FB2020_01, released Feb 12, 2020

Gene: Dmel\exd

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General Information
Symbol
Dmel\exd
Species
D. melanogaster
Name
extradenticle
Annotation Symbol
CG8933
Feature Type
protein_coding_gene
FlyBase ID
FBgn0000611
Gene Model Status
Current
Stock Availability
13 publicly available
Gene Snapshot
extradenticle (exd) encodes a homeodomain transcription factor that is imported to the nuclueus upon binding the product of hth. The protein complex hth-exd are Hox cofactors but also have additional Hox-independent functions such as in antennal development, proximal appendages patterning, and cell division promotion of the undifferentiated eye field. [Date last reviewed: 2019-03-21]
Other Summaries
Also Known As

Dpbx, lincRNA.S9404

Key Links
Genomic Location
Cytogenetic map
14A5-14A5
Sequence location
X:15,992,468..15,996,509 [+]
Recombination map

1-52

RefSeq locus
NC_004354 REGION:15992468..15996509
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
GO Summary Ribbons
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the TALE/PBX homeobox family. (P40427)
Protein Signatures (InterPro)
Molecular Function (GO)
[Detailed GO annotations]
Experimental Evidence
activating transcription factor binding; DNA binding; DNA-binding transcription factor activity; contributes_tocontributes_to DNA-binding transcription factor activity, RNA polymerase II-specific; enhancer binding; protein binding; protein heterodimerization activity; proximal promoter sequence-specific DNA binding; sequence-specific DNA binding; transcription coactivator activity; transcription factor binding
Predictions / Assertions
-
Summaries
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)
Transcription factor which acts with the selector homeodomain proteins altering the regulation of downstream target genes such as wingless (wg), teashirt (tsh) and decapentaplegic (dpp), thus affecting segmental identity. Delimits the eye field and prevent inappropriate eye development. Required for proper localization of chordotonal organs within the peripheral nervous system.
(UniProt, P40427)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
exd: extradenticle
Homozygous lethal; meso- and metathoracic segments resemble prothorax; first abdominal like posterior abdominal segments. In combination with Pc-like mutants shows abdominal transformations.
Summary (Interactive Fly)

TALE-class homeodomain transcription factor - cooperatively interacts with homeotic proteins to increase the specificity of homeotic protein binding to DNA - Engrailed cooperates directly with Extradenticle and Homothorax on a distinct class of homeodomain binding sites to repress - controls dendritic and axonal targeting of olfactory projection neurons in the brain - control adult muscle fiber identity

Gene Model and Products
Number of Transcripts
4
Number of Unique Polypeptides
1

Please see the GBrowse view of Dmel\exd or the JBrowse view of Dmel\exd 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
Comments on Gene Model

Gene model reviewed during 5.50

Low-frequency RNA-Seq exon junction(s) not annotated.

Annotated transcripts do not represent all supported alternative splices within 5' UTR.

Gene model reviewed during 5.46

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated.

Gene model reviewed during 5.52

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
exd-RA
FBtr0074135
NM_078636
2386
376
exd-RB
FBtr0074137
NM_167478
2347
376
exd-RD
FBtr0333322
NM_001272663
2930
376
exd-RE
FBtr0333323
NM_001272664
2883
376
Additional Transcript Data and Comments
Reported size (kB)

3.4, 2.8 (northern blot)

(Rauskolb et al., 1993)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
exd-PA
FBpp0073940
41.7
376
7.00
NP_523360
AAF48555
exd-PB
FBpp0073942
41.7
376
7.00
NP_727923
AAF48556
exd-PD
FBpp0305514
41.7
376
7.00
NP_001259592
AGB95434
exd-PE
FBpp0305515
41.7
376
7.00
NP_001259593
AGB95435
Polypeptides with Identical Sequences

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

376 aa isoforms: exd-PA, exd-PB, exd-PD, exd-PE
Additional Polypeptide Data and Comments
Reported size (kDa)

376 (aa); 41.7 (kD)

(Rauskolb et al., 1993)
Comments
External Data
Subunit Structure (UniProtKB)

Interacts with Ubx and hth.

(UniProt, P40427)
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\exd using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Gene Ontology (24 terms)
Molecular Function (11 terms)
Terms Based on Experimental Evidence (11 terms)
CV Term
Evidence
References
activating transcription factor binding
inferred from physical interaction with FLYBASE:Dfd; FB:FBgn0000439
(Li et al., 1999)
DNA binding
inferred from direct assay
(assigned by UniProt )
(Passner et al., 1999)
inferred from direct assay
(Stultz et al., 2006)
DNA-binding transcription factor activity
inferred from mutant phenotype
(assigned by UniProt )
(Rauskolb et al., 1993)
DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Li-Kroeger et al., 2008, Li et al., 1999)
contributes_to DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from physical interaction with FLYBASE:hth; FB:FBgn0001235
(Joshi et al., 2010)
inferred from physical interaction with FLYBASE:lab; FB:FBgn0002522
(Grieder et al., 1997)
enhancer binding
inferred from direct assay
(Stultz et al., 2012)
protein binding
inferred from physical interaction with UniProtKB:O46339
(assigned by UniProt )
(Kurant et al., 1998, Pai et al., 1998)
inferred from physical interaction with FLYBASE:hth; FB:FBgn0001235
(Rieckhof et al., 1997)
inferred from physical interaction with UniProtKB:P83949
(assigned by UniProt )
(Passner et al., 1999)
protein heterodimerization activity
inferred from physical interaction with FLYBASE:lab; FB:FBgn0002522
(Grieder et al., 1997)
proximal promoter sequence-specific DNA binding
inferred from direct assay
(Wang et al., 2017)
sequence-specific DNA binding
inferred from direct assay
(Ahn et al., 2011)
transcription coactivator activity
inferred from direct assay
(Li et al., 1999)
transcription factor binding
inferred from physical interaction with FLYBASE:hth; FB:FBgn0001235
(Joshi et al., 2010)
Terms Based on Predictions or Assertions (0 terms)
Biological Process (9 terms)
Terms Based on Experimental Evidence (8 terms)
CV Term
Evidence
References
brain development
inferred from mutant phenotype
(Nagao et al., 2000)
eye development
inferred from mutant phenotype
(assigned by UniProt )
(Pai et al., 1998)
oenocyte development
inferred from mutant phenotype
(Brodu et al., 2002)
peripheral nervous system development
inferred from mutant phenotype
(assigned by UniProt )
(Kurant et al., 1998)
positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Wang et al., 2017, Joshi et al., 2010)
inferred from mutant phenotype
(Grieder et al., 1997)
regulation of cell fate specification
inferred from mutant phenotype
(Bryantsev et al., 2012)
regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(assigned by UniProt )
(Rauskolb et al., 1993)
somatic muscle development
inferred from mutant phenotype
(Bryantsev et al., 2012)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
salivary gland boundary specification
traceable author statement
(Abrams et al., 2003, Bradley et al., 2001)
non-traceable author statement
(Nelson, 2003)
Cellular Component (4 terms)
Terms Based on Experimental Evidence (4 terms)
CV Term
Evidence
References
cytoplasm
inferred from direct assay
(Abu-Shaar and Mann, 1998, Rieckhof et al., 1997)
nucleus
inferred from direct assay
(Abu-Shaar and Mann, 1998, Azpiazu and Morata, 1998, Rieckhof et al., 1997)
RNA polymerase II transcription factor complex
inferred from physical interaction with FLYBASE:abd-A; FB:FBgn0000014, FLYBASE:hth; FB:FBgn0001235
(Wang et al., 2017)
transcription factor complex
inferred from physical interaction with FLYBASE:abd-A; FB:FBgn0000014
(Li-Kroeger et al., 2008)
inferred from physical interaction with FLYBASE:hth; FB:FBgn0001235
(Li-Kroeger et al., 2008)
inferred from direct assay
(assigned by UniProt )
(Rauskolb et al., 1993)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
cytoplasm
traceable author statement
(Azpiazu and Morata, 1998)
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
presumptive embryonic/larval central nervous system
(Rauskolb et al., 1993)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
visceral mesoderm | gradient
(Rauskolb et al., 1993)
embryonic stage 12
abdomen
(Rauskolb et al., 1993)
embryonic stage 12 -- 17
organism | gradient
(Rauskolb et al., 1993)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

(Fisher et al., 2012)
embryonic stage 4 -- 6
anlage in statu nascendi
(Fisher et al., 2012)
dorsal ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
foregut anlage in statu nascendi
(Fisher et al., 2012)
hindgut anlage in statu nascendi
(Fisher et al., 2012)
mesectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
mesoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
ventral ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
visual anlage in statu nascendi
(Fisher et al., 2012)
embryonic stage 7 -- 8
antennal anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
central brain anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
visual anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
anterior endoderm anlage
(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
foregut anlage
(Fisher et al., 2012)
head mesoderm
(Fisher et al., 2012)
hindgut anlage
(Fisher et al., 2012)
mesectoderm
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
posterior endoderm
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium

Comment: reported as ventral nerve cord anlage

(Fisher et al., 2012)
embryonic stage 9 -- 10
antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
visual primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)
anterior endoderm
(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
head mesoderm
(Fisher et al., 2012)
inclusive hindgut primordium
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
posterior endoderm
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
dorsal epidermis primordium
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic brain
(Fisher et al., 2012)
embryonic central brain neuron
(Fisher et al., 2012)
embryonic hindgut
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
embryonic/larval muscle system
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
embryonic/larval somatic muscle
(Fisher et al., 2012)
embryonic/larval visceral muscle
(Fisher et al., 2012)
lateral cord neuron
(Fisher et al., 2012)
organism | ubiquitous
(Fisher et al., 2012)
ventral nerve cord
(Fisher et al., 2012)
Additional Descriptive Data

exd is expressed at all stages of development. exd transcripts are present at high levels in early embryos except in the pole cells. Levels decline at gastrulation and then rise during germ band extension. From stage 12 on, transcript levels are modulated with high levels in head and thoracic regions and declining levels in abdominal regions. Levels are moderate in abdominal segments 1 and 2, lower in segments 3 and 4, and lowest in segments 5 and 6. Abdominal segments 7-9 have levels intermeditiate between those of segments 4 and 5. Expression is also observed in the CNS, visceral mesoderm, and somatic mesoderm.

(Rauskolb et al., 1993)

exd transcript levels are greatly reduced in the exd1 mutant.

(Rauskolb et al., 1993)

exd transcript levels are greatly reduced in the exd2 mutant.

(Rauskolb et al., 1993)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 15 -- 16
chordotonal organ
• nucleus
(Kurant et al., 2001)
presumptive embryonic/larval central nervous system
• nucleus
(Kurant et al., 2001)
ectoderm
• nucleus
(Kurant et al., 2001)
embryonic stage 10
parasegment 2 | ventral
• nucleus
(Ryoo and Mann, 1999)
third instar larval stage
embryonic/larval motor neuron | subset of larval thoracic neuromere
(Baek et al., 2013)
tracheal metamere 1
(Sato et al., 2008)
tracheal metamere 2
(Sato et al., 2008)
tracheal metamere 3
(Sato et al., 2008)
pupal stage
motor neuron | subset of thoracic neuromere
(Baek et al., 2013)
embryonic stage 11 -- 17
neuroblast NB5-6 of thorax
(Karlsson et al., 2010)
neuroblast NB5-6 of abdomen
(Karlsson et al., 2010)
embryonic stage 15
embryonic/larval central nervous system | variable
(Huang et al., 2003)
embryonic stage 12
bridge cell
(Merabet et al., 2005)
embryonic stage 13 -- 16
embryonic/larval Malpighian tubule | distal | restricted
(Zohar-Stoopel et al., 2014)
Additional Descriptive Data

exd is expressed in NB5-6 starting at embryonic stage 11 in abdominal and thoracic lineages, is expressed in all cells in the lineage at stage 13, and is maintained throughout the lineage in subsequent stages. It is also expressed in more anterior NB5-6 lineages.

(Karlsson et al., 2010)

Cells in the embryonic CNS anterior to the border of metathoracic segment (T3) and abdominal segment 1 express high levels of exd, largely in nuclei, wheras cells to the posterior of that boundary express lower levels of exd in the cytoplasm.

(Huang et al., 2003)
Marker for
 
Subcellular Localization
CV Term
Evidence
References
cytoplasm
inferred from direct assay
(Abu-Shaar and Mann, 1998, Rieckhof et al., 1997)
nucleus
inferred from direct assay
(Abu-Shaar and Mann, 1998, Azpiazu and Morata, 1998, Rieckhof et al., 1997)
RNA polymerase II transcription factor complex
inferred from physical interaction with FLYBASE:abd-A; FB:FBgn0000014, FLYBASE:hth; FB:FBgn0001235
(Wang et al., 2017)
transcription factor complex
inferred from physical interaction with FLYBASE:abd-A; FB:FBgn0000014
(Li-Kroeger et al., 2008)
inferred from physical interaction with FLYBASE:hth; FB:FBgn0001235
(Li-Kroeger et al., 2008)
inferred from direct assay
(assigned by UniProt )
(Rauskolb et al., 1993)
Expression Deduced from Reporters
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\exd in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • 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, and Transgenic Constructs
Classical and Insertion Alleles ( 11 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 54 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of exd
Transgenic constructs containing regulatory region of exd
Deletions and Duplications ( 10 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal | pupal stage, with Scer\GAL4pnr-MD237
lethal | recessive
lethal - all die before end of embryonic stage
lethal - all die before end of embryonic stage | recessive
lethal - all die before end of larval stage | recessive
lethal - all die before end of P-stage | recessive
lethal - all die during larval stage, with Scer\GAL4Mef2.PR
partially lethal, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
partially lethal, with Scer\GAL4tin.CΔ4
partially lethal - majority die | pupal stage, with Scer\GAL4pnr-MD237
Other Phenotypes
Allele
neuroanatomy defective
pair rule phenotype | embryonic stage
pair rule phenotype | ventral
short lived | heat sensitive, with Scer\GAL4tin.CΔ4
size defective, with Scer\GAL41151
some die during embryonic stage
some die during embryonic stage | recessive
some die during larval stage | recessive
some die during pharate adult stage | recessive
visible
visible, with Scer\GAL4Dll-md23
visible, with Scer\GAL4dpp.blk1
visible, with Scer\GAL4pnr-MD237
visible, with Scer\GAL4upd1-E132
visible | recessive | somatic clone
visible | somatic clone
wild-type
Phenotype manifest in
Allele
abdominal 1 lch5 neuron | embryonic stage
abdominal 2 lch5 neuron | embryonic stage
abdominal 3 lch5 neuron | embryonic stage
abdominal 4 lch5 neuron | embryonic stage
abdominal 5 lch5 neuron | embryonic stage
abdominal 6 lch5 neuron | embryonic stage
abdominal 7 lch5 neuron | embryonic stage
abdominal segment 1
abdominal sternite 2
abdominal sternite 3
abdominal sternite 4
abdominal sternite 5
abdominal tergite 1
abdominal tergite 2
abdominal tergite 3
abdominal tergite 4
adult abdominal segment 1
adult abdominal segment 1 | somatic clone
adult abdominal segment 2 | ectopic
adult abdominal segment 2 | somatic clone
adult abdominal segment 3 | somatic clone
adult abdominal segment 4 | somatic clone
adult abdominal segment 6
adult abdominal segment 6 | somatic clone
adult abdominal segment 7
adult abdominal segment 7 | somatic clone
adult abdominal segment 8
adult abdominal segment 9
adult head
adult head | somatic clone
adult heart, with Scer\GAL4HCH.Hand
adult optic lobe | precursor
antenna
arista
arista | somatic clone
Bolwig nerve
Bolwig organ
cervical connective
cuticle | pharate adult stage
dental sclerite
deutero/tritocerebral embryonic fiber tract founder cluster
embryonic/larval Malpighian tubule | embryonic stage
embryonic/larval oenocyte
embryonic/larval salivary gland | embryonic stage
embryonic abdominal segment 1
embryonic abdominal segment 1 | ventral
embryonic abdominal segment 2
embryonic abdominal segment 2 | ventral
embryonic abdominal segment 3 | ventral
embryonic abdominal segment 4 | ventral
embryonic abdominal segment 5 | ventral
embryonic abdominal segment | ventral
embryonic gnathal segment | ventral
embryonic head
embryonic mesothoracic segment | dorsal | posterior
embryonic mesothoracic segment | ventral
embryonic metathoracic segment
embryonic metathoracic segment | dorsal | posterior
embryonic metathoracic segment | ventral
embryonic stomatogastric nervous system
embryonic thorax | ventral
epipharyngeal sclerite
extracoxal depressor muscle | ectopic, with Scer\GAL41151
eye | ectopic
female genitalia
female genitalia | somatic clone
femur
femur, with Scer\GAL4Dll-md23
femur, with Scer\GAL4dpp.blk1
femur | distal, with Scer\GAL4dpp.blk1
femur | distal, with Scer\GAL4upd1-E132
flight muscle, with Scer\GAL41151
head | dorsal | posterior
head | ventral
head segment
hypostomal sclerite
lateral process
leg
leg, with Scer\GAL4Dll-md23
leg | distal | ectopic
leg | ectopic
leg | ectopic | medial
leg | ectopic | proximal
leg | ectopic | somatic clone
leg | proximal
leg | proximal | somatic clone
leg | somatic clone
leg sensillum | ectopic
leg sensillum | ectopic | somatic clone
macrochaeta & scutum
male genitalia
male genitalia | somatic clone
mandibular segment
median tooth
mesothoracic segment
mesothoracic tergum, with Scer\GAL4pnr-MD237
mesothoracic tergum & macrochaeta | ectopic | somatic clone
mesothoracic ventral denticle belt
metathoracic laterotergite
metathoracic segment
metathoracic ventral denticle belt
neuroblast NB6-4 of abdomen
neuroblast NB6-4 of thorax | ectopic
presumptive embryonic/larval peripheral nervous system
proboscis
proboscis | somatic clone
prothoracic dorsal trichome
prothoracic ventral denticle belt
sarcomere, with Scer\GAL4Mef2.PR
scutellum & chaeta | somatic clone
scutum | ectopic
subesophageal embryonic fiber tract founder cluster
supraesophageal embryonic fiber tract founder cluster
tarsal segment, with Scer\GAL4Dll-md23
tarsal segment | distal, with Scer\GAL4dpp.blk1
third segment of antenna
third segment of antenna | somatic clone
tibia, with Scer\GAL4dpp.blk1
tibia, with Scer\GAL4upd1-E132
tibia | distal, with Scer\GAL4dpp.blk1
tritocerebral commissure
trochanter
unguis, with Scer\GAL4Dll-md23
unguis | distal, with Scer\GAL4dpp.blk1
ventral thoracic disc, with Scer\GAL4Dll-md23
ventral thoracic disc, with Scer\GAL4upd1-E132
Z disc, with Scer\GAL4Mef2.PR
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\PBX2
13 of 15
Yes
Yes
Hsap\PBX1
12 of 15
No
Yes
 
Hsap\PBX3
12 of 15
No
Yes
Hsap\PBX4
8 of 15
No
Yes
 
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (4)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Pbx1
13 of 15
Yes
Yes
Mmus\Pbx2
13 of 15
Yes
Yes
Mmus\Pbx3
12 of 15
No
Yes
Mmus\Pbx4
7 of 15
No
Yes
Rattus norvegicus (Norway rat) (4)
Rnor\Pbx3
11 of 13
Yes
Yes
Rnor\Pbx2
10 of 13
No
Yes
Rnor\Pbx4
8 of 13
No
Yes
Rnor\Pbx1
4 of 13
No
Yes
Xenopus tropicalis (Western clawed frog) (3)
Xtro\pbx1
10 of 12
Yes
Yes
Xtro\pbx2
10 of 12
Yes
Yes
Xtro\pbx3
4 of 12
No
Yes
Danio rerio (Zebrafish) (7)
Drer\pbx1b
13 of 15
Yes
Yes
Drer\pbx4
13 of 15
Yes
Yes
Drer\pbx1a
8 of 15
No
Yes
Drer\pbx3b
8 of 15
No
Yes
Drer\pbx2
6 of 15
No
Yes
Drer\pbx3a
5 of 15
No
Yes
Drer\LOC100537845
3 of 15
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (4)
Cele\F31E3.2
7 of 15
Yes
Yes
Cele\ceh-20
6 of 15
No
Yes
Cele\ceh-40
5 of 15
No
Yes
Cele\ceh-60
2 of 15
No
Yes
Arabidopsis thaliana (thale-cress) (21)
Atha\BLH5
4 of 9
Yes
Yes
Atha\ATH1
3 of 9
No
Yes
Atha\BLH7
3 of 9
No
Yes
Atha\BEL10
2 of 9
No
No
Atha\BLH3
2 of 9
No
No
Atha\BLH11
2 of 9
No
No
Atha\KNAT3
2 of 9
No
Yes
Atha\RPL
2 of 9
No
No
Atha\BEL1
1 of 9
No
No
Atha\BLH1
1 of 9
No
No
Atha\BLH2
1 of 9
No
No
Atha\BLH4
1 of 9
No
No
Atha\BLH6
1 of 9
No
No
Atha\BLH8
1 of 9
No
No
Atha\KNAT1
1 of 9
No
No
Atha\KNAT2
1 of 9
No
No
Atha\KNAT4
1 of 9
No
No
Atha\KNAT5
1 of 9
No
No
Atha\KNAT6
1 of 9
No
No
Atha\KNAT7
1 of 9
No
No
Atha\STM
1 of 9
No
No
Saccharomyces cerevisiae (Brewer's yeast) (5)
Scer\CUP9
1 of 15
Yes
No
Scer\HMLALPHA2
1 of 15
Yes
No
Scer\HMRA2
1 of 15
Yes
No
Scer\MATALPHA2
1 of 15
Yes
No
Scer\TOS8
1 of 15
Yes
No
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190BIB )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Dmel\exd
Drosophila simulans
Dsim\GD17261
Drosophila sechellia
Dsec\GM11678
Drosophila erecta
Dere\GG17938
Drosophila yakuba
Dyak\GE17244
Drosophila ananassae
Dana\GF20172
Drosophila pseudoobscura pseudoobscura
Dpse\GA21419
Drosophila persimilis
Dper\GL10055
Drosophila willistoni
Dwil\GK20113
Drosophila virilis
Dvir\GJ15726
Drosophila virilis
Dvir\GJ15917
Drosophila mojavensis
Dmoj\GI16078
Drosophila grimshawi
Dgri\GH17639
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG0915076R )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17012775
Musca domestica
House fly
Mdoa\17008026
Musca domestica
House fly
Mdoa\17008027
Glossina morsitans
Tsetse fly
Gmor\GMOY011586
Glossina morsitans
Tsetse fly
Gmor\GMOY000609
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_14432
Mayetiola destructor
Hessian fly
Mdes\Mdes010135
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL012877
Anopheles darlingi
American malaria mosquito
Adar\ADAC002586
Anopheles gambiae
Malaria mosquito
Agam\exd
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ008039
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0MI0 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA007946
Danaus plexippus
Monarch butterfly
Dple\DPOGS208840-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL015430
Apis florea
Little honeybee
Aflo\2580670532
Apis florea
Little honeybee
Aflo\4598012980
Apis mellifera
Western honey bee
Amel\GB51904
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP25869
Bombus terrestris
Buff-tailed bumblebee
Bter\15215485
Linepithema humile
Argentine ant
Lhum\LH15134
Linepithema humile
Argentine ant
Lhum\LH15785
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:00039c
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG008684
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG003648
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_01649
Tribolium castaneum
Red flour beetle
Tcas\TC011311
Pediculus humanus
Human body louse
Phum\PHUM213170
Rhodnius prolixus
Kissing bug
Rpro\RPRC004192
Cimex lectularius
Bed bug
Clec\CLEC008620
Cimex lectularius
Bed bug
Clec\CLEC008621
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI009075
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_12397
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0MTY )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR009718
Strigamia maritima
European centipede
Smar\SMAR009720
Ixodes scapularis
Black-legged tick
Isca\ISCW023222
Ixodes scapularis
Black-legged tick
Isca\ISCW023208
Stegodyphus mimosarum
African social velvet spider
Smim\KFM73839.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM70119.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM70118.1
Stegodyphus mimosarum
African social velvet spider
Smim\KFM61484.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur07g02140
Tetranychus urticae
Two-spotted spider mite
Turt\tetur21g02040
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_347218
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G10TD )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Pbx_1
Strongylocentrotus purpuratus
Purple sea urchin
Spur\SPU_025503gn
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-Pbx
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000009075
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000013419
Gallus gallus
Domestic chicken
Ggal\PBX3
Gallus gallus
Domestic chicken
Ggal\PBX1
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (3)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
achi
1 of 10
hth
1 of 10
vis
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    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 v7.1 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?
    PBX2; PBX homeobox 2
    13 of 15
    176311
        PBX1; PBX homeobox 1
        12 of 15
        176310
           
          PBX3; PBX homeobox 3
          12 of 15
          176312
              PBX4; PBX homeobox 4
              8 of 15
              608127
                   
                  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
                  esyN Network Diagram
                  Show neighbor-neighbor interactions:
                  Select Layout:
                  Legend:
                  Protein
                  RNA
                  Selected Interactor(s)
                  Interactions Browser
                  View in Interactions Browser
                  Please see the Physical Interaction reports below for full details
                  protein-protein
                  Physical Interaction
                  Assay
                  References
                  exd - Abd-B
                  bimolecular fluorescence complementation, fluorescence microscopy, pull down, autoradiography, electrophoretic mobility shift assay
                  (Romero-Pozuelo et al., 2019, Rivas et al., 2013, Hudry et al., 2012)
                  exd - Antp
                  bimolecular fluorescence complementation, fluorescence microscopy, electrophoretic mobility shift assay, autoradiography
                  (Altamirano-Torres et al., 2018, Hudry et al., 2012, Ahn et al., 2011, Ryoo and Mann, 1999)
                  exd - CG32425
                  anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
                  (Rhee et al., 2014)
                  exd - CG34163
                  anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
                  (Rhee et al., 2014)
                  exd - CG9776
                  two hybrid array
                  (Shokri et al., 2019)
                  exd - CtBP
                  bimolecular fluorescence complementation, fluorescence microscopy, two hybrid array
                  (Shokri et al., 2019, Bischof et al., 2018)
                  exd - Dfd
                  nuclear magnetic resonance, electrophoretic mobility shift assay, autoradiography
                  (Maiti et al., 2019, Sánchez-Higueras et al., 2019, Joshi et al., 2010, Li et al., 1999, Grieder et al., 1997)
                  exd - Dll
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - Doc2
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - Dref
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - Poxm
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - Scr
                  electrophoretic mobility shift assay, autoradiography, x-ray crystallography, predetermined participant, bimolecular fluorescence complementation, fluorescence microscopy, anti tag coimmunoprecipitation, anti tag western blot, nuclear magnetic resonance
                  (Maiti et al., 2019, Hudry et al., 2012, Papadopoulos et al., 2012, Ahn et al., 2011, Joshi et al., 2010, Joshi et al., 2007, Ryoo and Mann, 1999)
                  exd - Su(H)
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - Tsen34
                  anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
                  (Rhee et al., 2014)
                  exd - Ubx
                  two hybrid, electrophoretic mobility shift assay, autoradiography, bimolecular fluorescence complementation, fluorescence microscopy, x-ray crystallography, predetermined participant
                  (Sánchez-Higueras et al., 2019, Foos et al., 2015, Bryantsev et al., 2012, Hudry et al., 2012, Ahn et al., 2011, Reed et al., 2010, Passner et al., 1999, Ryoo and Mann, 1999, Johnson et al., 1995, Chan et al., 1994)
                  exd - YL-1
                  two hybrid array
                  (Shokri et al., 2019)
                  exd - abd-A
                  electrophoretic mobility shift assay, autoradiography, bimolecular fluorescence complementation, fluorescence microscopy, two hybrid
                  (Duffraisse et al., 2014, Hudry et al., 2012, Hudry et al., 2011, Ryoo and Mann, 1999, Johnson et al., 1995)
                  exd - bap
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - bin
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - crm
                  two hybrid array
                  (Shokri et al., 2019)
                  exd - dsx
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Romero-Pozuelo et al., 2019)
                  exd - eve
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - ey
                  pull down, autoradiography
                  (Bessa et al., 2002)
                  exd - hth
                  electrophoretic mobility shift assay, autoradiography, bimolecular fluorescence complementation, fluorescence microscopy, two hybrid array, pull down, western blot, interologs mapping, inferred by curator, anti bait coimmunoprecipitation, anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
                  (Romero-Pozuelo et al., 2019, Shokri et al., 2019, Rhee et al., 2014, Bischof et al., 2013, Bryantsev et al., 2012, Hudry et al., 2012, Stultz et al., 2012, Hudry et al., 2011, Bessa et al., 2002, Abu-Shaar et al., 1999, Ryoo et al., 1999, Rieckhof et al., 1997)
                  exd - kni
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - lab
                  bimolecular fluorescence complementation, fluorescence microscopy, electrophoretic mobility shift assay, autoradiography
                  (Sánchez-Higueras et al., 2019, Hudry et al., 2012, Stultz et al., 2012, Ryoo et al., 1999, Grieder et al., 1997)
                  exd - lbe
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - mam
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - mid
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - nau
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - pnt
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - salm
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - slou
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - slp2
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - srp
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - tin
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  exd - tsh
                  pull down, autoradiography, bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018, Bessa et al., 2002)
                  exd - tup
                  bimolecular fluorescence complementation, fluorescence microscopy
                  (Bischof et al., 2018)
                  Summary of Genetic Interactions
                  esyN Network Diagram
                  esyN Network Key:
                  Suppression
                  Enhancement
                  View in Interactions Browser
                  Please look at the allele data for full details of the genetic interactions
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  Starting gene(s)
                  Interaction type
                  Interacting gene(s)
                  Reference
                  Antp
                  suppressible
                  exd
                  (Prince et al., 2008)
                  Antp|N
                  suppressible
                  exd
                  (Prince et al., 2008)
                  Dfd
                  enhanceable
                  exd
                  (Florence and McGinnis, 1998, Pinsonneault et al., 1997)
                  en
                  suppressible
                  exd
                  (Kobayashi et al., 2003)
                  Ubx
                  suppressible
                  exd|hth
                  (Sprecher et al., 2004)
                  External Data
                  Subunit Structure (UniProtKB)
                  Interacts with Ubx and hth.
                  (UniProt, P40427 )
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DroID - A comprehensive database of gene and protein interactions.
                  InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
                  MIST (genetic) - An integrated Molecular Interaction Database
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Pathways
                  Signaling Pathways (FlyBase)
                  Metabolic Pathways
                  External Data
                  Linkouts
                  Genomic Location and Detailed Mapping Data
                  Chromosome (arm)
                  X
                  Recombination map

                  1-52

                  Cytogenetic map
                  14A5-14A5
                  Sequence location
                  X:15,992,468..15,996,509 [+]
                  FlyBase Computed Cytological Location
                  Cytogenetic map
                  Evidence for location
                  14A5-14A5
                  Limits computationally determined from genome sequence between P{EP}Gβ13FEP1071 and P{EP}EP1458&P{EP}EP1522
                  Experimentally Determined Cytological Location
                  Cytogenetic map
                  Notes
                  References
                  14A-14A
                  (determined by in situ hybridisation)
                  (Flegel et al., 1993)
                  Experimentally Determined Recombination Data
                  Location

                  1-52

                  (FlyBase, 2016, Pinsonneault et al., 1997)

                  1-52.4

                  (Comeron et al., 2012)

                  1-54

                  (Tearle and Nusslein-Volhard, 1987)
                  Left of (cM)
                  Right of (cM)
                  Notes
                  Stocks and Reagents
                  Stocks (13)
                  Genomic Clones (10)
                   

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

                  cDNA Clones (335)
                   

                  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 sequences
                  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.

                  cDNA Clones, End Sequenced (ESTs)
                  BDGP DGC clones
                  Other clones
                  RNAi and Array Information
                  Linkouts
                  DRSC - Results frm RNAi screens
                  GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
                  Antibody Information
                  Laboratory Generated Antibodies
                   
                  Commercially Available Antibodies
                   
                  Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                  Other Information
                  Relationship to Other Genes
                  Source for database identify of

                  Source for identity of: exd CG8933

                  (Wang, 1999.11)
                  Source for database merge of

                  Source for merge of: exd anon- EST:fe1H3

                  (Bayraktaroglu, 2000.11.2)
                  Additional comments
                  Other Comments

                  exd is required for salivary gland formation.

                  (Henderson and Andrew, 2000)

                  exd mutants show patterning defects in the primary axonal scaffolds of the developing embryonic brain.

                  (Nagao et al., 2000)

                  The exd gene product's function is balanced by the activities of its NES and NLS. The hth protein alters this balance in favor of the NLS.

                  (Abu-Shaar et al., 1999)

                  The exd gene encodes a product that includes an NLS (nuclear localization signal), an NES (nuclear export signal) and a region required for hth-mediated nuclear localization. The hth gene product is required to overcome the influence of the exd NES (nuclear export signal), possibly by inducing a conformational change in the exd product. The exd and hth gene products interact in the cytoplasm.

                  (Abu-Shaar et al., 1999)

                  The subcellular location of exd gene product and its mammalian counterpart Hsap\PBX1 depends on nuclear export and import signals and is modulated by association with hth protein or its mammalian counterpart Hsap\PKNOX1.

                  (Berthelsen et al., 1999)

                  exd encodes a specificity cofactor for the trunk Hox genes. The control of exd subcellular localization is a mechanism to regulate Hox activity during development.

                  (Ryoo and Mann, 1999)

                  Scr negatively regulates hth, which is required for the nuclear localization of the exd gene product.

                  (Ryoo and Mann, 1999)

                  exd activity is required for the apparent stability of the hth protein.

                  (Abu-Shaar and Mann, 1998)

                  Subcellular localisation of exd is regulated by the BX-C genes and each BX-C gene can prevent or reduce nuclear translocation of exd to different extents. Ubx and abd-A require exd activity for their maintenance and functions.

                  (Azpiazu and Morata, 1998)

                  exd and hth are antennal-determining genes. hth is an antennal selector gene and Antp promotes leg development by repressing hth and consequently nuclear exd.

                  (Casares and Mann, 1998)

                  exd function and dpp/wg signalling are antagonistic and divide the leg into two mutually exclusive domains. In the proximal domain exd activity prevents cells from responding to dpp and wg. In the distal domain, exd function is suppressed by the dpp/wg response gene Dll, which prevents the nuclear transport of exd. The division of the limb into two antagonistic domains, as defined by exd function and hh signaling, may be a general feature of limb development.

                  (Gonzalez-Crespo et al., 1998)

                  hth acts upstream of exd and is required and sufficient for nuclear localisation of exd protein. hth and exd are both negative regulators of eye development.

                  (Pai et al., 1998)

                  exd protein localised to the nucleus is proposed to suppress tarsus development and activate arista development. In the mesodermal adepithelial cells of the leg imaginal discs, Scr protein is proposed to be required for the synthesis of a tarsus-inducer that when secreted acts on the ectoderm cells inhibiting nuclear accumulation of exd protein, such that tarsus determination is no longer suppressed and arista determination is no longer activated.

                  (Percival-Smith and Hayden, 1998)

                  Loss of exd activity is epistatic to loss of pb activity or loss of pb and Scr activity.

                  (Percival-Smith and Hayden, 1998)

                  Normal ph-p and ph-d expression depends on exd expression in vivo.

                  (Serrano and Maschat, 1998)

                  The nucleocytoplasmic localisation of exd protein is highly regulated throughout development.

                  (Aspland and White, 1997)

                  exd function is required for the autoactivation phase of Dfd expression in the posterior head. A minimal Dfd response element is regulated directly by the combination of Dfd and exd without the need for a specialised heterodimer-binding site for the two proteins.

                  (Pinsonneault et al., 1997)

                  hth is necessary for the nuclear localisation of exd protein.

                  (Rieckhof et al., 1997)

                  Although exd product affects wing development the presence of exd fails to modify the wing to haltere transformation caused by ectopic expression of either Ubx or abd-A.

                  (Casares et al., 1996)

                  Protein-DNA interactions are important for forming the lab-exd-DNA complex. Data suggests a model in which lab and exd bind DNA as a heterodimer in a head-to-tail orientation.

                  (Chan and Mann, 1996)

                  A 20bp oligonucleotide from the 5' region of Mmus\Hoxb1, a homolog of lab, is sufficient to direct an expression pattern in Drosophila very similar to endogenous lab. This expression requires lab abd exd function. In vitro DNA binding studies reveal that lab requires exd to bind DNA with high affinity.

                  (Chan et al., 1996)

                  Clonal analysis establishes a precise boundary for exd requirement along the proximal-distal axis of the leg at the proximal femur; exd is required in proximal, not in distal leg. exd and Dll are required in near complementary domains along the leg, with the exception of the trochanter and proximal femur where both gene functions are needed. exd is expressed in a normal pattern in the absence of hh function.

                  (Gonzalez-Crespo and Morata, 1996)

                  Both dpp and wg are required for the nuclear localization of exd protein in the endoderm. Ectopic expression of dpp and wg expands the domain of nuclear exd. The nuclear import of exd product correlates with the transcription of an exd-dependent reporter gene in the endoderm.

                  (Mann and Abu-Shaar, 1996)

                  Two trypotophan residues within the en EH2 domain are required for cooperativity with the exd gene product.

                  (Peltenburg and Murre, 1996)

                  Mosaic analysis reveals that exd is cell autonomously required and critically involved in adult morphogenesis, not only in the homeotic function, but also in several other developmental processes.

                  (Gonzalez-Crespo and Morata, 1995)

                  The physical association of exd protein with Ubx protein and other HOM proteins is studied using a yeast two-hybrid system. Highly selective interactions exist between exd and certain isoforms of Ubx and with the abd-A protein.

                  (Johnson et al., 1995)

                  exd is critically involved in adult morphogenesis, not only in the homeotic function, but also in several other developmental processes. In some regions of the adult exd mutant clones exhibit homeotic transformations similar to those produced by known homeotic mutations at Ubx, lab, ss or Antp. In other regions the lack of exd causes novel homeotic transformations producing ectopic eyes and legs. exd is also required for functions not normally associated with homeosis, such as the maintenance of the dorso-ventral pattern in the abdomen, the specification of subpatterns in adult appendages or the arrangement of bristles in the mesonotum and genitalia.

                  (Morata, 1995)

                  In vitro DNA binding studies indicate that Dfd product affinity for the 120bp Dfd epidermal element is markedly enhanced when exd protein is added to the binding reaction.

                  (Pinsonneault and McGinnis, 1995)

                  Virtually all of the P{r4/lacZ} expression in Drosophila depends on the same three conserved sequence elements involved in regulating expression in the mouse and lab expression. Expression in the head and the visceral mesoderm requires exd function.

                  (Popperl et al., 1995)

                  Uniform expression of exd+ suppresses the phenotype of exd null mutations. exd mosaics survive to adult stages but with reduced viability, exd does play an essential role in the development of adult structures. Loss of exd in the adult abdomen and in the head leads to homeotic transformations.

                  (Rauskolb et al., 1995)

                  Ubx protein directly regulates dpp expression. At least one other activity, possibly exd, is required in conjunction with Ubx for PS7-specific activation.

                  (Sun et al., 1995)

                  Cooperative interactions between Ubx protein and exd protein selectively increase the affinity of Ubx but not Antp for a particular DNA target.

                  (Chan et al., 1994)

                  exd is required for the appropriate regulation of at least some of the selector homeodomain proteins downstream target genes.

                  (Rauskolb and Wieschaus, 1994)

                  The exd protein raises the DNA binding specificity of Ubx and abd-A protein, but not that of Abd-B. The exd protein modulates the DNA binding activity of en to a different site. While a region N-terminal of the exd homeodomain is required for Ubx and abd-A cooperativity, en requires a domain C-terminal of the exd homeobox.

                  (van Dijk and Murre, 1994)

                  Isolated from a Drosophila adult cDNA library, using a fragment containing the human pbx1 homeobox as a probe, under low stringency conditions.

                  (Flegel et al., 1993)

                  An exd cDNA has been cloned and sequenced, and its expression pattern has been analysed.

                  (Flegel et al., 1993)

                  Mutations in exd cause homeotic transformations in both the embryo and the adult. The exd gene product acts with the selector homeodomain proteins as a DNA binding transcription factor, thereby altering their regulation of downstream target genes.

                  (Rauskolb et al., 1993)

                  exd alters segmental identity without altering the pattern of expression of homeotic genes. It may act by altering the target specificity of these genes.

                  (Peifer and Wieschaus, 1990)

                  exd mutants display mesothoracic and metathoracic segments like the prothorax, first abdominal segment is like the posterior abdomen.

                  (Wieschaus et al., 1984)
                  Origin and Etymology
                  Discoverer
                  Etymology
                  Identification
                  External Crossreferences and Linkouts ( 56 )
                  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.
                  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/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                  UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
                  Other crossreferences
                  BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                  Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                  Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                  Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                  Flygut - An atlas of the Drosophila adult midgut
                  GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
                  iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
                  InterPro - A database of protein families, domains and functional sites
                  KEGG Genes - Molecular building blocks of life in the genomic space.
                  modMine - A data warehouse for the modENCODE project
                  SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  DPiM - Drosophila Protein interaction map
                  DroID - A comprehensive database of gene and protein interactions.
                  DRSC - Results frm RNAi screens
                  Developmental Studies Hybridoma Bank - Monoclonal antibodies for use in research
                  FLIGHT - Cell culture data for RNAi and other high-throughput technologies
                  FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                  FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                  FlyMine - An integrated database for Drosophila genomics
                  Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                  InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
                  MIST (genetic) - An integrated Molecular Interaction Database
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Synonyms and Secondary IDs (16)
                  Reported As
                  Symbol Synonym
                  CG8933
                  (Kumar et al., 2016, Hosono et al., 2015, Ni et al., 2011.7.19, Neely et al., 2010, Keleman et al., 2009.8.5, Perkins et al., 2009)
                  DExd
                  (Galant and Carroll, 2002)
                  Dm-EXD
                  (Prpic et al., 2003)
                  Dpbx
                  (Flegel et al., 1993, Flegel, 1992.12.1)
                  EXD
                  (Altamirano-Torres et al., 2018, Blatti et al., 2015, Sivanantharajah and Percival-Smith, 2015, Bischof et al., 2013, Kim et al., 2010, Plaza et al., 2008, Percival-Smith et al., 2005, Rauskolb, 1993)
                  Exd
                  (Delker et al., 2019, Sánchez-Higueras et al., 2019, Zandvakili et al., 2019, Bischof et al., 2018, Powers and Srivastava, 2018, Chambers et al., 2017, Requena et al., 2017, Beh et al., 2016, Uhl et al., 2016, Zandvakili and Gebelein, 2016, Foos et al., 2015, Garaulet and Lai, 2015, Boekhoff-Falk and Eberl, 2014, Duffraisse et al., 2014, Hsiao et al., 2014, Makki et al., 2014, Spletter and Schnorrer, 2014, Wernet et al., 2014, Heffer et al., 2013, Sambrani et al., 2013, Wang et al., 2013, Agelopoulos et al., 2012, Fujioka et al., 2012, Hudry et al., 2012, Papadopoulos et al., 2012, Agrawal et al., 2011, Hudry et al., 2011, Lelli et al., 2011, Noro et al., 2011, Slattery et al., 2011, Reed et al., 2010, Uhl et al., 2010, Witt et al., 2010, Liu et al., 2009, Gebelein et al., 2008, Heffer and Pick, 2008, Karlsson et al., 2008, Moretti et al., 2008, Joshi et al., 2007, Joshi et al., 2007, Karlsson et al., 2007, Merabet et al., 2007, Merabet et al., 2007, Monier et al., 2007, Noro and Mann, 2007, Friedrich, 2006, Noro et al., 2006, Pueschel and Boekhoff-Falk, 2006, Bolinger and Boekhoff-Falk, 2005, Merabet et al., 2005, Mann and Carroll, 2002)
                  Pbx1
                  anon-EST:fe1H3
                  exd
                  (DeAguero et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Rastogi et al., 2018, Baril et al., 2017, Bielmeier et al., 2016, Bürglin and Affolter, 2016, Lycette et al., 2016, Camino et al., 2015, Schertel et al., 2015, Oas et al., 2014, Rogers et al., 2014, Sambrani et al., 2014, Shukla et al., 2014, Spletter and Schnorrer, 2014, Zohar-Stoopel et al., 2014, Baek et al., 2013, Corsetti and Azpiazu, 2013, Li et al., 2013, Bryantsev et al., 2012, Spokony and White, 2012.5.22, Stultz et al., 2012, Ahn et al., 2011, Kuzin et al., 2011, Merabet et al., 2011, Nègre et al., 2011, Saadaoui et al., 2011, Singh et al., 2011, Duncan et al., 2010, Joshi et al., 2010, Kannan et al., 2010, Karlsson et al., 2010, Popodi et al., 2010-, Venken et al., 2010, Venken et al., 2009, Venken et al., 2009, Coiffier et al., 2008, Li-Kroeger et al., 2008, Prince et al., 2008, Rogulja-Ortmann et al., 2008, Sato et al., 2008, Singh et al., 2008, Stultz et al., 2008, Aldaz et al., 2007, Ebacher et al., 2007, Mukherjee and Bürglin, 2007, Singh and Wook-Choi, 2007, Sprecher et al., 2007, Stevens and Mann, 2007, Chopra and Mishra, 2006, Legube et al., 2006, Stultz et al., 2006, Conant and Wagner, 2005, Merabet et al., 2005, Pearson et al., 2005, Grienenberger et al., 2003, Huang et al., 2003, Gim et al., 2001, Trachtulec and Forejt, 2001)
                  l(1)IV
                  lincRNA.S9404
                  (FlyBase Genome Annotators, 2014, Young et al., 2012)
                  td48
                  (Rauskolb et al., 1993)
                  Name Synonyms
                  Extradenticle
                  (Spletter and Schnorrer, 2014, Bischof et al., 2013, Corsetti and Azpiazu, 2013, Wang et al., 2013, Hudry et al., 2012, Ahn et al., 2011, Joshi et al., 2010, Kim et al., 2010, Reed et al., 2010, Witt et al., 2010, Liu et al., 2009, Venken et al., 2009, Gebelein et al., 2008, Li-Kroeger et al., 2008, Sato et al., 2008, Joshi et al., 2007, Monier et al., 2007, Noro and Mann, 2007, Friedrich, 2006, Noro et al., 2006, Pueschel and Boekhoff-Falk, 2006, Percival-Smith et al., 2005)
                  Pbx1-oncogene-like
                  anon-fast-evolving-1H3
                  extradenticle
                  (Powers and Srivastava, 2018, Wieschaus and Nüsslein-Volhard, 2016, Shukla et al., 2014, Baek et al., 2013, Bryantsev et al., 2012, Stultz et al., 2012, Kannan et al., 2010, Karlsson et al., 2010, Neely et al., 2010, Coiffier et al., 2008, Moretti et al., 2008, Stultz et al., 2008, Ebacher et al., 2007, Chopra and Mishra, 2006, Merabet et al., 2005, Grienenberger et al., 2003, Gim et al., 2001, Smith, 1995.8.11)
                  Secondary FlyBase IDs
                  • FBgn0010248
                  • FBgn0025242
                  Datasets (0)
                  Study focus (0)
                  Experimental Role
                  Project
                  Project Type
                  Title
                  References (385)