FB2020_06, released Dec 22, 2020

Gene: Dmel\fkh

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General Information
Symbol
Dmel\fkh
Species
D. melanogaster
Name
fork head
Annotation Symbol
CG10002
Feature Type
protein_coding_gene
FlyBase ID
FBgn0000659
Gene Model Status
Current
Stock Availability
14 publicly available
Gene Snapshot
fork head (fkh) encodes a winged-helix nuclear transcription factor most studied for its role in salivary gland formation, where it is required for salivary gland viability, invagination, and maintaining expression of other early-expressed salivary gland transcription factors. It works with the product of sage to activate expression of salivary gland specific gene products, such as secreted proteins and their modifying enzymes. [Date last reviewed: 2019-03-07]
Other Summaries
Also Known As

Secretion enhancer-binding protein 2, Sebp2

Key Links
Genomic Location
Cytogenetic map
98D5-98D5
Sequence location
3R:28,580,976..28,585,264 [-]
Recombination map

3-96

RefSeq locus
NT_033777 REGION:28580976..28585264
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 Ontology (GO) Annotations (26 terms)
Molecular Function (5 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Abrams et al., 2006)
enables RNA polymerase II transcription regulatory region sequence-specific DNA binding
inferred from direct assay
(Abrams et al., 2006, Lehmann and Korge, 1995)
Terms Based on Predictions or Assertions (4 terms)
CV Term
Evidence
References
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
enables protein domain specific binding
inferred from electronic annotation with InterPro:IPR013638
(assigned by InterPro )
(FlyBase Curators et al., 2004-)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
enables transcription factor binding
inferred from electronic annotation with InterPro:IPR013638
(assigned by InterPro )
(FlyBase Curators et al., 2004-)
Biological Process (18 terms)
Terms Based on Experimental Evidence (14 terms)
CV Term
Evidence
References
involved_in determination of adult lifespan
inferred from mutant phenotype
(Bolukbasi et al., 2017)
involved_in DNA endoreduplication
inferred from mutant phenotype
(Maruyama et al., 2011)
involved_in ecdysone-mediated induction of salivary gland cell autophagic cell death
inferred from direct assay
(Cao et al., 2007)
involved_in insulin receptor signaling pathway
inferred from direct assay
(Bolukbasi et al., 2017)
involved_in Malpighian tubule morphogenesis
inferred from mutant phenotype
(Maruyama et al., 2011)
involved_in negative regulation of apoptotic process
inferred from mutant phenotype
(Myat and Andrew, 2000)
involved_in negative regulation of cell growth
inferred from mutant phenotype
(Bülow et al., 2010)
involved_in negative regulation of multicellular organism growth
inferred from mutant phenotype
(Bülow et al., 2010)
involved_in negative regulation of programmed cell death
inferred from direct assay
(Cao et al., 2007)
involved_in neuroendocrine cell differentiation
inferred from mutant phenotype
(Miguel-Aliaga et al., 2008)
involved_in positive regulation of transcription by RNA polymerase II
inferred from direct assay
(Abrams et al., 2006)
involved_in regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Liu and Lehmann, 2008)
involved_in salivary gland development
inferred from mutant phenotype
(Maruyama et al., 2011, Abrams and Andrew, 2005)
involved_in salivary gland morphogenesis
inferred from mutant phenotype
(Myat and Andrew, 2000)
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
involved_in anatomical structure morphogenesis
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
involved_in cell differentiation
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
involved_in endoderm formation
traceable author statement
(Stainier, 2002)
involved_in negative regulation of transcription by RNA polymerase II
traceable author statement
(Bradley et al., 2001)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
Cellular Component (3 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Fox et al., 2013, Jattani et al., 2009, Lehmann and Korge, 1995, Weigel et al., 1989)
located_in polytene chromosome
inferred from direct assay
(Fox et al., 2013)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
located_in chromatin
inferred from biological aspect of ancestor with PANTHER:PTN000930906
(assigned by GO_Central )
(Gaudet et al., 2011)
Gene Group (FlyBase)
Pathway (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
FORK HEAD BOX TRANSCRIPTION FACTORS -
The forkhead box family of transcription factors are sequence-specific DNA binding proteins that regulate transcription. They are characterized by approximately 110 amino acid DNA-binding domain known as the forkhead or winged helix. (Adapted from FBrf0173142).
Pathway (FlyBase)
Insulin-like Receptor Signaling Pathway Core Components -
The Insulin-like Receptor (IR) signaling pathway in Drosophila is initiated by the binding of an insulin-like peptides to the Insulin-like receptor (InR). (Adapted from FBrf0232297, FBrf0230017 and FBrf0229989.)
Protein Function (UniProtKB)
Fkh promotes terminal as opposed to segmental development. In the absence of fkh, this developmental switch does not occur. The nuclear localization of the fkh protein suggest that fkh regulates the transcription of other, subordinate, genes.
(UniProt, P14734)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
fkh: fork head
Embryonic lethal. fkh+ appears to be required in the most anterior and posterior regions of the embryo; in amorphic mutants homeotic transformation effects the appearance of post-oral head structures in these terminal domains. The ectopic head structures are sometimes associated with thoracic structures anteriorly and anterior tail structures posteriorly. Thus fkh mutations produce transformations directed to the center involving structures normally found in different parasegments. Anteriorly, esophagus and proventriculus, derivatives of the ectodermal stomodaeum, are absent but not the pharynx or the hypopharyngeal organ, which resides on the anterior surface of the embryo; parts of the head skeleton are distorted; other parts apparently normal. Salivary glands absent. Posteriorly, anal pads and Malpighian tubules, derivatives of the ectodermal proctodaeum, are absent; replaced by anterior tail structures and post-oral head structures; supernumerary anal sensilla and dorsal hairs also present. Anterior and posterior fkh domains are beyond the regions controlled by ANTC and BXC, respectively; however ectopic expression of ANTC and BXC expression can occur in the fkh domains of fkh but not fkh+ embryos. No maternal effect. Homozygous fkh clones in adult cuticle completely normal in structure indicating no requirements for fkh+ in imaginal disk development. In situ hybridization reveals transcript in two terminal domains shortly before blastoderm cellularization occupying 5% embryonic length anteriorly and 15% posteriorly. fkh protein first detected in posterior domain at the end of syncytial blastoderm and the anterior domain at the beginning of cellularization. Expression continues in the derivatives of the ectodermal stomodaeum and proctodaeum throughout gastrulation. In addition fkh protein is detected in the midgut, the salivary glands, the central nervous system, and the yolk cells.
Summary (Interactive Fly)

winged-helix nuclear transcription factor - plays two roles in the formation of the embryonic salivary glands: an early role in promoting survival of the secretory cells, and a later role in secretory cell invagination works with bHLH protein Sage to activate expression of salivary gland specific gene products, such as secreted proteins and their modifying enzymes

Gene Model and Products
Number of Transcripts
5
Number of Unique Polypeptides
3

Please see the JBrowse view of Dmel\fkh 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 5.55

Gene model reviewed during 5.44

Double stop-codon suppression (UGA, UAA) postulated; FBrf0216884,

gene_with_stop_codon_read_through ; SO:0000697

Gene model reviewed during 5.40

Gene model reviewed during 6.02

Sequence Ontology: Class of Gene
gene_with_stop_codon_read_through
(Jungreis et al., 2011)
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
fkh-RA
FBtr0085321
NM_079818
3280
510
fkh-RB
FBtr0300259
NM_001170291
4030
510
fkh-RC
FBtr0330333
NM_001276109
4030
692
fkh-RD
FBtr0345227
NM_001300645
4289
510
fkh-RE
FBtr0346765
NM_001316532
4030
518
Additional Transcript Data and Comments
Reported size (kB)

4.2 (northern blot)

(Weigel et al., 1989)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
fkh-PA
FBpp0084690
54.3
510
8.71
NP_524542
AAF56798
fkh-PB
FBpp0289487
54.3
510
8.71
NP_001163762
ACZ95056
fkh-PC
FBpp0303365
74.4
692
8.54
NP_001263038
AGB96418
fkh-PD
FBpp0311419
54.3
510
8.71
NP_001287574
AHN57573
fkh-PE
FBpp0312360
55.2
518
8.71
NP_001303461
ALI30648
Polypeptides with Identical Sequences

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

510 aa isoforms: fkh-PA, fkh-PB, fkh-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

510 (aa); 54 (kD)

(Weigel et al., 1989)
Comments

fkh protein is used as a marker for the epithelium of the foregut and hindgut primordia.

(Maeda et al., 2007)
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
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 5
organism | 90-100% egg length

Comment: 95-100% egg length

(Weigel et al., 1989)
embryonic stage 4
organism | 90-100% egg length

Comment: 95-100% egg length

(Weigel et al., 1989)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10
inclusive hindgut primordium
(Wu and Lengyel, 1998)
embryonic stage 13
Malpighian tubule
(Wu and Lengyel, 1998)
embryonic hindgut
(Iwaki et al., 2001)
MP1 neuron
(Wheeler et al., 2006)
H-cell sib
(Wheeler et al., 2006)
VUM interneuron
(Wheeler et al., 2006)
embryonic stage 4
organism | 0-20% egg length

Comment: 0-15% egg length

(Weigel et al., 1989)
organism | terminal expression pattern
(Wu and Lengyel, 1998)
organism | 90-100% egg length

Comment: 95-100% egg length

(Wu and Lengyel, 1998)
organism | 0-10% egg length
(Wu and Lengyel, 1998)
embryonic stage 5
organism | 0-20% egg length

Comment: 0-15% egg length

(Weigel et al., 1989)
embryonic stage 15
hindgut
(Wu and Lengyel, 1998)
embryonic stage 6 -- 7
esophagus anlage
(De Velasco et al., 2004)
proventriculus anlage
(De Velasco et al., 2004)
embryonic stage 4 -- 6
anterior endoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
endoderm 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)
posterior ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
yolk nucleus
(Fisher et al., 2012)
embryonic stage 7 -- 8
anterior endoderm anlage
(Fisher et al., 2012)
foregut anlage
(Fisher et al., 2012)
hindgut anlage
(Fisher et al., 2012)
posterior endoderm
(Fisher et al., 2012)
yolk nucleus
(Fisher et al., 2012)
embryonic stage 9 -- 10
amnioserosa
(Fisher et al., 2012)
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)
foregut primordium
(Fisher et al., 2012)
inclusive hindgut primordium
(Fisher et al., 2012)
posterior endoderm
(Fisher et al., 2012)
salivary gland body primordium

Comment: reported as salivary gland body specific anlage

(Fisher et al., 2012)
yolk nucleus
(Fisher et al., 2012)
embryonic stage 11 -- 12
anterior midgut primordium
(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
embryonic central brain glial cell
(Fisher et al., 2012)
embryonic optic lobe primordium
(Fisher et al., 2012)
foregut primordium
(Fisher et al., 2012)
hindgut proper primordium
(Fisher et al., 2012)
large intestine primordium

Comment: reported as large intestine specific anlage

(Fisher et al., 2012)
lateral cord glial cell
(Fisher et al., 2012)
Malpighian tubule main body primordium
(Fisher et al., 2012)
Malpighian tubule primordium
(Fisher et al., 2012)
midline primordium
(Fisher et al., 2012)
midline ventral glial cell
(Fisher et al., 2012)
posterior midgut primordium
(Fisher et al., 2012)
proventriculus primordium
(Fisher et al., 2012)
rectum primordium

Comment: reported as rectum specific anlage

(Fisher et al., 2012)
salivary gland body primordium

Comment: reported as salivary gland primordium

(Fisher et al., 2012)
small intestine primordium

Comment: reported as small intestine specific anlage

(Fisher et al., 2012)
stomatogastric nervous system primordium
(Fisher et al., 2012)
yolk nucleus
(Fisher et al., 2012)
embryonic stage 13 -- 16
embryonic anal pad
(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic central brain glial cell
(Fisher et al., 2012)
embryonic central brain neuron
(Fisher et al., 2012)
embryonic esophagus
(Fisher et al., 2012)
embryonic foregut
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic hindgut
(Fisher et al., 2012)
embryonic large intestine
(Fisher et al., 2012)
embryonic main segment of Malpighian tubule
(Fisher et al., 2012)
embryonic Malpighian tubule
(Fisher et al., 2012)
embryonic proventriculus
(Fisher et al., 2012)
embryonic rectum
(Fisher et al., 2012)
embryonic small intestine
(Fisher et al., 2012)
embryonic stomatogastric nervous system
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
embryonic/larval salivary gland
(Fisher et al., 2012)
embryonic/larval salivary gland body
(Fisher et al., 2012)
ventral midline of embryo
(Fisher et al., 2012)
yolk nucleus
(Fisher et al., 2012)
adult stage
adult mushroom body | restricted
(Kobayashi et al., 2006)
embryonic stage 11
ventral midline of embryo | subset

Comment: late stage 11

(Wheeler et al., 2006)
embryonic stage 17
MP1 neuron
(Wheeler et al., 2006)
H-cell sib
(Wheeler et al., 2006)
VUM interneuron
(Wheeler et al., 2006)
median neuroblast MNB interneuron
(Wheeler et al., 2006)
embryonic stage
embryonic/larval proventriculus
(Hernández de Madrid and Casanova, 2018)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
late third instar larval stage
embryonic/larval salivary gland
(Cao et al., 2007)
Additional Descriptive Data

Expression assayed at stages 9, 11, 13, and 17. Expression may be continuous between assayed stages in some tissues.

(Wheeler et al., 2006)

In early embryos, fkh expression forms a posterior and an anterior cap. At later developmental stages fkh transcript expression is used to label the elongating malpighian tubules.

(Wu and Lengyel, 1998)

fkh transcripts are detected in blastoderm embryos in the terminal domains from 0-15% egg length and from 95-100% egg length.

(Weigel et al., 1989)
Marker for
 
embryonic Malpighian tubule
(Wu and Lengyel, 1998)
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 9
stomodeum
(Weigel et al., 1989)
embryonic stage 5
organism | 90-100% egg length

Comment: 94-100% egg length

(Weigel et al., 1989)
embryonic stage 12 -- 17
embryonic/larval salivary gland
(Weigel et al., 1989)
dissected tissue
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
embryonic/larval salivary gland
(Lehmann and Korge, 1996)
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 12 -- 17
presumptive embryonic/larval central nervous system | presumptive
(Weigel et al., 1989)
embryonic foregut

Comment: ectodermal cells

(Fuss et al., 2004)
embryonic stage 5
organism | 0-20% egg length

Comment: 0-13% egg length

(Weigel et al., 1989)
embryonic stage 9
yolk
(Weigel et al., 1989)
embryonic/larval anterior midgut
(Weigel et al., 1989)
embryonic stage 14 -- 15
keyhole structure | anterior
(Fuss et al., 2004)
embryonic stage 17
embryonic proventriculus inner layer
(Fuss et al., 2004)
embryonic/larval proventriculus
(Lechner et al., 2007)
embryonic stage 11 -- 12
anterior midgut primordium
(Weigel et al., 1990)
posterior midgut primordium
(Weigel et al., 1990)
stomodeal invagination
(Weigel et al., 1990)
hindgut proper primordium
(Weigel et al., 1990)
embryonic stage 12
salivary gland body primordium
(Weigel et al., 1990)
neuroblast | subset
(Maruyama et al., 2011)
embryonic stage 4 -- 5
organism | posterior
(Perkins et al., 1992)
embryonic stage
embryonic/larval dMP2 Ilp7 neuron
(Castellanos et al., 2013)
embryonic stage 10 -- 12
salivary gland body primordium
(Maruyama et al., 2011)
embryonic Malpighian tubule
(Maruyama et al., 2011)
embryonic stage 13 -- 15
embryonic/larval salivary gland
(Maruyama et al., 2011)
embryonic/larval Malpighian tubule
(Maruyama et al., 2011)
embryonic/larval proventriculus
(Maruyama et al., 2011)
embryonic stage 10 -- 11
stomodeum
(Maruyama et al., 2011)
embryonic stage 10
anterior midgut primordium
(Maruyama et al., 2011)
posterior midgut primordium
(Maruyama et al., 2011)
embryonic stage 14
embryonic/larval hindgut
(Maruyama et al., 2011)
embryonic stage 11 -- 14
yolk nucleus
(Maruyama et al., 2011)
embryonic stage 12 -- 13
embryonic foregut
(Lechner et al., 2007)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
late third instar larval stage
embryonic/larval salivary gland
(Cao et al., 2007)
prepupal stage P1 -- P3
embryonic/larval salivary gland
(Cao et al., 2007)
Additional Descriptive Data

fkh protein gradually disappears from the salivary gland nuclei, but remains strong in the imaginal rings that will later form the adult salivary gland.

(Cao et al., 2007)

fkh protein is used as a marker for the epithelium of the foregut and hindgut primordia.

(Maeda et al., 2007)

fkh protein is expressed in the ectodermal cells of the developing embryonic foregut. Expression stops sharply at the ectodermal/endodermal boundary.

(Fuss et al., 2004)

In csw mutant embryos, the posterior fkh domain is reduced posteriorly.

(Perkins et al., 1992)

fkh protein isdetected shortly after the onset of fkh transcription. It is firstdetected in the posterior domain at the end of the syncytial blastodermstage and in the anterior domain at the beginning of the cellularblastoderm stage. By the end of cellular blastoderm, the expressiondomains cover 0-13% egg length and 94-100% egg length. Double stainingwith the ftz antibody indicates that the posterior domain covers theregion posterior to parasegment 15. Subsequently, the anterior domain ofexpression expands such that by germ band elongation it covers theanterior midgut and stomodeal primordia. Yolk nuclei also stain. Later, itis expressed in the developing salivary glands and in the CNS.

(Weigel et al., 1989)
Marker for
 
embryonic foregut
(Maeda et al., 2007)
embryonic hindgut
(Maeda et al., 2007)
embryonic proventriculus inner layer
(Fuss et al., 2004)
Subcellular Localization
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Fox et al., 2013, Jattani et al., 2009, Lehmann and Korge, 1995, Weigel et al., 1989)
located_in polytene chromosome
inferred from direct assay
(Fox et al., 2013)
Expression Deduced from Reporters
Reporter: P{fkh-GAL4.14-3}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 8 -- 17
embryonic hindgut
(Johansen et al., 2003)
embryonic stage
embryonic/larval midgut | restricted
(Fuss and Hoch, 1998)
embryonic/larval hindgut
(Fuss et al., 2001, Fuss and Hoch, 1998)
embryonic/larval salivary gland
(Fuss and Hoch, 1998)
Reporter: P{fkh-GAL4.H}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10 -- 13
salivary gland body primordium
(Henderson and Andrew, 2000)
embryonic stage 13 -- 17
presumptive embryonic salivary gland
(Henderson and Andrew, 2000)
Reporter: P{fkh-lacZ.250}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 11
parasegment 2
(Zhou et al., 2001)
organism | segmentally repeated
(Gebelein et al., 2002, Zhou et al., 2001)
salivary gland body primordium
(Zhou et al., 2001)
embryonic stage 10
parasegment 2
(Ryoo and Mann, 1999)
third instar larval stage
prothoracic leg disc | restricted

Comment: absent in other thoracic discs

(Ryoo and Mann, 1999)
prothoracic trochanter | precursor
(Ryoo and Mann, 1999)
Reporter: P{GMR16E04-GAL4}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 11 -- 17
embryonic hindgut
(Hernández et al., 2015)
embryonic stage 15 -- larval stage
embryonic/larval hindgut
(Hernández et al., 2015)
embryonic stage 13 -- 16
embryonic/larval midgut
(Hernández et al., 2015)
larval stage
embryonic/larval salivary gland
(Mehta et al., 2020)
Reporter: P{GMR16H06-GAL4}
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
anterior optic tubercle | intense | restricted
(Jenett and Svirskas, 2012.9.24)
embryonic stage 11 -- 17
presumptive embryonic/larval central nervous system | faint
(Hernández et al., 2015)
Reporter: P{GMR17D02-GAL4}
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult mushroom body | faint | restricted
(Jenett and Svirskas, 2012.9.24)
embryonic stage 9 -- 12
early stomodeal invagination
(Hernández et al., 2015)
hindgut proper primordium
(Hernández et al., 2015)
embryonic stage 13 -- 17
embryonic hindgut
(Hernández et al., 2015)
presumptive embryonic/larval central nervous system | faint
(Hernández et al., 2015)
embryonic stage 13 -- 16
embryonic Malpighian tubule
(Hernández et al., 2015)
Reporter: P{GMR18A08-GAL4}
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult antennal lobe | restricted
(Jenett and Svirskas, 2012.9.24)
embryonic stage 11 -- 17
embryonic hindgut
(Hernández et al., 2015)
embryonic stage 15 -- 16
embryonic Malpighian tubule
(Hernández et al., 2015)
embryonic stage 11 -- 15
embryonic stomatogastric nervous system
(Hernández et al., 2015)
Reporter: P{lacZ}fkhF223
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 11
neuroblast MP2
(Kim et al., 2007)
embryonic stage 11 -- 16
dMP2 neuron
(Kim et al., 2007)
vMP2 neuron
(Kim et al., 2007)
Reporter: P{NXl.lacZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | posterior
(Weigel et al., 1990)
embryonic stage 12 -- 13
anterior midgut primordium
(Weigel et al., 1990)
posterior midgut primordium
(Weigel et al., 1990)
foregut primordium
(Weigel et al., 1990)
hindgut proper primordium
(Weigel et al., 1990)
salivary gland body primordium
(Weigel et al., 1990)
Malpighian tubule primordium
(Weigel et al., 1990)
embryonic stage 13
anal pad primordium
(Weigel et al., 1990)
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\fkh 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) 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 ( 18 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 16 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of fkh
Transgenic constructs containing regulatory region of fkh
Deletions and Duplications ( 18 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal | recessive
lethal - all die before end of embryonic stage | recessive
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
decreased cell death
decreased cell death | embryonic stage
decreased cell size | larval stage | somatic clone, with Scer\GAL4Act5C.PP
increased cell death
increased cell size | conditional, with Scer\GAL4Act5C.PP
neuroanatomy defective
small body | larval stage, with Scer\GAL4ppl.PP
some die during embryonic stage | recessive
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4pnr-MD237
Phenotype manifest in
Allele
adult dMP2 Ilp7 neuron
anterior midgut primordium
cephalopharyngeal skeleton
cercus
embryo | gap
embryo | posterior
embryonic/larval anterior midgut | embryonic stage
embryonic/larval esophagus | embryonic stage
embryonic/larval foregut
embryonic/larval foregut | embryonic stage
embryonic/larval hindgut | embryonic stage
embryonic/larval proventriculus | embryonic stage
embryonic/larval salivary gland
embryonic/larval salivary gland, with Scer\GAL4Tub.PU
embryonic/larval salivary gland | embryonic stage
embryonic brain
embryonic brain & ganglion mother cell
embryonic central brain
embryonic central brain glial cell
embryonic head
embryonic hindgut
embryonic primordium of adult salivary gland
embryonic stomatogastric nervous system
embryonic visceral muscle
external sensory organ | ectopic
hemocyte
labial disc
longitudinal visceral muscle primordium
macrophage
Malpighian tubule
Malpighian tubule primordium
mesothoracic tergum, with Scer\GAL4pnr-MD237
pars intercerebralis primordium
salivary gland
salivary gland common duct primordium
somatic mesoderm, with Scer\GAL4twi.PGa
stomatogastric nervous system
visceral mesoderm derivative, with Scer\GAL4twi.PGa
wing, with Scer\GAL4Bx-MS1096
Orthologs
Downloads
Download All DIOPT Orthologs
Download All OrthoDB Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (29)
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\FOXA2
12 of 15
Yes
Yes
Hsap\FOXA1
10 of 15
No
Yes
Hsap\FOXA3
7 of 15
No
Yes
Hsap\FOXB1
1 of 15
No
No
Hsap\FOXB2
1 of 15
No
No
Hsap\FOXC1
1 of 15
No
No
Hsap\FOXC2
1 of 15
No
No
Hsap\FOXD1
1 of 15
No
No
Hsap\FOXD2
1 of 15
No
No
Hsap\FOXD3
1 of 15
No
No
1  
Hsap\FOXD4
1 of 15
No
No
Hsap\FOXD4L1
1 of 15
No
No
Hsap\FOXD4L3
1 of 15
No
No
Hsap\FOXD4L4
1 of 15
No
No
Hsap\FOXD4L5
1 of 15
No
No
Hsap\FOXD4L6
1 of 15
No
No
Hsap\FOXE1
1 of 15
No
No
Hsap\FOXE3
1 of 15
No
No
Hsap\FOXF1
1 of 15
No
No
Hsap\FOXF2
1 of 15
No
No
Hsap\FOXG1
1 of 15
No
No
Hsap\FOXI1
1 of 15
No
No
Hsap\FOXI2
1 of 15
No
No
Hsap\FOXI3
1 of 15
No
No
Hsap\FOXJ1
1 of 15
No
No
Hsap\FOXL1
1 of 15
No
No
Hsap\FOXL2
1 of 15
No
No
Hsap\FOXQ1
1 of 15
No
No
Hsap\FOXS1
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (25)
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\Foxa2
11 of 15
Yes
Yes
Mmus\Foxa1
10 of 15
No
Yes
Mmus\Foxa3
7 of 15
No
Yes
Mmus\Foxb1
1 of 15
No
No
Mmus\Foxb2
1 of 15
No
No
Mmus\Foxc1
1 of 15
No
No
Mmus\Foxc2
1 of 15
No
No
Mmus\Foxd1
1 of 15
No
No
Mmus\Foxd2
1 of 15
No
No
Mmus\Foxd3
1 of 15
No
No
Mmus\Foxd4
1 of 15
No
No
Mmus\Foxe1
1 of 15
No
No
Mmus\Foxe3
1 of 15
No
No
Mmus\Foxf1
1 of 15
No
No
Mmus\Foxf2
1 of 15
No
No
Mmus\Foxg1
1 of 15
No
No
Mmus\Foxi1
1 of 15
No
No
Mmus\Foxi2
1 of 15
No
No
Mmus\Foxi3
1 of 15
No
No
Mmus\Foxj1
1 of 15
No
No
Mmus\Foxl1
1 of 15
No
No
Mmus\Foxl2
1 of 15
No
No
Mmus\Foxq1
1 of 15
No
No
Mmus\Foxs1
1 of 15
No
No
Mmus\Gm5294
1 of 15
No
No
Rattus norvegicus (Norway rat) (25)
Rnor\Foxa2
11 of 13
Yes
Yes
Rnor\Foxa3
7 of 13
No
Yes
Rnor\Foxa1
6 of 13
No
Yes
Rnor\Foxb1
1 of 13
No
No
Rnor\Foxb2
1 of 13
No
No
Rnor\Foxc1
1 of 13
No
No
Rnor\Foxc2
1 of 13
No
No
Rnor\Foxd1
1 of 13
No
No
Rnor\Foxd2
1 of 13
No
No
Rnor\Foxd3
1 of 13
No
No
Rnor\Foxd4
1 of 13
No
No
Rnor\Foxe1
1 of 13
No
No
Rnor\Foxe3
1 of 13
No
No
Rnor\Foxf1
1 of 13
No
No
Rnor\Foxf2
1 of 13
No
Yes
Rnor\Foxg1
1 of 13
No
No
Rnor\Foxi1
1 of 13
No
No
Rnor\Foxi2
1 of 13
No
No
Rnor\Foxi3
1 of 13
No
No
Rnor\Foxj1
1 of 13
No
No
Rnor\Foxl1
1 of 13
No
No
Rnor\Foxl2
1 of 13
No
No
Rnor\Foxq1
1 of 13
No
No
Rnor\Foxs1
1 of 13
No
No
Rnor\LOC680273
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (27)
Xtro\foxa2
11 of 12
Yes
Yes
Xtro\foxa1
10 of 12
No
Yes
Xtro\foxa4
10 of 12
No
Yes
Xtro\foxb1
1 of 12
No
No
Xtro\foxb2
1 of 12
No
No
Xtro\foxc1
1 of 12
No
No
Xtro\foxc2
1 of 12
No
No
Xtro\foxd1
1 of 12
No
No
Xtro\foxd2
1 of 12
No
No
Xtro\foxd3
1 of 12
No
No
Xtro\foxd4l1.1
1 of 12
No
No
Xtro\foxd4l1.2
1 of 12
No
No
Xtro\foxe1
1 of 12
No
No
Xtro\foxe3
1 of 12
No
No
Xtro\foxf1
1 of 12
No
No
Xtro\foxf2
1 of 12
No
No
Xtro\foxg1
1 of 12
No
No
Xtro\foxi1
1 of 12
No
No
Xtro\foxi2
1 of 12
No
No
Xtro\foxi4.1
1 of 12
No
No
Xtro\foxi4.2
1 of 12
No
No
Xtro\foxj1.2
1 of 12
No
No
Xtro\foxj1
1 of 12
No
No
Xtro\foxl1
1 of 12
No
No
Xtro\foxl2
1 of 12
No
No
Xtro\foxq1
1 of 12
No
No
Xtro\wi2-2373i1.2
1 of 12
No
No
Danio rerio (Zebrafish) (40)
Drer\foxa2
13 of 15
Yes
Yes
Drer\foxa1
9 of 15
No
Yes
Drer\foxa3
9 of 15
No
Yes
Drer\foxa
3 of 15
No
Yes
Drer\foxb1a
1 of 15
No
No
Drer\foxb1b
1 of 15
No
No
Drer\foxb2
1 of 15
No
No
Drer\foxc1a
1 of 15
No
No
Drer\foxc1b
1 of 15
No
No
Drer\foxd1
1 of 15
No
No
Drer\foxd2
1 of 15
No
No
Drer\foxd3
1 of 15
No
No
Drer\foxd5
1 of 15
No
No
Drer\foxe1
1 of 15
No
No
Drer\foxe3
1 of 15
No
No
Drer\foxf1
1 of 15
No
No
Drer\foxf2a
1 of 15
No
No
Drer\foxf2b
1 of 15
No
No
Drer\foxg1a
1 of 15
No
No
Drer\foxg1b
1 of 15
No
No
Drer\foxg1c
1 of 15
No
No
Drer\foxg1d
1 of 15
No
No
Drer\foxi1
1 of 15
No
No
Drer\foxi2
1 of 15
No
No
Drer\foxi3a
1 of 15
No
No
Drer\foxi3b
1 of 15
No
No
Drer\foxj1a
1 of 15
No
No
Drer\foxj1b
1 of 15
No
No
Drer\foxj2
1 of 15
No
No
Drer\foxl1
1 of 15
No
No
Drer\foxl2a
1 of 15
No
No
Drer\foxl2b
1 of 15
No
No
Drer\foxo6a
1 of 15
No
No
Drer\foxq1a
1 of 15
No
No
Drer\foxq1b
1 of 15
No
No
Drer\foxq2
1 of 15
No
No
Drer\si:ch211-145o7.3
1 of 15
No
No
Drer\si:dkey-7e14.3
1 of 15
No
No
Drer\zgc:162612
1 of 15
No
No
Drer\zgc:194189
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (10)
Cele\pha-4
7 of 15
Yes
Yes
Cele\fkh-2
1 of 15
No
No
Cele\fkh-6
1 of 15
No
No
Cele\fkh-8
1 of 15
No
No
Cele\fkh-9
1 of 15
No
No
Cele\fkh-10
1 of 15
No
No
Cele\let-381
1 of 15
No
No
Cele\lin-31
1 of 15
No
No
Cele\pes-1
1 of 15
No
No
Cele\unc-130
1 of 15
No
No
Arabidopsis thaliana (thale-cress) (0)
No records found.
Saccharomyces cerevisiae (Brewer's yeast) (4)
Scer\FKH1
3 of 15
Yes
No
Scer\HCM1
3 of 15
Yes
No
Scer\FKH2
2 of 15
No
No
Scer\FHL1
1 of 15
No
No
Schizosaccharomyces pombe (Fission yeast) (4)
Spom\mei4
3 of 12
Yes
No
Spom\fhl1
2 of 12
No
No
Spom\sep1
2 of 12
No
No
Spom\fkh2
1 of 12
No
No
Ortholog(s) in Drosophila Species (via OrthoDB v9.1) ( EOG091909A1 )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00011329
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00013343
Drosophila suzukii
Spotted wing Drosophila
Dsuz\DS10_00013551
Drosophila simulans
Dsim\GD18028
Drosophila sechellia
Dsec\GM16286
Drosophila erecta
Dere\GG12065
Drosophila yakuba
Dyak\GE10507
Drosophila ananassae
Dana\GF16437
Drosophila pseudoobscura pseudoobscura
Dpse\GA10002
Drosophila persimilis
Dper\GL23472
Drosophila willistoni
Dwil\GK10997
Drosophila virilis
Dvir\fkh
Drosophila mojavensis
Dmoj\GI22864
Drosophila grimshawi
Dgri\GH18716
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG091507ZY )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Mdoa\17025006
Glossina morsitans
Tsetse fly
Gmor\GMOY004775
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_01124
Lucilia cuprina
Australian sheep blowfly
Lcup\FF38_11190
Mayetiola destructor
Hessian fly
Mdes\Mdes014235
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL003163
Aedes aegypti
Yellow fever mosquito
Aaeg\AAEL003173
Anopheles darlingi
American malaria mosquito
Adar\ADAC009607
Anopheles gambiae
Malaria mosquito
Agam\AGAP001671
Culex quinquefasciatus
Southern house mosquito
Cqui\CPIJ001843
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W08HZ )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Bmor\BGIBMGA005101
Danaus plexippus
Monarch butterfly
Dple\DPOGS200192-PA
Heliconius melpomene
Postman butterfly
Hmel\HMEL017967
Apis florea
Little honeybee
Aflo\5239739604
Apis mellifera
Western honey bee
Amel\GB53404
Bombus impatiens
Common eastern bumble bee
Bimp\BIMP15851
Bombus terrestris
Buff-tailed bumblebee
Bter\15207928
Linepithema humile
Argentine ant
Lhum\LH11086
Megachile rotundata
Alfalfa leafcutting bee
Mrot\MROTU:002d7b
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG003582
Nasonia vitripennis
Parasitic wasp
Nvit\Nasvi2EG036450
Dendroctonus ponderosae
Mountain pine beetle
Dpod\YQE_04282
Tribolium castaneum
Red flour beetle
Tcas\fkh
Pediculus humanus
Human body louse
Phum\PHUM546090
Rhodnius prolixus
Kissing bug
Rpro\RPRC000089
Cimex lectularius
Bed bug
Clec\CLEC001132
Acyrthosiphon pisum
Pea aphid
Apim\ACYPI52843
Zootermopsis nevadensis
Nevada dampwood termite
Znev\Znev_13477
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0C7Q )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Smar\SMAR005037
Ixodes scapularis
Black-legged tick
Isca\ISCW006922
Stegodyphus mimosarum
African social velvet spider
Smim\KFM75273.1
Tetranychus urticae
Two-spotted spider mite
Turt\tetur14g02480
Daphnia pulex
Water flea
Dpux\DAPPUDRAFT_325321
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( EOG091G0HW9 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-FoxL1
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-FoxC
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-FoxA
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-FoxD
Strongylocentrotus purpuratus
Purple sea urchin
Spur\Sp-FoxD_1
Ciona intestinalis
Vase tunicate
Cint\foxc
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000023488
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000003394
Ciona intestinalis
Vase tunicate
Cint\ENSCING00000022098
Ciona intestinalis
Vase tunicate
Cint\foxd-a/b
Ciona intestinalis
Vase tunicate
Cint\foxE
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000012644
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000028153
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000025873
Gallus gallus
Domestic chicken
Ggal\FOXC2
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000022935
Gallus gallus
Domestic chicken
Ggal\ENSGALG00000026252
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (16)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
croc
3 of 10
fd19B
3 of 10
fd59A
3 of 10
fd96Ca
3 of 10
fd96Cb
3 of 10
fd102C
3 of 10
FoxL1
3 of 10
slp1
3 of 10
bin
2 of 10
CG32006
2 of 10
fd3F
2 of 10
slp2
2 of 10
FoxK
1 of 10
foxo
1 of 10
FoxP
1 of 10
jumu
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 v8.0 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?
    FOXA2; forkhead box A2
    12 of 15
    600288
        FOXA1; forkhead box A1
        10 of 15
        602294
            FOXA3; forkhead box A3
            7 of 15
            602295
                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
                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
                byn
                enhanceable
                fkh
                (Kusch and Reuter, 1999)
                External Data
                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 (protein-protein) - An integrated Molecular Interaction Database
                Pathways
                Signaling Pathways (FlyBase)
                Insulin-like Receptor Signaling Pathway Core Components -
                The Insulin-like Receptor (IR) signaling pathway in Drosophila is initiated by the binding of an insulin-like peptides to the Insulin-like receptor (InR). (Adapted from FBrf0232297, FBrf0230017 and FBrf0229989.)
                Metabolic Pathways
                External Data
                Linkouts
                KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
                Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                Genomic Location and Detailed Mapping Data
                Chromosome (arm)
                3R
                Recombination map

                3-96

                Cytogenetic map
                98D5-98D5
                Sequence location
                3R:28,580,976..28,585,264 [-]
                FlyBase Computed Cytological Location
                Cytogenetic map
                Evidence for location
                98D5-98D5
                Limits computationally determined from genome sequence between P{PZ}l(3)0648706487 and P{EP}EP3390EP3390
                Experimentally Determined Cytological Location
                Cytogenetic map
                Notes
                References
                98D2-98D3
                (determined by in situ hybridisation)
                (Ranz et al., 1997)
                29E-29E
                (determined by in situ hybridisation)
                (Zhong et al., 1993)
                98D1-98E1
                (determined by in situ hybridisation)
                (Weigel and Jäckle, 1990, Weigel et al., 1989)
                98D2-98D3
                (Tearle and Nusslein-Volhard, 1987)
                Location based on the common breakpoints of several translocations.
                (Tearle and Nusslein-Volhard, 1987)
                Experimentally Determined Recombination Data
                Location

                3-96

                (FlyBase, 2016)

                3-93.8

                (Comeron et al., 2012)

                3-95

                (Tearle and Nusslein-Volhard, 1987)
                Left of (cM)
                Right of (cM)
                Notes
                Stocks and Reagents
                Stocks (14)
                Genomic Clones (16)
                 

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

                cDNA Clones (51)
                 

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

                cDNA clones, fully sequenced
                BDGP DGC clones
                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
                 

                polyclonal

                (Bülow et al., 2010, Weigel et al., 1989)
                Commercially Available Antibodies
                 
                Other Information
                Relationship to Other Genes
                Source for database identify of
                Source for database merge of
                Additional comments
                Other Comments

                Under fed conditions, fkh has a negative effect on growth. Under conditions of starvation, reduction of fkh levels attenuates the size reduction associated with these conditions.

                (Bülow et al., 2010)

                fkh protects the salivary glands from hormone-induced death until a stage-specific, hormone-induced loss of the protein earmarks the tissue for destruction in response to future hormone exposure.

                (Cao et al., 2007)

                Downregulation of fkh at puparium formation is necessary for proper repression of Sgs4.

                (Renault et al., 2001)

                fkh has at least two roles in the formation of embryonic salivary glands; an early role in promoting survival of secretory cells and a later role in secretory cell invagination, specifically in the constriction of the apical surface membrane.

                (Myat and Andrew, 2000)

                byn, supported by fkh, mediates the early specification of the caudal visceral mesoderm along with zfh1.

                (Kusch and Reuter, 1999)

                The amino terminal of the Scr homeodomain is necessary for the specific activation of the fkh 37bp fkh250 element in vivo.

                (Ryoo and Mann, 1999)

                In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes.

                (Kliman and Eyre-Walker, 1998)

                Mutants are isolated in an EMS mutagenesis screen to identify zygotic mutations affecting germ cell migration at discrete points during embryogenesis: mutants exhibit gap pattern defects.

                (Moore et al., 1998)

                cad acts in hindgut development through fog, fkh and wg, but does not play a role in activating tll, hkb, byn and bowl which are also required for proper hindgut development. cad, fkh, byn and wg constitute a conserved constellation of genes that plays a required role in gastrulation and gut development.

                (Wu and Lengyel, 1998)

                hh, wg and dpp are required for the establishment of signaling centres that coordinate morphogenesis in the hindgut epithelium. Activation of these genes in the developing hindgut and foregut requires fkh.

                (Hoch and Pankratz, 1996)

                The distinction between pregland and preduct cells is made by the combination of two spatially separated negative regulatory steps: the Egfr signaling pathway represses fkh in the preduct cells and fkh represses duct specific genes in the pregland cells. trh is a duct-specific gene activator and is one of the targets of fkh repression.

                (Kuo et al., 1996)

                fkh protein strongly interacts with the proximal element of Sgs3 and it regulates Sgs3 tissue-specific expression. fkh is expressed in the appropriate tissue and is bound to many loci including the Sgs genes on polytene chromosomes.

                (Mach et al., 1996)

                fkh is required for stomodeal nervous system development.

                (Gonzalez-Gaitan and Jäckle, 1995)

                The organisation of the tail region of the embryo is documented from studies of cuticular markers enabling a more direct comparison between homologous structures on the embryo and larval cuticle.

                (Kuhn et al., 1995)

                In salivary gland development the activity of fkh prevents the expression of duct-specific cells, and in preduct cells the spi group signalling pathway prevents the expression of gland specific markers. fkh is repressed by spi group signalling, which is strongest in the most ventral cells of the epidermis.

                (Kuo et al., 1995)

                Identified by virtue of their products binding the Sgs4 regulatory region in a mobility shift assay.

                (Lehmann and Korge, 1995)

                fkh and cnc homeotic genes are required for the maintenance of segment polarity gene expression in the foregut. fkh is required for the continued expression of the oesopharyngeal domains of wg and hh expression past stage 10.

                (Mohler et al., 1995)

                Scr is required to activate fkh expression, which then acts to maintain its own expression.

                (Zhou and Beckendorf, 1995)

                trx gene product binds within an 8.4kb regulatory region that directs fkh expression in several embryonic tissues.

                (Kuzin et al., 1994)

                The limits of the mesoderm primordium do not depend on tll or fkh.

                (Reuter and Leptin, 1994)

                Co-crystal structure of the HNF-3/fork head DNA-recognition motif resembles histone H5.

                (Clark et al., 1993)

                The analogous expression patterns of fkh and rat hepatocyte nuclear factor 3, together with similar findings for rat hepatocyte nuclear factor 4 and Hnf4, suggest evolutionary relationships between a group of genes involved in gut formation in invertebrates and mammals.

                (Zhong et al., 1993)

                Seven genes isolated by low stringency hybridization to a probe containing coding sequences for fkh domain conserved in the rodent hepatocyte enriched nuclear transcription factor. All seven genes include a 110 amino acid conserved sequence, overall amino acid identity varies between 45% and 62%, and two are accounted for by the two slp genes slp1 and slp2. The remaining five are fd59A, fd64A, fd78E, fd96Ca and fd96Cb.

                (Hacker et al., 1992)

                fkh is highly homologous to the rat hepatocyte nuclear factor-3 gene family.

                (Lai et al., 1991)

                Zygotically active locus involved in the terminal developmental program in the embryo.

                (Strecker et al., 1991)

                The effect of fkh on hb and ftz expression has been studied.

                (Casanova, 1990)

                fkh mutants exhibit transformation of nonsegmental terminal regions into segmental derivatives, especially deletions of hindgut and Malpighian tubules.

                (Gaul and Jäckle, 1990)

                A sequence comparison between fkh and the rat HNF-3A gene suggests the existence of a new class of transcription factors that is conserved between Drosophila and mammals. The name "fork head" domain is suggested for the characteristic DNA binding motif.

                (Weigel and Jäckle, 1990)

                A number of cis regulatory elements within the genomic DNA of the fkh gene have been identified.

                (Weigel et al., 1990)

                The molecular identification, structure and expression pattern of the fkh gene product demonstrate that it is localized in the nucleus and may act as a transcriptional regulator.

                (Weigel et al., 1989)

                fkh is required in the very anterior and posterior ectodermal regions of the embryo. Inactive fkh causes homeotic transformations, the appearance of post-oral head structures in the terminal domains. There is no morphological evidence for the expression of fkh in the cells of the female germ line. Mitotic recombination demonstrates that fkh is not required in a cell autonomous manner for normal development of the imaginal discs.

                (Jurgens and Weigel, 1988)

                fkh mutants display a forked head skeleton and no anal pads.

                (Jurgens et al., 1984)

                Embryonic lethal. fkh+ appears to be required in the most anterior and posterior regions of the embryo; in amorphic mutants homeotic transformation effects the appearance of post-oral head structures in these terminal domains. The ectopic head structures are sometimes associated with thoracic structures anteriorly and anterior tail structures posteriorly. Thus fkh mutations produce transformations directed to the center involving structures normally found in different parasegments. Anteriorly, esophagus and proventriculus, derivatives of the ectodermal stomodaeum, are absent but not the pharynx or the hypopharyngeal organ, which resides on the anterior surface of the embryo; parts of the head skeleton are distorted; other parts apparently normal. Salivary glands absent. Posteriorly, anal pads and Malpighian tubules, derivatives of the ectodermal proctodaeum, are absent; replaced by anterior tail structures and post-oral head structures; supernumerary anal sensilla and dorsal hairs also present. Anterior and posterior fkh domains are beyond the regions controlled by ANTC and BXC, respectively; however ectopic expression of ANTC and BXC expression can occur in the fkh domains of fkh but not fkh+ embryos. No maternal effect. Homozygous fkh clones in adult cuticle completely normal in structure indicating no requirements for fkh+ in imaginal disk development. In situ hybridization reveals transcript in two terminal domains shortly before blastoderm cellularization occupying 5% embryonic length anteriorly and 15% posteriorly. fkh protein first detected in posterior domain at the end of syncytial blastoderm and the anterior domain at the beginning of cellularization. Expression continues in the derivatives of the ectodermal stomodaeum and proctodaeum throughout gastrulation. In addition fkh protein is detected in the midgut, the salivary glands, the central nervous system and the yolk cells.

                Origin and Etymology
                Discoverer
                Etymology
                Identification
                External Crossreferences and Linkouts ( 59 )
                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.
                DroID - A comprehensive database of gene and protein interactions.
                DRSC - Results frm RNAi screens
                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.
                KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.
                MIST (protein-protein) - An integrated Molecular Interaction Database
                Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                Synonyms and Secondary IDs (16)
                Reported As
                Symbol Synonym
                CG10002
                (Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009.3.31)
                Dm-FoxA
                (Heingård et al., 2019)
                Dmfkh
                (Mazet et al., 2003)
                FKH
                (Varma et al., 2014, Kolesnikov and Beckendorf, 2005)
                Fkh
                (Naidu et al., 2020, Babski et al., 2019, Gomez-Lamarca et al., 2018, Castellanos et al., 2013, Busser et al., 2012, Crombach et al., 2012, Alic et al., 2011, Nedelsky et al., 2010, Jattani et al., 2009, Noyes et al., 2008, Chandraratna et al., 2007, Thummel, 2007, Abrams et al., 2006, Jafar-Nejad and Bellen, 2004, Kusch et al., 2000)
                Sebp2
                (Lehmann and Korge, 1995)
                dFoxA
                (Kanao et al., 2010)
                fkh
                (Hung et al., 2020, Sun et al., 2020, Babski et al., 2019, Krmpot et al., 2019, Ray et al., 2019, Shokri et al., 2019, Yang et al., 2019, Campbell et al., 2018, Davie et al., 2018, Garner et al., 2018, Hernández de Madrid and Casanova, 2018, Lan et al., 2018, Rajpurohit et al., 2018, Rastogi et al., 2018, Segal et al., 2018, Bolukbasi et al., 2017, Chung et al., 2017, Karaiskos et al., 2017, Bielmeier et al., 2016, Hernández et al., 2015, Schertel et al., 2015, Sivanantharajah and Percival-Smith, 2015, Archbold et al., 2014, Aleksic et al., 2013, Burlin and Tillib, 2013, Combs and Eisen, 2013, Fox et al., 2013, Knowles and Biggin, 2013, Li and Gilmour, 2013, Merabet and Hudry, 2013, Rousso et al., 2013, Saunders et al., 2013, Spokony and White, 2013.5.10, Hudry et al., 2012, Nikulova et al., 2012, Papadopoulos et al., 2012, Fowlkes et al., 2011, Jungreis et al., 2011, Kaplan et al., 2011, Lelli et al., 2011, Maruyama et al., 2011, Weake et al., 2011, Bülow et al., 2010, Cook and Cook, 2010.5.7, Ismat et al., 2010, Joshi et al., 2010, Kazemian et al., 2010, Kong et al., 2010, Kong et al., 2010, Wheeler et al., 2009, Wolfstetter et al., 2009, Cook et al., 2008.7.11, Juven-Gershon et al., 2008, Liu and Lehmann, 2008, Maruyama et al., 2008, Miguel-Aliaga et al., 2008, Segal et al., 2008, Srinivasan et al., 2008, Wheeler et al., 2008, Xu et al., 2008, Cao et al., 2007, Christensen et al., 2007.10.29, de Velasco et al., 2007, Harris and Beckendorf, 2007, Joshi et al., 2007, Lechner et al., 2007, Lehmann et al., 2007, Ryakhovskiy and Tillib, 2007, Ryakhovskiy and Tillib, 2007, Sprecher et al., 2007, Thummel, 2007, Kobayashi et al., 2006, Moran and Jimenez, 2006, Noro et al., 2006, Wheeler et al., 2006, Brown and Feder, 2005, Burgler and Macdonald, 2005, Copley, 2005, Lee and Frasch, 2004, Wang et al., 2004, Williams et al., 2004, Gim et al., 2001, Myat et al., 2000, Sugimura et al., 2000, Lehmann and Korge, 1996)
                Name Synonyms
                FORK HEAD
                (Myat and Andrew, 1999)
                Fork Head
                (Cao et al., 2007, Thummel, 2007, Keung et al., 2004)
                Fork head
                (Castellanos et al., 2013, Abrams et al., 2006)
                Fork head, which is crucial for survival, is enriched in the MT transcriptome, and its down regulation in
                (Ojha and Tapadia, 2020)
                Forkhead
                (Chambers et al., 2017, Varma et al., 2014, Castellanos et al., 2013, Busser et al., 2012, Crombach et al., 2012, Alic et al., 2011, Chandraratna et al., 2007, Gim et al., 2001, Holland, 2000)
                Secretion enhancer-binding protein 2
                (Lehmann and Korge, 1995)
                fork head
                (Papadopoulos et al., 2019, Burlin and Tillib, 2013, Merabet and Hudry, 2013, Rousso et al., 2013, Biehs et al., 2010, Chung et al., 2009, Wolfstetter et al., 2009, Liu and Lehmann, 2008, Wheeler et al., 2008, Joshi et al., 2007, Ryakhovskiy and Tillib, 2007, Chan et al., 2005, Takiya et al., 2003)
                forkhead
                (Wieschaus and Nüsslein-Volhard, 2016, Hudry et al., 2012, Lelli et al., 2011, Joshi et al., 2010, Coiffier et al., 2008, Juven-Gershon et al., 2008, Joshi et al., 2007, Schwartz and Pirrotta, 2007, Moran and Jimenez, 2006, Noro et al., 2006, Papatsenko et al., 2006, Pearson et al., 2005, Lee and Frasch, 2004, Sugimura et al., 2000)
                Secondary FlyBase IDs
                • FBgn0015292
                Datasets (0)
                Study focus (0)
                Experimental Role
                Project
                Project Type
                Title
                References (326)