FB2025_05 , released December 11, 2025
Gene: Dmel\tsh
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General Information
Symbol
Dmel\tsh
Species
D. melanogaster
Name
teashirt
Annotation Symbol
CG1374
Feature Type
protein_coding_gene
FlyBase ID
FBgn0003866
Gene Model Status
Current
Stock Availability
49 publicly available
Gene Summary
Homeotic protein that acts downstream of Arm in the Wg cascade during embryogenesis to determine segment identity throughout the entire trunk. Acts cooperatively with other trunk homeotic proteins to repress head homeotic genes and therefore repress head segmental identity. Necessary, in combination with Scr, for the formation of the prothoracic segment. Promotes eye development in the dorsal region of the eye disk and suppresses eye development in the ventral region in combination with Wg-signaling and several early dorso-ventral eye patterning genes. Required for proper development of proximal leg segments. Has differential functions along the dorso-ventral axs of the antennal and leg disks. May play a role in wing hinge development. Possible involvement in chromatin structure for modulation of transcription. Binds DNA and can act as both a transcriptional repressor and activator. Positively regulates its own expression as well as that of Dll. Negatively regulates the expression of mod. Required for Wg-mediated transcriptional repression of Ubx in the midgut. Also represses transcription of lab in the midgut and is necessary for the proper formation of anterior and central midgut structures. Tiptop (tio) and teashirt (tsh) have, on the whole, common activities. Tio and tsh repress each other's expression and tsh has a crucial role for trunk patterning that is in part masked by ectopic expression of tiptop. Both genes share a common activity required for the activation of Ser and svb and the maintenance of en and wg. (UniProt, P22265)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

ae

Key Links
Genomic Location
Cytogenetic map
40A5-40A5
Sequence location
Recombination map
2-54
RefSeq locus
NT_033779 REGION:21828593..21837011
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
Gene Ontology (GO) Annotations (27 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables DNA binding
inferred from direct assay
(Alexandre et al., 1996)
enables DNA-binding transcription repressor activity, RNA polymerase II-specific
inferred from direct assay
(Alexandre et al., 1996)
inferred from mutant phenotype
(Alexandre et al., 1996)
enables protein binding
inferred from physical interaction with UniProtKB:P18824
(Gallet et al., 1998)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables DNA binding
inferred from biological aspect of ancestor with PANTHER:PTN000281113
(GO Reference Genome Project, 2011-)
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from biological aspect of ancestor with PANTHER:PTN000281113
(GO Reference Genome Project, 2011-)
Biological Process (21 terms)
Terms Based on Experimental Evidence (19 terms)
CV Term
Evidence
References
involved_in compound eye development
inferred from mutant phenotype
(Bessa and Casares, 2005)
involved_in dorsal/ventral pattern formation, imaginal disc
inferred from mutant phenotype
(Singh et al., 2004)
involved_in epidermis morphogenesis
inferred from mutant phenotype
(Laugier et al., 2005)
involved_in eye-antennal disc development
inferred from mutant phenotype
(Singh et al., 2004)
involved_in head involution
inferred from mutant phenotype
(Laugier et al., 2005)
involved_in imaginal disc-derived wing morphogenesis
inferred from mutant phenotype
(Dworkin and Gibson, 2006)
involved_in leg disc proximal/distal pattern formation
inferred from mutant phenotype
(Wu and Cohen, 2000, Erkner et al., 1999)
involved_in Malpighian tubule stellate cell differentiation
inferred from expression pattern
(Laugier et al., 2005, Liu et al., 1999)
inferred from mutant phenotype
(Denholm et al., 2013)
involved_in midgut development
inferred from mutant phenotype
(Mathies et al., 1994)
involved_in negative regulation of DNA-templated transcription
inferred from mutant phenotype
(Soanes et al., 2001)
involved_in negative regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Bessa et al., 2009, Alexandre et al., 1996)
inferred from mutant phenotype
(Soanes et al., 2001, Roder et al., 1992)
involved_in positive regulation of cell proliferation involved in compound eye morphogenesis
inferred from mutant phenotype
(Peng et al., 2009)
involved_in positive regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Peng et al., 2009)
involved_in regulation of transcription by RNA polymerase II
inferred from mutant phenotype
(Roder et al., 1992)
involved_in specification of segmental identity, head
inferred from mutant phenotype
(Laugier et al., 2005)
involved_in specification of segmental identity, thorax
inferred from mutant phenotype
(Roder et al., 1992)
involved_in specification of segmental identity, trunk
inferred from mutant phenotype
(Laugier et al., 2005, Roder et al., 1992)
inferred from mutant phenotype
(Gallet et al., 1998, de Zulueta et al., 1994)
involved_in wing disc proximal/distal pattern formation
inferred from mutant phenotype
(Soanes et al., 2001)
involved_in Wnt signaling pathway
inferred from mutant phenotype
(Gallet et al., 1998)
inferred from physical interaction with UniProtKB:P18824
(Gallet et al., 1998)
Terms Based on Predictions or Assertions (3 terms)
CV Term
Evidence
References
involved_in negative regulation of salivary gland boundary specification
traceable author statement
(Abrams et al., 2003, Bradley et al., 2001)
involved_in regulation of gene expression
inferred from electronic annotation with InterPro:IPR027008
(InterPro Project Members, 2004-)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN000281113
(GO Reference Genome Project, 2011-)
Cellular Component (2 terms)
Terms Based on Experimental Evidence (2 terms)
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Gallet et al., 1998)
located_in nucleus
inferred from direct assay
(Gallet et al., 1998)
inferred from direct assay
(Bessa et al., 2009, Alexandre et al., 1996)
Terms Based on Predictions or Assertions (1 term)
CV Term
Evidence
References
is_active_in nucleus
inferred from biological aspect of ancestor with PANTHER:PTN000281113
(GO Reference Genome Project, 2011-)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the teashirt C2H2-type zinc-finger protein family. (P22265)
Protein Signatures (InterPro)
Summaries
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).
Protein Function (UniProtKB)
Homeotic protein that acts downstream of Arm in the Wg cascade during embryogenesis to determine segment identity throughout the entire trunk. Acts cooperatively with other trunk homeotic proteins to repress head homeotic genes and therefore repress head segmental identity. Necessary, in combination with Scr, for the formation of the prothoracic segment. Promotes eye development in the dorsal region of the eye disk and suppresses eye development in the ventral region in combination with Wg-signaling and several early dorso-ventral eye patterning genes. Required for proper development of proximal leg segments. Has differential functions along the dorso-ventral axs of the antennal and leg disks. May play a role in wing hinge development. Possible involvement in chromatin structure for modulation of transcription. Binds DNA and can act as both a transcriptional repressor and activator. Positively regulates its own expression as well as that of Dll. Negatively regulates the expression of mod. Required for Wg-mediated transcriptional repression of Ubx in the midgut. Also represses transcription of lab in the midgut and is necessary for the proper formation of anterior and central midgut structures. Tiptop (tio) and teashirt (tsh) have, on the whole, common activities. Tio and tsh repress each other's expression and tsh has a crucial role for trunk patterning that is in part masked by ectopic expression of tiptop. Both genes share a common activity required for the activation of Ser and svb and the maintenance of en and wg.
(UniProt, P22265)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
*ae: aeroplane
Wings spread, balancers drooping. Overlaps wild type. RK3.
Summary (Interactive Fly)

transcription factor - zinc finger - required globally for segmental identity throughout the entire trunk - required for identity of the anterior prothorax - required for the subdivision of midgut mesoderm acting in partnership with the homeotics - Restricted teashirt expression confers eye-specific responsiveness to Dpp and Wg signals during eye specification

Gene Model and Products
Number of Transcripts
1
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\tsh 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 P22265)

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 5.44

Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
tsh-RA
FBtr0085906
NM_078891
4732
954
Additional Transcript Data and Comments
Reported size (kB)

8.5, 8.1, 5.4, 5.0 (northern blot)

(Fasano et al., 1991)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
tsh-PA
FBpp0085261
100.6
954
8.64
P22265
NP_523615
AAF57236
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)

993 (aa); 106 (kD)

(Fasano et al., 1991)
Comments

Several isoforms of tsh protein are observed on gels. The smaller 116kD form was determined to be a hypophosphorylated form.The phosphorylated forms have apparent molecular weights of 120~130kD.

(Gallet et al., 1999)

The tsh protein has three separated Zn fingers of the HX5H type which are also found in the hb, human PRDII-BF1,Pep, and Su(var)3-7 proteins. Outside of the Zn-finger region, no clear regions of similarity were observed to other proteins.

(Fasano et al., 1991)
External Data
Subunit Structure (UniProtKB)

Binds arm.

(UniProt, P22265)
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

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

-0.48

Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
head
(Roder et al., 1992)
presumptive embryonic/larval central nervous system
(Fasano et al., 1991)
embryonic midgut constriction 2
(Fasano et al., 1991)
embryonic/larval anus
(Fasano et al., 1991)
epidermis of parasegment 3 -- parasegment 13
(Fasano et al., 1991)
embryonic stage 13
mesothoracic segment
(Roder et al., 1992)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
embryonic/larval pharynx
(Fasano et al., 1991)
somatic trunk mesoderm
(Fasano et al., 1991)
visceral trunk mesoderm
(Fasano et al., 1991)
blastoderm stage
parasegment 3 -- parasegment 13
(Roder et al., 1992)
embryonic stage 4 -- 6
anlage in statu nascendi
(Fisher et al., 2012)
dorsal ectoderm anlage
(Fisher et al., 2012)
organism | gap expression pattern
(Fisher et al., 2012)
trunk mesoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
ventral ectoderm anlage
(Fisher et al., 2012)
embryonic stage 7 -- 8
amnioserosa anlage
(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
organism | gap expression pattern
(Fisher et al., 2012)
trunk mesoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic cycle 14A
organism | 40-60% egg length

Comment: reference states 45-60% egg length

(Fasano et al., 1991)
embryonic stage 9 -- 10
dorsal ectoderm
(Fisher et al., 2012)
organism | gap expression pattern
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
embryonic stage 11 -- 12
dorsal epidermis primordium
(Fisher et al., 2012)
ventral epidermis primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 13
abdominal segment 1 -- abdominal segment 8
(Roder et al., 1992)
embryonic stage 13 -- 16
embryonic dorsal epidermis
(Fisher et al., 2012)
embryonic midgut chamber
(Fisher et al., 2012)
embryonic ventral epidermis
(Fisher et al., 2012)
embryonic/larval midgut
(Fisher et al., 2012)
larval nervous system
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
early third instar larval stage
leg disc | restricted
(Patel et al., 2007)
wandering third instar larval stage
eye disc anterior to the morphogenetic furrow
(Datta et al., 2009)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Fasano et al., 1991)
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
wandering third instar larval stage
eye disc
(Datta et al., 2009)
Additional Descriptive Data

tsh transcript is expressed in the eye disc in the region anterior to the morphogenetic furrow. tsh and tio transcripts are expressed at similar levels in the eye disc.

(Datta et al., 2009)

tsh is expressed in the outer ring of leg discs. It is co-expressed with the disco genes in many of these cells but not in the dorsal (stalk) region.

(Patel et al., 2007)
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 8
parasegment 3 -- parasegment 14
(Bachiller et al., 1994)
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 10
embryonic thoracic segment
(Pages and Kerridge, 2000)
embryonic abdominal segment 1 -- embryonic abdominal segment 8
(Pages and Kerridge, 2000)
embryonic stage 11
epidermis of parasegment 3
(McCormick et al., 1995)
mesoderm of parasegment 3
(McCormick et al., 1995)
epidermis of parasegment 4
(McCormick et al., 1995)
mesoderm of parasegment 4
(McCormick et al., 1995)
epidermis of parasegment 5
(McCormick et al., 1995)
mesoderm of parasegment 5
(McCormick et al., 1995)
larval ventral nerve cord | restricted

Comment: parasegments 3-13

(McCormick et al., 1995)
embryonic stage 13 -- 17
embryonic Malpighian tubule Type II cell
(Denholm et al., 2013)
embryonic stage 15
embryonic Malpighian tubule | restricted
(Denholm et al., 2003)
embryonic stage 16
embryonic/larval Malpighian tubule Type II cell
(Zohar-Stoopel et al., 2014)
first instar larval stage
Malpighian tubule Type II cell
(Tapadia and Gautam, 2011)
second instar larval stage
wing disc
(Wu and Cohen, 2002)
second instar larval stage -- third instar larval stage
eye disc
(Bessa et al., 2002)
third instar larval stage
wing disc
(Wu and Cohen, 2002)
eye disc | anterior
(Bessa et al., 2009)
leg disc | proximal
(Bessa et al., 2009)
wing disc | proximal
(Bessa et al., 2009)
embryonic/larval Malpighian tubule Type II cell
(Denholm et al., 2013)
early third instar larval stage
eye-antennal disc | restricted
(Luque and Milan, 2007)
wandering third instar larval stage
eye disc anterior to the morphogenetic furrow | restricted
(Bessa et al., 2002)
mesothoracic tergum primordium
(Nusinow et al., 2008)
wing hinge primordium
(Nusinow et al., 2008)
adult stage
adult Malpighian tubule Type II cell
(Denholm et al., 2013)
Additional Descriptive Data

tsh protein is present in the anterior part of the eye disc and in the proximal domains of the leg and wing discs in third instar larvae. The expression domain overlaps that of tio.

(Bessa et al., 2009)

It appears that the hypophosphorylated form of tsh protein is largely cytoplasmic while the phosphorylated forms are nuclear.

(Nusinow et al., 2008, Gallet et al., 1999)

Expression of the tsh protein was detected in stage 15 embryonic Malpighian tubule stellate cells.

(Denholm et al., 2003)

tsh protein is expressed in all cells of the second instar eye disc; expression levels are lower in the posterior disc. By late third instar, tsh protein is expressed in the eye disc anterior to the morphogenetic furrow, but not in the anterior-most part of the eye disc, where it contacts the antennal disc.

(Bessa et al., 2002)

tsh protein is expressed in the presumptive notum region of the wing disc and repressed in the presumptive wing region

(Wu and Cohen, 2002)

At stages 5-8, the faster, nonphosphorylated tsh protein is detected in the region of the first three thoracic and the first abdominal segments as a diffuse signal within the nuclei and cytoplasm of all cells. Between stages 9 and 11, the larger form becomes more apparent. tsh protein becomes highly concentrated in nuclei in the posterior part of each segment. Anteriorly in the segments, it is found both in the nuclei and the cytoplasm and is still diffuse. In later stages, the larger forms are more predominant and staining is high in all nuclei.

(Gallet et al., 1999)

tsh expression is specific to the trunk region of the developing embryo and is responsive to changes in homeotic gene expression.

(Bachiller et al., 1994)
Marker for
 
Malpighian tubule Type II cell
(Tapadia and Gautam, 2011)
embryonic/larval Malpighian tubule Type II cell
(Zohar-Stoopel et al., 2014)
Subcellular Localization
CV Term
Evidence
References
located_in cytoplasm
inferred from direct assay
(Gallet et al., 1998)
located_in nucleus
inferred from direct assay
(Gallet et al., 1998)
inferred from direct assay
(Bessa et al., 2009, Alexandre et al., 1996)
Expression Deduced from Reporters
Reporter: P{A92}tsh1
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
visceral trunk mesoderm
(Fasano et al., 1991)
somatic trunk mesoderm
(Fasano et al., 1991)
central nervous system
(Fasano et al., 1991)
embryonic/larval anus
(Fasano et al., 1991)
embryonic stage
presumptive embryonic/larval peripheral nervous system
(Fasano et al., 1991)
embryonic midgut constriction 2
(Fasano et al., 1991)
embryonic/larval pharynx
(Fasano et al., 1991)
epidermis of parasegment 3 -- parasegment 13
(Fasano et al., 1991)
larval stage
embryonic/larval midgut
(Lemaitre and Coen, 1991)
imaginal disc
(Lemaitre and Coen, 1991)
larval brain
(Lemaitre and Coen, 1991)
embryonic/larval cuticle
(Lemaitre and Coen, 1991)
wing disc | restricted
(Zecca and Struhl, 2002)
wing hinge | presumptive
(Zecca and Struhl, 2002)
adult stage
axillary sclerite
(Casares and Mann, 2000)
pleural wing sclerite
(Casares and Mann, 2000)
Reporter: P{GawB}tshc724
Stage
Tissue/Position (including subcellular localization)
Reference
larval stage
Malpighian tubule Type II cell
(Sozen et al., 1997)
adult stage
Malpighian tubule Type II cell
(Sozen et al., 1997)
initial segment of Malpighian tubule | restricted
(Sozen et al., 1997)
embryonic stage 15
embryonic Malpighian tubule | restricted
(Denholm et al., 2003)
embryonic stage 17
embryonic/larval Malpighian tubule Type II cell
(Gautam and Tapadia, 2010)
embryonic/larval salivary gland
(Gautam and Tapadia, 2010)
larval stage
Malpighian tubule Type II cell
(Sozen et al., 1997)
first instar larval stage
embryonic/larval Malpighian tubule Type II cell
(Gautam and Tapadia, 2010)
embryonic/larval salivary gland
(Gautam and Tapadia, 2010)
larval visceral muscle cell
(Gautam and Tapadia, 2010)
adult stage
initial segment of Malpighian tubule
(Sozen et al., 1997)
Malpighian tubule Type II cell
(Sozen et al., 1997)
adult Malpighian tubule stellate cell of main segment
(Xu et al., 2023, Cabrero et al., 2020)
adult Malpighian tubule Type II cell
(Rosay et al., 1997)
adult Malpighian tubule bar-shaped cell
(Rosay et al., 1997)
Reporter: P{GawB}tshGal4-1
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
peripodial epithelium
(Shiga et al., 1996)
wing hinge | presumptive
(Shiga et al., 1996)
wing disc | restricted
(Tomoyasu et al., 1998, Shiga et al., 1996)
mesothoracic tergum primordium
(Tomoyasu et al., 1998)
Reporter: P{GawB}tshmd621
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage | male
larval abdominal segment 8 of male genital disc
(Petzoldt et al., 2012)
early third instar larval stage -- prepupal stage P4
abdominal segment 8 of male genital disc
(Speder et al., 2006)
wandering third instar larval stage
wing hinge primordium
(Peterson and O'Connor, 2013, Pallavi and Shashidhara, 2003)
mesothoracic tergum primordium
(Peterson and O'Connor, 2013, Pallavi and Shashidhara, 2003)
peripodial epithelium | restricted of wing disc
(Pallavi and Shashidhara, 2003)
adult stage
adult fruitless aSP-g (female) neuron
(Yu et al., 2010)
peripodial epithelium | restricted of wing disc | restricted
(Gibson et al., 2002)
Reporter: P{lacW}A(2)-6
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
(Sun et al., 1995)
third instar larval stage
eye disc | restricted
(Sun et al., 1995)
wing disc | restricted
(Sun et al., 1995)
leg disc | restricted
(Sun et al., 1995)
antennal disc | restricted
(Sun et al., 1995)
Reporter: P{lacW}tshB4-2-12
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 11
ectoderm of abdominal segment
(Bier et al., 1989)
ectoderm of thoracic segment
(Bier et al., 1989)
embryonic stage 11 -- 17
epidermis of prothoracic segment
(Hartenstein and Jan, 1992)
epidermis of abdominal segment 1 -- abdominal segment 9
(Hartenstein and Jan, 1992)
epidermis of mesothoracic segment
(Hartenstein and Jan, 1992)
embryonic ventral nerve cord of mesothoracic segment
(Hartenstein and Jan, 1992)
epidermis of metathoracic segment
(Hartenstein and Jan, 1992)
embryonic ventral nerve cord of prothoracic segment
(Hartenstein and Jan, 1992)
embryonic ventral nerve cord of metathoracic segment
(Hartenstein and Jan, 1992)
abdominal segment 1 -- abdominal segment 9 of embryonic ventral nerve cord
(Hartenstein and Jan, 1992)
embryonic stage 15
larval ventral nerve cord
(Bier et al., 1989)
Reporter: P{PZ}tsh04319
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
embryonic Malpighian tubule | restricted
(Denholm et al., 2003)
presumptive embryonic/larval central nervous system
(Jackle and Janning, 1998.8.10)
embryonic/larval midgut
(Jackle and Janning, 1998.8.10)
embryonic/larval anterior midgut
(Jackle and Janning, 1998.8.10)
epidermis
(Jackle and Janning, 1998.8.10)
Malpighian tubule Type II cell
(Denholm et al., 2003)
embryonic/larval pharynx
(Jackle and Janning, 1998.8.10)
wandering third instar larval stage
imaginal disc
(Meister and Braun, 1995.10)
embryonic/larval midgut
(Meister and Braun, 1995.10)
Malpighian tubule
(Meister and Braun, 1995.10)
adult stage
adult Malpighian tubule Type II cell
(Wang et al., 2004)
Reporter: P{UpP65L}tshmd621-lexA
Stage
Tissue/Position (including subcellular localization)
Reference
first instar larval stage -- third instar larval stage
neuron of larval ventral nerve cord
(Eschbach et al., 2021)
adult stage
neuron of adult ventral nerve cord
(Sherer et al., 2020)
High-Throughput Expression Data
Associated Tools

JBrowse - Visual display of RNA-Seq signals

View Dmel\tsh in JBrowse
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 48 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 41 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of tsh
Transgenic constructs containing regulatory region of tsh
Aberrations (Deficiencies and Duplications) ( 14 )
Variants
Variant Molecular Consequences
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal (with Df(2L)305)
lethal (with Df(2L)BSC150)
lethal | heat sensitive
lethal | recessive
lethal - all die before end of pupal stage, with Scer\GAL4dpp.blk1
lethal - all die during embryonic stage, with Scer\GAL448Y
lethal - all die during embryonic stage | recessive
partially lethal, with Scer\GAL4tsh-c724
viable
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4tin.CΔ4
viable (with Df(2L)BSC102)
viable (with tsh8)
viable (with tshrAO135)
Other Phenotypes
Allele
abnormal cell shape | somatic clone, with Scer\GAL4αTub84B.PZ
abnormal developmental rate, with Scer\GAL4dpp.blk1
abnormal mitotic cell cycle | recessive
abnormal planar polarity (with Df(2L)305)
flightless
flightless | recessive
increased body size, with Scer\GAL4dpp.blk1
increased cell number | larval stage | somatic clone, with Scer\GAL4αTub84B.PZ
short lived, with Scer\GAL4tsh-c724
short lived | heat sensitive, with Scer\GAL4tin.CΔ4
some die during embryonic stage, with Scer\GAL448Y
some die during embryonic stage | recessive
some die during pupal stage, with Scer\GAL4dpp.blk1
some die during pupal stage, with Scer\GAL4tsh-c724
visible
visible, with Scer\GAL4ap-md544
visible, with Scer\GAL4bi-omb-Gal4
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4cb26
visible, with Scer\GAL4cb41
visible, with Scer\GAL4dpp.blk1
visible, with Scer\GAL4dpp.PS
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4GMR.PF
visible, with Scer\GAL4Ser.PGF
visible, with Scer\GAL4tsh-c724
visible | dominant
visible | homeotic
visible | homeotic, with Scer\GAL469B
visible | recessive
visible | somatic clone
visible | somatic clone, with Scer\GAL4Act5C.PI
visible | somatic clone, with Scer\GAL4Ubx.PdC
Phenotype manifest in
Allele
adult abdomen, with Scer\GAL4tsh-c724
adult head
adult Malpighian tubule, with Scer\GAL4tsh-c724
adult Malpighian tubule Type II cell, with Scer\GAL4tsh-c724
antenna & eye | ectopic
antenna | somatic clone, with Scer\GAL4Act5C.PI
antennal disc, with Scer\GAL4dpp.PS
antennal disc | ectopic | third instar larval stage, with Scer\GAL4cb26
antennal disc | ectopic | third instar larval stage, with Scer\GAL4cb41
antennal disc | ectopic | third instar larval stage, with Scer\GAL4dpp.PS
antennal disc | ectopic | third instar larval stage, with Scer\GAL4Ser.PGF
antennal disc | somatic clone, with Scer\GAL4FRT.Act5C
antennal disc | somatic clone | ventral, with Scer\GAL4FRT.Act5C
anterior scutellar bristle
arista
copper cell, with Scer\GAL448Y
coxa
coxa (with Df(2L)305)
coxa (with tshHD1)
coxa (with tshmd621)
denticle, with Scer\GAL469B
denticle belt & embryonic prothoracic segment
denticle row 1
denticle row 2
denticle row 3
denticle row 4
denticle row 5 | increased number
embryo | dorsal closure stage, with Scer\GAL469B
embryonic/first instar larval cuticle
embryonic/first instar larval cuticle, with Scer\GAL469B
embryonic/first instar larval cuticle, with Scer\GAL4hs.PB
embryonic/larval cuticle
embryonic antennal segment, with Scer\GAL469B
embryonic head, with Scer\GAL469B
embryonic labial segment, with Scer\GAL469B
embryonic labial segment | ectopic
embryonic maxillary segment, with Scer\GAL469B
embryonic midgut constriction 1
embryonic midgut constriction 2
embryonic prothoracic segment
embryonic prothoracic segment | ectopic, with Scer\GAL469B
eye, with Scer\GAL4arm.PS, Scer\GAL4ey.PH
eye, with Scer\GAL4dpp.blk1
eye, with Scer\GAL4ey.PH
eye | dorsal, with Scer\GAL4bi-omb-Gal4
eye | dorsal | somatic clone
eye | dorsal | somatic clone, with Scer\GAL4Act5C.PI
eye | ectopic
eye | ectopic, with Scer\GAL4Bx-MS1096
eye | ectopic, with Scer\GAL4cb26
eye | ectopic, with Scer\GAL4cb41
eye | ectopic, with Scer\GAL4dpp.blk1
eye | ectopic, with Scer\GAL4dpp.PS
eye | ectopic, with Scer\GAL4MD705
eye | ectopic, with Scer\GAL4Ser.PGF
eye | ectopic | somatic clone, with Scer\GAL4Act5C.PI
eye | somatic clone, with Scer\GAL4Act5C.PI
eye | somatic clone | ventral
eye | somatic clone | ventral, with Scer\GAL4Act5C.PI
eye | ventral, with Scer\GAL4bi-omb-Gal4
eye-antennal disc, with Scer\GAL4dpp.blk1
eye disc, with Scer\GAL4bi-omb-Gal4
eye disc, with Scer\GAL4ey.PH
eye disc | somatic clone, with Scer\GAL4αTub84B.PZ
eye disc | somatic clone | ventral, with Scer\GAL4Act5C.PI
eye photoreceptor cell, with Scer\GAL4GMR.PF
femur
femur (with tshHD1)
femur (with tshmd621)
femur | proximal (with Df(2L)305)
femur | somatic clone, with Scer\GAL4Act5C.PP
femur | somatic clone, with Scer\GAL4VP16.arm
haltere
haltere sclerite
haltere sclerite (with tsh8.1)
haltere sclerite (with tshae-1)
head & eye | ectopic
interommatidial bristle | ectopic, with Scer\GAL4Bx-MS1096
Keilin organ
labial segment | heat sensitive
larval abdomen
larval abdominal segment 1
larval antennal segment & denticle | ectopic, with Scer\GAL469B
larval head, with Scer\GAL469B
larval labial segment, with Scer\GAL469B
larval maxillary segment & denticle | ectopic, with Scer\GAL469B
larval mesothoracic segment
larval thorax
leg
leg (with tshHD1)
leg (with tshmd621)
leg & cuticle | dorsal | somatic clone, with Scer\GAL4Act5C.PP
leg & cuticle | dorsal | somatic clone, with Scer\GAL4VP16.arm
leg & cuticle | somatic clone, with Scer\GAL4Act5C.PP
leg & cuticle | somatic clone, with Scer\GAL4VP16.arm
leg & cuticle | ventral | somatic clone, with Scer\GAL4Act5C.PP
leg & cuticle | ventral | somatic clone, with Scer\GAL4VP16.arm
leg | ectopic
leg disc | somatic clone, with Scer\GAL4Act5C.PP
leg disc | somatic clone, with Scer\GAL4VP16.arm
maxillary palp
maxillary segment | heat sensitive
mesothoracic coxal bristle on hairy island
mesothoracic coxal bristle on hairy island (with tshHD1)
mesothoracic coxal bristle on hairy island (with tshmd621)
mesothoracic coxal trichoid sensillum
mesothoracic coxal trichoid sensillum (with tshHD1)
mesothoracic coxal trichoid sensillum (with tshmd621)
mesothoracic femoral bract | somatic clone, with Scer\GAL4Act5C.PP
mesothoracic femoral bract | somatic clone, with Scer\GAL4VP16.arm
metaphase & condensed nuclear chromosome
metathoracic coxal trichoid sensillum
metathoracic coxal trichoid sensillum (with tshHD1)
metathoracic coxal trichoid sensillum (with tshmd621)
metathoracic femoral bract | somatic clone, with Scer\GAL4Act5C.PP
metathoracic femoral bract | somatic clone, with Scer\GAL4VP16.arm
mitotic anaphase
mitotic cell cycle
mitotic metaphase
photoreceptor neuron | ectopic, with Scer\GAL4Bx-MS1096
photoreceptor neuron | ectopic, with Scer\GAL4hs.PB
photoreceptor neuron | ectopic, with Scer\GAL4MD705
photoreceptor neuron | somatic clone, with Scer\GAL4Act5C.PI
procephalic segment | heat sensitive
prothoracic coxal bristle on hairy island
prothoracic coxal bristle on hairy island (with tshHD1)
prothoracic coxal bristle on hairy island (with tshmd621)
prothoracic coxal trichoid sensillum
prothoracic coxal trichoid sensillum (with tshHD1)
prothoracic coxal trichoid sensillum (with tshmd621)
prothoracic femoral bract | somatic clone, with Scer\GAL4Act5C.PP
prothoracic femoral bract | somatic clone, with Scer\GAL4VP16.arm
prothoracic segment
T1 beard, with Scer\GAL469B
T1 beard | ectopic, with Scer\GAL469B
tibia | somatic clone, with Scer\GAL4Act5C.PP
tibia | somatic clone, with Scer\GAL4VP16.arm
trochanter
trochanter (with Df(2L)305)
trochanter (with tshHD1)
trochanter (with tshmd621)
ventral denticle belt
ventral denticle belt, with Scer\GAL469B
ventral denticle belt | heat sensitive
ventral denticle belt | larval stage
vertical cleft (with tsh8.1)
vertical cleft (with tshae-1)
wing
wing | somatic clone, with Scer\GAL4Ubx.PdC
wing blade dorsal compartment, with Scer\GAL4ap-md544
wing blade dorsal compartment, with Scer\GAL4Bx-MS1096
wing disc, with Scer\GAL4bi-omb-Gal4
wing disc, with Scer\GAL4dpp.blk1
wing disc, with Scer\GAL4sd-SG29.1
wing hinge
wing hinge (with tsh8.1)
wing hinge (with tshae-1)
wing hinge | ectopic, with Scer\GAL4ap-md544
wing hinge | ectopic, with Scer\GAL4Bx-MS1096
wing hinge | ectopic, with Scer\GAL4Ubx.PdC
wing margin
wing sclerite
wing sclerite (with tsh8.1)
wing sclerite (with tshae-1)
wing sclerite | ectopic, with Scer\GAL4ap-md544
wing sclerite | ectopic, with Scer\GAL4Bx-MS1096
wing sclerite | ectopic, with Scer\GAL4Ubx.PdC
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
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Homo sapiens (Human) (3)
Hsap\TSHZ2
6 of 14
Yes
Yes
Hsap\TSHZ1
5 of 14
No
Yes
Hsap\TSHZ3
5 of 14
No
Yes
Model Organism Orthologs (via DIOPT v9.1)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
Domainoid
eggNOG
Hieranoid
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Complementation?
Transgene?
Rattus norvegicus (Norway rat) (3)
Rnor\Tshz3
5 of 14
Yes
Yes
Rnor\Tshz1
4 of 14
No
Yes
Rnor\Tshz2
4 of 14
No
Yes
Mus musculus (laboratory mouse) (3)
Mmus\Tshz2
6 of 14
Yes
Yes
Mmus\Tshz1
5 of 14
No
Yes
Mmus\Tshz3
5 of 14
No
Yes
Xenopus tropicalis (Western clawed frog) (4)
Xtro\tshz1
3 of 13
Yes
No
Xtro\tshz2
3 of 13
Yes
Yes
Xtro\tshz3
3 of 13
Yes
Yes
Xtro\shq1
1 of 13
No
No
Danio rerio (Zebrafish) (4)
Drer\tshz2
6 of 14
Yes
Yes
Drer\tshz3a
6 of 14
Yes
Yes
Drer\tshz1
5 of 14
No
Yes
Drer\tshz3b
5 of 14
No
Yes
Caenorhabditis elegans (Nematode, roundworm) (0)
Anopheles gambiae (African malaria mosquito) (1)
Agam\AgaP_AGAP009028
4 of 12
Yes
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:tsh. Refer to their site for version information.
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v9.1)
Gene Symbol
Score
Source
Compara
Domainoid
eggNOG
Homologene
Inparanoid
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
SonicParanoid
Alignment
Drosophila melanogaster (Fruit fly) (1)
tio
8 of 13
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 5 )
    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?
    TSHZ2; teashirt zinc finger homeobox 2
    6 of 14
    614118
        TSHZ1; teashirt zinc finger homeobox 1
        5 of 14
        614427
          TSHZ3; teashirt zinc finger homeobox 3
          5 of 14
          614119
              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
              protein-protein
              Physical Interaction
              Assay
              References
              tsh - Abd-B
              bimolecular fluorescence complementation, fluorescence microscopy
              (Baëza et al., 2015)
              tsh - Antp
              bimolecular fluorescence complementation, fluorescence microscopy, pull down, autoradiography
              (Baëza et al., 2015, Taghli-Lamallem et al., 2007)
              tsh - CG15098
              anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
              (Rhee et al., 2014)
              tsh - CtBP
              anti tag coimmunoprecipitation, anti tag western blot
              (Datta et al., 2011)
              tsh - Ppt1
              anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
              (Rhee et al., 2014)
              tsh - Sat
              anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
              (Rhee et al., 2014)
              tsh - Scr
              bimolecular fluorescence complementation, fluorescence microscopy, pull down, autoradiography, two hybrid
              (Baëza et al., 2015, Taghli-Lamallem et al., 2007)
              tsh - Ubx
              bimolecular fluorescence complementation, fluorescence microscopy
              (Bischof et al., 2018, Baëza et al., 2015)
              tsh - abd-A
              bimolecular fluorescence complementation, fluorescence microscopy
              (Bischof et al., 2018, Baëza et al., 2015, Hudry et al., 2011)
              tsh - arm
              pull down, autoradiography, anti bait coimmunoprecipitation, western blot
              (Gallet et al., 1998)
              tsh - cindr
              anti tag coimmunoprecipitation, peptide massfingerprinting, experimental knowledge based
              (Rhee et al., 2014)
              tsh - exd
              bimolecular fluorescence complementation, fluorescence microscopy, pull down, autoradiography
              (Bischof et al., 2018, Bessa et al., 2002)
              tsh - hth
              pull down, autoradiography, anti tag coimmunoprecipitation, anti tag western blot, anti bait coimmunoprecipitation, western blot
              (Datta et al., 2011, Bessa et al., 2002)
              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
              tsh
              suppressible
              Axn
              (Bessa and Casares, 2005)
              tsh
              enhanceable
              brk
              (Saller et al., 2002)
              tsh|Dfd
              enhanceable
              disco
              (Robertson et al., 2004)
              tsh
              suppressible
              hth
              (Singh et al., 2002)
              tsh
              enhanceable
              spen
              (Wiellette et al., 1999)
              tsh
              suppressible
              tio
              (Denholm et al., 2013, Laugier et al., 2005)
              tsh
              suppressible
              wg
              (Singh et al., 2002)
              Starting gene(s)
              Interaction type
              Interacting gene(s)
              Reference
              Antp
              enhanceable
              tsh
              (Bhojwani et al., 1997)
              arm
              suppressible
              tsh
              (Gallet et al., 1998)
              brk
              enhanceable
              tsh
              (Saller et al., 2002)
              ci
              suppressible
              tsh
              (Angelats et al., 2002)
              Delta
              enhanceable
              tsh
              (Singh et al., 2004)
              Dfd
              enhanceable
              tsh
              (Robertson et al., 2004)
              gt
              suppressible
              tsh
              (Sanders et al., 2008)
              hng1
              suppressible
              tsh
              (Shukla et al., 2014)
              Ser
              suppressible
              tsh
              (Singh et al., 2004)
              spen
              suppressible
              tsh
              (Wiellette et al., 1999)
              External Data
              Subunit Structure (UniProtKB)
              Binds arm.
              (UniProt, P22265 )
              Linkouts
              BioGRID - A database of protein and genetic interactions.
              DroID - A comprehensive database of gene and protein interactions.
              MIST (genetic) - An integrated Molecular Interaction Database
              MIST (protein-protein) - An integrated Molecular Interaction Database
              Pathways
              Signaling Pathways (FlyBase)
              Metabolic Pathways
              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.
              SignaLink - A signaling pathway resource with multi-layered regulatory networks.
              Class of Gene
              Genomic Location and Detailed Mapping Data
              Chromosome (arm)
              2L
              Recombination map
              2-54
              Cytogenetic map
              40A5-40A5
              Sequence location
              FlyBase Computed Cytological Location
              Cytogenetic map
              Evidence for location
              40A5-40A5
              Limits computationally determined from genome sequence between P{lacW}Df31k05815 and P{PZ}cul-202074&P{lacW}l(2)k08110k08110
              Experimentally Determined Cytological Location
              Cytogenetic map
              Notes
              References
              40A1-40A4
              (determined by in situ hybridization)
              (Spradling et al., 1999)
              40A1-40A4
              (BDGP Project Members, 1994-1999)
              40A-40C
              (determined by in situ hybridization)
              (Hartenstein and Jan, 1992)
              40A-40A
              (determined by in situ hybridization)
              (Fasano et al., 1991)
              Experimentally Determined Recombination Data
              Location

              2-54

              (FlyBase, 2016)

              2-54.6

              (Comeron et al., 2012, Soanes and Bell, 1999)

              2-55.8

              Left of (cM)
              (Soanes and Bell, 1999)
              Right of (cM)
              (Soanes and Bell, 1999)
              Notes

              Map position determined using tshae-l.

              (Soanes and Bell, 1999)
              Stocks and Reagents
              Stocks (49)
              Genomic Clones (16)
               

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

              cDNA Clones (42)
               

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

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

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

              cDNA Clones, End Sequenced (ESTs)
              BDGP DGC clones
              Other clones
              RNAi and Array Information
              Linkouts
              DRSC - Results frm RNAi screens
              Antibody Information
              Laboratory Generated Antibodies
               
              Commercially Available Antibodies
               
              Cell Line Information
              Publicly Available Cell Lines
               
                Other Stable Cell Lines
                 
                  Other Comments

                  tsh is required in undifferentiated cells for eye specification, but its expression must later be turned off to allow retinal differentiation.

                  (Bessa and Casares, 2005)

                  dsRNA made from templates generated with primers directed against this gene and injected into embryos on the dorsal side causes a phenotype at the larval stage. These larvae have no T1 denticle beard and also show a reduced number of denticle rows in the abdominal segments compared to wild type.

                  (Laugier et al., 2005)

                  tsh and tio repress each other's expression.

                  (Laugier et al., 2005)

                  The primary signal for repression of tsh expression during wing development is wg.

                  (Zirin and Mann, 2004)

                  Different combinations of the proteins ci, tsh and arm appear to be employed for the specification of naked cuticle at distinct positions both along the anterior posterior axis and within individual trunk segments.

                  (Angelats et al., 2002)

                  ci, tsh and arm may act as a complex.

                  (Angelats et al., 2002)

                  tsh is required for thr formation of proximal leg segments, but has no role in boundary formation.

                  (Wu and Cohen, 2000)

                  tsh contributes to the differences in cell-cell adhesion between proximal and distal leg cells.

                  (Erkner et al., 1999)

                  wg signaling promotes the phosphorylation and the nuclear accumulation of tsh protein.

                  (Gallet et al., 1999)

                  spen cooperates with Antp and tsh to repress head-like sclerites in the thorax and promote thoracic identity.

                  (Wiellette et al., 1999)

                  tsh protein binds to, and requires, arm protein for the naked cell-fate choice in the larval trunk.

                  (Gallet et al., 1998)

                  tsh and ey induce the expression of each other, acting in a gene network that functions to specify eye identity.

                  (Pan and Rubin, 1998)

                  Targeted expression of tsh in imaginal discs is sufficient to induce ectopic eye formation in non-eye tissues. tsh functions with the other eye specification genes ey, so and dac to specify eye identity.

                  (Pan and Rubin, 1998)

                  tsh specifies adult head segments by repressing Antp expression.

                  (Bhojwani et al., 1997)

                  mod is a genetic target of tsh. A biological role of tsh is to repress mod expression in the ectoderm and this negative control is performed independently of Scr. tsh protein binds to specific DNA sequences within a 5' mod regulatory region that reproduces tsh-dependent expression in the ectoderm.

                  (Alexandre et al., 1996)

                  P{wAR}4-3 localizes to the 3' end of the tsh gene.

                  (Kerridge, 1996.1.31)

                  tsh expression in the visceral mesoderm is required for the development of anterior and central midgut structures and is regulated by homeotic genes and dpp and wg.

                  (Mathies et al., 1995)

                  tsh may act as a cofactor for homeotic function, acting in parallel to Antp in epidermal enhancer activity in the thorax while playing an antagonistic role to Ubx in somatic mesoderm.

                  (McCormick et al., 1995)

                  A homeotic response element (HOMRE) has been identified and characterized within the tsh gene. Different homeotic proteins control tsh enhancer activity in different tissues.

                  (McCormick et al., 1995)

                  Expression pattern of Ecol\lacZ from silencer detector insertion P{lacW}A(2)-6 reflects that of tsh.

                  (Sun et al., 1995)

                  Salivary gland induction by Scr in the trunk is limited by tsh.

                  (Andrew et al., 1994)

                  tsh is necessary for proper formation of anterior and central midgut structures. Antp, Ubx, abd-A, dpp and wg are required for proper tsh expression. The control of tsh by Ubx, abd-A and probably also by Antp is mediated by secreted signaling molecules.

                  (Mathies et al., 1994)

                  tsh is a target gene of the homeotic selector genes and is regulated by exd.

                  (Rauskolb and Wieschaus, 1994)

                  Ectopic expression of dpp eliminates Scr and Antp expression, attenuating abd-A expression, inducing Ubx, dpp, wg and tsh expression in the visceral mesoderm and inducing lab expression in the apposing endoderm. The result is failure of all of the morphogenetic events except formation of midgut constriction 2.

                  (Staehling-Hampton and Hoffmann, 1994)

                  tsh acts in a partially redundant way with certain homeotic complex proteins but co-operates with them for the establishment of specific segment types. The tsh product and homeotic complex proteins regulate common sets of downstream target genes.

                  (de Zulueta et al., 1994)

                  In the epidermis tsh is expressed in a parasegmental and then a segmental register whereas in the CNS its expression remains parasegmental. The maintenance of tsh expression in the posterior compartments of trunk segments is dependent on the common function of Antp and the bithorax complex genes. Scr and lab are expressed ectopically in embryos deficient for tsh and Antp. Antp and Ubx act independently of tsh for the determination of trunk identity, although tsh is essential for the complete function of the trunk homeotic genes.

                  (Roder et al., 1992)

                  tsh is required for normal development of the cells giving rise to the labial prothoracic region, ventral trunk region and anal opening.

                  (Fasano et al., 1991)

                  Mutants have outheld wings.

                  Relationship to Other Genes
                  Source for database merge of

                  Source for merge of: tsh ae

                  (Soanes and Bell, 2000)
                  Additional comments

                  Complementation tests between tshae-l and various other tsh alleles define a distinct complementation group though molecular analysis places a tshae-l lesion very close to the 3' end of the tsh transcription unit, between the 3' end of tsh and previously defined regulatory sequences.

                  (Soanes and Bell, 1999)
                  Nomenclature History
                  Source for database identify of

                  Source for identity of: tsh CG1374

                  (Deng, 1999.11)
                  Nomenclature comments
                  Etymology
                  Synonyms and Secondary IDs (13)
                  Reported As
                  Symbol Synonym
                  CG1374
                  (National Institute of Genetics Fly Stocks, 2022-, Meadows, 2017.9.14, Neuert et al., 2017, Datta et al., 2011, Ni et al., 2011.7.19, Perkins et al., 2009)
                  T shirt
                  (Chinnadurai, 2003)
                  TSH
                  (Harvey and Hariharan, 2012)
                  Tsh
                  (Gou et al., 2023, Gracia-Latorre et al., 2022, Gogia et al., 2021, Chen and Desplan, 2020, Ruiz-Losada et al., 2018, Davis and Rebay, 2017, Apitz and Salecker, 2016, Fallahi et al., 2016, Baëza et al., 2015, Irvine and Harvey, 2015, Zohar-Stoopel et al., 2014, Enderle and McNeill, 2013, Wang et al., 2013, Weasner and Kumar, 2013, Weavers and Skaer, 2013, Baker and Firth, 2011, Datta et al., 2011, Halder and Johnson, 2011, Hudry et al., 2011, Lynch and Wagner, 2011, Papadopoulos et al., 2011, Zhao et al., 2011, Benítez et al., 2009, Campbell et al., 2009, Gambetta et al., 2009, McKay et al., 2009, Urbano et al., 2009, Ayala and Bach, 2008, Datta and Kumar, 2008, Nusinow et al., 2008, Rafel and Milán, 2008, Lan et al., 2007, Luque and Milan, 2007, Zirin and Mann, 2007, Venken et al., 2006)
                  ae
                  (Soanes and Bell, 1999, Lindsley and Zimm, 1992)
                  ael
                  (Soanes and Bell, 1999)
                  l(2)04319
                  l(2)B4-2-12
                  tsh
                  (Gemeinhardt et al., 2025, Ricolo et al., 2025, Ko et al., 2024, Pollex et al., 2024, Voutyraki et al., 2023, Zhang et al., 2023, Clarembaux-Badell et al., 2022, Deshpande et al., 2022, Dow et al., 2022, Messina et al., 2022, National Institute of Genetics Fly Stocks, 2022-, Winant et al., 2022, Casares and McGregor, 2021, Dow et al., 2021, Loganathan et al., 2021, Loker et al., 2021, Poliacikova et al., 2021, Winbush and Singh, 2021, Xie et al., 2021, Jonusaite et al., 2020, Overton et al., 2020, Wang and Spradling, 2020, Gaspar et al., 2019, Tiwari et al., 2019, Baker et al., 2018, Bischof et al., 2018, Palliyil et al., 2018, Sander et al., 2018, Yeung et al., 2018, Zhu et al., 2018, Karaiskos et al., 2017, Lee et al., 2017, Neto et al., 2017, Neuert et al., 2017, Transgenic RNAi Project members, 2017-, Zhu et al., 2017, Zouaz et al., 2017, Carrasco-Rando et al., 2016, Neto et al., 2016, Seyres et al., 2016, Shlyueva et al., 2016, Testa and Dworkin, 2016, Voutev and Mann, 2016, Doggett et al., 2015, Schertel et al., 2015, Ugrankar et al., 2015, Orsi et al., 2014, Piñeiro et al., 2014, Shukla et al., 2014, Bateman et al., 2013, Denholm et al., 2013, Jin et al., 2013, Merabet and Hudry, 2013, Moran et al., 2013, Schaub and Frasch, 2013, Southall et al., 2013, Berni et al., 2012, Foronda et al., 2012, Hudry et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Nfonsam et al., 2012, Brockmann et al., 2011, Cherbas et al., 2011, Datta et al., 2011, Datta et al., 2011, Tapadia and Gautam, 2011, Zhang et al., 2011, Campbell et al., 2010, Cunha et al., 2010, Estella and Mann, 2010, Lopes and Casares, 2010, Oros et al., 2010, Rubinstein et al., 2010, Ayroles et al., 2009, Bao and Friedrich, 2009, Bessa et al., 2009, Datta et al., 2009, Dichtel-Danjoy et al., 2009, Fang et al., 2009, Firth and Baker, 2009, Gambetta et al., 2009, Grieder et al., 2009, Peng et al., 2009, Perea et al., 2009, Salzer and Kumar, 2009, Anderson and Kumar, 2008, Chen et al., 2008, Coiffier et al., 2008, Oktaba et al., 2008, Sanders et al., 2008, Takacs et al., 2008, Terriente et al., 2008, Kumar and Datta, 2007, Kumar and Luhur, 2007, Kumar and Salzer, 2007, Merabet et al., 2007, Minidorff et al., 2007, Patel et al., 2007, Peng et al., 2007, Singh et al., 2007, Taghli-Lamallem et al., 2007, Tanaka et al., 2007, Zirin and Mann, 2007, Culi et al., 2006, Dworkin and Gibson, 2006, Dworkin and Gibson., 2006, Friedrich, 2006, Jordan et al., 2006, Kim et al., 2006, Noro et al., 2006, Venken, 2006, Akimoto et al., 2005, Pütz et al., 2005, Roederer et al., 2005, Grad et al., 2004, Wang et al., 2004, Chauvet et al., 2000)
                  Name Synonyms
                  T-shirt
                  (Foronda et al., 2009)
                  Teashirt
                  (Choubey et al., 2020, Harvey et al., 2013, Wang et al., 2013, Datta et al., 2011, Halder and Johnson, 2011, Hudry et al., 2011, Itasaki and Hoppler, 2010, Peng et al., 2009, Urbano et al., 2009, Ayala and Bach, 2008, Dougherty et al., 2008, Ishihara and Shibata, 2008, Rafel and Milán, 2008, Luque and Milan, 2007, Taghli-Lamallem et al., 2007, Zirin and Mann, 2007)
                  aeroplane
                  (Lindsley and Zimm, 1992)
                  teashirt
                  (Cohen et al., 2020, Neto et al., 2016, Baëza et al., 2015, Piñeiro et al., 2014, Shukla et al., 2014, Bateman et al., 2013, Denholm, 2013, Merabet and Hudry, 2013, Southall et al., 2013, Weavers and Skaer, 2013, Hudry et al., 2012, Brockmann et al., 2011, Datta et al., 2011, Estella and Mann, 2010, Gautam and Tapadia, 2010, Lopes and Casares, 2010, Oros et al., 2010, Bao and Friedrich, 2009, Bessa et al., 2009, Datta et al., 2009, Dichtel-Danjoy et al., 2009, Gambetta et al., 2009, Coiffier et al., 2008, Oktaba et al., 2008, Takacs et al., 2008, Kumar and Datta, 2007, Kumar and Luhur, 2007, Kumar and Salzer, 2007, Patel et al., 2007, Tanaka et al., 2007, Zirin and Mann, 2007, Dworkin and Gibson, 2006, Friedrich, 2006, Jordan et al., 2006, Noro et al., 2006, Sandmann et al., 2006, Akimoto et al., 2005, Pearson et al., 2005, Roederer et al., 2005, Wang et al., 2004, Chauvet et al., 2000)
                  Secondary FlyBase IDs
                  • FBgn0000058
                  • FBgn0010570
                  • FBgn0013543
                  Datasets (0)
                  Study focus (0)
                  Experimental Role
                  Project
                  Project Type
                  Title
                  Study result (0)
                  Result
                  Result Type
                  Title
                  External Crossreferences and Linkouts ( 40 )
                  Sequence Crossreferences
                  NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
                  GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
                  GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
                  RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
                  UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
                  UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
                  Other crossreferences
                  AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
                  BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
                  DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
                  EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
                  FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
                  FlyMine - An integrated database for Drosophila genomics
                  InterPro - A database of protein families, domains and functional sites
                  KEGG Genes - Molecular building blocks of life in the genomic space.
                  MARRVEL_MODEL - MARRVEL (model organism gene)
                  Linkouts
                  BioGRID - A database of protein and genetic interactions.
                  Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
                  DroID - A comprehensive database of gene and protein interactions.
                  DRSC - Results frm RNAi screens
                  Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
                  FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
                  FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
                  Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
                  Flygut - An atlas of the Drosophila adult midgut
                  Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
                  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
                  MIST (protein-protein) - An integrated Molecular Interaction Database
                  Reactome - An open-source, open access, manually curated and peer-reviewed pathway database.
                  SignaLink - A signaling pathway resource with multi-layered regulatory networks.
                  References (411)