - An integrated database for Drosophila genomics

FBgn0015904

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

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

dme:Dmel_CG10571

MARRVEL_MODEL - MARRVEL (model organism gene)

39439

Linkouts

BioGRID - A database of protein and genetic interactions.

64788

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

FBgn0015904

DroID - A comprehensive database of gene and protein interactions.

FBgn0015904

DRSC - Results frm RNAi screens

FBgn0015904

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

ara_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0015904

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0015904

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

CG10571

Flygut - An atlas of the Drosophila adult midgut

FBgn0015904

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC031040

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

Interactive Fly

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

39439

References (243)

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

-0.71

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