FB2024_03 , released June 25, 2024
Gene: Dmel\how
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General Information
Symbol
Dmel\how
Species
D. melanogaster
Name
held out wings
Annotation Symbol
CG10293
Feature Type
protein_coding_gene
FlyBase ID
FBgn0264491
Gene Model Status
Current
Stock Availability
22 publicly available
Gene Summary
held out wings (how) encodes an RNA-binding protein highly expressed in the mesoderm and tendon cells. It is spliced into two isoforms: how(L) and how(S). how(L) induces RNA destabilization, whereas How(S) stabilizes the RNA targets. [Date last reviewed: 2019-03-07] (FlyBase Gene Snapshot)
All Summaries
Also Known As

who, struthio, Held Out Wing, l(3)j5B5, sci

Key Links
Genomic Location
Cytogenetic map
94A1-94A1
Sequence location
Recombination map
3-74
RefSeq locus
NT_033777 REGION:22042191..22079321
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
Gene Ontology (GO) Annotations (27 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables mRNA 3'-UTR binding
inferred from direct assay
(Israeli et al., 2007, Toledano-Katchalski et al., 2007)
enables mRNA 5'-UTR binding
inferred from direct assay
(Graindorge et al., 2013)
enables mRNA binding
inferred from direct assay
(Volohonsky et al., 2007, Di Fruscio et al., 1998)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables mRNA binding
inferred from biological aspect of ancestor with PANTHER:PTN000129548
(GO Reference Genome Project, 2011-)
enables RNA binding
inferred from electronic annotation with InterPro:IPR004088, InterPro:IPR036612
(InterPro Project Members, 2004-)
Biological Process (21 terms)
Terms Based on Experimental Evidence (19 terms)
CV Term
Evidence
References
involved_in apposition of dorsal and ventral imaginal disc-derived wing surfaces
inferred from mutant phenotype
(Walsh and Brown, 1998)
involved_in axon ensheathment
inferred from mutant phenotype
(Edenfeld et al., 2006)
involved_in cell adhesion
inferred from mutant phenotype
(Lo and Frasch, 1997)
involved_in cell migration
inferred from mutant phenotype
(Baehrecke, 1997)
involved_in germ-line stem cell population maintenance
inferred from mutant phenotype
(Monk et al., 2010)
involved_in glial cell migration
inferred from mutant phenotype
(Edenfeld et al., 2006)
involved_in mesoderm development
inferred from expression pattern
(Furlong et al., 2001)
involved_in mesodermal cell migration
inferred from mutant phenotype
(Toledano-Katchalski et al., 2007)
involved_in muscle attachment
inferred from mutant phenotype
(Baehrecke, 1997)
involved_in muscle cell fate determination
inferred from direct assay
(Fyrberg et al., 1997)
involved_in muscle organ development
inferred from mutant phenotype
(Baehrecke, 1997, Zaffran et al., 1997)
involved_in negative regulation of RNA export from nucleus
inferred from direct assay
(Graindorge et al., 2013)
involved_in oenocyte development
inferred from mutant phenotype
(Edenfeld et al., 2006)
involved_in regulation of alternative mRNA splicing, via spliceosome
inferred from mutant phenotype
(Volohonsky et al., 2007)
inferred from high throughput mutant phenotype
(Park et al., 2004)
involved_in regulation of embryonic development
inferred from mutant phenotype
(Walsh and Brown, 1998)
involved_in regulation of heart contraction
inferred from mutant phenotype
(Zaffran et al., 1997)
involved_in sarcomere organization
inferred from mutant phenotype
(Schnorrer et al., 2010)
involved_in somatic muscle development
inferred from mutant phenotype
(Baehrecke, 1997)
involved_in spermatogonial cell division
inferred from mutant phenotype
(Monk et al., 2010)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
involved_in cell differentiation
traceable author statement
(Bokel and Brown, 2002)
involved_in regulation of mRNA splicing, via spliceosome
inferred from biological aspect of ancestor with PANTHER:PTN000129548
(GO Reference Genome Project, 2011-)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Rodrigues et al., 2012)
located_in nucleus
inferred from direct assay
(Rodrigues et al., 2012, Di Fruscio et al., 1998)
inferred from direct assay
(Zaffran et al., 1997)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000129548
(GO Reference Genome Project, 2011-)
located_in nucleus
inferred by curator from GO:0000381
(Park et al., 2004)
Gene Group (FlyBase)
Protein Family (UniProt)
-
Protein Signatures (InterPro)
Summaries
Gene Snapshot
held out wings (how) encodes an RNA-binding protein highly expressed in the mesoderm and tendon cells. It is spliced into two isoforms: how(L) and how(S). how(L) induces RNA destabilization, whereas How(S) stabilizes the RNA targets. [Date last reviewed: 2019-03-07]
Protein Function (UniProtKB)
RNA-binding protein involved in muscle development and dosage compensation (PubMed:23788626, PubMed:9118803, PubMed:9169854, PubMed:9344542). Vital role in steroid regulation of muscle development and to control heart rate (PubMed:9169854). Required during embryogenesis, in late stages of somatic muscle development, for myotube migration and during metamorphosis for muscle reorganization (PubMed:9118803). Required for integrin-mediated cell-adhesion in wing blade (PubMed:9344542). Together with Sxl, acts as an inhibitor of dosage compensation in females by preventing production of msl-2 protein, an essential component of the MSL complex (PubMed:23788626). Specifically binds to the 5'-UTR of msl-2 transcripts and cooperates with Sxl to promote nuclear retention of msl-2 mRNAs (PubMed:23788626).
(UniProt, O01367)
Summary (Interactive Fly)

KH motif - putative RNA-binding protein - functions in the migration and attachment of somatic muscle cells - targets Dpp mRNA - cell divisions in the embryonic mesoderm are repressed via posttranscriptional regulation of string/cdc25 by How - repression of the Drosophila midkine and pleiotrophin homolog miple by HOW is essential for correct mesoderm spreading

Gene Model and Products
Number of Transcripts
6
Number of Unique Polypeptides
5

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

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

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry O01367)

If you don't see a structure in the viewer, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

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

Experimentally Determined Structures
Crossreferences
Comments on Gene Model

Annotated transcripts do not represent all possible combinations of alternative exons and/or alternative promoters.

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

Gene model reviewed during 5.47

Sequence Ontology: Class of Gene
gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
how-RA
FBtr0084177
NM_079723
3964
405
how-RB
FBtr0084178
NM_169992
1759
375
how-RC
FBtr0100514
NM_001032032
2181
380
how-RD
FBtr0301401
NM_001170212
3884
405
how-RE
FBtr0301402
NM_001170213
3103
404
how-RF
FBtr0335195
NM_001275893
4003
418
Additional Transcript Data and Comments
Reported size (kB)

4.0 (northern blot)

(Baehrecke, 1997)

4.5, 4.0, 3.5 (northern blot)

(Fyrberg et al., 1997)

4.4, 3.6 (northern blot)

(Zaffran et al., 1997)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
how-PA
FBpp0083575
44.3
405
8.10
O01367
NP_524447
AAF55952
how-PB
FBpp0083576
40.9
375
6.96
NP_732695
AAN13901
how-PC
FBpp0099954
41.5
380
7.02
NP_001027203
AAZ52538
how-PD
FBpp0290615
44.3
405
8.10
O01367
NP_001163683
ACZ94979
how-PE
FBpp0290616
44.2
404
8.10
E1JIT0
NP_001163684
ACZ94980
how-PF
FBpp0307182
45.6
418
8.00
A0A0B4KHV5
NP_001262822
AGB96202
Polypeptides with Identical Sequences

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

405 aa isoforms: how-PA, how-PD
Additional Polypeptide Data and Comments
Reported size (kDa)

405 (aa)

(Baehrecke, 1997)

404 (aa)

(Fyrberg et al., 1997)

404 (aa); 44 (kD predicted)

(Zaffran et al., 1997)
Comments

how protein binds homopolymeric RNA (poly(A)) in vitro. A Myc-epitope-tagged how protein expressed in HeLa cells migrates with a molecular mass of about 55 kD.

(Di Fruscio et al., 1998)
External Data
Subunit Structure (UniProtKB)

Homodimer (PubMed:23788626). Interacts with Sxl; promoting nuclear retention of msl-2 transcripts (PubMed:23788626).

(UniProt, O01367)
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\how 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
Testis-specificity index

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

-1.32

Transcript Expression
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
mesoderm
(Fyrberg et al., 1997)
myoblast of embryonic/larval muscle system
(Baehrecke, 1997)
embryonic stage 1 -- 3
organism

Comment: maternally deposited

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

Comment: anlage in statu nascendi

(Fisher et al., 2012)
head mesoderm 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)
embryonic stage 6
mesoderm anlage
(Fyrberg et al., 1997, Zaffran et al., 1997)
ventral furrow
(Fyrberg et al., 1997)
embryonic stage 6 -- 17
mesoderm
(Baehrecke, 1997)
embryonic stage 7 -- 8
head mesoderm
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
embryonic stage 9 -- 10
head mesoderm
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
embryonic stage 10
mesoderm
(Toledano-Katchalski et al., 2007)
embryonic stage 11 -- 12
fat body primordium

Comment: reported as fat body specific anlage

(Fisher et al., 2012)
head mesoderm
(Fisher et al., 2012)
organism | faint | ubiquitous
(Fisher et al., 2012)
pericardial cell primordium

Comment: reported as pericardial cell specific anlage

(Fisher et al., 2012)
somatic muscle primordium
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
visceral muscle primordium
(Fisher et al., 2012)
embryonic stage 12
visceral mesoderm
(Baehrecke, 1997)
somatic mesoderm
(Baehrecke, 1997)
embryonic stage 13 -- 16
larval foregut atrium
(Fisher et al., 2012)
dorsal pharyngeal muscle cell
(Fisher et al., 2012)
embryonic dorsal apodeme
(Fisher et al., 2012)
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic head epidermis
(Fisher et al., 2012)
embryonic ventral apodeme
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval muscle system
(Fisher et al., 2012)
embryonic/larval visceral muscle cell
(Fisher et al., 2012)
embryonic stage 14
pharyngeal muscle cell
(Baehrecke, 1997)
embryonic stage 14 -- 17
embryonic/larval heart
(Baehrecke, 1997)
muscle attachment site
(Baehrecke, 1997)
embryonic stage 15 -- 17
embryonic/larval somatic muscle cell
(Zaffran et al., 1997)
late embryonic stage
embryonic/larval visceral muscle cell
(Fyrberg et al., 1997)
embryonic/larval somatic muscle cell
(Fyrberg et al., 1997)
pharyngeal muscle cell
(Fyrberg et al., 1997)
embryonic/larval dorsal vessel
(Fyrberg et al., 1997)
pupal stage
muscle attachment site
(Baehrecke, 1997)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Fyrberg et al., 1997)
embryonic stage & pupal stage
(Fyrberg et al., 1997)
embryonic stage -- adult stage
(Fyrberg et al., 1997, Zaffran et al., 1997)
embryonic stage 1 -- 4
organism
(Zaffran et al., 1997)
embryonic stage 6 -- 7
ventral furrow
(Zaffran et al., 1997)
embryonic stage 12
mesoderm
(Zaffran et al., 1997)
embryonic stage 14 -- 17
muscle attachment site
(Zaffran et al., 1997)
embryonic stage 15 -- 17
embryonic/larval dorsal vessel
(Zaffran et al., 1997)
embryonic/larval aorta
(Zaffran et al., 1997)
pharyngeal muscle cell
(Zaffran et al., 1997)
embryonic/larval visceral muscle cell
(Zaffran et al., 1997)
prepupal stage P1
imaginal disc
(Fortier et al., 2006)
gut
(Fortier et al., 2006)
fat body
(Fortier et al., 2006)
salivary gland
(Fortier et al., 2006)
adult stage | female
(Zaffran et al., 1997)
Additional Descriptive Data

how is expressed in the embryonic mesoderm at the time of mesoderm spreading (stages 7-9). Only the "long" isoform is detected at this time.

(Toledano-Katchalski et al., 2007)

how transcripts are detected in imaginal disc, gut, fat body and salivary gland at 0hr pupae on northern blots and reduced levels are seen in the gut and salivary glands at the 12 hour prepupal stage. Transcripts are not observed in the nervous system.

(Fortier et al., 2006)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 6 -- 7
cephalic furrow
(Zaffran et al., 1997)
ventral furrow
(Zaffran et al., 1997)
mesoderm anlage
(Zaffran et al., 1997)
head mesoderm
(Zaffran et al., 1997)
embryonic stage 8 -- 17
mesoderm
(Zaffran et al., 1997)
embryonic stage 14
embryonic/larval hindgut muscle cell
(Zaffran et al., 1997)
cardioblast
(Zaffran et al., 1997)
embryonic stage 14 -- 17
embryonic/larval visceral muscle cell
(Zaffran et al., 1997)
muscle attachment site
(Zaffran et al., 1997)
pharyngeal muscle cell
(Zaffran et al., 1997)
embryonic/larval somatic muscle cell
(Zaffran et al., 1997)
embryonic stage 15 -- 17
embryonic/larval aorta
(Zaffran et al., 1997)
embryonic/larval dorsal vessel
(Zaffran et al., 1997)
larval stage
imaginal disc
(Zaffran et al., 1997)
visceral muscle fiber of embryonic/larval muscle system
(Zaffran et al., 1997)
adepithelial cell
(Zaffran et al., 1997)
adult stage
tendon cell
(Zaffran et al., 1997)
muscle attachment site
(Zaffran et al., 1997)
adult stage & oogenesis
stalk follicle cell
(Zaffran et al., 1997)
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 7 -- 9
mesoderm

Comment: long isoform

(Toledano-Katchalski et al., 2007)
embryonic stage 16
midline glial cell
(Reuveny et al., 2009)
PNS glial cell
(Reuveny et al., 2009)
embryonic/larval CNS perineurial glial cell
(Reuveny et al., 2009)
embryonic/larval CNS subperineurial glial cell
(Reuveny et al., 2009)
cardioblast
(Harpaz et al., 2013)
prepupal stage P1
imaginal disc
(Fortier et al., 2006)
gut
(Fortier et al., 2006)
fat body
(Fortier et al., 2006)
salivary gland
(Fortier et al., 2006)
prepupal stage P1 -- P4
central nervous system
(Fortier et al., 2006)
adult stage | male
male germline stem cell
(Monk et al., 2010)
primary gonial cell
(Monk et al., 2010)
spermatogonial cyst
(Monk et al., 2010)
hub cell
(Monk et al., 2010)
male germline cyst
(Monk et al., 2010)
oogenesis
female germline stem cell
(Monk et al., 2011)
cystoblast
(Monk et al., 2011)
germarium cap cell
(Monk et al., 2011)
terminal filament
(Monk et al., 2011)
stalk follicle cell
(Monk et al., 2011)
mass spectroscopy
Stage
Tissue/Position (including subcellular localization)
Reference
adult stage
adult head
• membrane
(Aradska et al., 2015)
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 6 -- 10
organism

Comment: long isoform

(Toledano-Katchalski et al., 2007)
embryonic stage 16
organism

Comment: long and short isoforms

(Toledano-Katchalski et al., 2007)
Additional Descriptive Data

how protein is expressed in female germline stem cells and in the cystoblast. how expression is downregulated by the 2-cell stage in the female germarium. In the male, how is detected in 2 cell cysts. how expression is complementary to bam::GFP . How protein is also detected in some somatic cells of the ovary including the terminal filament cells, cap cells, and interfollicle cells.

(Monk et al., 2011)

how protein expression was detected in vas-expressing GSCs, their immediate daughter cells (gonialblasts), and 2-cell spermatogonial cysts. Expression is downregulated at the 4-cell stage. how protein expression is also detected in somatic hub cells and in the nuclei of mature cyst cells but not in somatic stem cells.

(Monk et al., 2010)

A GFPFBtr0091350:pb-XRhow fusion protein localizes to the nucleus of mouse NIH 3T3 and Drosophila S2 cells.

(Reuveny et al., 2009, Di Fruscio et al., 1998)

how is expressed in the embryonic mesoderm at the time of mesoderm spreading (stages 7-9). Only the "long" isoform is detected at this time while long and short isoforms are detected at later embryonic stages.

(Toledano-Katchalski et al., 2007)

how protein is detected by immunolocalization in dissected tissues in imaginal disc, gut, fat body and salivary gland at 0hr pupae. It is also seen in the CNS.

(Fortier et al., 2006)

how protein is first detected in the cells of the ventral furrow and the staining extends beyond the cephalic furrow. In the early stages of gastrulation, staining is mostly in the ventral furrow but is also seen in head mesoderm and around the cephalic furrow. Later, expression becomes restricted to myogenic cells. Expression is observed in precursors of the somatic muscles, pharyngeal muscles, visceral muscles and cardiac cells. Expression is also seen in muscle attachment sites. In larvae, expression is observed in the nuclei of muscle fibers and in adepithelial cells of the imaginal discs. In adults, expression occurs in the tendon cells that attach the flight muscle fibers to the dorsal cuticle of the thorax. During oogenesis, expression is observed in follicular stalk cells that join the different egg chambers in the ovary.

(Zaffran et al., 1997)
Marker for
 
midline glial cell
(Harpaz et al., 2013)
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Rodrigues et al., 2012)
located_in nucleus
inferred from direct assay
(Rodrigues et al., 2012, Di Fruscio et al., 1998)
inferred from direct assay
(Zaffran et al., 1997)
Expression Deduced from Reporters
Reporter: P{GawB}how24B
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10 -- 17
mesoderm
(Brand, 1997.6.30)
embryonic stage 11 -- 17
mesoderm
(Luo et al., 1994)
third instar larval stage
haltere disc
(Brand, 1997.6.30)
wing disc
(Brand, 1997.6.30)
imaginal disc
(Brand, 1997.6.30)
leg disc
(Brand, 1997.6.30)
eye-antennal disc
(Brand, 1997.6.30)
late embryonic stage
somatic muscle cell
(Staehling-Hampton et al., 1994)
embryonic stage
embryonic/larval somatic muscle cell
(Schuster et al., 1996)
mesoderm
(DeFalco et al., 2003, Brand and Perrimon, 1993)
embryonic/larval hypodermal muscle cell | subset
(Rodrigues et al., 2012)
embryonic/larval glial cell | subset
(Rodrigues et al., 2012)
embryonic/larval CNS subperineurial glial cell
(Rodrigues et al., 2012)
mesoderm anlage
(Kim et al., 2007)
embryonic somatic muscle cell
(Kim et al., 2007)
visceral mesoderm
(Kim et al., 2007)
embryonic stage 6
mesoderm anlage
(Zaffran et al., 1997)
embryonic stage 10 -- 17
mesoderm
(Marquez et al., 2001)
embryonic stage 11
mesoderm
(Allen et al., 2002, Brand and Perrimon, 1993)
embryonic stage 12
mesoderm
(Lee and Frasch, 2000, Zaffran et al., 1997)
embryonic stage 13 -- 16
mesoderm
(Jenkins et al., 2003)
embryonic stage 13 -- 17
embryonic/larval muscle system
(Brand and Perrimon, 1993)
embryonic stage 14 -- 17
muscle attachment site
(Zaffran et al., 1997)
embryonic stage 15 -- 17
embryonic/larval somatic muscle cell
(Zaffran et al., 1997)
pharyngeal muscle cell
(Zaffran et al., 1997)
embryonic/larval dorsal vessel
(Zaffran et al., 1997)
embryonic/larval visceral muscle cell
(Zaffran et al., 1997)
embryonic/larval aorta
(Zaffran et al., 1997)
larval stage
embryonic/larval somatic muscle cell
(Schuster et al., 1996)
embryonic/larval glial cell | subset
(Rodrigues et al., 2012)
embryonic/larval neuron
(Rodrigues et al., 2012)
embryonic/larval central nervous system | restricted
(Suster et al., 2003)
symmetrical commissure | subset of embryonic/larval brain | restricted
(Suster et al., 2003)
first instar larval stage -- second instar larval stage
tendon cell
(Subramanian et al., 2003)
muscle cell
(Subramanian et al., 2003)
third instar larval stage
visceral muscle fiber

Comment: Ascribed to all muscle fibers.

(Nitabach et al., 2006)
embryonic/larval somatic muscle cell
(van der Plas et al., 2006)
wandering third instar larval stage
circular visceral muscle fiber
(Jiang and Edgar, 2009)
embryonic/larval midgut longitudinal visceral muscle cell
(Jiang and Edgar, 2009)
P-stage
embryonic/larval dorsal vessel
(Monier et al., 2005)
adult dorsal vessel
(Monier et al., 2005)
adult stage
adult somatic muscle cell
(Wu et al., 2010)
adult visceral muscle cell
(Xu et al., 2011, Buchon et al., 2010)
adult heart
(Qian et al., 2008)
adult midgut
(Ferveur et al., 1997)
crop
(Ferveur et al., 1997)
adult Malpighian tubule
(Ferveur et al., 1997)
adult fat body
(Ferveur et al., 1997)
Reporter: P{lacW}B1-93F
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
larval ventral nerve cord | medial
(Hartenstein and Jan, 1992)
embryonic/larval midgut | restricted
(Hartenstein and Jan, 1992)
embryonic stage 6
mesoderm anlage
(Zaffran et al., 1997)
embryonic stage 11 -- 17
cardioblast
(Hartenstein and Jan, 1992)
embryonic/larval pericardial cell
(Hartenstein and Jan, 1992)
visceral mesoderm of embryonic/larval digestive system
(Hartenstein and Jan, 1992)
embryonic/larval hypodermal muscle cell | presumptive
(Hartenstein and Jan, 1992)
embryonic stage 12
mesoderm
(Zaffran et al., 1997)
embryonic stage 14 -- 17
muscle attachment site
(Zaffran et al., 1997)
embryonic stage 15 -- 17
visceral muscle cell
(Zaffran et al., 1997)
pharyngeal muscle cell
(Zaffran et al., 1997)
embryonic/larval aorta
(Zaffran et al., 1997)
somatic muscle cell
(Zaffran et al., 1997)
embryonic/larval dorsal vessel
(Zaffran et al., 1997)
oogenesis
stalk follicle cell
(Grammont and Irvine, 2001)
Reporter: P{lArB}how1A122
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
mesoderm
(Perrimon et al., 1991)
embryonic stage 4 -- 17
mesoderm
(Perrimon et al., 1991)
embryonic stage 6
mesoderm anlage
(Zaffran et al., 1997)
embryonic stage 12
mesoderm
(Zaffran et al., 1997)
embryonic stage 14 -- 17
muscle attachment site
(Zaffran et al., 1997)
embryonic stage 15 -- 17
pharyngeal muscle cell
(Zaffran et al., 1997)
embryonic/larval somatic muscle cell
(Zaffran et al., 1997)
embryonic/larval visceral muscle cell
(Zaffran et al., 1997)
embryonic/larval dorsal vessel
(Zaffran et al., 1997)
embryonic/larval aorta
(Zaffran et al., 1997)
embryonic stage 17
embryonic/larval dorsal vessel
(Perrimon et al., 1991)
embryonic/larval muscle system
(Perrimon et al., 1991)
adult stage
muscle attachment site
(Zaffran et al., 1997)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\how in JBrowse
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
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Images
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 44 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 13 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of how
Transgenic constructs containing regulatory region of how
Aberrations (Deficiencies and Duplications) ( 11 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal, with Scer\GAL4Mef2.PR
lethal, with Scer\GAL4pnr-MD237
lethal (with Df(3R)e-N19)
lethal | female limited, with Scer\GAL4dome-PG14
lethal | recessive
lethal - all die before end of larval stage | recessive
lethal - all die before end of pupal stage (with howr4)
lethal - all die before end of pupal stage (with howr17)
lethal - all die before end of pupal stage | recessive
lethal - all die during embryonic stage | recessive
lethal - all die during larval stage, with Scer\GAL4Mef2.PR
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4tin.CΔ4
Sterility
Allele
female sterile, with Scer\GAL4VP16.otu
fertile
Other Phenotypes
Allele
abnormal cell cycle | germline clone
abnormal flight | recessive | somatic clone
abnormal mitotic cell cycle | germline clone | pre-blastoderm stage
flightless
increased cell number, with Scer\GAL4VP16.nanos.UTR
increased occurrence of cell division | embryonic stage
some die during embryonic stage | recessive
some die during larval stage | recessive
some die during pupal stage (with howr4)
some die during pupal stage (with howr17)
some die during pupal stage | recessive
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4nub-AC-62
visible, with Scer\GAL4sd.PU
visible, with Scer\GAL4sd-SG29.1
visible | recessive
visible | recessive | somatic clone
visible | somatic clone
Phenotype manifest in
Allele
anterior wing margin, with Scer\GAL4sd.PU
cardial cell, with Scer\GAL4twi.PG
embryo | dorsal closure stage
embryonic/first instar larval cuticle | dorsal
embryonic/first instar larval cuticle | posterior
embryonic/larval dorsal vessel | embryonic stage
embryonic/larval oenocyte
embryonic/larval somatic muscle cell
embryonic pericardial cell, with Scer\GAL4twi.PG
embryonic somatic muscle cell, with Scer\GAL4twi.PG
germline cyst, with Scer\GAL4VP16.nanos.UTR
germline cyst | germline clone (with howr4)
germline cyst | germline clone (with howr17)
head
male germline cell | germline clone
male germline stem cell, with Scer\GAL4VP16.nanos.UTR
male germline stem cell | germline clone (with howr4)
male germline stem cell | germline clone (with howr17)
mesoderm
perineurial glial cell
PNS glial cell
sarcomere, with Scer\GAL4Mef2.PR
spermatocyte cyst | germline clone
spermatocyte fusome | germline clone
spermatogonial cyst, with Scer\GAL4VP16.nanos.UTR
spermatogonial cyst | germline clone
spermatogonium, with Scer\GAL4VP16.nanos.UTR
tendon cell
testis, with Scer\GAL4VP16.nanos.UTR
testis | germline clone (with howr4)
testis | germline clone (with howr17)
wing
wing, with Scer\GAL4nub-AC-62
wing, with Scer\GAL4sd.PU
wing, with Scer\GAL4sd-SG29.1
wing | somatic clone
wing vein | ectopic, with Scer\GAL4Bx-MS1096
Z disc, with Scer\GAL4Mef2.PR
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (7)
Hsap\QKI
11 of 14
Yes
Yes
Hsap\KHDRBS1
2 of 14
No
No
Hsap\KHDRBS2
2 of 14
No
No
Hsap\KHDRBS3
2 of 14
No
No
Hsap\SF1
2 of 14
No
No
Hsap\DYNLT2
1 of 14
No
No
Hsap\FUBP3
1 of 14
No
No
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (5)
Rnor\Qk
10 of 14
Yes
Yes
Rnor\Khdrbs1
2 of 14
No
No
Rnor\Khdrbs2
2 of 14
No
No
Rnor\Khdrbs3
2 of 14
No
No
Rnor\Sf1
2 of 14
No
No
Mus musculus (laboratory mouse) (6)
Mmus\Qk
9 of 14
Yes
Yes
Mmus\Khdrbs1
2 of 14
No
No
Mmus\Khdrbs2
2 of 14
No
No
Mmus\Khdrbs3
2 of 14
No
No
Mmus\Sf1
2 of 14
No
No
Mmus\Ms4a14
1 of 14
No
No
Xenopus tropicalis (Western clawed frog) (5)
Xtro\qki
6 of 13
Yes
Yes
Xtro\sf1
2 of 13
No
No
Xtro\khdrbs2
1 of 13
No
No
Xtro\khdrbs3
1 of 13
No
No
Xtro\serhl2
1 of 13
No
No
Danio rerio (Zebrafish) (10)
Drer\qkia
12 of 14
Yes
Yes
Drer\qki2
10 of 14
No
Yes
Drer\qkib
8 of 14
No
Yes
Drer\khdrbs1a
2 of 14
No
No
Drer\khdrbs1b
2 of 14
No
No
Drer\khdrbs2
2 of 14
No
No
Drer\sf1
2 of 14
No
No
Drer\khdrbs3
1 of 14
No
No
Drer\LOC110440170
1 of 14
No
Yes
Drer\map7d2b
1 of 14
No
No
Caenorhabditis elegans (Nematode, roundworm) (12)
Cele\asd-2
11 of 14
Yes
Yes
Cele\gld-1
8 of 14
No
Yes
Cele\K07H8.9
5 of 14
No
Yes
Cele\Y57G11C.36
4 of 14
No
Yes
Cele\B0280.17
3 of 14
No
Yes
Cele\F54D1.1
3 of 14
No
Yes
Cele\Y69A2AR.32
3 of 14
No
Yes
Cele\sfa-1
2 of 14
No
No
Cele\fubl-3
1 of 14
No
No
Cele\fubl-4
1 of 14
No
No
Cele\vamp-7
1 of 14
No
No
Cele\ZK899.6
1 of 14
No
Yes
Anopheles gambiae (African malaria mosquito) (5)
Agam\AgaP_AGAP008975
7 of 12
Yes
Yes
Agam\AgaP_AGAP003356
3 of 12
No
No
Agam\AgaP_AGAP000049
2 of 12
No
No
Agam\AgaP_AGAP003355
2 of 12
No
No
Agam\AgaP_AGAP012103
1 of 12
No
No
Arabidopsis thaliana (thale-cress) (7)
Atha\AT2G38610
9 of 13
Yes
Yes
Atha\AT3G08620
9 of 13
Yes
Yes
Atha\AT4G26480
9 of 13
Yes
Yes
Atha\AT5G56140
9 of 13
Yes
Yes
Atha\AT1G09660
8 of 13
No
Yes
Atha\AT3G32940
2 of 13
No
No
Atha\AT5G51300
2 of 13
No
No
Saccharomyces cerevisiae (Brewer's yeast) (1)
Scer\MSL5
3 of 13
Yes
No
Schizosaccharomyces pombe (Fission yeast) (1)
Spom\bpb1
2 of 12
Yes
No
Escherichia coli (enterobacterium) (0)
Other Organism Orthologs (via OrthoDB)
Data provided directly from OrthoDB:how. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (9)
CG3927
6 of 13
CG4021
6 of 13
nsr
6 of 13
qkr54B
6 of 13
qkr58E-1
6 of 13
qkr58E-2
6 of 13
qkr58E-3
6 of 13
CG10384
5 of 13
SF1
5 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    how18
    ameliorates  oculopharyngeal muscular dystrophy
    modeled by Hsap\PABPN117ala.Act88F
    (Chartier et al., 2015)
    Disease Associations of Human Orthologs (via DIOPT v9.1 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    QKI; QKI, KH domain containing RNA binding
    11 of 14
    609590
        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:
        Show/hide secondary interactors 
        (data from AllianceMine provided by esyN)
        Select Layout:
        Legend:
        Protein
        RNA
        Selected Interactor(s)
        Other Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please see the Physical Interaction reports below for full details
        RNA-protein
        Physical Interaction
        Assay
        References
        how - CG31638
        pull down, anti tag western blot
        (Toledano-Katchalski et al., 2007)
        how - Grasp65
        anti tag coimmunoprecipitation, molecular weight estimation by staining, anti bait coimmunoprecipitation, primer specific pcr
        (Giuliani et al., 2014)
        how - Hrb98DE
        anti bait coimmunoprecipitation, reverse transcription pcr
        (Ji and Tulin, 2016)
        how - Nrx-IV
        anti tag coimmunoprecipitation, molecular weight estimation by staining, anti bait coimmunoprecipitation, primer specific pcr, pull down, anti tag western blot
        (Giuliani et al., 2014, Edenfeld et al., 2006)
        how - bam
        anti bait coimmunoprecipitation, quantitative reverse transcription pcr
        (Monk et al., 2010)
        how - dpp
        pull down, anti tag western blot
        (Israeli et al., 2007)
        how - fal
        pull down, anti tag western blot
        (Toledano-Katchalski et al., 2007)
        how - lap
        pull down, anti tag western blot
        (Toledano-Katchalski et al., 2007)
        how - miple1
        pull down, anti tag western blot
        (Toledano-Katchalski et al., 2007)
        how - msl-2
        pull down, western blot, peptide massfingerprinting
        (Graindorge et al., 2013)
        how - sr
        pull down, western blot, anti tag western blot
        (Nir et al., 2012, Israeli et al., 2007, Volohonsky et al., 2007, Nabel-Rosen et al., 2005, Nabel-Rosen et al., 1999)
        how - stg
        anti bait coimmunoprecipitation, quantitative reverse transcription pcr, pull down, anti tag western blot
        (Monk et al., 2010, Nabel-Rosen et al., 2005)
        protein-protein
        Physical Interaction
        Assay
        References
        how
        anti tag coimmunoprecipitation, anti tag western blot
        (Nir et al., 2012)
        how - Hrb27C
        anti bait coimmunoprecipitation, western blot
        (Szostak et al., 2018)
        how - Sxl
        anti bait coimmunoprecipitation, western blot, pull down, peptide massfingerprinting, molecular weight estimation by staining
        (Graindorge et al., 2013)
        how - crn
        anti tag coimmunoprecipitation, western blot
        (Edenfeld et al., 2006)
        Summary of Genetic Interactions
        esyN Network Diagram
        Show/hide secondary interactors 
        (data from AllianceMine provided by esyN)
        esyN Network Key:
        Suppression
        Enhancement
        Other Interaction Browsers
        View in FlyBase Interactions Browser   View in MIST tool (external link)
        Please look at the allele data for full details of the genetic interactions
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        how
        suppressible
        crn
        (Edenfeld et al., 2006)
        how
        enhanceable
        crn
        (Edenfeld et al., 2006)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        bam
        suppressible
        how
        (Monk et al., 2010)
        bam
        enhanceable
        how
        (Monk et al., 2010)
        ftz
        suppressible
        how
        (Kankel et al., 2004)
        ftz
        enhanceable
        how
        (Kankel et al., 2004)
        if
        enhanceable
        how
        (Walsh and Brown, 1998)
        External Data
        Subunit Structure (UniProtKB)
        Homodimer (PubMed:23788626). Interacts with Sxl; promoting nuclear retention of msl-2 transcripts (PubMed:23788626).
        (UniProt, O01367 )
        Linkouts
        DroID - A comprehensive database of gene and protein interactions.
        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)
        3R
        Recombination map
        3-74
        Cytogenetic map
        94A1-94A1
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        94A1-94A1
        Limits computationally determined from genome sequence between P{PZ}tsl00617 and P{lacW}howj5B5&P{EP}EP738
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        93F-93F
        (Baehrecke, 1997.1.16, Fyrberg, 1997.1.24, Zaffran et al., 1997, Zaffran, 1996.9.24)
        94A1-94A2
        (determined by in situ hybridisation)
        (Spradling et al., 1999)
        87F3-87F10
        (determined by in situ hybridisation)
        (Szeged Stock Center, 1998-, Deak et al., 1997)
        93F-93F
        (determined by in situ hybridisation)
        (Fyrberg et al., 1997, Zaffran et al., 1995)
        93F13-93F13
        (determined by in situ hybridisation)
        (Baehrecke, 1997)
        94A1-94A2
        (BDGP Project Members, 1994-1999)
        Experimentally Determined Recombination Data
        Location

        3-74

        (FlyBase, 2016)

        3-76.2

        (Comeron et al., 2012)

        3-

        (Fyrberg, 1997.1.24, Prout et al., 1997, Zaffran, 1996.9.24)
        Left of (cM)
        Right of (cM)
        Notes

        Located on 3R.

        (Prout et al., 1997)
        Stocks and Reagents
        Stocks (22)
        Genomic Clones (33)
        cDNA Clones (256)
         

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

        cDNA clones, fully sequenced
        BDGP DGC clones
        Other clones
          Drosophila Genomics Resource Center cDNA clones

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

          cDNA Clones, End Sequenced (ESTs)
          BDGP DGC clones
          Other clones
          RNAi and Array Information
          Linkouts
          Antibody Information
          Laboratory Generated Antibodies
           

          polyclonal

          (Graindorge et al., 2013, Nir et al., 2012, Zaffran et al., 1997)
          Commercially Available Antibodies
           
          Cell Line Information
          Other Comments

          The balance between two distinct forms of how protein maintains low levels of sr protein at the precursor stage in future tendon cells and high levels in the mature tendon.

          (Nabel-Rosen et al., 1999)

          Identification: Genetic screen for mutations that produce blisters in somatic wing clones.

          (Walsh and Brown, 1998)

          how encodes a member of the KH RNA binding protein family that functions during muscle development.

          (Baehrecke, 1997)

          Cloning and partial characterisation of how, defined by the Scer\GAL4 24B transposant. Four cDNAs (2-9, 4-2, D-11 and D-5) that contain a KH domain are isolated and sequence compared to how : a family of proteins having closely related KH domains is discovered.

          (Fyrberg et al., 1997)

          how appears to regulate some aspects of integrin functioning.

          (Lo and Frasch, 1997)

          Identification: Genetic screen for autosomal mutations that produce blisters in somatic wing clones. 1 allele of how has been isolated.

          (Prout et al., 1997)

          how function is required late in embryogenesis to control heart rate and muscular activity.

          (Zaffran et al., 1997)
          Relationship to Other Genes
          Source for database merge of

          Source for merge of: how anon- EST:Liang-2.39

          (FlyBase Genome Annotators, 2005)

          Source for merge of: how l(3)j5B5

          (Spradling et al., 1999)

          Source for merge of: how sci

          (Brown et al., 2000)

          Source for merge of: how l(3)S090417 l(3)S053606

          (Link et al., 2007)
          Additional comments
          Nomenclature History
          Source for database identify of

          Source for identity of: how CG10293

          (Zheng, 1999.11)
          Nomenclature comments
          Etymology
          Synonyms and Secondary IDs (34)
          Reported As
          Symbol Synonym
          0904/17
          (Deak et al., 1997)
          24B
          (Johnson et al., 2011, Taylor, 2006)
          CG10293
          (Haye et al., 2014, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Quinones-Coello, 2007, Whitfield, 2000.4.28)
          HOW
          (Ko and Martin, 2020, Szostak et al., 2018, König and Shcherbata, 2013, Lim et al., 2012, Patel et al., 2012, Monk et al., 2010, Reuveny et al., 2009, Elhanany and Volk, 2008, Reuveny and Volk, 2007, Sofola et al., 2007, Volk and Reuveny, 2007, Nabel-Rosen et al., 2004)
          How
          (Berry et al., 2022, Dubey et al., 2020, Ji and Tulin, 2016, Dutta et al., 2015, Ryu et al., 2011, Hime et al., 2007, Nabel-Rosen and Volk, 2003)
          KH93F
          P62
          (Mushegian et al., 1998)
          SZ1
          (Zaffran et al., 1995)
          Who/How
          (Di Fruscio et al., 1999, Di Fruscio et al., 1998)
          anon-EST:Liang-2.39
          (Ashburner, 2004.6.30)
          clone 2.39
          (Liang and Biggin, 1998)
          how
          (Burghardt et al., 2023, Kao et al., 2021, Aboukilila et al., 2020, Overton et al., 2020, Aw et al., 2017, Bonneaud et al., 2017, Nie et al., 2017, Zhang et al., 2016, Chartier et al., 2015, Duff et al., 2015, Hsu et al., 2015, Sîrbu et al., 2015, Haye et al., 2014, Graindorge et al., 2013, Harpaz et al., 2013, Kwon et al., 2013, Japanese National Institute of Genetics, 2012.5.21, Nir et al., 2012, Rodrigues et al., 2012, Kazemian et al., 2011, Monk et al., 2011, Park et al., 2011, Xu et al., 2011, Bulchand et al., 2010, Carmon et al., 2010, Schnorrer et al., 2010, Christensen et al., 2009.5.6, Christensen et al., 2009.5.6, Dworkin et al., 2009, Hazelett et al., 2009, Reuveny et al., 2009, Schulze et al., 2009, Christensen et al., 2008.6.11, Medioni et al., 2008, Assa-Kunik et al., 2007, Beltran et al., 2007, Buszczak et al., 2007, Dietzl et al., 2007, Georlette et al., 2007, Israeli et al., 2007, Junion et al., 2007, Link et al., 2007, Minidorff et al., 2007, Quinones-Coello, 2007, Reuveny and Volk, 2007, Sofola et al., 2007, Toledano-Katchalski et al., 2007, Volohonsky et al., 2007, Zhang et al., 2007, Edenfeld et al., 2006, Fortier et al., 2006, Mack et al., 2006, Sandmann et al., 2006, Nabel-Rosen et al., 2002, Gim et al., 2001)
          l(3)S053606
          l(3)S090417
          l(3)j5B5
          (Meister and Braun, 1995.10, BDGP Project Members, 1994-1999)
          l(3)j5D5
          l(3)s2612
          qkr
          (Fyrberg et al., 1997)
          qkr93F
          (Fyrberg et al., 1997, Vernet and Artzt, 1997)
          sci
          (Walsh and Brown, 1998)
          stru
          (Lo and Frasch, 1997)
          who
          (Volk and Rosen, 1999, Fyrberg et al., 1998, Baehrecke, 1997, Baehrecke, 1997.1.16, Sweeney et al., 1997, Vernet and Artzt, 1997)
          Name Synonyms
          Held Out Wing
          (Ko and Martin, 2020, Reuveny et al., 2009, Reuveny and Volk, 2007)
          Held out wings
          (König and Shcherbata, 2013, Gim et al., 2001)
          Held-out wings
          (Monk et al., 2010)
          KH motif at 93F
          held out wings
          (Carmon et al., 2010, Schulze et al., 2009, Medioni et al., 2008, Link et al., 2007, Fortier et al., 2006, Mack et al., 2006, Burnette et al., 2005, Zaffran et al., 1997)
          lethal (3) S053606
          lethal (3) S090417
          quaking related, 93F
          (Fyrberg et al., 1997)
          scorpion
          (Walsh and Brown, 1998)
          struthio
          (Carmon et al., 2010, MacIntyre et al., 2007, Lo, 1997.5.7, Lo, 1997.5.7, Prout et al., 1997)
          wings held out
          wings held-out
          (Baehrecke, 1997)
          Secondary FlyBase IDs
          • FBgn0028068
          • FBgn0082187
          • FBgn0017397
          • FBgn0011379
          • FBgn0011385
          • FBgn0015249
          • FBgn0020779
          • FBgn0025731
          • FBgn0025956
          Datasets (0)
          Study focus (0)
          Experimental Role
          Project
          Project Type
          Title
          Study result (0)
          Result
          Result Type
          Title
          External Crossreferences and Linkouts ( 85 )
          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/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
          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
          AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
          BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
          DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
          EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
          FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
          InterPro - A database of protein families, domains and functional sites
          KEGG Genes - Molecular building blocks of life in the genomic space.
          MARRVEL_MODEL - MARRVEL (model organism gene)
          Linkouts
          Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
          DroID - A comprehensive database of gene and protein interactions.
          Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
          FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
          FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
          Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
          iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
          Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
          MIST (genetic) - An integrated Molecular Interaction Database
          MIST (protein-protein) - An integrated Molecular Interaction Database
          References (185)