FlyBase Gene Report: Dmel\shn

FB2026_01 , released March 12, 2026

FB2026_01 , released March 12, 2026

Gene: Dmel\shn

General Information

Symbol

Dmel\shn

Species

D. melanogaster

Name

schnurri

Annotation Symbol

CG7734

Feature Type

protein_coding_gene

FlyBase ID

FBgn0003396

Gene Model Status

Stock Availability

144 publicly available

Gene Summary

schnurri (shn) encodes a zinc finger C2H2 transcription factor involved in Dpp signaling. It contributes to multiple processes including ectoderm and midgut development, wing disc patterning and morphogenesis of Malpighian tubules and dendrites. [Date last reviewed: 2019-06-13] (FlyBase Gene Snapshot)

All Summaries

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

Also Known As

quo, l(2)04738, EP2359

Key Links

Genomic Location

Cytogenetic map

Sequence location

2R:11,159,086..11,217,394 [+]

Recombination map

2-63

RefSeq locus

NT_033778 REGION:11159086..11217394

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 (23 terms)

Molecular Function (6 terms)

Terms Based on Experimental Evidence (4 terms)

CV Term

Evidence

References

enables DNA-binding transcription factor activity, RNA polymerase II-specific

inferred from direct assay

(Dai et al., 2000)

enables RNA polymerase II-specific DNA-binding transcription factor binding

inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648

(Dai et al., 2000)

enables transcription coactivator activity

inferred from direct assay

(Dai et al., 2000)

enables transcription corepressor activity

inferred from direct assay

(Yao et al., 2006, Muller et al., 2003)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

enables nucleic acid binding

inferred from electronic annotation with InterPro:IPR003604

(InterPro Project Members, 2004-)

enables zinc ion binding

inferred from electronic annotation with InterPro:IPR003604

(InterPro Project Members, 2004-)

Biological Process (15 terms)

Terms Based on Experimental Evidence (14 terms)

CV Term

Evidence

References

involved_in BMP signaling pathway

inferred from direct assay

(Yao et al., 2006, Muller et al., 2003, Dai et al., 2000)

inferred from mutant phenotype

(Zhang et al., 2025, Torres-Vazquez et al., 2001, Dai et al., 2000)

inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490

(Arora et al., 1995, Staehling-Hampton et al., 1995)

involved_in dendrite morphogenesis

inferred from mutant phenotype

(Iyer et al., 2013)

involved_in dorsal closure

inferred from mutant phenotype

(Torres-Vazquez et al., 2001)

involved_in dorsal/ventral pattern formation

inferred from mutant phenotype

(Torres-Vazquez et al., 2001, Arora et al., 1995)

involved_in ectoderm development

inferred from mutant phenotype

(Grieder et al., 1995)

involved_in imaginal disc-derived wing morphogenesis

inferred from mutant phenotype

(Torres-Vazquez et al., 2000)

inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490

(Torres-Vazquez et al., 2000)

involved_in imaginal disc-derived wing vein morphogenesis

inferred from mutant phenotype

(Torres-Vazquez et al., 2000)

involved_in Malpighian tubule morphogenesis

inferred from mutant phenotype

(Jack and Myette, 1999)

involved_in mesoderm development

inferred from genetic interaction with FLYBASE:dpp; FB:FBgn0000490

(Arora et al., 1995)

involved_in midgut development

inferred from mutant phenotype

(Grieder et al., 1995)

involved_in negative regulation of transcription by RNA polymerase II

inferred from direct assay

(Yao et al., 2006, Muller et al., 2003)

inferred from mutant phenotype

(Torres-Vazquez et al., 2001, Torres-Vazquez et al., 2000)

involved_in positive regulation of transcription by RNA polymerase II

inferred from direct assay

(Yao et al., 2006, Dai et al., 2000)

inferred from mutant phenotype

(Torres-Vazquez et al., 2001, Torres-Vazquez et al., 2000)

involved_in wing disc anterior/posterior pattern formation

inferred from mutant phenotype

(Torres-Vazquez et al., 2000)

involved_in wing disc dorsal/ventral pattern formation

inferred from genetic interaction with FLYBASE:Bx; FB:FBgn0265598,FLYBASE:ap; FB:FBgn0267978

(Bejarano et al., 2008)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

involved_in negative regulation of salivary gland boundary specification

traceable author statement

(Bradley et al., 2001)

Cellular Component (2 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

is_active_in nucleus

inferred from direct assay

(Ell et al., 2024, Dai et al., 2000)

part_of RNA polymerase II transcription repressor complex

inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648,FLYBASE:Med; FB:FBgn0288966

(Yao et al., 2006, Muller et al., 2003)

Terms Based on Predictions or Assertions (0 terms)

Gene Group (FlyBase)

C2H2 ZINC FINGER TRANSCRIPTION FACTORS

Pathway (FlyBase)

BMP SIGNALING PATHWAY CORE COMPONENTS

Protein Family (UniProt)

-

Protein Signatures (InterPro)

Summaries

Gene Snapshot

schnurri (shn) encodes a zinc finger C2H2 transcription factor involved in Dpp signaling. It contributes to multiple processes including ectoderm and midgut development, wing disc patterning and morphogenesis of Malpighian tubules and dendrites. [Date last reviewed: 2019-06-13]

Gene Group (FlyBase)

C2H2 ZINC FINGER TRANSCRIPTION FACTORS -: Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys₂His₂ zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).

Pathway (FlyBase)

BMP SIGNALING PATHWAY CORE COMPONENTS -: The Bone Morphogenetic Protein (BMP) signaling pathway is one of two branches of Transforming Growth Factor-β family signaling in Drosophila. The binding of a BMP family dimer to a heterodimeric serine/threonine kinase receptor complex results in the phosphorylation of Mad, a member of the Smad family. Mad forms a complex with the co-Smad, Med. This complex translocates into the nucleus and regulates the transcription of target genes in concert with other nuclear cofactors. (Adapted from FBrf0236482.)

Protein Function (UniProtKB)

Transcription cofactor involved in dpp/BMP TGF-beta signal-induced transcriptional regulation downstream of the receptor serine/threonine kinases tkv/thickvein and put/punt (PubMed:11071761, PubMed:11071762, PubMed:11290303, PubMed:15296719, PubMed:7774017, PubMed:7774018). Recruited to DNA enhancer elements by association with the SMAD protein Mad-Med complex (PubMed:15296719, PubMed:16109720). Mad-Med acts as a transcriptional activator but recruits shn/shnurri to enhance its activity or switch to transcriptional repression (PubMed:11071761, PubMed:16109720). When recruited by Mad-Med can regulate dpp-responsive genes by directly activating their expression or by repressing expression of the transcriptional repressor brk/brinker (PubMed:11290303, PubMed:12705870, PubMed:15296719, PubMed:16109720). Also involved in repression of other key developmental genes including bam and gsb (PubMed:15296719). Required for dpp-signaling induced repression of brk and induction of sna during embryonic development (PubMed:11290303). Required for transcriptional regulation of dpp-responsive genes in the mesoderm and endoderm during embryonic gut development; required for the proper differentiation of dorsal ectodermal cells (PubMed:11071761, PubMed:7774017, PubMed:7774018). Not required for early embryo development (PubMed:11290303). Required for expression of dpp-responsive genes involved in patterning of the wing imaginal disc (PubMed:11071762). Required for dpp-responsive cell proliferation and anterior-posterior patterning in the wing (PubMed:11071762). Required for vein formation in the developing wing (PubMed:11071762, PubMed:38344065, PubMed:7588072, PubMed:7774018). Required in somatic cyst cells of the testes for regulation of germ cell proliferation (PubMed:9334286). Prevents JNK-induced apoptosis during embryogenesis by repressing expression of the pro-apoptotic protein rpr/reaper; required for survival of dorsal edge cells during dorsal closure (PubMed:25307481). Required for tissue maintenance after DNA damage, probably by inducing expression of the growth factor Pvf1, which stimulates the macrophage-like activity of hemocytes (PubMed:22275438). ACTIVITY REGULATION: DNA binding, specifically of the enhancer region of the brk gene, is activated by dpp-signaling (PubMed:15296719). Dpp-signaling induces phosphorylation of Mad by dpp-specific receptor serine/threonine kinase complexes, which in turn induces interaction of Mad with Med and nuclear localization of the Mad-Med complex, where it can bind the enhancer regions of dpp-responsive genes and recruit shn (Probable).

(UniProt, A1Z8F4)

Phenotypic Description (Red Book; Lindsley and Zimm 1992)

shn: schnurri

Embryonic lethal. Embryos lack dorsal hypoderm. Internal organs appear normal and extruded through the open dorsal side of the embryo. Ventral hypoderm contracted.

Summary (Interactive Fly)

transcription factor - zinc finger - target of Dpp - the activity of the Mad/Schnurri complex is modulated in a promoter specific fashion to active or repress gene transcription - corepressor Schnurri protects epithelial cells from JNK-induced apoptosis in drosophila embryos.

Interactive Fly

Gene Model and Products

Number of Transcripts

6

Number of Unique Polypeptides

3

Please see the JBrowse view of Dmel\shn 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 A1Z8F4)

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

Stop-codon suppression (UGA) postulated; FBrf0216884

Gene model reviewed during 5.44

Gene model reviewed during 5.49

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

FBtr0088098

8940

2532

FBtr0088099

9552

2532

FBtr0088100

9026

2532

FBtr0088101

9886

2577

FBtr0330619

9011

2532

FBtr0330620

9886

2587

Additional Transcript Data and Comments

Reported size (kB)

9.5 (northern blot)

(Arora et al., 1995)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

UniProt

RefSeq ID

GenBank

FBpp0087200

271.9

2532

9.12

FBpp0087201

271.9

2532

9.12

FBpp0087202

271.9

2532

9.12

FBpp0089118

276.8

2577

9.27

FBpp0303469

271.9

2532

9.12

FBpp0303470

278.0

2587

9.26

Polypeptides with Identical Sequences

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

2532 aa isoforms: shn-PA, shn-PB, shn-PC, shn-PE

Additional Polypeptide Data and Comments

Reported size (kDa)

2578, 2527 (aa)

(Arora et al., 1995)

2529 (aa)

(Grieder et al., 1995)

2527 (aa)

(Staehling-Hampton et al., 1995)

Comments

External Data

Subunit Structure (UniProtKB)

May form higher order homomultimers (PubMed:11071761). Associates with the Mad-Med complex; association is dependent on the sequence of Mad-Med bound DNA (PubMed:15296719, PubMed:16109720). Recruitment by DNA associated Mad-Med requires C2H2-type zinc-finger domains 5 and 7 (PubMed:15296719). Interacts (via the region upstream of C2H2-type zinc-finger 3) with Mad (via MH2 or linker domains); the interaction is direct (PubMed:11071761).

(UniProt, A1Z8F4)

Domain

Possesses at least 2 DNA-binding domains (DBD1 and DBD2) (PubMed:11071761). DBD1, made up of C2H2-type zinc-fingers 1 and 2, binds DNA with a consensus sequence of 5'-GGGN(6)CCC-3' and an optimal sequence of 5'-GGGA[AT]CGTTCCC-3' (PubMed:11071761). DBD2, made up of C2H2-type zinc-fingers 3 and 4, binds DNA with a consensus sequence of 5'-GGGN(5)CCC-3' and an optimal sequence of 5'-GGGGA[AC][AT]TCCC-3' (PubMed:11071761). C2H2-type zinc-finger domains 1, 2, 3 and 4 are involved in vein formation in the developing wing (PubMed:38344065).

The C-terminal zinc-finger domains are required for recruitment to the brk gene enhancer region by Mad-Med and repression of the brk gene; C2H2-type zinc-finger domains 5 and 7 are essential while C2H2-type zinc-finger domain 6 is dispensable.

A region between C2H2-type zinc-finger domains 4 and 5 has inherent ability to repress transcription and is required to repress transcription of brk.

(UniProt, A1Z8F4)

Crossreferences

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

Linkouts

Sequences Consistent with the Gene Model

Nucleotide / Polypeptide Records

Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\shn 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).

-1.79

Transcript Expression

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 1 -- 3

Comment: maternally deposited

(Fisher et al., 2012)

blastoderm stage

organism | dorsal

(Grieder et al., 1995, Staehling-Hampton et al., 1995)

embryonic stage 4 -- 5

organism | dorsal

(Arora et al., 1995)

embryonic stage 4 -- 6

central brain anlage in statu nascendi

(Fisher et al., 2012)

clypeo-labral anlage in statu nascendi

(Fisher et al., 2012)

dorsal ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

dorsal head epidermis anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

embryonic stage 5

mesoderm | presumptive

(Arora et al., 1995)

(Grieder et al., 1995)

(Arora et al., 1995)

embryonic stage 7 -- 8

clypeo-labral anlage

(Fisher et al., 2012)

dorsal ectoderm

(Fisher et al., 2012)

ventral ectoderm

(Fisher et al., 2012)

embryonic stage 7 -- 10

(Staehling-Hampton et al., 1995)

embryonic stage 8 -- 10

(Arora et al., 1995, Grieder et al., 1995)

embryonic/larval anterior midgut

(Arora et al., 1995)

ectoderm | dorsal

(Arora et al., 1995)

embryonic/larval posterior midgut

(Arora et al., 1995)

embryonic stage 9 -- 10

clypeo-labral primordium

(Fisher et al., 2012)

dorsal ectoderm

(Fisher et al., 2012)

ventral ectoderm

(Fisher et al., 2012)

embryonic stage 10 -- 12

endoderm of embryonic/larval midgut

(Grieder et al., 1995)

embryonic stage 11 -- 12

clypeo-labral primordium

(Fisher et al., 2012)

dorsal epidermis primordium

(Fisher et al., 2012)

ventral epidermis primordium

(Fisher et al., 2012)

embryonic stage 12

parasegment 4 of endoderm

(Staehling-Hampton et al., 1995)

parasegment 7 of endoderm

(Staehling-Hampton et al., 1995)

embryonic stage 13 -- 16

embryonic dorsal epidermis

(Fisher et al., 2012)

embryonic hindgut

(Fisher et al., 2012)

embryonic ventral epidermis

(Fisher et al., 2012)

embryonic/larval midgut

(Fisher et al., 2012)

(Fisher et al., 2012)

embryonic stage 13 -- 17

(Staehling-Hampton et al., 1995)

embryonic stage 14

of visceral trunk mesoderm & parasegment 6 & parasegment 7

(Arora et al., 1995)

(Arora et al., 1995)

embryonic/larval pharynx

(Arora et al., 1995)

(Organista et al., 2015)

(Arora et al., 1995)

adult stage & oogenesis

(Arora et al., 1995)

(Arora et al., 1995)

(Arora et al., 1995)

Additional Descriptive Data

Marker for

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 14

(Staehling-Hampton et al., 1995)

embryonic/larval foregut

(Staehling-Hampton et al., 1995)

Additional Descriptive Data

shn protein is detected in the endoderm and in the foregut of germ band retracted embryos.

(Staehling-Hampton et al., 1995)

Marker for

Subcellular Localization

CV Term

Evidence

References

is_active_in nucleus

inferred from direct assay

(Ell et al., 2024, Dai et al., 2000)

part_of RNA polymerase II transcription repressor complex

inferred from physical interaction with FLYBASE:Mad; FB:FBgn0011648,FLYBASE:Med; FB:FBgn0288966

(Yao et al., 2006, Muller et al., 2003)

Expression Deduced from Reporters

Reporter: P{PZ}shn⁰⁴⁷³⁸

Stage

Tissue/Position (including subcellular localization)

Reference

wandering third instar larval stage

(Meister and Braun, 1995.10)

embryonic/larval fat body

(Meister and Braun, 1995.10)

embryonic/larval midgut | restricted

(Meister and Braun, 1995.10)

Reporter: PBac{GAL4D,EYFP}shn¹⁶⁶

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 12

(Horn et al., 2003)

embryonic stage 16

embryonic/larval gut

(Horn et al., 2003)

Reporter: TI{TI}shn^HA

Stage

Tissue/Position (including subcellular localization)

Reference

third instar larval stage

larval glial cell

(Vuilleumier et al., 2022)

High-Throughput Expression Data

Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\shn in JBrowse

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

View exonic expression by developmental stage for Dmel\shn

View exonic expression by tissue for Dmel\shn

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

FBgn0003396

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

36171/tissue=All

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

FBgn0003396

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0003396

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

FBgn0003396

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

CG7734

Flygut - An atlas of the Drosophila adult midgut

FBgn0003396

Images

Image Based Resources

Alleles, Insertions, Transgenic Constructs, and Aberrations

Classical and Insertion Alleles ( 89 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 25 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of shn

Transgenic constructs containing regulatory region of shn

Aberrations (Deficiencies and Duplications) ( 17 )

Inferred from experimentation ( 17 )

Gene not disrupted in

(Kania et al., 1995, BDGP Project Members, 1994-1999)

(Chase and Baker, 1995)

(Bieser et al., 2019)

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

Gene partially disrupted in

(Ruan et al., 2017)

Inferred from location ( 12 )

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

shn¹

lethal | heat sensitive

shn²

lethal | recessive

lethal - all die during embryonic stage

lethal - all die during embryonic stage | recessive

viable, with Scer\GAL4^pnr-MD237

shn^GD1644

Sterility

Allele

shn^zf1.2.4.5

Other Phenotypes

Allele

abnormal circadian rhythm, with Scer\GAL4^P2.4.Pdf

abnormal circadian rhythm, with Scer\GAL4^tim.PE

shn^UAS.cMa

abnormal circadian rhythm | heat sensitive, with Scer\GAL4^P2.4.Pdf, Scer\GAL80^Pdf.PS

shn^UAS.cMa

abnormal circadian rhythm | heat sensitive, with Scer\GAL4^tim.PE, Scer\GAL80^Pdf.PS

shn⁷⁵⁶

abnormal cytokinesis | somatic clone | spermatogenesis

abnormal locomotor behavior | adult stage, with Scer\GAL4^ple.PF

shn^KK101278

abnormal memory

shn^umnitza

abnormal neuroanatomy

abnormal neuroanatomy, with Scer\GAL4²²¹

abnormal neuroanatomy, with Scer\GAL4^ppk.1.9

abnormal neuroanatomy (with shn¹)

shn^k04412

abnormal neuroanatomy (with shn^k04412)

shn¹

abnormal neuroanatomy | larval stage, with Scer\GAL4^elav-C155

shn^EP644

abnormal proboscis extension reflex, with Scer\GAL4^ple.PF

shn^KK101278

decreased cell size | somatic clone | spermatogenesis

increased cell number | somatic clone | spermatogenesis

increased occurrence of cell division | somatic clone | spermatogenesis

some die during embryonic stage

some die during embryonic stage | recessive

visible, with Scer\GAL4^en-e16E

shn^EP2359

visible, with Scer\GAL4^ey.PH

shn^EP2359

visible, with Scer\GAL4^hs.2sev

shn^EP2359

visible, with Scer\GAL4^sca-537.4

shn^EP2359

visible | adult stage

Phenotype manifest in

Allele

abdominal anterior ventral multidendritic neuron vdaa, with Scer\GAL4^ppk.1.9

anterior fascicle & synapse, with Scer\GAL4^elav-C155

shn^EP644

chordotonal organ | lateral

double row of wing sensilla | cell autonomous | somatic clone

double row of wing sensilla | ectopic | somatic clone

shn²

embryo | dorsal

shn⁰⁴⁷³⁸

embryo | dorsal closure stage

embryonic/first instar larval cuticle

embryonic/larval crystal cell

shn¹

embryonic/larval midgut | embryonic stage

shn¹

embryonic abdominal dorsal sensory organ precursor cluster

embryonic abdominal dorsal sensory organ precursor cluster (with shn¹)

shn^k04412

embryonic abdominal dorsal sensory organ precursor cluster (with shn^k04412)

shn¹

embryonic abdominal lateral sensory organ precursor cluster

embryonic abdominal lateral sensory organ precursor cluster (with shn¹)

shn^k04412

embryonic abdominal lateral sensory organ precursor cluster (with shn^k04412)

shn¹

embryonic dorsal epidermis

embryonic epidermis

embryonic epidermis | dorsal

shn⁰⁴⁷³⁸

embryonic midgut constriction

shn¹

epidermis | dorsal

shn²

eye, with Scer\GAL4^ey.PH

shn^EP2359

eye, with Scer\GAL4^hs.2sev

shn^EP2359

eye | somatic clone

shn^56M

shn^k16131

female germline stem cell | germline clone

shn⁰⁴⁷³⁸

shn⁰⁴⁷³⁸

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

larval dorsal multidendritic neuron ddaD, with Scer\GAL4²²¹

male germline cyst | somatic clone | spermatogenesis

Malpighian tubule

shn¹

ommatidium, with Scer\GAL4^hs.2sev

shn^EP2359

photoreceptor neuron | somatic clone

shn^56M

presumptive embryonic/larval peripheral nervous system

scutellar bristle, with Scer\GAL4^sca-537.4

shn^EP2359

triple row of wing sensilla | cell autonomous | somatic clone

ventral denticle belt

visceral trunk mesoderm

shn¹

wing | somatic clone

shn¹

wing blade anterior compartment | somatic clone

shn²

shn¹

wing vein, with Scer\GAL4^en-e16E

shn^EP2359

wing vein | cell autonomous | somatic clone

shn²

wing vein | cell non-autonomous | ectopic | somatic clone

shn²

wing vein | ectopic

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) (29)

6 of 14

Yes

Yes

1

5 of 14

No

Yes

5 of 14

No

Yes

1 of 14

No

No

1

1 of 14

No

No

1 of 14

No

No

1 of 14

No

Yes

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1

1 of 14

No

No

1 of 14

No

No

1 of 14

No

Yes

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

Yes

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

1 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) (3)

5 of 14

Yes

Yes

5 of 14

Yes

Yes

5 of 14

Yes

Yes

Mus musculus (laboratory mouse) (3)

6 of 14

Yes

Yes

5 of 14

No

Yes

5 of 14

No

Yes

Xenopus tropicalis (Western clawed frog) (12)

5 of 13

Yes

Yes

4 of 13

No

Yes

4 of 13

No

Yes

Xtro\LOC100486443

1 of 13

No

No

Xtro\LOC101734875

1 of 13

No

No

Xtro\LOC116411624

1 of 13

No

No

Xtro\LOC116411645

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

1 of 13

No

No

Danio rerio (Zebrafish) (5)

5 of 14

Yes

Yes

5 of 14

Yes

Yes

4 of 14

No

Yes

3 of 14

No

Yes

2 of 14

No

Yes

Caenorhabditis elegans (Nematode, roundworm) (4)

5 of 14

Yes

Yes

1 of 14

No

No

1 of 14

No

No

1 of 14

No

No

Anopheles gambiae (African malaria mosquito) (7)

Agam\AgaP_AGAP007640

9 of 12

Yes

Yes

Agam\AgaP_AGAP001555

1 of 12

No

No

Agam\AgaP_AGAP006592

1 of 12

No

No

Agam\AgaP_AGAP008232

1 of 12

No

No

Agam\AgaP_AGAP011262

1 of 12

No

No

Agam\AgaP_AGAP011674

1 of 12

No

No

Agam\AgaP_AGAP012741

1 of 12

No

No

Arabidopsis thaliana (thale-cress) (0)

Saccharomyces cerevisiae (Brewer's yeast) (0)

Schizosaccharomyces pombe (Fission yeast) (0)

Escherichia coli (enterobacterium) (0)

Other Organism Orthologs (via OrthoDB)

Data provided directly from OrthoDB:shn. 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) (6)

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

1 of 13

Human Disease Associations

FlyBase Human Disease Model Reports

juvenile polyposis syndrome, TGF-β/BMP signaling pathway

Disease Ontology (DO) Annotations

Models Based on Experimental Evidence ( 1 )

Allele

Disease

Evidence

References

model of juvenile polyposis syndrome

CEA

(Zhou et al., 2021, Zhou and Boutros, 2020)

CEA with puc^E69

(Zhou et al., 2021)

Potential Models Based on Orthology ( 1 )

Human Ortholog

Disease

Evidence

References

HIVEP2; HIVEP zinc finger 2

model of autosomal dominant intellectual developmental disorder 43

IEA

(FlyBase, 2019-)

Modifiers Based on Experimental Evidence ( 2 )

Allele

Disease

Interaction

References

exacerbates tauopathy

modeled by Hsap\MAPT^UAS.cWa

(Feuillette et al., 2020)

model of juvenile polyposis syndrome

is ameliorated by BacA\p35^lexAop.cEa

(Zhou et al., 2021)

is ameliorated by hid^Pe05016

(Zhou et al., 2021)

is exacerbated by puc^E69

(Zhou et al., 2021, Zhou and Boutros, 2020)

is ameliorated by bsk^DN.UAS

(Zhou and Boutros, 2020)

is ameliorated by kay^RNAi.UAS

(Zhou and Boutros, 2020)

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?

HIVEP1; HIVEP zinc finger 1

6 of 14

HIVEP2; HIVEP zinc finger 2

5 of 14

INTELLECTUAL DEVELOPMENTAL DISORDER, AUTOSOMAL DOMINANT 43; MRD43

autosomal dominant intellectual developmental disorder 43

HIVEP3; HIVEP zinc finger 3

5 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

(Yao 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

electrophoretic mobility shift assay, autoradiography

(Chen et al., 2008)

protein-protein

Physical Interaction

Assay

References

anti tag coimmunoprecipitation, anti tag western blot

(Cai and Laughon, 2009, Yao et al., 2008)

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

(Dai et al., 2000)

anti tag coimmunoprecipitation, anti tag western blot

(Cai and Laughon, 2009)

anti tag coimmunoprecipitation, anti tag western blot

(Cai and Laughon, 2009)

anti tag coimmunoprecipitation, anti tag western blot

(Cai and Laughon, 2009)

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

suppressible

(Rusten et al., 2002, Torres-Vazquez et al., 2001, Marty et al., 2000)

suppressible

(Beckwith et al., 2013)

suppressible

(Tseng and Hariharan, 2002)

enhanceable

(Toraason et al., 2024)

suppressible

(Chao et al., 2003)

suppressible

(Beckwith et al., 2013)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

suppressible

(Bejarano et al., 2008)

suppressible

(Torres-Vazquez et al., 2001)

enhanceable

(Bejarano et al., 2008)

suppressible

(Bejarano et al., 2008)

suppressible

(Lowery et al., 2010)

suppressible

(Krupp et al., 2005)

suppressible

(Horsfield et al., 1998)

suppressible

(Bangi and Wharton, 2006, Torres-Vazquez et al., 2000)

enhanceable

(George and Terracol, 1997)

suppressible

(Bangi and Wharton, 2006)

enhanceable

(Gregory et al., 2007)

suppressible

(Beckwith et al., 2013)

enhanceable

(Pena-Rangel et al., 2002)

enhanceable

(Staehling-Hampton et al., 1995)

suppressible

(Hoodless et al., 1996)

External Data

Subunit Structure (UniProtKB)

May form higher order homomultimers (PubMed:11071761). Associates with the Mad-Med complex; association is dependent on the sequence of Mad-Med bound DNA (PubMed:15296719, PubMed:16109720). Recruitment by DNA associated Mad-Med requires C2H2-type zinc-finger domains 5 and 7 (PubMed:15296719). Interacts (via the region upstream of C2H2-type zinc-finger 3) with Mad (via MH2 or linker domains); the interaction is direct (PubMed:11071761).

(UniProt, A1Z8F4 )

Linkouts

BioGRID - A database of protein and genetic interactions.

61969

DroID - A comprehensive database of gene and protein interactions.

FBgn0003396

MIST (genetic) - An integrated Molecular Interaction Database

36171

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

36171

Pathways

Signaling Pathways (FlyBase)

BMP SIGNALING PATHWAY CORE COMPONENTS

Metabolic Pathways

FlyBase

External Links

External Data

Linkouts

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q0E9C5

Class of Gene

gene_with_stop_codon_read_through

(Jungreis et al., 2011)

protein_coding_gene

Genomic Location and Detailed Mapping Data

Chromosome (arm)

2R

Recombination map

2-63

Cytogenetic map

Sequence location

2R:11,159,086..11,217,394 [+]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

47D5-47D7

; Limits computationally determined from genome sequence between P{EP}shn^EP2359&P{EP}shn^EP644 and P{PZ}rM547

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

47D5-47D6

(determined by in situ hybridisation)

(Abdelilah-Seyfried et al., 2000)

47E1-47E2

(determined by in situ hybridisation)

(Spradling et al., 1999, Perrimon et al., 1996)

47D3-47D6

(determined by in situ hybridisation)

(Kania et al., 1995)

47D3-47D6

47D5--6 47D7--8 47E1--2

(BDGP Project Members, 1994-1999)

Experimentally Determined Recombination Data

Location

2-63

(FlyBase, 2016)

2-64.0

(Comeron et al., 2012)

2-62

(Tearle and Nusslein-Volhard, 1987)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (144)

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

12085

P{VT014973-GAL4}attP2

12617

P{VT014974-GAL4}attP2

13253

P{VT014962-GAL4}attP2

14775

P{VT014977-GAL4}attP2

14851

P{VT014961-GAL4}attP2

15073

P{VT014966-GAL4}attP2

15074

P{VT014969-GAL4}attP2

15453

P{VT014968-GAL4}attP2

15546

P{GMR85F12-GAL4}attP2

11068

16288

P{GMR85F06-lexA}attP40

v105643

P{KK101278}VIE-260B

More stocks available...

Genomic Clones (57)

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

cDNA Clones (39)

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

IP16071

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.

FBgn0003396

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

Other clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0003396

Antibody Information

Laboratory Generated Antibodies

polyclonal

(von Saucken et al., 2025, Staehling-Hampton et al., 1995)

Commercially Available Antibodies

Cell Line Information

Publicly Available Cell Lines

Other Stable Cell Lines

Other Comments

DNA-protein interactions: genome-wide binding profile assayed for shn protein in 2-3 hr embryos; see BDTNP1_TFBS_shn collection report.

(MacArthur et al., 2009)

shn has an essential role in dpp signal transduction in the wing.

(Torres-Vazquez et al., 2000)

One of five genes identified as encoding downstream components of the dpp signalling cascade which is necessary for blocking salivary gland gene activation by Scr in the dorsal region of parasegment 2.

(Henderson et al., 1999)

Loss of function alleles of tkv, put, Mad, Med and shn suppress the CycE^JP mutant eye phenotype in combination with dpp^d-ho.

(Horsfield et al., 1998)

shn and put are required to limit transient amplification of germ cells. Mosaic analysis demonstrates shn and put act within somatic cyst cells that surround germ cells, rather than in germ cells. Thus a cyst-cell-derived signal restricts germ cell proliferation and this signal is initiated by input from a member of the TGF-β superfamily. Thus, a signal relay regulates progression through the germline stem cell lineage.

(Matunis et al., 1997)

dpp receptors put and tkv, or the shn transcription factor, are autonomously required for cell proliferation in the entire developing wing.

(Burke and Basler, 1996)

shn mutations enhance the weak dpp mutations.

(Arora et al., 1995)

Genetic interactions and the strikingly similar phenotypes in shn and dpp receptors encoded by tkv and put suggest the shn plays a downstream role in dpp signalling.

(Arora et al., 1995)

shn is required for the proper development of the midgut and dorsal ectoderm. shn may represent a direct target of the dpp signalling pathway and respond to its stimulation by the activation or repression of dpp responsive genes.

(Grieder et al., 1995)

shn represents the first identified downstream component of the signal transduction pathway used by dpp and its receptors.

(Staehling-Hampton et al., 1995)

shn function is critical for cells to respond properly to dpp.

(Staehling-Hampton et al., 1995)

Relationship to Other Genes

Source for database merge of

Source for merge of: shn BEST:SD06302

(FlyBase, 1996-)

Additional comments

Nomenclature History

Source for database identify of

Nomenclature comments

Etymology

Synonyms and Secondary IDs (14)

Reported As

Symbol Synonym

BEST:SD06302

CG7734

(National Institute of Genetics Fly Stocks, 2022-, Schwartz et al., 2016, Lin et al., 2015, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Perkins et al., 2009, Quinones-Coello, 2007, Molnar et al., 2006, Dubnau et al., 2003, Kraut et al., 2001, Kraut et al., 2001)

EP(2)2359

(Pena-Rangel et al., 2002)

EP2359

(Abdelilah-Seyfried et al., 2000)

SD06302

(Pena-Rangel et al., 2002)

SHN

(Pyrowolakis et al., 2017, Dresch et al., 2016, Blatti et al., 2015, Restrepo et al., 2014, Cheng et al., 2013, MacArthur et al., 2009)

Schnurri

(Van Bortle et al., 2015)

l(2)04738

(Russell et al., 1998, Perrimon et al., 1996)

quo

(Poeck et al., 1997, Kania et al., 1995, Poeck et al., 1995)

schnurri

(Berger et al., 2008, Edwards et al., 2006, Staudt et al., 2005)

shn

(Alizada et al., 2025, Collins et al., 2024, Zhang et al., 2024, Ding et al., 2022, Vuilleumier et al., 2022, Guzman et al., 2021, McParland et al., 2021, Ruiz-Losada et al., 2021, Winbush and Singh, 2021, Aboukilila et al., 2020, Berndt et al., 2020, Feuillette et al., 2020, Geigges et al., 2020, Mahmud et al., 2020, Port et al., 2020, Robles-Murguia et al., 2020, Bieser et al., 2019, Mortensen et al., 2018, Karaiskos et al., 2017, Transgenic RNAi Project members, 2017-, Upadhyay et al., 2017, Clandinin and Owens, 2016-, Sandler and Stathopoulos, 2016, Baëza et al., 2015, Beckwith and Ceriani, 2015, Charbonnier et al., 2015, Gene Disruption Project members, 2015-, Morozova et al., 2015, Nadimpalli et al., 2015, Organista et al., 2015, Schertel et al., 2015, Svendsen et al., 2015, Van Bortle et al., 2015, Ashwal-Fluss et al., 2014, Beira et al., 2014, Esteves et al., 2014, Jiang and Singh, 2014, Taylor et al., 2014, Beckwith et al., 2013, Curtis et al., 2013, Enuameh et al., 2013, Gafner et al., 2013, Iyer et al., 2013, Li and Gilmour, 2013, Saunders et al., 2013, Terriente-Felix et al., 2013, Kelsey et al., 2012, Kvon et al., 2012, Spokony, 2012.12.12, Abruzzi et al., 2011, Bhattacharya and Baker, 2011, Johnson et al., 2011, Jungreis et al., 2011, Li et al., 2011, Nègre et al., 2011, Campos et al., 2010, Cook et al., 2010.2.12, Lowery et al., 2010, Vuilleumier et al., 2010, Cai and Laughon, 2009, Firth and Baker, 2009, Telonis-Scott et al., 2009, Chen et al., 2008, Christensen et al., 2008.9.3, Christensen et al., 2008.10.20, McClure and Schubiger, 2008, Yao et al., 2008, Beltran et al., 2007, Christensen and Cook, 2007.10.29, Christensen et al., 2007.10.29, Christensen et al., 2007.10.29, Fernández et al., 2007, Minidorff et al., 2007, Quinones-Coello, 2007, Walsh and Carroll, 2007, Bangi and Wharton, 2006, Chen and Schupbach, 2006, Chertemps et al., 2006, Dworkin and Gibson, 2006, Dworkin and Gibson., 2006, Friedman and Perrimon, 2006, Molnar et al., 2006, Yao et al., 2006, Stathopoulos and Levine, 2005, Staudt et al., 2005, Xie et al., 2005, Pyrowolakis et al., 2004)

Name Synonyms

Schnurri

(Upadhyay et al., 2017, Charbonnier et al., 2015, Van Bortle et al., 2015, Li et al., 2014, Peterson and O'Connor, 2013, Kvon et al., 2012, Zoller and Schulz, 2012, Harris and Ashe, 2011, Nègre et al., 2011, Vuilleumier et al., 2010, Cai and Laughon, 2009, MacArthur et al., 2009, Campbell and Moser, 2008, Warrior et al., 2008, Fernández et al., 2007, Walsh and Carroll, 2007)

quo vadis

schnurri

(Merino et al., 2022, Merrill et al., 2022, Wieschaus and Nüsslein-Volhard, 2016, Beckwith and Ceriani, 2015, Svendsen et al., 2015, Peterson and O'Connor, 2013, Winbush et al., 2012, Bhattacharya and Baker, 2011, Campos et al., 2010, Krejcí et al., 2009, Bejarano et al., 2008, McClure and Schubiger, 2008, Gregory et al., 2007, Chertemps et al., 2006, Dworkin and Gibson, 2006, Edwards et al., 2006, Ganguly-Fitzgerald et al., 2006, Staudt et al., 2005)

umnitza

(Berger et al., 2008)

Secondary FlyBase IDs

FBgn0010584
FBgn0013727
FBgn0043359

Datasets (2)

Study focus (2)

Experimental Role

Project

Project Type

Title

bait_protein

genome binding

ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.

bait_protein

modENCODE_regulation_TFs

genome binding

Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.

Study result (0)

Result

Result Type

Title

External Crossreferences and Linkouts ( 91 )

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.

36171

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.

A1Z8F4

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

A1Z8F4

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.

A1Z8F4

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0003396

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

36171/tissue=All

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

FBgn0003396

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

FBgn0003396

FlyMine - An integrated database for Drosophila genomics

FBgn0003396

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

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

dme:Dmel_CG7734

MARRVEL_MODEL - MARRVEL (model organism gene)

36171

Linkouts

BioGRID - A database of protein and genetic interactions.

61969

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

FBgn0003396

DroID - A comprehensive database of gene and protein interactions.

FBgn0003396

DRSC - Results frm RNAi screens

FBgn0003396

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

FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array

FBgn0003396

FlyCyc Genes - Genes from a BioCyc PGDB for Dmel

FBGN0003396

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

CG7734

Flygut - An atlas of the Drosophila adult midgut

FBgn0003396

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

TC009542

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

Interactive Fly

MIST (genetic) - An integrated Molecular Interaction Database

36171

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

36171

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q0E9C5

References (334)