FlyBase Gene Report: Dmel\shrb

FB2025_05 , released December 11, 2025

FB2025_05 , released December 11, 2025

Gene: Dmel\shrb

General Information

Symbol

Dmel\shrb

Species

D. melanogaster

Name

shrub

Annotation Symbol

CG8055

Feature Type

protein_coding_gene

FlyBase ID

FBgn0086656

Gene Model Status

Stock Availability

17 publicly available

Gene Summary

shrub (shrb) encodes Shrub encodes Vps32/Snf7, a filament-forming subunit of the ESCRT-III complex involved in inward membrane budding. It regulates multiple cellular processes involving ESCRTs activity including multivesicular body biogenesis, cytokinesis, membrane repair and regulation of membrane associated signaling. [Date last reviewed: 2019-03-14] (FlyBase Gene Snapshot)

All Summaries

Gene Snapshot
Alliance
Auto summary
Gene Group
UniProtKB
Interactive Fly

Also Known As

Vps32, ESCRT-III, vps32/snf7, ESCRT, Snf7

Key Links

Genomic Location

Cytogenetic map

Sequence location

2R:9,142,146..9,143,914 [-]

Recombination map

2-60

RefSeq locus

NT_033778 REGION:9142146..9143914

Sequence

Get Decorated FASTA

Genomic Maps

Other Genome Views

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

Function

Gene Ontology (GO) Annotations (22 terms)

Molecular Function (1 term)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

enables protein binding

inferred from physical interaction with FLYBASE:borr; FB:FBgn0032105

(Capalbo et al., 2012)

Terms Based on Predictions or Assertions (0 terms)

Biological Process (12 terms)

Terms Based on Experimental Evidence (8 terms)

CV Term

Evidence

References

involved_in autophagy

inferred from mutant phenotype

(Rusten et al., 2007)

involved_in dendrite morphogenesis

inferred from mutant phenotype

(Gao et al., 1999)

involved_in endosome transport via multivesicular body sorting pathway

inferred from mutant phenotype

(Hori et al., 2011, Vaccari et al., 2009)

inferred from mutant phenotype

(Baeumers et al., 2020)

involved_in female germ-line stem cell asymmetric division

inferred from mutant phenotype

(Eikenes et al., 2015, Matias et al., 2015)

involved_in mitotic cytokinesis

inferred from mutant phenotype

(Eikenes et al., 2015, Matias et al., 2015)

involved_in neuron remodeling

inferred from mutant phenotype

(Loncle et al., 2015, Issman-Zecharya and Schuldiner, 2014)

involved_in symbiont entry into host cell

inferred from high throughput mutant phenotype

(Hodgson et al., 2019)

involved_in ubiquitin-dependent protein catabolic process via the multivesicular body sorting pathway

inferred from mutant phenotype

(Vaccari et al., 2009)

Terms Based on Predictions or Assertions (4 terms)

CV Term

Evidence

References

involved_in late endosome to vacuole transport via multivesicular body sorting pathway

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

involved_in nuclear membrane reassembly

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

involved_in vacuolar transport

inferred from electronic annotation with InterPro:IPR005024

(InterPro Project Members, 2004-)

involved_in vesicle budding from membrane

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

Cellular Component (9 terms)

Terms Based on Experimental Evidence (6 terms)

CV Term

Evidence

References

located_in contractile ring

inferred from direct assay

(Eikenes et al., 2015)

located_in fusome

inferred from direct assay

(Eikenes et al., 2015, Matias et al., 2015)

located_in midbody

inferred from direct assay

(Eikenes et al., 2015, Matias et al., 2015, Capalbo et al., 2012)

located_in multivesicular body

inferred from direct assay

(Hori et al., 2011)

located_in neuronal cell body

inferred from direct assay

(Loncle et al., 2015)

located_in proximal dendrite

inferred from direct assay

(Loncle et al., 2015)

Terms Based on Predictions or Assertions (4 terms)

CV Term

Evidence

References

is_active_in cytoplasmic side of plasma membrane

inferred from biological aspect of ancestor with PANTHER:PTN000509454

(GO Reference Genome Project, 2011-)

part_of ESCRT III complex

inferred from sequence or structural similarity with SGD:S000004015

(Capalbo et al., 2012, Vaccari et al., 2009)

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

is_active_in multivesicular body

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

is_active_in nuclear envelope

inferred from biological aspect of ancestor with PANTHER:PTN000509386

(GO Reference Genome Project, 2011-)

Gene Group (FlyBase)

ESCRT-III COMPLEX

Protein Family (UniProt)

Belongs to the SNF7 family. (Q8T0Q4)

Protein Signatures (InterPro)

Snf7 family

Summaries

Gene Snapshot

shrub (shrb) encodes Shrub encodes Vps32/Snf7, a filament-forming subunit of the ESCRT-III complex involved in inward membrane budding. It regulates multiple cellular processes involving ESCRTs activity including multivesicular body biogenesis, cytokinesis, membrane repair and regulation of membrane associated signaling. [Date last reviewed: 2019-03-14]

Gene Group (FlyBase)

ESCRT-III COMPLEX -: The ESCRT (Endosomal Sorting Complexes Required for Transport) machinery is made up of five complexes, ESCRT-0 to -III and the Vps4-Vta1 ATPase complex, which are sequentially recruited to the cytosolic face of the membrane, and mediate cargo recognition, membrane budding and fission. ESCRT-III is a dynamic complex, that assembles on endosomes. The components of the complex are less well defined, but it has four core subunits. It assembles into a filament around the neck of the budding vesicle. ESCRT-III mediates the recruitment of deubiquitinases and, together with the Vps4-Vta1 complex, which it recruits, mediates membrane scission. (Adapted from PMID:22361144 and FBrf0209668).

Protein Function (UniProtKB)

Probable core polymerisation component of the endosomal sorting required for transport (ESCRT) III complex involved in multiple cellular processes requiring the outward bending of membranes, including vesicle budding, membrane repair and cytokinesis (Probable). The ESCRT pathway involves 4 complexes (ESCRT-0, -I, -II and -III) that sequentially assemble on the cytoplasmic side of membranes and induce membrane remodeling, budding and scission. As part of the ESCRT-III complex, involved in the budding of intraluminal vesicles (ILVs) into endosomes to form multivesicular bodies (MVBs), which target their contents for degradation via the endolysosomal pathway (PubMed:16713958). Involved in regulation of signal transduction pathways, including the Notch and BMP/decapentaplegic (dpp) pathways, by sequestering the intracellular domains of activated receptors into ILVs, isolating them from the cytoplasm and targeting them for lysosomal degradation (PubMed:22162134, PubMed:25804739). Involved in targeting ubiquitilated proteins, such as mono-ubiquitilanated N/Notch, to MVBs for degradation (PubMed:22162134, PubMed:33660341). Plays a role in wing development by regulating Notch signaling (PubMed:22162134). Involved in abscission of germline cells during oogenesis (PubMed:25647097). Involved in spermiogenesis (PubMed:33660341). Required for efficient cytoplasmic isolation and abscission during cytokinesis of epithelial sensory organ precursor cells (PubMed:37226981). May be involved in septate junction remodeling and maintenance (PubMed:37226981). ACTIVITY REGULATION: May be regulated by aurB/Aurora kinase B-dependent phosphorylation.

(UniProt, Q8T0Q4)

Summary (Interactive Fly)

vacuolar sorting protein - required for Endosomal Sorting Complexes Required for Transport (ESCRT) machinery - ESCRT III component - regulation of dendritic branching - completion of membrane abscission during female germline stem cell division

Interactive Fly

Gene Model and Products

Number of Transcripts

1

Number of Unique Polypeptides

1

Please see the JBrowse view of Dmel\shrb 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 Q8T0Q4)

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

PDB - An information portal to biological macromolecular structures

5J45
5VO5

Comments on Gene Model

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

Gene model reviewed during 5.49

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

FBtr0088623

1267

226

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

FBpp0087704

25.4

226

4.42

Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments

Reported size (kDa)

Comments

External Data

Subunit Structure (UniProtKB)

Homopolymer; forms elongated striated filaments of uniform 10nm width (PubMed:27452459). Monomers interact in a staggered arrangement mediated by complementary charged electrostatic surfaces (PubMed:27452459). Interacts with l(2)gd1 (via DM14 domains 1 and 3); the interaction is direct and blocks access to the surface involved in homopolymerization (PubMed:22389409, PubMed:27452459, PubMed:28564595). This interaction may be required for the ESCRT-III complex role in multivesicular body formation (PubMed:22389409).

(UniProt, Q8T0Q4)

Crossreferences

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

Snf7 family

PDB - An information portal to biological macromolecular structures

5J45
5VO5

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\shrb using the Feature Mapper tool.

External Data

Crossreferences

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

shrb_1

Linkouts

Expression Data

Testis-specificity index

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

-0.91

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 4 -- 6

(Fisher et al., 2012)

embryonic stage 7 -- 8

organism | ubiquitous

(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

organism | ubiquitous

(Fisher et al., 2012)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

mass spectroscopy

Stage

Tissue/Position (including subcellular localization)

Reference

• membrane

(Aradska et al., 2015)

day 7 of adulthood -- day 49 of adulthood

(Cammarato et al., 2011)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Evidence

References

located_in contractile ring

inferred from direct assay

(Eikenes et al., 2015)

located_in fusome

inferred from direct assay

(Eikenes et al., 2015, Matias et al., 2015)

located_in midbody

inferred from direct assay

(Eikenes et al., 2015, Matias et al., 2015, Capalbo et al., 2012)

located_in multivesicular body

inferred from direct assay

(Hori et al., 2011)

located_in neuronal cell body

inferred from direct assay

(Loncle et al., 2015)

located_in proximal dendrite

inferred from direct assay

(Loncle et al., 2015)

Expression Deduced from Reporters

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\shrb in JBrowse

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

View exonic expression by developmental stage for Dmel\shrb

View exonic expression by tissue for Dmel\shrb

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

FBgn0086656

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

35933/tissue=All

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

FBgn0086656

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0086656

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

FBgn0086656

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

CG8055

Flygut - An atlas of the Drosophila adult midgut

FBgn0086656

Images

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 9 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 35 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of shrb

Transgenic constructs containing regulatory region of shrb

Aberrations (Deficiencies and Duplications) ( 3 )

Inferred from experimentation ( 3 )

Gene disrupted in

(Kahsai and Cook, 2018, Kahsai et al., 2016)

(Sweeney et al., 2006)

(Sweeney et al., 2006)

Inferred from location ( 5 )

Variants

Variant Molecular Consequences

Alleles Representing Disease-Implicated Variants

Phenotypes

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

Lethality

Allele

cell lethal | somatic clone | third instar larval stage

shrb⁴

lethal (with Df(2R)BSC279)

shrb^k11201

lethal | recessive

shrb^k11201

lethal - all die before end of first instar larval stage, with shrb⁴

shrb^{cDNA.LAP.EGFP}

lethal - all die before end of first instar larval stage, with shrb^{cDNA.LAP.EGFP}

shrb⁴

lethal - all die before end of larval stage, with shrb⁴

shrb^{cDNA.3xGly.EGFP}

lethal - all die before end of larval stage, with shrb^{cDNA.3xGly.EGFP}

shrb⁴

lethal - all die before end of P-stage, with Scer\GAL4^dome-PG14

shrb^KK108557

lethal - all die before end of second instar larval stage, with shrb⁴

lethal - all die before end of second instar larval stage, with shrb^D79K

shrb⁴

lethal - all die before end of second instar larval stage, with shrb^E40K

shrb⁴

lethal - all die before end of second instar larval stage, with shrb^E86K

shrb⁴

lethal - all die during embryonic stage

shrb⁴

partially lethal - majority die | paternal effect, with Scer\GAL4^{VP16.nanos.UTR}

shrb^EY05194

Sterility

Allele

shrb^EY05194

fertile, with shrb⁴

fertile, with shrb^{+tCH322-47O20}

shrb⁴

fertile, with shrb^{cDNA.N.Tag:MYC}

shrb⁴

fertile, with shrb^R59E.Tag:MYC

shrb⁴

fertile, with shrb^R70E.Tag:MYC

shrb⁴

fertile, with shrb^WT.cDNA

shrb⁴

Other Phenotypes

Allele

abnormal cell number | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

abnormal cytokinesis, with Scer\GAL4^nanos.PG

shrb^HMS01767

abnormal cytokinesis | dominant

abnormal immune response | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

abnormal meiotic cell cycle | oogenesis

shrb^GFP.KI

abnormal mitotic cell cycle | P-stage

shrb^GFP.KI

abnormal mitotic cell cycle | P-stage, with Scer\GAL4^sca.PU

shrb^HMS01767

abnormal neuroanatomy, with Scer\GAL4²²¹

abnormal neuroanatomy, with Scer\GAL4^Tab2-201Y

shrb^UAS.GFP

abnormal neuroanatomy | cell autonomous | pupal stage | somatic clone

shrb⁴

abnormal neuroanatomy | embryonic stage

shrb⁴

abnormal neuroanatomy | P-stage, with Scer\GAL4^ppk.1.9

shrb^UAS.GFP

abnormal neuroanatomy | P-stage, with Scer\GAL4^ppk.PU

shrb^UAS.GFP

abnormal neuroanatomy | P-stage | somatic clone

shrb^G5

abnormal neuroanatomy | pupal stage, with Scer\GAL4^ppk.1.9

shrb^RNAi.UAS

abnormal neuroanatomy | pupal stage, with Scer\GAL4^smid-C161

shrb^RNAi.UAS

abnormal neuroanatomy | pupal stage, with Scer\GAL4^Tab2-201Y

shrb^RNAi.UAS

increased cell death, with Scer\GAL4^GMR.PF

shrb^UAS.GFP

increased cell number | larval stage | somatic clone

shrb^G5

increased cell number | third instar larval stage, with Scer\GAL4^{E(spl)m6-BFM.PF}

shrb^KK108557

increased occurrence of cell division | cell non-autonomous | third instar larval stage

shrb^G5

increased occurrence of cell division | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

increased size | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

majority die during embryonic stage | paternal effect, with Scer\GAL4^{VP16.nanos.UTR}

short lived, with Scer\GAL4^MESO.dome, Scer\GAL80^elav.PU

shrb^KK108557

some die during embryonic stage

shrb⁴

Phenotype manifest in

Allele

abdominal dorsal multidendritic neuron | P-stage | somatic clone

shrb^G5

abdominal lateral adult muscle precursor cell | third instar larval stage, with Scer\GAL4^{E(spl)m6-BFM.PF}

shrb^KK108557

adult mushroom body gamma-lobe | pupal stage, with Scer\GAL4^Tab2-201Y

shrb^RNAi.UAS

axon | pupal stage, with Scer\GAL4^Tab2-201Y

shrb^RNAi.UAS

cystic bulge | decreased number, with Scer\GAL4^{VP16.nanos.UTR}

dendrite | cell autonomous | pupal stage | somatic clone

shrb⁴

dendrite | dorsal | embryonic stage

shrb⁴

dendrite | P-stage, with Scer\GAL4^ppk.1.9

shrb^UAS.GFP

dendrite | pupal stage, with Scer\GAL4^ppk.1.9

shrb^RNAi.UAS

dendrite | pupal stage, with Scer\GAL4^smid-C161

shrb^RNAi.UAS

dendritic arborizing neuron | embryonic stage

shrb⁴

dendritic arborizing neuron | embryonic stage (with Df(2R)Np5)

shrb⁴

dendritic arborizing neuron | embryonic stage (with In(2LR)w45-32n)

shrb⁴

dendritic tree | P-stage, with Scer\GAL4^ppk.PU

shrb^UAS.GFP

dendritic tree | P-stage | somatic clone

shrb^G5

dorsal multidendritic neuron ddaE & dendritic tree | ectopic, with Scer\GAL4²²¹

shrb^RNAi.UAS

dorsal multidendritic neuron ddaF & dendritic tree | ectopic, with Scer\GAL4²²¹

shrb^RNAi.UAS

egg chamber, with Scer\GAL4^nanos.PG

egg chamber, with Scer\GAL4^nanos.PG, Scer\GAL80^bam.PM

shrb^HMS01767

embryonic/larval crystal cell | increased number | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

embryonic/larval plasmatocyte | increased number | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

endosome | embryonic stage

shrb⁴

endosome | P-stage | somatic clone

shrb^G5

endosome | somatic clone | third instar larval stage

shrb^O3

external sensory organ precursor cell | P-stage

shrb^GFP.KI

external sensory organ precursor cell | P-stage, with Scer\GAL4^sca.PU

shrb^HMS01767

extracellular vesicle | decreased number | larval stage, with Scer\GAL4^VGlut1-OK371

shrb^HMS01767

extracellular vesicle | decreased number | larval stage, with Spyo\Cas9^VGlut1-OK371

shrb^U6.gRNAx2

extracellular vesicle | decreased number | larval stage, with Spyo\Cas9^wor.PC

shrb^U6.gRNAx2

eye, with Scer\GAL4^GMR.PF

shrb^UAS.GFP

eye disc | cell non-autonomous | somatic clone

shrb^G5

eye disc | somatic clone | third instar larval stage

shrb^O3

female germline cell

female germline cell, with Scer\GAL4^nanos.PG

female germline cell, with Scer\GAL4^nanos.PG, Scer\GAL80^bam.PM

shrb^HMS01767

female germline cyst

shrb^O3

female germline stem cell

female germline stem cell, with Scer\GAL4^nanos.PG

shrb^HMS01767

female germline stem cell | germline clone

shrb^G5

gamma Kenyon cell, with Scer\GAL4^Tab2-201Y

shrb^UAS.GFP

shrb^O3

intracellular vesicle | larval stage, with Scer\GAL4^VGlut1-OK371

shrb^HMS01767

intracellular vesicle | larval stage, with Spyo\Cas9^VGlut1-OK371

shrb^U6.gRNAx2

intracellular vesicle | larval stage, with Spyo\Cas9^wor.PC

shrb^U6.gRNAx2

lamellocyte | increased number | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

lamellocyte | increased number | larval stage | somatic clone

shrb^G5

larval dorsal multidendritic neuron | embryonic stage 16

shrb⁴

larval dorsal multidendritic neuron | embryonic stage 16 (with Df(2R)Np5)

shrb⁴

larval dorsal multidendritic neuron | embryonic stage 16 (with In(2LR)w45-32n)

shrb⁴

larval dorsal multidendritic neuron | embryonic stage 17

shrb⁴

larval dorsal multidendritic neuron | embryonic stage 17 (with Df(2R)Np5)

shrb⁴

larval dorsal multidendritic neuron | embryonic stage 17 (with In(2LR)w45-32n)

shrb⁴

larval dorsal multidendritic neuron ddaA | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaB | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaC | cell autonomous | pupal stage | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaC | P-stage, with Scer\GAL4^ppk.1.9

shrb^UAS.GFP

larval dorsal multidendritic neuron ddaC | P-stage, with Scer\GAL4^ppk.PU

shrb^UAS.GFP

larval dorsal multidendritic neuron ddaC | P-stage | somatic clone

shrb^G5

larval dorsal multidendritic neuron ddaC | pupal stage, with Scer\GAL4^ppk.1.9

shrb^RNAi.UAS

larval dorsal multidendritic neuron ddaC | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaD | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaE, with Scer\GAL4²²¹

shrb^UAS.GFP

larval dorsal multidendritic neuron ddaE | somatic clone

shrb⁴

larval dorsal multidendritic neuron ddaF | somatic clone

shrb⁴

larval multidendritic class I neuron | pupal stage, with Scer\GAL4^smid-C161

shrb^RNAi.UAS

larval tracheal system | third instar larval stage, with Scer\GAL4^Cg.PA

shrb^UAS.GFP

larval tracheal system | third instar larval stage, with Scer\GAL4^ppl.PP

shrb^UAS.GFP

larval tracheocyte | embryonic stage

shrb⁴

midbody | P-stage, with Scer\GAL4^sca.PU

shrb^HMS01767

NMJ bouton | increased number | larval stage, with Scer\GAL4^VGlut1-OK371

shrb^HMS01767

NMJ bouton | increased number | larval stage, with Spyo\Cas9^VGlut1-OK371

shrb^U6.gRNAx2

NMJ bouton | increased number | larval stage, with Spyo\Cas9^wor.PC

shrb^U6.gRNAx2

nucleus | increased number | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

primary lobe of embryonic/larval lymph gland | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

secondary lobe of embryonic/larval lymph gland | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

spermatozoon | absent, with Scer\GAL4^{VP16.nanos.UTR}

tertiary lobe of embryonic/larval lymph gland | larval stage, with Scer\GAL4^dome-PG14

shrb^KK108557

testis, with Scer\GAL4^{VP16.nanos.UTR}

type Ib terminal bouton | larval stage, with Spyo\Cas9^wor.PC

shrb^U6.gRNAx2

waste bag | absent, with Scer\GAL4^{VP16.nanos.UTR}

wing disc | somatic clone

shrb⁴

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) (7)

14 of 14

Yes

Yes

6

12 of 14

No

Yes

2

12 of 14

No

Yes

3

8 of 14

No

Yes

2 of 14

No

No

1 of 14

No

No

1 of 14

No

No

Model Organism Orthologs (via DIOPT v9.1)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

Rattus norvegicus (Norway rat) (7)

14 of 14

Yes

Yes

12 of 14

No

Yes

11 of 14

No

Yes

2 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

Mus musculus (laboratory mouse) (7)

13 of 14

Yes

Yes

2

12 of 14

No

Yes

1

2 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

Xenopus tropicalis (Western clawed frog) (6)

11 of 13

Yes

Yes

6 of 13

No

No

6 of 13

No

Yes

2 of 13

No

No

2 of 13

No

No

1 of 13

No

No

Danio rerio (Zebrafish) (8)

14 of 14

Yes

Yes

12 of 14

No

Yes

12 of 14

No

Yes

2 of 14

No

No

2 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

Caenorhabditis elegans (Nematode, roundworm) (6)

13 of 14

Yes

Yes

7 of 14

No

Yes

2 of 14

No

No

2 of 14

No

No

1 of 14

No

No

1 of 14

No

Yes

Anopheles gambiae (African malaria mosquito) (4)

Agam\AgaP_AGAP005316

12 of 12

Yes

Yes

Agam\AgaP_AGAP005100

2 of 12

No

No

Agam\AgaP_AGAP006229

1 of 12

No

No

Agam\AgaP_AGAP012459

1 of 12

No

No

Arabidopsis thaliana (thale-cress) (9)

10 of 13

Yes

Yes

10 of 13

Yes

Yes

2 of 13

No

No

2 of 13

No

No

1 of 13

No

No

1 of 13

No

No

1 of 13

No

No

1 of 13

No

No

1 of 13

No

No

Saccharomyces cerevisiae (Brewer's yeast) (4)

11 of 13

Yes

Yes

2 of 13

No

No

1 of 13

No

No

1 of 13

No

No

Schizosaccharomyces pombe (Fission yeast) (4)

12 of 12

Yes

Yes

Spom\SPCC162.06c

2 of 12

No

No

Spom\SPBC1604.18c

1 of 12

No

No

1 of 12

No

No

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:shrb. 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) (3)

3 of 13

2 of 13

2 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 ( 1 )

Human Ortholog

Disease

Evidence

References

CHMP4B; charged multivesicular body protein 4B

model of cataract 31 multiple types

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 2 )

Allele

Disease

Interaction

References

ameliorates fragile X syndrome

modeled by Fmr1^Δ50M

(Vita and Broadie, 2017)

exacerbates Huntington's disease

modeled by Hsap\HTT^GMR.Q120

(Rusten et al., 2007)

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?

CHMP4B; charged multivesicular body protein 4B

14 of 14

CATARACT 31, MULTIPLE TYPES; CTRCT31

cataract 31 multiple types

CHMP4A; charged multivesicular body protein 4A

12 of 14

CHMP4C; charged multivesicular body protein 4C

12 of 14

CHMP4BP1; charged multivesicular body protein 4B pseudogene 1

8 of 14

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.

Dmel gene

Ortholog showing functional complementation

Supporting References

(Baeumers et al., 2022)

(Sweeney et al., 2006)

(Sweeney et al., 2006)

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

anti bait coimmunoprecipitation, quantitative reverse transcription pcr

(Vita and Broadie, 2017)

protein-protein

Physical Interaction

Assay

References

anti tag coimmunoprecipitation, anti tag western blot, pull down, autoradiography

(Lie-Jensen et al., 2019, Eikenes et al., 2015)

anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based

(Rhee et al., 2014)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

anti tag coimmunoprecipitation, peptide massfingerprinting

(Kwon et al., 2013)

anti tag coimmunoprecipitation, anti tag western blot

(Mathieu et al., 2022)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

experimental knowledge based

(Guruharsha et al., 2011)

affinity technology, Identification by mass spectrometry, pull down, autoradiography

(Capalbo et al., 2012)

anti tag coimmunoprecipitation, peptide massfingerprinting

(Kwon et al., 2013)

anti tag coimmunoprecipitation, anti tag western blot, x-ray crystallography, predetermined participant, surface plasmon resonance, pull down, western blot

(McMillan et al., 2017, Troost et al., 2012)

(Loncle et al., 2015)

anti tag coimmunoprecipitation, Identification by mass spectrometry

(Vinayagam et al., 2016)

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

enhanceable

(Martin et al., 2022)

suppressible

(Shravage et al., 2013)

suppressible

(Matias et al., 2015)

suppressible

(Bruelle et al., 2023)

suppressible

(Matias et al., 2015)

enhanceable

(Matias et al., 2015)

enhanceable

(Martin et al., 2022)

suppressible

(Mathieu et al., 2022)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

enhanceable

(Martin et al., 2022)

enhanceable

(Martin et al., 2022)

suppressible

(Tjota et al., 2011)

suppressible

(Uytterhoeven et al., 2011)

enhanceable

(Sen et al., 2013)

suppressible

(Mathieu et al., 2022)

suppressible

(Bäumers et al., 2019)

suppressible

(Bäumers et al., 2019)

External Data

Subunit Structure (UniProtKB)

Homopolymer; forms elongated striated filaments of uniform 10nm width (PubMed:27452459). Monomers interact in a staggered arrangement mediated by complementary charged electrostatic surfaces (PubMed:27452459). Interacts with l(2)gd1 (via DM14 domains 1 and 3); the interaction is direct and blocks access to the surface involved in homopolymerization (PubMed:22389409, PubMed:27452459, PubMed:28564595). This interaction may be required for the ESCRT-III complex role in multivesicular body formation (PubMed:22389409).

(UniProt, Q8T0Q4 )

Linkouts

BioGRID - A database of protein and genetic interactions.

61771

DroID - A comprehensive database of gene and protein interactions.

FBgn0086656

MIST (genetic) - An integrated Molecular Interaction Database

35933

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

35933

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.

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

Class of Gene

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

2R

Recombination map

2-60

Cytogenetic map

Sequence location

2R:9,142,146..9,143,914 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

45A12-45A12

Limits computationally determined from genome sequence between P{lacW}CG8788^k04512&P{lacW}l(2)k13412^k13412 and P{PZ}Pkn⁰⁶⁷³⁶

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

Determined by deficiency mapping (details unspecified).

(Gao et al., 1999)

44F-45E

(Gao, 1999.11.4, Gao et al., 1999)

49C-50D

(Gao et al., 1999)

Experimentally Determined Recombination Data

Location

2-60

(FlyBase, 2016)

2-58.2

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (17)

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

606716

w^*; P{sqh-sfGFP.shrb}21

606436

w^*; PBac{U6-shrb.gRNAx2}VK00027

32559

w^*; P{UAS-shrb-GFP}2; P{NIG.4699R}3

606717

P{sStMCA}10, w^*; P{sqh-sfGFP.shrb}31

38305

y¹ v¹; P{TRiP.HMS01767}attP40

11016

y¹ w^67c23; P{lacW}shrb^k11201/CyO

19705

y¹ w^67c23; P{EPgy2}shrb^EY05194/CyO

14372

y¹; P{SUPor-P}shrb^KG01481/SM6a; ry⁵⁰⁶

102787

y¹ w^67c23 ; P{lacW}shrb^k11201 / CyO

v106823

P{KK108557}VIE-260B

v341830

P{hsFLP}1, y¹ w¹¹¹⁸; P{HD_CFD00982}attP40/CyO-GFP

v341963

P{hsFLP}1, y¹ w¹¹¹⁸; P{HD_CFD01267}attP40/CyO-GFP

More stocks available...

Genomic Clones (14)

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

cDNA Clones (63)

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

GH13992

Other clones

DPiM_GH13992

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.

FBgn0086656

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0086656

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Bäumers et al., 2019)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

shrb is required cell autonomously for dendrite pruning in sensory neurons.

(Loncle et al., 2015)

Mutants show aberrant endosome morphology, fail to degrade ubiquitinated cargoes, and a degree of endosomal accumulation of the signaling receptors.

(Vaccari and Bilder, 2006)

dsRNA made from templates generated with primers directed against Vps32 that is transfected into S2 treated with Listeria monocytogenes reveals Vps32 to be involved in Listeria monocytogenes entry, intracellular growth, and vacuolar escape.

(Cheng et al., 2005)

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

(Boutros et al., 2004)

Area matching Drosophila EST AA264253. This EST has sequence similarity to SNF7 genes.

(Davies, 2001.3.15)

shrb has a function in dendritic development.

(Gao et al., 1999)

"44F--45E" was stated as revision.

(Gao, 1999.11.4, Gao et al., 1999)

Relationship to Other Genes

Source for database merge of

Source for merge of: shrb l(2)k11201

(Sweeney et al., 2006)

Source for merge of: shrb Vps32

(Sweeney et al., 2006, Sweeney et al., 2006)

Source for merge of: CG8055 anon-WO0118547.180

(FlyBase, 1996-)

Additional comments

A 31 nucleotide deletion in the coding region of shrb⁴ confirms that shrb is Vps32.

(Sweeney et al., 2006)

Source for merge of CG8055 anon-WO0118547.180 was sequence comparison ( date:051113 ).

(FlyBase, 1996-)

Nomenclature History

Source for database identify of

Source for identity of: Vps32 CG8055

(Vaccari, 2006.3.1)

Nomenclature comments

Etymology

Synonyms and Secondary IDs (16)

Reported As

Symbol Synonym

CG8055

(National Institute of Genetics Fly Stocks, 2022-, Laffafian and Tepass, 2019, Gupta et al., 2014, Lee et al., 2014, Cammarato et al., 2011, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Vaccari et al., 2009, Philips et al., 2008, Sepp et al., 2008, Gandhi et al., 2007, Irion and St Johnston, 2007, Stuart et al., 2007, Sweeney et al., 2006, Sweeney et al., 2006, Cheng et al., 2005)

CHMP4C

(Capalbo et al., 2019)

DmSNF2

(Winter and Hauser, 2006)

Dvps32

(Gupta et al., 2014)

ESCRT

(Hounjet and Vooijs, 2021, Carnes et al., 2015)

ESCRT-III

(Tognon et al., 2014, El Amine et al., 2013, Juhász et al., 2008, Rusten et al., 2007)

Shrb

(Linnemannstöns et al., 2022, Laffafian and Tepass, 2019, Wang et al., 2018, Capalbo et al., 2012, Giansanti et al., 2012, Tokusumi et al., 2012, Lone et al., 2010, Michelet et al., 2010)

Shrub

(González-Méndez et al., 2020, Warecki et al., 2020, Aradhya et al., 2015, Copf, 2015, Parchure et al., 2015, Baron, 2012)

Snf7

(Gohel et al., 2020, Vaccari et al., 2009)

Snf7/Vps32

(Vaccari and Bilder, 2006)

Vps32

(Ray et al., 2024, Wang et al., 2022, Hodgson et al., 2019, Sheng et al., 2016, Kanamori et al., 2015, Li and Blissard, 2015, Dong et al., 2014, Szatmári et al., 2014, Zhang et al., 2014, Shravage et al., 2013, Chittaranjan et al., 2009, Vaccari et al., 2009, Juhász et al., 2008, Sweeney et al., 2006, Sweeney et al., 2006)

anon-WO0118547.180

shrub

(Mathieu and Huynh, 2024, Ricolo et al., 2021, Arii et al., 2018, Salazar and Yamamoto, 2018, Hayashi and Dong, 2017, Vita and Broadie, 2017, Palmer et al., 2015, Dong et al., 2014, Sen et al., 2013, Sweeney et al., 2006)

vps32

(Wang et al., 2021, Khezri and Rusten, 2019, Jiang et al., 2018, Mukherjee et al., 2016, Tjota et al., 2011, Uytterhoeven et al., 2011, Saj et al., 2010, Vaccari et al., 2009, Irion and St Johnston, 2007, Link et al., 2007, Rusten et al., 2007)

vps32/snf7

(Irion and St Johnston, 2007, Vaccari, 2006.3.1)

Name Synonyms

Shrub

(Xiao et al., 2025, Sachan et al., 2024, Papagiannouli, 2022, Reiff et al., 2021, Aradhya and Jagla, 2020, Jiang et al., 2019, Lie-Jensen et al., 2019, Daniel et al., 2018, McMillan et al., 2017, Aradhya et al., 2015, Eikenes et al., 2015, West et al., 2015, Issman-Zecharya and Schuldiner, 2014, Puram and Bonni, 2013, Guruharsha et al., 2012)

Vps32

(Vaccari, 2006.3.1)

Secondary FlyBase IDs

FBgn0033385
FBgn0028565
FBgn0062188

Datasets (0)

Study focus (0)

Experimental Role

Project

Project Type

Title

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 46 )

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.

35933

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.

Q8T0Q4

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

Q8T0Q4

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.

Q8T0Q4

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0086656

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

35933/tissue=All

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

FBgn0086656

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

FBgn0086656

FlyMine - An integrated database for Drosophila genomics

FBgn0086656

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

Snf7 family

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

dme:Dmel_CG8055

MARRVEL_MODEL - MARRVEL (model organism gene)

35933

PDB - An information portal to biological macromolecular structures

5J45
5VO5

Linkouts

BioGRID - A database of protein and genetic interactions.

61771

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

FBgn0086656

DroID - A comprehensive database of gene and protein interactions.

FBgn0086656

DRSC - Results frm RNAi screens

FBgn0086656

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

shrb_1

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0086656

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0086656

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

CG8055

Flygut - An atlas of the Drosophila adult midgut

FBgn0086656

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

FBgn0086656

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

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

Interactive Fly

KEGG Pathways - A collection of manually drawn pathway maps representing knowledge of molecular interaction, reaction and relation networks.

MIST (genetic) - An integrated Molecular Interaction Database

35933

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

35933

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

References (186)