FlyBase Gene Report: Dmel\bel

General Information

Symbol

Dmel\bel

Species

D. melanogaster

Name

belle

Annotation Symbol

CG9748

Feature Type

protein_coding_gene

FlyBase ID

FBgn0263231

Gene Model Status

Current

Stock Availability

18 publicly available

Enzyme Name (EC)

RNA helicase (3.6.4.13)

Gene Summary

belle (bel) encodes a DEAD-box RNA helicase that functions both as positive and negative translation regulator. It is essential for embryo viability and fertility, and is required for the maintenance of male germ-line stem cells. [Date last reviewed: 2019-05-16] (FlyBase Gene Snapshot)

All Summaries

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

Also Known As

cap, ms(3)neo30, l(3)85Ac, L3, psg9

Key Links

Genomic Location

Cytogenetic map

85A5-85A5

Sequence location

3R:8,654,975..8,660,761 [-]

Recombination map

3-48

RefSeq locus

NT_033777 REGION:8654975..8660761

Sequence

Get Decorated FASTA

Genomic Maps

JBrowse

Other Genome Views

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

Function

Gene Ontology (GO) Annotations (23 terms)

Molecular Function (4 terms)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables RNA helicase activity

inferred from mutant phenotype

(Johnstone et al., 2005)

Terms Based on Predictions or Assertions (4 terms)

CV Term

Evidence

References

enables ATP binding

inferred from electronic annotation with InterPro:IPR011545

(InterPro Project Members, 2004-)

enables mRNA binding

inferred from biological aspect of ancestor with PANTHER:PTN002776867

(GO Reference Genome Project, 2011-)

enables nucleic acid binding

inferred from electronic annotation with InterPro:IPR011545

(InterPro Project Members, 2004-)

enables RNA helicase activity

inferred from sequence or structural similarity with SGD:S000005730

(Lasko, 2000)

inferred from biological aspect of ancestor with PANTHER:PTN002774595

(GO Reference Genome Project, 2011-)

inferred from electronic annotation with InterPro:IPR014014

(InterPro Project Members, 2004-)

Biological Process (12 terms)

Terms Based on Experimental Evidence (9 terms)

CV Term

Evidence

References

involved_in autophagic cell death

inferred from mutant phenotype

(Ihry and Bashirullah, 2014)

involved_in instar larval development

inferred from mutant phenotype

(Johnstone et al., 2005)

involved_in male germ-line stem cell population maintenance

inferred from mutant phenotype

(Kotov et al., 2016)

involved_in mitotic sister chromatid segregation

inferred from mutant phenotype

(Pek and Kai, 2011)

involved_in oogenesis

inferred from mutant phenotype

(Johnstone et al., 2005)

involved_in regulation of gene expression

inferred from mutant phenotype

(Poulton et al., 2011)

involved_in regulatory ncRNA-mediated post-transcriptional gene silencing

inferred from mutant phenotype

(Ulvila et al., 2006)

involved_in response to ecdysone

inferred from mutant phenotype

(Ihry and Bashirullah, 2014)

involved_in spermatogenesis

inferred from mutant phenotype

(Johnstone et al., 2005)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

involved_in cell differentiation

inferred from biological aspect of ancestor with PANTHER:PTN002774652

(GO Reference Genome Project, 2011-)

involved_in gamete generation

inferred from biological aspect of ancestor with PANTHER:PTN002774652

(GO Reference Genome Project, 2011-)

involved_in negative regulation of gene expression

inferred from biological aspect of ancestor with PANTHER:PTN002774722

(GO Reference Genome Project, 2011-)

Cellular Component (7 terms)

Terms Based on Experimental Evidence (5 terms)

CV Term

Evidence

References

located_in cytoplasm

inferred from direct assay

(Ambrus et al., 2007)

located_in cytosol

inferred from high throughput direct assay

(Lye et al., 2014)

located_in neuronal cell body

inferred from direct assay

(Rolls et al., 2007)

located_in P granule

inferred from direct assay

(Johnstone et al., 2005)

part_of RISC complex

inferred from direct assay

(Ulvila et al., 2006)

Terms Based on Predictions or Assertions (3 terms)

CV Term

Evidence

References

is_active_in mitochondrion

inferred from sequence or structural similarity with UniProtKB:O00571

(Gene Ontology Curators, 2002-)

is_active_in nucleus

inferred from biological aspect of ancestor with PANTHER:PTN002774649

(GO Reference Genome Project, 2011-)

is_active_in P granule

inferred from biological aspect of ancestor with PANTHER:PTN002774723

(GO Reference Genome Project, 2011-)

Gene Group (FlyBase)

DEAD-BOX RNA HELICASES

Protein Family (UniProt)

Belongs to the DEAD box helicase family. DDX3/DED1 subfamily. (Q9VHP0)

Protein Signatures (InterPro)

Catalytic Activity (EC/Rhea)

RNA helicase activity

ATP + H2O = ADP + phosphate + H(+) (3.6.4.13)

Summaries

Gene Snapshot

Gene Group (FlyBase)

DEAD-BOX RNA HELICASES -: DEAD-box RNA helicases belong to helicase superfamily 2 (SF2). They are characterized by the presence of nine conserved helicase motifs, including the DEAD sequence in motif II. (Adapted from PMID:21509200.)

Protein Function (UniProtKB)

ATP-dependent RNA helicase that is essential and required for cellular function, larval growth, and for male and female fertility. Also required for RNA interference (RNAi), double-stranded RNA induces potent and specific gene silencing, by acting downstream of dsRNA internalization. RNAi is mediated by the RNA-induced silencing complex (RISC), a sequence-specific, multicomponent nuclease that destroys or silences messenger RNAs homologous to the silencing trigger.

(UniProt, Q9VHP0)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

bel: belle (M.T. Fuller)

Recessive larval lethal. Homozygous null individuals hatch, but remain as first instar larvae until their death several days later.

Summary (Interactive Fly)

helicase that unwinds RNA - translational repressor - regulates expression of transgenes, essential for male germline stem cell maintenance and division - suppresses the effects of ectopic Wg signaling, regulates ecdysone-triggered transcriptional cascades - regulates the temporal pattern of Notch signaling in Drosophila follicle cells, regulates chromosome segregation

Interactive Fly

Gene Model and Products

Number of Transcripts

Number of Unique Polypeptides

Please see the JBrowse view of Dmel\bel 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 Q9VHP0)

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

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

Gene model reviewed during 5.47

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

bel-RA

FBtr0081888

NM_080522

3010

798

bel-RB

FBtr0334319

NM_001275450

3970

801

Additional Transcript Data and Comments

Reported size (kB)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

bel-PA

FBpp0081374

85.1

798

7.59

FBpp0306433

85.4

801

7.86

A0A0B4KGU4

NP_001262379

AGB95761

Polypeptides with Identical Sequences

None of the polypeptides share 100% sequence identity.

Additional Polypeptide Data and Comments

Reported size (kDa)

Comments

External Data

Crossreferences

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

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

UniProt

Mapped Features

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

External Data

Crossreferences

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

bel_1

Linkouts

Expression Data

Testis-specificity index

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

1.83

Transcript Expression

dissected tissue

Stage

Tissue/Position (including subcellular localization)

Reference

wandering third instar larval stage

embryonic/larval salivary gland

Comment: from -12 hr APF

(Ihry and Bashirullah, 2014)

prepupal stage P1 -- P4

embryonic/larval salivary gland

Comment: peak at P4

(Ihry and Bashirullah, 2014)

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

organism | ubiquitous

Comment: maternally deposited

(Fisher et al., 2012)

embryonic stage 4 -- 6

organism | ubiquitous

(Fisher et al., 2012)

primordial germ cell

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | ubiquitous

(Fisher et al., 2012)

primordial germ cell

(Fisher et al., 2012)

embryonic stage 9 -- 10

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 11 -- 12

organism | ubiquitous

(Fisher et al., 2012)

embryonic stage 13 -- 16

germline cell

(Fisher et al., 2012)

organism | ubiquitous

(Fisher et al., 2012)

Additional Descriptive Data

Expression increases after the prepupal (+10 hr APF) ecdysone pulse.

(Ihry and Bashirullah, 2014)

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

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

adult stage | female

ovary

(Pek and Kai, 2011)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in cytoplasm

inferred from direct assay

(Ambrus et al., 2007)

located_in cytosol

inferred from high throughput direct assay

(Lye et al., 2014)

located_in neuronal cell body

inferred from direct assay

(Rolls et al., 2007)

located_in P granule

inferred from direct assay

(Johnstone et al., 2005)

part_of RISC complex

inferred from direct assay

(Ulvila et al., 2006)

Expression Deduced from Reporters

Reporter: PBac{754.P.FSVS-0}bel^CPTI002755

Stage

Tissue/Position (including subcellular localization)

Reference

adult stage

medulla

(Knowles-Barley, 2011.8.24)

lamina

(Knowles-Barley, 2011.8.24)

adult brain cell body rind

(Knowles-Barley, 2011.8.24)

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\bel in JBrowse

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

View exonic expression by developmental stage for Dmel\bel

View exonic expression by tissue for Dmel\bel

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

FBgn0263231

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

45826/tissue=All

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

FBgn0263231

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0000171

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

FBgn0263231

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

CG9748

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 ( 44 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of bel

Transgenic constructs containing regulatory region of bel

Aberrations (Deficiencies and Duplications) ( 21 )

Inferred from experimentation ( 21 )

Gene not disrupted in

Df(3R)D6

(Bender et al., 1987)

Df(3R)D8

(Jones and Rawls, 1988)

Df(3R)dsx11

(Jones and Rawls, 1988)

Df(3R)p5

(Jones and Rawls, 1988)

Df(3R)V2

(Jones and Rawls, 1988)

Inferred from location ( 7 )

Variants

Variant Molecular Consequences

Alleles Representing Disease-Implicated Variants

Phenotypes

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

Lethality

Allele

lethal, with bel^S047110/bel^neo30

bel^{E460A.Tag:FLAG}

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

bel^GD1324

lethal, with Scer\GAL4^Act5C.PU

bel^GD1324

lethal, with Scer\GAL4^Bx-MS1096

bel^GD1324

lethal, with Scer\GAL4^GMR.PF, Scer\GAL4^ey.PH

bel^GD1324

lethal, with Scer\GAL4^pnr-MD237

bel^GD1324

lethal (with bel^L4740)

bel^S047110

lethal (with bel^neo30), with bel^{E460A.Tag:FLAG}

bel^S047110

lethal (with bel^S047110)

bel^L4740

lethal (with bel^S047110), with bel^{E460A.Tag:FLAG}

bel^neo30

lethal | pharate adult stage (with Df(3R)Exel6149)

bel^psg9

lethal | pupal stage (with Df(3R)Exel6149)

bel^psg9

lethal | recessive

lethal - all die before end of larval stage | recessive

lethal - all die during larval stage, with Scer\GAL4^Mef2.PR

bel^GD1324

viable

viable, with bel^S047110/bel^neo30

viable (with bel^neo30)

bel^S047110

viable (with bel^neo30), with bel^{D624A.Tag:FLAG}

bel^S047110

viable (with bel^neo30), with bel^{L51A.Tag:FLAG}

bel^S047110

viable (with bel^neo30), with bel^{Q595A.Tag:FLAG}

bel^S047110

viable (with bel^neo30), with bel^Tag:FLAG

bel^S047110

viable (with bel^neo30), with bel^{V321M.Tag:FLAG}

bel^S047110

viable (with bel^neo30), with bel^ΔC.Tag:FLAG

bel^S047110

viable (with bel^neo30), with bel^ΔN.Tag:FLAG

bel^S047110

viable (with bel^neo30), with bel^{ΔNC.Tag:FLAG}

bel^S047110

viable (with bel^S047110)

bel^neo30

viable (with bel^S047110), with bel^{D624A.Tag:FLAG}

bel^neo30

viable (with bel^S047110), with bel^{L51A.Tag:FLAG}

bel^neo30

viable (with bel^S047110), with bel^{Q595A.Tag:FLAG}

bel^neo30

viable (with bel^S047110), with bel^Tag:FLAG

bel^neo30

viable (with bel^S047110), with bel^{V321M.Tag:FLAG}

bel^neo30

viable (with bel^S047110), with bel^ΔC.Tag:FLAG

bel^neo30

viable (with bel^S047110), with bel^ΔN.Tag:FLAG

bel^neo30

viable (with bel^S047110), with bel^{ΔNC.Tag:FLAG}

bel^neo30

Sterility

Allele

female fertile

bel^neo30

female sterile

bel^neo30

female sterile (with bel⁶)

bel^neo30

female sterile (with bel^neo30)

bel⁶

fertile

bel^EY08943

male semi-sterile, with Scer\GAL4^nanos.PG

bel^GD1324

male sterile

bel^neo30

male sterile | recessive

Other Phenotypes

Allele

abnormal cytokinesis

bel^neo30

abnormal cytokinesis (with Df(3L)p¹⁶)

bel^neo30

abnormal cytokinesis (with Df(3L)p¹⁹)

bel^neo30

abnormal developmental rate | recessive

bel⁵

abnormal meiotic cell cycle

bel^neo30

abnormal meiotic cell cycle (with Df(3L)p¹⁶)

bel^neo30

abnormal meiotic cell cycle (with Df(3L)p¹⁹)

bel^neo30

abnormal mitotic cell cycle | somatic clone

bel⁶

abnormal size | adult stage

bel⁶

atrophy | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

atrophy | third instar larval stage (with bel⁶)

bel^neo30

atrophy | third instar larval stage (with bel^neo30)

bel⁶

decreased cell number | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

decreased cell number | adult stage (with bel⁶)

bel^neo30

decreased cell number | adult stage (with bel^EY08943)

bel^neo30

decreased cell number | adult stage (with bel^neo30)

decreased cell number | third instar larval stage (with bel⁶)

bel^neo30

decreased cell number | third instar larval stage (with bel^neo30)

bel⁶

increased cell death | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

increased cell death | adult stage (with bel⁶)

bel^neo30

increased cell death | adult stage (with bel^neo30)

bel⁶

increased cell number | adult stage, with Scer\GAL4^NP1624

bel^GD1324

increased cell number | adult stage (with bel⁶)

bel^neo30

increased cell number | adult stage (with bel^neo30)

bel⁶

increased cell size | adult stage (with bel^EY08943)

bel^neo30

increased cell size | adult stage (with bel^neo30)

bel^EY08943

increased cell size | third instar larval stage (with bel⁶)

bel^neo30

increased cell size | third instar larval stage (with bel^neo30)

bel⁶

some die during embryonic stage, with Dcr-2^UAS.cDa, Scer\GAL4^elav.PLu

bel^GD1324

some die during larval stage | recessive

Phenotype manifest in

Allele

cyst progenitor cell | adult stage | increased number (with bel⁶)

bel^neo30

cyst progenitor cell | adult stage | increased number (with bel^neo30)

bel⁶

dorsal appendage (with bel²)

bel^neo30

dorsal appendage (with bel⁶)

bel^neo30

dorsal appendage (with bel^neo30)

egg chorion

bel^neo30

egg chorion (with bel²)

bel^neo30

egg chorion (with bel⁶)

bel^neo30

egg chorion (with bel^neo30)

embryonic/larval salivary gland | prepupal stage (with Df(3R)Exel6149)

bel^psg9

embryonic/larval salivary gland | pupal stage (with Df(3R)Exel6149)

bel^psg9

eye disc morphogenetic furrow | somatic clone

bel^L4740

follicle cell | somatic clone

bel^S074407

germinal proliferation center hub | adult stage (with bel⁶)

bel^neo30

germinal proliferation center hub | adult stage (with bel^neo30)

bel⁶

male germline cell | adult stage

bel⁶

male germline cell | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

male germline cell | adult stage, with Scer\GAL4^NP1624

bel^GD1324

male germline cell | adult stage (with bel⁶)

bel^neo30

male germline cell | adult stage (with bel^EY08943)

bel^neo30

male germline cell | adult stage (with bel^neo30)

male germline cell | adult stage | increased number, with Scer\GAL4^NP1624

bel^GD1324

male germline stem cell | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

male germline stem cell | adult stage (with bel⁶)

bel^neo30

male germline stem cell | adult stage (with bel^EY08943)

bel^neo30

male germline stem cell | adult stage (with bel^neo30)

male germline stem cell | third instar larval stage (with bel⁶)

bel^neo30

male germline stem cell | third instar larval stage (with bel^neo30)

bel⁶

meiotic cell cycle

bel^neo30

Nebenkern

bel^neo30

Nebenkern (with Df(3L)p¹⁶)

bel^neo30

Nebenkern (with Df(3L)p¹⁹)

bel^neo30

onion stage spermatid & nucleus

bel^neo30

onion stage spermatid & nucleus (with Df(3L)p¹⁶)

bel^neo30

onion stage spermatid & nucleus (with Df(3L)p¹⁹)

bel^neo30

oocyte | germline clone

spermatid

bel^neo30

spermatocyte | adult stage (with bel⁶)

bel^neo30

spermatocyte | adult stage (with bel^neo30)

bel⁶

spermatocyte cell spectrosome | adult stage (with bel⁶)

bel^neo30

spermatocyte cell spectrosome | adult stage (with bel^neo30)

bel⁶

spermatocyte fusome, with Scer\GAL4^nanos.PG

bel^GD1324

spermatocyte fusome | adult stage, with Scer\GAL4^NP1624

bel^GD1324

spermatocyte fusome | adult stage (with bel⁶)

bel^neo30

spermatocyte fusome | adult stage (with bel^neo30)

bel⁶

spermatogonium | adult stage (with bel⁶)

bel^neo30

spermatogonium | adult stage (with bel^EY08943)

bel^neo30

spermatogonium | adult stage (with bel^neo30)

spermatogonium | third instar larval stage (with bel⁶)

bel^neo30

spermatogonium | third instar larval stage (with bel^neo30)

bel⁶

testis | adult stage

bel⁶

testis | adult stage, with Scer\GAL4^nanos.PG

bel^GD1324

testis | third instar larval stage (with bel⁶)

bel^neo30

testis | third instar larval stage (with bel^neo30)

bel⁶

Orthologs

Downloads

Download All DIOPT Orthologs

Human Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Homo sapiens (Human) (36)

Hsap\DDX3X

12 of 14

Yes

Hsap\DDX3Y

11 of 14

Yes

5 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (24)

Rnor\Ddx3x

12 of 14

Yes

Rnor\Ddx3y

12 of 14

Yes

Rnor\Ddx3

11 of 14

Yes

5 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Mus musculus (laboratory mouse) (26)

Mmus\Ddx3x

13 of 14

Yes

Mmus\Ddx3y

12 of 14

Yes

Mmus\D1Pas1

11 of 14

Yes

5 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Xenopus tropicalis (Western clawed frog) (14)

Xtro\ddx3x

7 of 13

Yes

3 of 13

2 of 13

2 of 13

2 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Danio rerio (Zebrafish) (20)

Drer\ddx3xa

12 of 14

Yes

Drer\ddx3xb

12 of 14

Yes

4 of 14

2 of 14

2 of 14

2 of 14

2 of 14

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Caenorhabditis elegans (Nematode, roundworm) (34)

Cele\laf-1

13 of 14

Yes

Cele\vbh-1

10 of 14

Yes

Cele\glh-1

5 of 14

Yes

Cele\glh-2

4 of 14

Yes

Cele\glh-3

4 of 14

Yes

Cele\mut-14

3 of 14

Yes

Cele\ddx-17

2 of 14

Cele\F55F8.2

2 of 14

Cele\glh-4

2 of 14

Yes

Cele\H20J04.4

2 of 14

Cele\rde-12

2 of 14

Yes

2 of 14

2 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Cele\smut-1

1 of 14

Yes

1 of 14

1 of 14

1 of 14

1 of 14

1 of 14

Yes

Anopheles gambiae (African malaria mosquito) (28)

Agam\AgaP_AGAP003047

10 of 12

Yes

5 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

2 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Arabidopsis thaliana (thale-cress) (54)

Atha\AT2G42520

13 of 13

Yes

Atha\AT3G58510

13 of 13

Yes

Atha\AT3G58570

13 of 13

Yes

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

Yes

Atha\emb1138

2 of 13

Atha\PDE340

2 of 13

Yes

2 of 13

2 of 13

2 of 13

2 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Yes

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Saccharomyces cerevisiae (Brewer's yeast) (22)

Scer\DED1

12 of 13

Yes

Scer\DBP1

11 of 13

Yes

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Schizosaccharomyces pombe (Fission yeast) (13)

Spom\sum3

11 of 12

Yes

2 of 12

2 of 12

2 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

1 of 12

Yes

Spom\SPBC691.04

1 of 12

Escherichia coli (enterobacterium) (3)

Ecol\rhlE

5 of 11

Yes

Ecol\deaD

2 of 11

Ecol\rhlB

2 of 11

Yes

Other Organism Orthologs (via OrthoDB)

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

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v9.1)

Gene Symbol

Score

Source

Alignment

Drosophila melanogaster (Fruit fly) (31)

vas

9 of 13

CG10077

5 of 13

Rm62

5 of 13

abs

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

3 of 13

ais

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

2 of 13

pit

2 of 13

Prp5

2 of 13

pths

2 of 13

Rs1

2 of 13

solo

2 of 13

Dbp73D

1 of 13

Ddx56

1 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

azoospermia, Sertoli-cell-only syndrome, DDX3-related

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 3 )

Allele

Disease

Evidence

References

bel^GD1324

model of Sertoli cell-only syndrome

CEA

(Kotov et al., 2016)

bel^neo30

model of Sertoli cell-only syndrome

CEA with bel⁶

(Kotov et al., 2016)

bel⁶

model of Sertoli cell-only syndrome

CEA with bel^neo30

(Kotov et al., 2016)

Potential Models Based on Orthology ( 0 )

Human Ortholog

Disease

Evidence

References

Modifiers Based on Experimental Evidence ( 10 )

Allele

Disease

Interaction

References

bel^S074407

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel^GL00057

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel^GL00205

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

exacerbates C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis

modeled by Zzzz\GGGGCC^58.UAS.EGFP

(Cheng et al., 2019)

ameliorates amyotrophic lateral sclerosis type 10

modeled by Hsap\TARDBP^{M337V.UAS.Tag:MYC}

(Markmiller et al., 2018)

ameliorates C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis

modeled by Zzzz\poly-GA^{80.UAS.Tag:FLAG}

(Markmiller et al., 2018)

bel^GD1324

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel^EY08943

exacerbates C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis

modeled by Zzzz\GGGGCC^58.UAS.EGFP

(Cheng et al., 2019)

bel^cap-1

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel⁶

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel⁵

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

bel^JF02884

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

exacerbates C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis

modeled by Zzzz\GGGGCC^58.UAS.EGFP

(Cheng et al., 2019)

ameliorates C9orf72 frontotemporal dementia and/or amyotrophic lateral sclerosis

modeled by Zzzz\poly-GA^{80.UAS.Tag:FLAG}

(Markmiller et al., 2018)

bel^L4740

ameliorates fragile X-associated tremor/ataxia syndrome

modeled by Zzzz\CGG^90.UAS.EGFP

(Linsalata et al., 2019)

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?

DDX3X; DEAD-box helicase 3 X-linked

12 of 14

300160

INTELLECTUAL DEVELOPMENTAL DISORDER, X-LINKED, SYNDROMIC, SNIJDERS BLOK TYPE; MRXSSB

DDX3Y; DEAD-box helicase 3 Y-linked

11 of 14

400010

DDX4; DEAD-box helicase 4

5 of 14

605281

Functional Complementation Data

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

Interactions

Summary of Physical Interactions

Interaction Browsers

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

Please see the Physical Interaction reports below for full details

RNA-protein

Physical Interaction

Assay

References

bel - Caf1-55

clip-seq

(Kotov et al., 2020)

bel - CycB

anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Liao et al., 2019)

bel - Eip74EF

anti tag coimmunoprecipitation, primer specific pcr

(Ihry et al., 2012)

bel - Fs(2)Ket

clip-seq

(Kotov et al., 2020)

bel - Hsp83

anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Götze et al., 2017)

bel - Not1

clip-seq

(Kotov et al., 2020)

bel - Rho1

clip-seq

(Kotov et al., 2020)

bel - Rpn7

clip-seq

(Kotov et al., 2020)

bel - br

anti tag coimmunoprecipitation, primer specific pcr

(Ihry et al., 2012)

bel - eIF4EHP

anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Yarunin et al., 2011)

bel - hpRNA:CR46342

anti tag coimmunoprecipitation, northern blot

(Zhou et al., 2008)

bel - mir-ban

anti tag coimmunoprecipitation, northern blot

(Zhou et al., 2008)

bel - nanos

pull down, Identification by mass spectrometry, western blot, anti tag coimmunoprecipitation, quantitative reverse transcription pcr

(Götze et al., 2017)

protein-protein

Physical Interaction

Assay

References

bel - AGO1

anti tag coimmunoprecipitation, western blot

(Zhou et al., 2008)

bel - AGO2

anti bait coimmunoprecipitation, western blot, anti tag coimmunoprecipitation

(Pek and Kai, 2011, Zhou et al., 2008)

bel - Cap-D2

anti bait coimmunoprecipitation, western blot

(Pek and Kai, 2011)

bel - Fmr1

anti tag coimmunoprecipitation, western blot

(Zhou et al., 2008)

bel - Hecw

pull down, Identification by mass spectrometry

(Fajner et al., 2021)

bel - Hrb27C

anti tag coimmunoprecipitation, anti tag western blot, Identification by mass spectrometry

(Bansal et al., 2020)

bel - SmD1

anti tag coimmunoprecipitation, western blot, anti tag western blot

(Xiong et al., 2013)

bel - Zn72D

anti tag coimmunoprecipitation, western blot, Identification by mass spectrometry

(Worringer et al., 2009)

bel - barr

anti bait coimmunoprecipitation, western blot

(Pek and Kai, 2011)

bel - cnn

anti bait coimmunoprecipitation, peptide massfingerprinting

(Habermann et al., 2012)

bel - cry

anti tag coimmunoprecipitation, Identification by mass spectrometry, western blot

(Cusumano et al., 2019)

bel - eIF4E1

pull down, western blot

(Yarunin et al., 2011)

bel - miso

anti tag coimmunoprecipitation, western blot, anti tag western blot

(Li et al., 2024)

bel - shep

anti tag coimmunoprecipitation, Identification by mass spectrometry

(Olesnicky et al., 2018)

bel - vas

anti tag coimmunoprecipitation, western blot, Identification by mass spectrometry

(Durdevic and Ephrussi, 2019)

bel - vig

anti tag coimmunoprecipitation, western blot

(Zhou et al., 2008)

Summary of Genetic Interactions

Interaction Browsers

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

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

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

bel

suppressible

CycB

(Kotov et al., 2016)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

E2f1

suppressible

bel

(Ambrus et al., 2009, Ambrus et al., 2007)

suppressible

bel

(Zhou et al., 2008)

External Data

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0263231

MIST (genetic) - An integrated Molecular Interaction Database

45826

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

45826

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

Neutrophil degranulation

Class of Gene

gene

gene_with_edited_transcript

(Graveley et al., 2011)

nuclear_gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

Recombination map

3-48

Cytogenetic map

85A5-85A5

Sequence location

3R:8,654,975..8,660,761 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

85A5-85A5

Left limit from in situ hybridisation (FBrf0067338) Right limit from in situ hybridisation (FBrf0067338); Limits computationally determined from genome sequence between P{EP}EP833 and P{lacW}l(3)L4740^L4740

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

85A-85A

(Naora, 2000.2.21)

85A4-85A5

(determined by in situ hybridisation)

(Spradling et al., 1999)

85A-85A

(determined by in situ hybridisation)

(Szeged Stock Center, 1998-)

85A5-85A6

(BDGP Project Members, 1994-1999)

Experimentally Determined Recombination Data

Location

3-48

(FlyBase, 2016)

3-44.2

(Sapiro et al., 2013)

3-45.4

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (18)

Genomic Clones (11)

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

cDNA Clones (56)

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

RE28061

Other clones

FI21109

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.

FBgn0263231

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Götze et al., 2017, Kotov et al., 2016, Pek and Kai, 2011)

monoclonal

(Na et al., 2016)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

bel is required autonomously in the male germline for germ-line stem cell maintenance.

(Kotov et al., 2016)

Identified as a potential component of the hh signalling pathway in a genome-wide RNAi screen. dsRNA made from templates generated with primers directed affects the extent of expression of a hh signaling reporter construct in Clone 8 cells.

(Nybakken et al., 2005)

Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days.

(Claridge-Chang et al., 2001)

Postmeiotic differentiation defect.

(Castrillon et al., 1993)

Sequence analysis suggests that the bel gene product is a member of a family of putative helicases.

(Deuring et al., 1989)

Recessive larval lethal. Homozygous null individuals hatch, but remain as first instar larvae until their death several days later.

Relationship to Other Genes

Source for database merge of

Source for merge of: cap CG9748

(Misra, 2000.8.9)

Source for merge of: bel cap

(Johnstone et al., 2001)

Source for merge of: bel anon-85Ab

(FlyBase, 1996-)

Source for merge of: bel l(3)S047110 l(3)L4740

(Poulton et al., 2011)

Additional comments

Source for merge of bel anon-85Ab was sequence comparison ( date:021125 ).

(FlyBase, 1996-)

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (24)

Reported As

Symbol Synonym

BEL

(Li et al., 2024, Rees et al., 2011, Johnstone et al., 2001)

BELLE

(Cusumano et al., 2019)

Bel

(Bansal et al., 2020, Xiong et al., 2013, Puri et al., 2008)

Belle

(Linsalata et al., 2019)

DDX3

(Fortuna et al., 2021)

DmRH6

(Boudet et al., 2001)

(Bender et al., 1987)

anon-85Ab

bel

(Saha et al., 2023, Dalton et al., 2022, Bilder et al., 2021, Ibaraki et al., 2021, Sureka and Mishra, 2021, Ellison et al., 2020, Kotov et al., 2020, Robinson et al., 2020, Taschuk et al., 2020, Cheng et al., 2019, Fic et al., 2019, Fingerhut et al., 2019, Markmiller et al., 2018, Hsu and Drummond-Barbosa, 2017, Brewer-Jensen et al., 2016, Gene Disruption Project members, 2016-, Jenny and Basler, 2016, Kotov et al., 2016, Lee et al., 2016, Lo et al., 2016, Na et al., 2016, Aradska et al., 2015, Kwong et al., 2015, Ashwal-Fluss et al., 2014, He et al., 2014, Moy et al., 2014, Kwon et al., 2013, Schertel et al., 2013, Thomae et al., 2013, Bradley et al., 2012, Butchar et al., 2012, Habermann et al., 2012, Miles et al., 2012, Friedman et al., 2011, Poulton et al., 2011, Yarunin et al., 2011, Ambrus and Frolov, 2010, Müller et al., 2010, Worringer et al., 2009, Poulton and Deng, 2008, Zhou et al., 2008, Akdemir et al., 2007, Ambrus et al., 2007, Buszczak et al., 2007, Dietzl et al., 2007, Link et al., 2007, Quinones-Coello, 2007, Rathke et al., 2007, Rolls et al., 2007, Syrzycka et al., 2007, Cermelli et al., 2006, Ulvila et al., 2006, Nybakken et al., 2005)

belle

(Xu et al., 2020, Doiguchi et al., 2016, Ambrus et al., 2009, Link et al., 2007)

cap

(Spradling et al., 1999, BDGP Project Members, 1994-1999, Castrillon et al., 1993)

cg9748

(Lasko, 2000)

l(3)85Ac

(Jones and Rawls, 1988)

l(3)L3

l(3)L4740

l(3)S047110

l(3)S097074

(Link et al., 2007)

ms(3)Pneo85A

ms(3)neo30

(Fuller, 1993)

psg9

(Ihry and Bashirullah, 2014, Sapiro et al., 2013)

Name Synonyms

Belle

(Linsalata et al., 2019, Jenny and Basler, 2016, Kibanov et al., 2012, Shimada et al., 2011, Worringer et al., 2009, Poulton and Deng, 2008, Ambrus et al., 2007, Nashchekin et al., 2006, Ulvila et al., 2006)

belle

(Fortuna et al., 2021, Kotov et al., 2020, Liao et al., 2019, Lo et al., 2016, Ihry and Bashirullah, 2014, Hoffmann et al., 2013, Ihry et al., 2012, Miles et al., 2012, Pek and Kai, 2011, Poulton et al., 2011, Yarunin et al., 2011, Ambrus and Frolov, 2010, Ulvila et al., 2010, Zhou et al., 2008, Link et al., 2007, Link et al., 2007, Rathke et al., 2007, Cermelli et al., 2006)

capon

lethal (3) L4740

lethal (3) S047110

Secondary FlyBase IDs

FBgn0000171
FBgn0002312
FBgn0011214
FBgn0037604
FBgn0041772
FBgn0028143
FBgn0024557

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 56 )

Sequence Crossreferences

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

45826

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.

Q9VHP0

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

Q9VHP0

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.

Q9VHP0

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0263231

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

45826/tissue=All

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

FBgn0263231

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

FBgn0263231

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

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

dme:Dmel_CG9748

MARRVEL_MODEL - MARRVEL (model organism gene)

45826

Linkouts

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

FBgn0263231

DroID - A comprehensive database of gene and protein interactions.

FBgn0263231

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

bel_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0000171

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0263231

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

CG9748

FlyMet - A comprehensive tissue-specific metabolomics resource for Drosophila.

FBgn0263231

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

TC013328

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

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

45826

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

45826

Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.

Neutrophil degranulation

References (168)

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

1.83

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