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Gene Dmel\twi

General Information
Symbol	Dmel\twi	Species	D. melanogaster
Name	twist	Annotation symbol	CG2956
Feature type	protein_coding_gene	FlyBase ID	FBgn0003900
Created / Updated	2005-09-09/2005-09-09
Genomic Location
Chromosome (arm)	2R	Recombination map	2-100
Cytogenetic map	59C2-59C2	Sequence location	2R:18,933,631..18,935,849 [+]
Map ( GBrowse )	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene twist is referred to in FlyBase by the symbol twi (CG2956, FBgn0003900). It has the cytological map location 59C2. Its sequence location is 2R:18933631..18935849. Its molecular function is described as: specific RNA polymerase II transcription factor activity; transcription activator activity; DNA binding; specific transcriptional repressor activity. It is involved in the biological processes described with 16 unique terms, many of which group under: anatomical structure development; organ development; mesoderm morphogenesis; regulation of metabolic process; mesoderm development; transcription from RNA polymerase II promoter; positive regulation of nucleobase, nucleoside, nucleotide and nucleic acid metabolic process; gland morphogenesis; dorsal/ventral axis specification; mesodermal cell fate specification; ventral furrow formation; cell-cell signaling; muscle development; organ morphogenesis; axis specification. 54 alleles are reported. The phenotypes of these alleles are annotated with 33 unique terms, many of which group under: organ system; primordium; adult segment; adult mesothoracic segment; late extended germ band embryo; germ layer; portion of tissue; adult somatic muscle; embryonic/larval tracheal system; nervous system. It has 2 annotated transcripts and 2 annotated polypeptides.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	twi: twist The wild-type allele of twi is involved in the establishment of germ layers. Mutants are embryonic lethals (zygotic), partially dorsalized, and without mesodermal differentiation. A normal blastoderm is formed; at gastrulation, no ventral furrow is visible, but the endoderm invaginates, a cephalic furrow is formed, and the germband elongated. The embryo is twisted or coiled in the egg case, often with posterior side up. There are few mesodermally derived internal tissues. Some embryos fail to make a properly differentiated cuticle, although 40-100% make normal ectodermal derivatives (Simpson, 1983). There is no maternal effect in germline chimeras. twi/+ heterozygous embryos have delayed ventral furrow formation. The TSP of the twi gene is around gastrulation (Thisse et al., 1987).
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 59C2-59C2   Limits computationally determined from genome sequence between P{PZ}Nup21410444 and P{PZ}l(2)0649606496&P{lacW}l(2)k09913k09913
Experimentally Determined Cytological Location
Cytogenetic map Notes References 59C3-59D2 (determined by in situ hybridisation)   (Thisse et al., 1988) 59C3-59D2 (determined by in situ hybridisation)   (Thisse et al., 1987)
Experimentally Determined Recombination Data
Location	2-100 (Tearle and Nusslein-Volhard, 1987)
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References Gene Order (overall orientation not stated) References Overall orientation not stated: anon-59Ca? twi? anon-59Cb? anon-EB8? anon-59Cc? anon-EB8? anon-59Cd? anon-59Ce? anon-59Cg? anon-59Cf? anon-59Ch? anon-59Cg? anon-59Ci? anon-59Ck? anon-59Cj? anon-59Ck? (Thisse et al., 1987)
Gene Model & Products
Please see the GBrowse view of Dmel\twi for information on other features
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) twi-RA FBtr0071953  NM_079092   2098   490 twi-RB FBtr0100130  NM_001038878   1880   490
Additional Transcript Data & Comments
Reported size (kB)	1.8 (northern blot) (Thisse et al., 1987) 1.9 (longest cDNA) (Thisse et al., 1988)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein twi-PA   FBpp0071864   54.4   490   6.59     NP_523816   AAF46941 twi-PB   FBpp0099476   54.4   490   6.59     NP_001033967   ABC66041
Additional Polypeptide Data & Comments
Reported size (kD)	490 (aa) (Thisse et al., 1988)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0003900
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Twist (IPR015789) Basic helix-loop-helix dimerisation region bHLH (IPR001092) Helix-loop-helix DNA-binding (IPR011598) TRANSFAC - Eukaryotic transcription factors, their genomic binding sites, and DNA-binding profiles • R01716 R01717 R01718 R01719 R01720 R01721 R01722 R03460 R03461 R03462 T01517
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE003459 AAF46941   AY075213 AAL68080   AY118454 AAM49823   BF485882   BF500249   BF501764   BT025192   CO306060   CO306897   CO308924   CO309640   CO317866   CO319927   EC066714   EC237874   S59100   X12506 CAA31024   X14569 CAA32707
UniProtKB/Swiss-Prot	P10627
UniProtKB/TrEMBL
Maps to
Does NOT map to
Identified with	AT15089 AT17310 (BDGP Project Members, 2000-) D1125 (BDGP Project Members, 1994-1999)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References point mutation twi[v50] 2R:18,934,069..18,934,069 evidence=experimental na_change=C18934069G pr_change=P22A|twi-PA,P22A|twi-PB reported_na_change=CCC?GCC reported_pr_change=P22A (Baylies and Bate, 1996) point mutation twi[RY50] 2R:18,934,754..18,934,754 comment=The difference between the authors reported amino acid acid substitution position and the FlyBase report is due to the use of a different initation codon. evidence=experimental na_change=A18934754T pr_change=Q250L|twi-PA,Q250L|twi-PB reported_na_change=CAG?CTG reported_pr_change=Q251L (Baylies and Bate, 1996) protein binding site twi-protein_bind-1 2R:18,932,786..18,932,796 bound_moiety=dl-XP evidence=experimental (Bergman et al., 2004, Thisse et al., 1991) protein binding site twi-protein_bind-10 2R:18,933,564..18,933,577 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-11 2R:18,933,819..18,933,831 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-12 2R:18,933,588..18,933,607 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-13 2R:18,933,547..18,933,557 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-14 2R:18,933,755..18,933,765 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-15 2R:18,933,531..18,933,541 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-16 2R:18,932,745..18,932,764 bound_moiety=dl-XP evidence=experimental (Bergman et al., 2004, Thisse et al., 1991) protein binding site twi-protein_bind-17 2R:18,933,536..18,933,557 evidence=experimental (Pan et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-2 2R:18,933,337..18,933,365 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-3 2R:18,933,771..18,933,786 evidence=experimental (Pan et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-4 2R:18,933,665..18,933,675 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-5 2R:18,932,732..18,932,742 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-6 2R:18,933,577..18,933,587 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-7 2R:18,933,244..18,933,258 evidence=experimental (Bergman et al., 2004, Pan et al., 1991) protein binding site twi-protein_bind-8 2R:18,933,730..18,933,744 evidence=experimental (Pan et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-9 2R:18,933,268..18,933,307 evidence=experimental (Pan et al., 1991, Bergman et al., 2004) protein binding site twi-protein_bind-18 2R:18,932,536..18,932,546 bound_moiety=dl-XP comment=TD2 evidence=experimental (Jiang et al., 1991) protein binding site twi-protein_bind-19 2R:18,932,845..18,932,855 bound_moiety=dl-XP comment=TD3 evidence=experimental (Jiang et al., 1991) protein binding site twi-protein_bind-20 2R:18,933,531..18,933,620 bound_moiety=dl-XP evidence=experimental (Pan et al., 1991) protein binding site twi-protein_bind-21 2R:18,931,531..18,931,540 bound_moiety=dl-XP comment=TD4 evidence=experimental (Jiang et al., 1991) protein binding site twi-protein_bind-22 2R:18,933,548..18,933,557 bound_moiety=dl-XP comment=TD5 evidence=experimental (Jiang et al., 1991) protein binding site twi-protein_bind-23 2R:18,933,733..18,933,738 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) protein binding site twi-protein_bind-24 2R:18,933,754..18,933,759 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) protein binding site twi-protein_bind-25 2R:18,933,774..18,933,779 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) protein binding site twi-protein_bind-26 2R:18,933,821..18,933,826 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) regulatory region twi-reg_element-1 2R:18,932,020..18,934,085 comment=Directs expression in a pattern very similar to wild-type twi expression. evidence=experimental (Pan et al., 1991) regulatory region twi-reg_element-2 2R:18,932,428..18,932,864 comment=DE (distal element) evidence=experimental linked_to=HindIII-EcoNI_rfrag (Jiang et al., 1991) regulatory region twi-reg_element-3 2R:18,933,311..18,933,578 comment=Ventral activator region, contains dl protein binding sites evidence=experimental (Pan et al., 1991) regulatory region twi-reg_element-4 2R:18,933,407..18,933,668 comment=PE (proximal element) evidence=experimental linked_to=EcoRI-XhoI_rfrag (Jiang et al., 1991) regulatory region twi-reg_element-5 2R:18,933,642..18,933,794 comment=element z, contains z protein binding sites evidence=experimental (Pan et al., 1991) rescue fragment twi[+t9.2] 2R:18,930,658..18,939,884 comment=Reported as a HaeIII-HaeIII fragment containing 3.0kb of 5' and 6.2kb of 3' sequences; position of restriction fragment on reference sequence inferred by FlyBase curator. evidence=experimental linked_to=HaeIII-HaeIII_rfrag (Thisse et al., 1991)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 1426
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  embryonic stage | 2-10 hr   (Thisse et al., 1987)  embryonic stage | cellular blastoderm  ectoderm | presumptive (Leptin, 1991)  embryonic stage | stage 6  mesoderm | presumptive (Thisse et al., 1987)  embryonic stage | stage 8  mesoderm (Thisse et al., 1987)  embryonic stage | cellular blastoderm  mesoderm | presumptive (Leptin, 1991)  embryonic stage | stage 5   (Thisse et al., 1987)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  embryonic stage | stage 7-10  mesoderm (Thisse et al., 1988)  embryonic stage | cellular blastoderm  ectoderm | presumptive (Leptin, 1991)  embryonic stage | stage 6  mesoderm | presumptive (Thisse et al., 1988)  embryonic stage | stage 12-14  larval visceral muscle (Thisse et al., 1988)  embryonic stage | stage 6  mesectoderm (Thisse et al., 1988)  embryonic stage | cellular blastoderm  mesoderm | presumptive (Leptin, 1991)  embryonic stage-pupal stage  mesothoracic tergum & myoblast (Sudarsan et al., 2001)  embryonic stage | stage 12-14  larval somatic muscle (Thisse et al., 1988)  embryonic stage | stage 5   (Thisse et al., 1988)  embryonic stage | stage 7-8  anterior midgut primordium (Thisse et al., 1988)
Marker for
Subcellular Localization
CV Term	nucleus (Thisse et al., 1988)
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003900 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • FBgn0003900 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0003900 BDGP expression data • Show all BDGP original images, annotations and embryonic microarray data for twi FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for twi Stage(s) 4-6 Stage(s) 7-8 Stage(s) 9-10 Stage(s) 11-12 Stage(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | recessive twiunspecified lethal | recessive | conditional ts twiRY50 lethal | embryonic stage | recessive twi1 twiRY50 twid50 twiey50 twiey53 twiey55 twiunspecified Other Phenotypes Allele wild-type twi+t9.2 twiT10 twiT11 twiT40 twiT48 twiT64 twiT71 twiT72 visible | recessive twiRY50 twiv50 flight behavior defective, with Scer\GAL4Mef2.PR twiGD1553 neuroanatomy defective twidsRNA.cIa Phenotype manifest in Allele embryonic somatic muscle, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 visceral muscle primordium, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval pericardial cell, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval heart, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval somatic muscle | ectopic, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 Malpighian tubule twi1 twi4 anterior midgut inclusive primordium twi1 twiRY50 twiey53 posterior midgut inclusive primordium twi1 twiRY50 twiey53 visceral mesoderm & embryo twi1 twiRY50 twiey53 endoderm twi1 twiHH07 visceral mesoderm twi1 twiunspecified embryonic/larval somatic muscle twi1 twiv50 embryonic central nervous system twi1 mesoderm twi1 mesoderm & embryo twi1 central nervous system twi1 ventral furrow twi1 twidsRNA.cGa twiunspecified embryonic/larval dorsal branch twi1 embryonic/larval dorsal trunk twi1 mesothoracic extracoxal depressor muscle 66 twi1 twiv50 embryonic salivary gland, with Scer\GAL4twi.PG twi2x.Scer\UAS longitudinal visceral muscle primordium, with Scer\GAL4twi.PG twi2x.Scer\UAS embryonic somatic muscle, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS visceral muscle primordium, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval pericardial cell, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval heart, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval somatic muscle | ectopic, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic somatic muscle | ectopic, with Scer\GAL4twi.PB twi2x.Scer\UAS twiScer\UAS.cBa principal midgut epithelial cell twi3 dorsal medial muscle | conditional ts twiRY50 oblique muscle | conditional ts twiRY50 wing | conditional ts twiRY50 dorsal medial muscle twiRY50 twiv50 mesothoracic extracoxal depressor muscle 66 | conditional ts twiRY50 skeletal muscle of abdomen | conditional ts twiRY50 indirect flight muscle | conditional ts twiRY50 direct flight muscle | conditional ts twiRY50 embryonic/larval somatic muscle | conditional ts twiRY50 visceral muscle, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval dorsal vessel & embryo, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval pericardial cell, with Scer\GAL4twi.PB twiScer\UAS.cBa cardioblast, with Scer\GAL4twi.PB twiScer\UAS.cBa visceral mesoderm, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval heart, with Scer\GAL4twi.PB twiScer\UAS.cBa abdominal ventral transverse muscle 1 founder cell, with Scer\GAL4twi.PB twiScer\UAS.cBa abdominal lateral oblique muscle 1 founder cell, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic epidermis, with Scer\GAL4da.G32 twiScer\UAS.cBa mesoderm, with Scer\GAL4twi.PG twiScer\UAS.cBa myoblast & pupa, with Scer\GAL41151 twiScer\UAS.cBa indirect flight muscle, with Scer\GAL41151 twiScer\UAS.cBa longitudinal visceral muscle primordium twiey53 circular visceral muscle fibers twiey53 embryo | ventral twiunspecified oblique muscle twiv50 wing twiv50 skeletal muscle of abdomen twiv50 indirect flight muscle twiv50 direct flight muscle twiv50 wing, with Scer\GAL4Mef2.PR twiGD1553
Classical Alleles ( 20 )
For All Classical Alleles Show
Allele of twi Class Mutagen Stocks Known lesion twi1 amorph, loss of function, hypomorph ethyl methanesulfonate 2 -- twi2 ethyl methanesulfonate 0 -- twi3 ethyl methanesulfonate 2 -- twi4 ethyl methanesulfonate 0 -- twi5 ethyl methanesulfonate 0 -- twi6 ethyl methanesulfonate 0 -- twi7 ethyl methanesulfonate 0 -- twi8 ethyl methanesulfonate 0 -- twiEY53R1 0 -- twiHH07 0 -- twiRY50 hypomorph X ray 0 Yes twid50 ethyl methanesulfonate 0 Yes twiey50 amorph ethyl methanesulfonate 0 Yes twiey53 amorph ethyl methanesulfonate 0 Yes twiey55 ethyl methanesulfonate 0 Yes twiey62 0 -- twiey63 ethyl methanesulfonate 0 -- twiry51 X ray 0 -- twiunspecified 0 -- twiv50 0 Yes
Alleles Carried on Transgenic Constructs ( 34 )
For All Alleles Carried on Transgenic Constructs Show
Allele of twi Class Mutagen Stocks Known lesion twi+t9.2 in vitro construct | deletion 0 Yes twi155P in vitro construct | deletion 0 Yes twi2x.CMV in vitro construct | coding region fusion in vitro construct | regulatory fusion 0 Yes twi2x.Scer\UAS in vitro construct | coding region fusion in vitro construct | regulatory fusion 0 Yes twi354.Act5C in vitro construct | regulatory fusion 0 Yes twi434P in vitro construct | deletion 0 Yes twi51P in vitro construct | deletion 0 Yes twi79P in vitro construct | deletion 0 Yes twi99P in vitro construct | deletion 0 Yes twiAct5C.PS in vitro construct | regulatory fusion 0 Yes twiCMV.cCa in vitro construct | regulatory fusion 0 Yes twiGD1553 in vitro construct | RNAi in vitro construct | regulatory fusion 2 Yes twiHsp83.bcd.3'UTR in vitro construct | regulatory fusion in vitro construct | coding region fusion 0 Yes twiScer\UAS.cBa in vitro construct | regulatory fusion 0 Yes twiT10 in vitro construct | deletion 0 Yes twiT11 in vitro construct | deletion 0 Yes twiT135 in vitro construct | genomic fragment 0 Yes twiT36 in vitro construct | deletion 0 Yes twiT40 in vitro construct | deletion 0 Yes twiT48 in vitro construct | deletion 0 Yes twiT64 in vitro construct | deletion 0 Yes twiT71 in vitro construct | deletion 0 Yes twiT72 in vitro construct | deletion 0 Yes twicRa in vitro construct | other 0 Yes twidsRNA.cBa in vitro construct | RNAi 0 Yes twidsRNA.cGa in vitro construct | RNAi 0 Yes twidsRNA.cIa in vitro construct | RNAi 0 Yes twidsRNA.cMa in vitro construct | RNAi 0 Yes