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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
Gene Model Status	Current	Stock availability	8 publicly available
Genomic Location
Chromosome (arm)	2R	Recombination map	2-100
Cytogenetic map	59C2-59C2	Sequence location	2R:18,933,631..18,935,849 [+]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents Beta	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 Dmel\twi (CG2956, FBgn0003900). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: RNA polymerase II distal enhancer sequence-specific DNA binding transcription factor activity; protein heterodimerization activity; protein homodimerization activity; DNA binding. There is experimental evidence that it is involved in the biological process: mesodermal cell fate commitment; muscle fiber development; Malpighian tubule morphogenesis; positive regulation of transcription from RNA polymerase II promoter; regulation of striated muscle tissue development; negative regulation of gene expression; actomyosin structure organization; salivary gland morphogenesis; mesoderm development. 58 alleles are reported. The phenotypes of these alleles are annotated with: organ system; portion of tissue; multicellular structure; adult segment; late extended germ band embryo; non-connected developing system; extended germ band embryo; organ system subdivision; endoderm derivative; indirect flight muscle. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: Myc-type, basic helix-loop-helix (bHLH) domain; Twist-related. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of very high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Little or no expression detected in any larval or adult organs/tissues. Comments on Affy2 ProbeSet: ProbeSet 1631682_at completely aligns to an exonic region common to each of the 2 FlyBase-annotated transcript isoforms of twi. Gene sequence location is 2R:18933631..18935849.
User Contributed Data
	Beta version, feedback welcome
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).
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2013_03
FB2013_02
All updates	Click here to see a list of all updates to this record from FB2010_08 and on.
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
Gene Model & Products
Please see the GBrowse view of Dmel\twi for information on other features To submit a correction to a gene model please use the Contact FlyBase form
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)	1.9 (longest cDNA) (Thisse et al., 1988) 1.8 (northern blot) (Thisse et al., 1987)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) 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 (kDa)	490 (aa) (Thisse et al., 1988)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Twist-related (IPR015789) Myc-type, basic helix-loop-helix (bHLH) domain (IPR011598)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE013599 AAF46941   AE013599 ABC66041   AY075213 AAL68080   AY118454 AAM49823   BF485882   BF500249   BF501764   BT025192   CO306060   CO306897   CO308924   CO309640   CO317866   CO319927   EC066714   EC237874   GH960150   S59100   X12506 CAA31024   X14569 CAA32707
UniProtKB/Swiss-Prot	P10627
UniProtKB/TrEMBL
Mapped Features
Mapped Features have been reorganized, please see this article for details. Additional mapped features and mutations can be found on GBrowse or related reports. Type Symbol & Location Additional Notes References protein binding site FBsf0000160677 2R:18,932,786..18,932,796 bound_moiety=dl-XP evidence=experimental (Bergman et al., 2005, Thisse et al., 1991) protein binding site FBsf0000160678 2R:18,933,564..18,933,577 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000160679 2R:18,933,819..18,933,831 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000160680 2R:18,933,588..18,933,607 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000160681 2R:18,933,547..18,933,557 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2005) protein binding site FBsf0000160682 2R:18,933,755..18,933,765 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000159787 2R:18,933,531..18,933,541 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2005) protein binding site FBsf0000159788 2R:18,932,745..18,932,764 bound_moiety=dl-XP evidence=experimental (Bergman et al., 2005, Thisse et al., 1991) protein binding site FBsf0000159789 2R:18,933,536..18,933,557 evidence=experimental (Pan et al., 1991, Bergman et al., 2005) protein binding site FBsf0000159790 2R:18,933,337..18,933,365 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000159791 2R:18,933,771..18,933,786 evidence=experimental (Pan et al., 1991, Bergman et al., 2005) protein binding site FBsf0000159792 2R:18,933,665..18,933,675 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2005) protein binding site FBsf0000160683 2R:18,932,732..18,932,742 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2005) protein binding site FBsf0000160684 2R:18,933,577..18,933,587 bound_moiety=dl-XP evidence=experimental (Thisse et al., 1991, Bergman et al., 2005) protein binding site FBsf0000160685 2R:18,933,244..18,933,258 evidence=experimental (Bergman et al., 2005, Pan et al., 1991) protein binding site FBsf0000160686 2R:18,933,730..18,933,744 evidence=experimental (Pan et al., 1991, Bergman et al., 2005) protein binding site FBsf0000160687 2R:18,933,268..18,933,307 evidence=experimental (Pan et al., 1991, Bergman et al., 2005) protein binding site FBsf0000159872 2R:18,932,536..18,932,546 bound_moiety=dl-XP comment=TD2 evidence=experimental (Jiang et al., 1991) protein binding site FBsf0000159873 2R:18,932,845..18,932,855 bound_moiety=dl-XP comment=TD3 evidence=experimental (Jiang et al., 1991) protein binding site FBsf0000159874 2R:18,933,531..18,933,620 bound_moiety=dl-XP evidence=experimental (Pan et al., 1991) protein binding site FBsf0000159876 2R:18,931,531..18,931,540 bound_moiety=dl-XP comment=TD4 evidence=experimental (Jiang et al., 1991) protein binding site FBsf0000159877 2R:18,933,548..18,933,557 bound_moiety=dl-XP comment=TD5 evidence=experimental (Jiang et al., 1991) protein binding site FBsf0000159878 2R:18,933,733..18,933,738 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) protein binding site FBsf0000159880 2R:18,933,754..18,933,759 evidence=experimental bound_moiety=z-XP (Pan et al., 1991) protein binding site FBsf0000159879 2R:18,933,774..18,933,779 bound_moiety=z-XP evidence=experimental (Pan et al., 1991) protein binding site FBsf0000159881 2R:18,933,821..18,933,826 evidence=experimental bound_moiety=z-XP (Pan et al., 1991) regulatory region FBsf0000435053 2R:18,933,408..18,934,003 evidence=Reporter construct (in vivo) (Jiang et al., 1991, Halfon, 2012.6.19) regulatory region FBsf0000435204 2R:18,932,428..18,933,663 evidence=Reporter construct (in vivo) (Jiang and Levine, 1993, Halfon, 2012.6.19) regulatory region FBsf0000435269 2R:18,932,428..18,933,842 (Papatsenko and Levine, 2005, Halfon, 2012.6.19) regulatory region FBsf0000435286 2R:18,932,001..18,932,927 evidence=Reporter construct (in vivo) (Pan et al., 1994, Halfon, 2012.6.19) regulatory region FBsf0000161217 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 FBsf0000161218 2R:18,932,428..18,932,864 comment=DE (distal element) evidence=experimental linked_to=HindIII-EcoNI_rfrag (Jiang et al., 1991) regulatory region FBsf0000161219 2R:18,933,311..18,933,578 comment=Ventral activator region, contains dl protein binding sites evidence=experimental (Pan et al., 1991) regulatory region FBsf0000161220 2R:18,933,407..18,933,668 comment=PE (proximal element) evidence=experimental linked_to=EcoRI-XhoI_rfrag (Jiang et al., 1991) regulatory region FBsf0000161221 2R:18,933,642..18,933,794 comment=element z, contains z protein binding sites evidence=experimental (Pan et al., 1991) regulatory region FBsf0000435591 2R:18,933,313..18,933,795 evidence=Reporter construct (in vivo) (Pan et al., 1991, Halfon, 2012.6.19) regulatory region FBsf0000435919 2R:18,933,190..18,933,581 evidence=Reporter construct (in vivo) (Pan et al., 1991, Halfon, 2012.6.19) regulatory region FBsf0000436568 2R:18,932,606..18,934,003 evidence=Reporter construct (in vivo) (Jiang et al., 1991, Halfon, 2012.6.19)
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 4 -- 5   mesoderm anlage   (Ou et al., 2002) embryonic stage 4 -- 6   anterior endoderm anlage   (Fisher et al., 2012) Comment:anlage in statu nascendi mesoderm anlage   (Fisher et al., 2012) Comment:anlage in statu nascendi trunk mesoderm anlage   (Fisher et al., 2012) Comment:anlage in statu nascendi embryonic stage 5   mesoderm | presumptive   (Leptin, 1991) ectoderm | presumptive   (Leptin, 1991) mesoderm anlage   (Pan et al., 1991) embryonic stage 6 -- 7   proventriculus anlage   (De Velasco et al., 2004) ventral furrow   (De Velasco et al., 2004) embryonic stage 7 -- 8   anterior endoderm anlage   (Fisher et al., 2012) head mesoderm   (Fisher et al., 2012) trunk mesoderm   (Fisher et al., 2012) embryonic stage 9 -- 10   head mesoderm   (Fisher et al., 2012) trunk mesoderm   (Fisher et al., 2012) embryonic stage 11 -- 12   dorsal pharyngeal muscle primordium   (Fisher et al., 2012) head mesoderm   (Fisher et al., 2012) longitudinal visceral muscle primordium   (Fisher et al., 2012) somatic muscle primordium   (Fisher et al., 2012) trunk mesoderm   (Fisher et al., 2012) visceral muscle primordium   (Fisher et al., 2012) embryonic stage 13 -- 16   adult muscle precursor primordium   (Fisher et al., 2012) embryonic esophagus   (Fisher et al., 2012) embryonic foregut   (Fisher et al., 2012) embryonic hindgut   (Fisher et al., 2012) esophageal visceral muscle   (Fisher et al., 2012) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage 5 -- 13     (Thisse et al., 1987) Comment:reference states 2-10 hr AEL radioisotope in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage 5   organism | ventral compartment   (Thisse et al., 1987) embryonic stage 6   mesoderm | presumptive   (Thisse et al., 1987) embryonic stage 8   mesoderm   (Thisse et al., 1987)
Additional Descriptive Data
	The 1.8 kb twi transcript is expressed at high levels between 2 to 6 hr after egg laying, with a peak during gastrulation. The levels of twi transcript are low betwen 8 and 10 hr, and undetectable thereafter. (Thisse et al., 1987) Both twi and sna transcript are first detectable during nuclear cycle 12 as a diffuse band half the width of the presumptive mesoderm. Although the expression patterns of sna and twi transcript and protein are similar early in embryogenesis, there are some subtle differences. At mid-cellularization, the sna transcript and protein expression boundaries sharply delimit the presumptive mesoderm. At the same time, twi transcript and protein is expressed in a gradient that extends past the sna expression zone into the presumptive ectoderm. The twi transcript and protein expression pattern gets sharper later, during gastrulation. twi protein is expressed in the mesoderm throughout germ band extension, whereas the sna protein product disappears from the mesoderm partway through germ band extension, and appears in neurectodermal cells which might be neuroblasts. (Leptin, 1991) twi expression is detected in a limited number of ventral cells in stage 5 embryos. The ventral furrow forms shortly thereafter, and twi transcript accumulates in the invaginated cells of the ventral furrow (presumptive mesodermal cells). At stage 8, twi is expressed in the mesodermal layer of the germ band. (Thisse et al., 1987)
Marker for
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage -- pupal stage   mesothoracic tergum & myoblast   (Sudarsan et al., 2001) embryonic stage 4   organism | dorsal   (James et al., 2002) embryonic stage 5   ectoderm | presumptive   (Leptin, 1991) mesoderm | presumptive   (Leptin, 1991) organism | ventral   (Thisse et al., 1988) embryonic stage 6   mesoderm | presumptive   (Thisse et al., 1988) mesectoderm   (Thisse et al., 1988) embryonic stage 7 -- 8   anterior midgut primordium   (Thisse et al., 1988) embryonic stage 7 -- 10   mesoderm   (Thisse et al., 1988) embryonic stage 10 -- 11   hindgut visceral muscle primordium   (San Martin and Bate, 2001) embryonic stage 12 -- 14   embryonic/larval somatic muscle   (Thisse et al., 1988) embryonic/larval visceral muscle   (Thisse et al., 1988)
Additional Descriptive Data
	twi is a nuclear protein (Leptin, 1991, Thisse et al., 1988) Protein can be detected in myoblasts in the presumptive and larval wing disc. (Sudarsan et al., 2001) During stage 5 of embryogenesis, the twi protein is expressed ventrally, extending past the anterior and posterior poles. During stage 6, twi protein is seen in the presumptive mesodermal cells of the invaginating ventral furrow, in endodermal cells in the anterior and posterior regions of the embryo, and in the ectodermal cells of the anterior midgut primordium. The mesodermal cells forming the ventral furrow, as well as the four adjoining mesectodermal cells express twi protein at stage 7. At stage 7, twi protein is no longer detected in the cells which will form the posterior midgut rudiment, although labeling is still seen in the anterior midgut rudiment. twi protein becomes restricted to the mesoderm starting at stage 9. In stage 11 embryos, only a few mesodermal cells in each segment express twi protein. At stage 12, a subset of the cells of the somatopleura and the splanchnopleura (mesodermally derived cells which will give rise to the somatic and visceral musculature, respectively) are labeled by anti-twi antibody, and by stage 14, only the splanchnopleura labeling is visible. (Thisse et al., 1988)
Marker for
Subcellular Localization (GO Cellular Component)
CV term Evidence References
cytoplasm inferred from direct assay (Tipping et al., 2010) nucleus inferred from direct assay (Thisse et al., 1988)
Expression Deduced from Reporters
Reporter: P{GAL4-twi.2xPE} Stage Tissue/Position (including subcellular localization) Reference embryonic stage 5   organism | ventral   (Schubiger, 1997.3.14) Reporter: P{GAL4-twi.B} Stage Tissue/Position (including subcellular localization) Reference embryonic stage   mesoderm   (Jenkins et al., 2003) mesoderm   (Staehling-Hampton et al., 1994) embryonic Malpighian tubule | restricted   (Denholm et al., 2003) Reporter: P{GAL4-twi.G} Stage Tissue/Position (including subcellular localization) Reference embryonic stage   mesoderm   (Greig and Akam, 1993, Cheng et al., 1998) Reporter: P{twi-βgal-1.4a} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage -- pupal stage   mesothoracic tergum & myoblast   (Sudarsan et al., 2001) embryonic stage 4   mesoderm anlage   (Thisse et al., 1991) organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   posterior midgut primordium   (Thisse et al., 1991) anterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) pupal stage   adult muscle precursor cell   (Roy and VijayRaghavan, 1997) Reporter: P{twi-βgal-1.4t} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage 4   organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   anterior midgut primordium   (Thisse et al., 1991) posterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) Reporter: P{twi-βgal-3.0a} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage 4   organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   anterior midgut primordium   (Thisse et al., 1991) posterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) Reporter: P{twi-βgal-3.0t} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage 4   mesoderm anlage   (Thisse et al., 1991) organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   posterior midgut primordium   (Thisse et al., 1991) anterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) Reporter: P{twi-βgal-5.0a} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage 4   organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   anterior midgut primordium   (Thisse et al., 1991) posterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) Reporter: P{twi-βgal-5.0t} Stage Tissue/Position (including subcellular localization) Reference embryonic stage | late   embryonic/larval visceral muscle   (Thisse et al., 1991) embryonic/larval somatic muscle   (Thisse et al., 1991) embryonic stage 4   mesoderm anlage   (Thisse et al., 1991) organism | ventral   (Thisse et al., 1991) gastrula stage   endoderm | anterior   (Thisse et al., 1991) ventral furrow   (Thisse et al., 1991) extended germ band stage   posterior midgut primordium   (Thisse et al., 1991) anterior midgut primordium   (Thisse et al., 1991) embryonic stage 9 -- 17   mesoderm   (Thisse et al., 1991) Reporter: P{twi-GAL4.Ga} Stage Tissue/Position (including subcellular localization) Reference embryonic stage   mesoderm   (Riechmann et al., 1997) Reporter: P{twi-lacZ.P} Stage Tissue/Position (including subcellular localization) Reference embryonic stage 5   organism | ventral   (Jiang and Levine, 1993)
High-Throughput Expression Data
Associated Tools
Reference	See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes.
FlyAtlas Anatomy Microarray
FlyAtlas Anatomical Expression Data (FlyAtlas-RNA.adult) (FlyAtlas-RNA.larva)    Styles Linear Logarithmic Heatmap Back-to-back    Scales gene max expression Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Little or no expression detected in any larval or adult organs/tissues. [download data (TSV)] Guide to FlyAtlas expression level colors   No expression (0 - 9.999)   Low expression (10 - 99.999)   Moderate expression (100 - 499.999)   High level expression (500 - 999.999)   Very high expression (>999.999) Linear, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate   High  Linear, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate   High   Very high  Linear, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  Very high  log, scaled to maximum expression level Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate  log, scaled to Moderate expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate   High  log, scaled to High level expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate   High   Very high  log, scaled to Very high expression Tissue   Expression Level Larval Central Nervous System    8.9 Larval Midgut    11.4 Larval Hindgut    76.1 Larval Malpighian Tubules    9.3 Larval Fat Body    14.4 Larval Salivary Gland    7.6 Larval Trachea    5.975 Larval Carcass    28.625 Adult Head    10.5 Adult Eye    1.575 Adult Brain    4.8 Adult Thoracic-Abdominal Ganglion    10 Adult Crop    51.4 Adult Midgut    6.1 Adult Hindgut    67.6 Adult Malpighian Tubules    6.7 Adult Fat Body    10.2 Adult Salivary Gland    12.5 Adult Heart    15.05 Adult VirginFemale Spermatheca    19.7 Adult InseminatedFemale Spermatheca    21.9 Adult Ovary    2.8 Adult Testis    94.1 Adult Male Accessory Gland    82.9 Adult Carcass    15.2 Expression Level Scale  None   Low   Moderate   High   Very high  Heatmap Tissue   Expression Level Larval Central Nervous System     Larval Midgut     Larval Hindgut     Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea     Larval Carcass     Adult Head     Adult Eye     Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop     Adult Midgut     Adult Hindgut     Adult Malpighian Tubules     Adult Fat Body     Adult Salivary Gland     Adult Heart     Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis     Adult Male Accessory Gland     Adult Carcass     FlyAtlas Organ/Tissue Expression, larval vs. adult Larval Expression Level Tissue Adult Expression Level   NA  Head    no informative data   NA  Eye    1.575   NA  Brain    4.8   8.9  Central Nervous System    NA   NA  Thoracic-Abdominal Ganglion    no informative data   NA  Crop    51.4   no informative data  Midgut    6.1   76.1  Hindgut    67.6   9.3  Malpighian Tubules    6.7   no informative data  Fat Body    no informative data   7.6  Salivary Gland    no informative data   NA  Heart    no informative data   5.975  Trachea    NA   NA  VirginFemale Spermatheca    no informative data   NA  InseminatedFemale Spermatheca    no informative data   NA  Ovary    2.8   NA  Testis    94.1   NA  Male Accessory Gland    82.9   no informative data  Carcass    no informative data FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
modENCODE Anatomy RNA-Seq
modENCODE Tissue Expression Data (modENCODE_mRNA-Seq_tissues)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate  Linear, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Extremely high  log, scaled to maximum expression level Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to Moderate expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Tissue   Expression Level imaginal disc, larvae L3 wandering    10 central nervous system, larvae L3    1 central nervous system, pupae P8    1 head, virgin 1-day female    0 head, virgin 4-day female    0 head, virgin 20-day female    0 head, mated 1-day female    1 head, mated 4-day female    0 head, mated 20-day female    0 head, mated 1-day male    0 head, mated 4-day male    0 head, mated 20-day male    0 salivary gland, larvae L3 wandering    0 salivary gland, white prepupae    1 digestive system, larvae L3 wandering    1 digestive system, 1-day adult    1 digestive system, 4-day adult    1 digestive system, 20-day adult    1 fat body, larvae L3 wandering    0 fat body, white prepupae    2 fat body, pupae P8    3 carcass, larvae L3 wandering    5 carcass, 1-day adult    1 carcass, 4-day adult    1 carcass, 20-day adult    1 ovary, virgin 4-day female    0 ovary, mated 4-day female    0 testis, mated 4-day male    19 accessory gland, mated 4-day male    4 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Tissue   Expression Level imaginal disc, larvae L3 wandering     central nervous system, larvae L3     central nervous system, pupae P8     head, virgin 1-day female     head, virgin 4-day female     head, virgin 20-day female     head, mated 1-day female     head, mated 4-day female     head, mated 20-day female     head, mated 1-day male     head, mated 4-day male     head, mated 20-day male     salivary gland, larvae L3 wandering     salivary gland, white prepupae     digestive system, larvae L3 wandering     digestive system, 1-day adult     digestive system, 4-day adult     digestive system, 20-day adult     fat body, larvae L3 wandering     fat body, white prepupae     fat body, pupae P8     carcass, larvae L3 wandering     carcass, 1-day adult     carcass, 4-day adult     carcass, 20-day adult     ovary, virgin 4-day female     ovary, mated 4-day female     testis, mated 4-day male     accessory gland, mated 4-day male
modENCODE Development RNA-Seq
modENCODE Temporal Expression Data (modENCODE_mRNA-Seq_U)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of very high expression to a trough of very low expression. Peak expression observed within 00-06 hour embryonic stages. [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    150 embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr  (150) embryo 04-06hr  (69) embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr  (150) embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    150 embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Extremely high  log, scaled to maximum expression level Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    150 embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr  (150) embryo 04-06hr  (69) embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr  (150) embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    9 embryo 02-04hr    150 embryo 04-06hr    69 embryo 06-08hr    23 embryo 08-10hr    19 embryo 10-12hr    13 embryo 12-14hr    9 embryo 14-16hr    9 embryo 16-18hr    4 embryo 18-20hr    4 embryo 20-22hr    3 embryo 22-24hr    3 larva L1    3 larva L2    1 larva L3 12hr old    1 larva L3 puffstage 1-2    2 larva L3 puffstage 3-6    4 larva L3 puffstage 7-9    5 white prepupae new    6 white prepupae 12hr    11 white prepupae 24hr    10 pupae 2d postWPP    7 pupae 3d postWPP    6 pupae 4d postWPP    9 adult male 01day    10 adult male 05day    13 adult male 30day    18 adult female 01day    2 adult female 05day    0 adult female 30day    0 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Developmental Stage   Expression Level embryo 00-02hr     embryo 02-04hr     embryo 04-06hr     embryo 06-08hr     embryo 08-10hr     embryo 10-12hr     embryo 12-14hr     embryo 14-16hr     embryo 16-18hr     embryo 18-20hr     embryo 20-22hr     embryo 22-24hr     larva L1     larva L2     larva L3 12hr old     larva L3 puffstage 1-2     larva L3 puffstage 3-6     larva L3 puffstage 7-9     white prepupae new     white prepupae 12hr     white prepupae 24hr     pupae 2d postWPP     pupae 3d postWPP     pupae 4d postWPP     adult male 01day     adult male 05day     adult male 30day     adult female 01day     adult female 05day     adult female 30day     modENCODE Temporal Expression Data (Graveley et al., 2011)
modENCODE Cell Lines RNA-Seq
modENCODE Cell Line Expression Data (modENCODE_mRNA-Seq_cell.A) (modENCODE_mRNA-Seq_cell.B)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1    128 wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1    114 CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Moderate expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4  (84) embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3  (98) leg disc CME L1  (128) wing disc CME-W2    0 wing disc ML-DmD8  (70) wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1  (114) CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1  (128) wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1  (114) CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1    128 wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1    114 CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Extremely high  log, scaled to maximum expression level Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1    128 wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1    114 CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Moderate expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4  (84) embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3  (98) leg disc CME L1  (128) wing disc CME-W2    0 wing disc ML-DmD8  (70) wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1  (114) CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1  128 wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1  114 CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Cell Line   Expression Level Schneider line 2 S2R+    0 Schneider line 2 Sg4    84 embryonic 1182-4H    0 embryonic GM2    0 embryonic Kc167    0 embryonic S1    0 embryonic S3    98 leg disc CME L1    128 wing disc CME-W2    0 wing disc ML-DmD8    70 wing disc ML-DmD9    0 wing disc ML-DmD16-c3    0 wing disc ML-DmD21    0 wing disc ML-DmD32    0 haltere disc ML-DmD17-c3    0 eye-antennal disc ML-DmD11    0 antennal disc ML-DmD20-c5    0 mixed discs ML-DmD4-c1    114 CNS ML-DmBG1-c1    0 CNS ML-DmBG2-c2    0 tumorous blood cells mbn2    0 ovary fGS/OSS    0 ovary OSC    0 ovary OSS    1 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Cell Line   Expression Level Schneider line 2 S2R+     Schneider line 2 Sg4     embryonic 1182-4H     embryonic GM2     embryonic Kc167     embryonic S1     embryonic S3     leg disc CME L1     wing disc CME-W2     wing disc ML-DmD8     wing disc ML-DmD9     wing disc ML-DmD16-c3     wing disc ML-DmD21     wing disc ML-DmD32     haltere disc ML-DmD17-c3     eye-antennal disc ML-DmD11     antennal disc ML-DmD20-c5     mixed discs ML-DmD4-c1     CNS ML-DmBG1-c1     CNS ML-DmBG2-c2     tumorous blood cells mbn2     ovary fGS/OSS     ovary OSC     ovary OSS
modENCODE Treatments RNA-Seq
modENCODE Treatment Expression Data (modENCODE_mRNA-Seq_treatments)    Styles Linear Logarithmic Heatmap    Scales gene max expression Moderate expression bin max High expression bin max Extremely high expression bin max [download data (TSV)] Guide to modENCODE expression level colors   No/Extremely low expression (0 - 0)   Very low expression (1 - 3)   Low expression (4 - 10)   Moderate expression (11 - 25)   Moderately high expression (26 - 50)   High expression (51 - 100)   Very high expression (101 - 1000)   Extremely high expression (>1000) Linear, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate  Linear, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Extremely high  log, scaled to maximum expression level Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate  log, scaled to Moderate expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate   Moderately high  log, scaled to High expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Treatment   Expression Level extended cold, 4-day adult    1 cold shock, 4-day adult    2 heat shock, 4-day adult    1 Cadmium 50 mM 6 hrs, larvae L3    1 Cadmium 50 mM 12 hrs, larvae L3    1 Cadmium 50 mM 48 hrs, 4-day adult    1 Cadmium 100 mM 48 hrs, 4-day adult    2 Copper 0.5 mM 12 hrs, larvae L3    3 Copper 15 mM 48 hrs, 4-day adult    3 Zinc 5 mM 12 hrs, larvae L3    1 Zinc 4.5 mM 48 hrs, 4-day adult    4 Ethanol 2.5% 3 hrs, larvae L3    2 Ethanol 5% 3 hrs, larvae L3    0 Ethanol 10% 3 hrs, larvae L3    1 Caffeine 1.5 mg/ml 4 hrs, larvae L3    1 Caffeine 2.5 mg/ml 48 hrs, 4-day adult    1 Caffeine 25 mg/ml 48 hrs, 4-day adult    3 Paraquat 5 mM 48 hrs, 4-day adult    3 Paraquat 10 mM 48 hrs, 4-day adult    3 Rotenone 2 μg 12 hrs, larvae L3    0 Rotenone 8 μg 12 hrs, larvae L3    2 Expression Level Scale  Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  Heatmap Treatment   Expression Level extended cold, 4-day adult     cold shock, 4-day adult     heat shock, 4-day adult     Cadmium 50 mM 6 hrs, larvae L3     Cadmium 50 mM 12 hrs, larvae L3     Cadmium 50 mM 48 hrs, 4-day adult     Cadmium 100 mM 48 hrs, 4-day adult     Copper 0.5 mM 12 hrs, larvae L3     Copper 15 mM 48 hrs, 4-day adult     Zinc 5 mM 12 hrs, larvae L3     Zinc 4.5 mM 48 hrs, 4-day adult     Ethanol 2.5% 3 hrs, larvae L3     Ethanol 5% 3 hrs, larvae L3     Ethanol 10% 3 hrs, larvae L3     Caffeine 1.5 mg/ml 4 hrs, larvae L3     Caffeine 2.5 mg/ml 48 hrs, 4-day adult     Caffeine 25 mg/ml 48 hrs, 4-day adult     Paraquat 5 mM 48 hrs, 4-day adult     Paraquat 10 mM 48 hrs, 4-day adult     Rotenone 2 μg 12 hrs, larvae L3     Rotenone 8 μg 12 hrs, larvae L3
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_31
External Data & Images
Linkouts	BDGP expression data • AT15089 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003900 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG2956-RA FlyExpress - Embryonic expression images (BDGP data) • Show all FlyExpress processed images for twi Stages(s) 4-6 Stages(s) 7-8 Stages(s) 9-10 Stages(s) 11-12 Stages(s) 13-16
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | embryonic stage | recessive twi1 twid50 twiey50 twiey53 twiey55 twiRY50 twiunspecified lethal | recessive twiunspecified lethal | recessive | heat sensitive twiRY50 viable, with Scer\GAL4pnr-MD237 twiGD1553 Other Phenotypes Allele flight defective, with Scer\GAL4Mef2.PR twiGD1553 neuroanatomy defective twidsRNA.cIa visible | recessive twiRY50 twiv50 wild-type twi+t9.2 twiT10 twiT11 twiT40 twiT48 twiT64 twiT71 twiT72 Phenotype manifest in Allele abdominal lateral oblique muscle 1 founder cell, with Scer\GAL4twi.PB twiScer\UAS.cBa abdominal ventral transverse muscle 1 founder cell, with Scer\GAL4twi.PB twiScer\UAS.cBa anterior midgut primordium twi1 twiRY50 twiey53 cardioblast, with Scer\GAL4twi.PB twiScer\UAS.cBa central nervous system twi1 circular visceral muscle fiber twiey53 corpus cardiacum primordium twi1 corpus cardiacum primordium, with Scer\GAL4twi.2PE twiScer\UAS.cBa direct flight muscle twiv50 direct flight muscle | heat sensitive twiRY50 dorsal medial muscle twiRY50 twiv50 dorsal medial muscle | heat sensitive twiRY50 embryo | ventral twiunspecified embryonic/larval dorsal branch twi1 embryonic/larval dorsal trunk twi1 embryonic/larval dorsal vessel | embryonic stage, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval heart, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval heart, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval heart, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval pericardial cell, with Scer\GAL4twi.PB twiScer\UAS.cBa embryonic/larval pericardial cell, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval pericardial cell, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval somatic muscle twi1 twiv50 embryonic/larval somatic muscle | ectopic, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic/larval somatic muscle | ectopic, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic/larval somatic muscle | heat sensitive twiRY50 embryonic epidermis, with Scer\GAL4da.G32 twiScer\UAS.cBa embryonic somatic muscle, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS embryonic somatic muscle, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 embryonic somatic muscle | ectopic, with Scer\GAL4twi.PB twi2x.Scer\UAS twiScer\UAS.cBa endoderm twi1 twiHH07 indirect flight muscle twiv50 indirect flight muscle, with Scer\GAL41151 twiScer\UAS.cBa indirect flight muscle | heat sensitive twiRY50 longitudinal visceral muscle primordium twiey53 longitudinal visceral muscle primordium, with Scer\GAL4twi.PG twi2x.Scer\UAS Malpighian tubule twi1 twi4 mesoderm twi1 mesoderm, with Scer\GAL4twi.PG twiScer\UAS.cBa mesothoracic extracoxal depressor muscle 66 twi1 twiv50 mesothoracic extracoxal depressor muscle 66 | heat sensitive twiRY50 myoblast | P-stage, with Scer\GAL41151 twiScer\UAS.cBa oblique muscle twiv50 oblique muscle | heat sensitive twiRY50 posterior midgut primordium twi1 twiRY50 twiey53 presumptive embryonic/larval central nervous system twi1 presumptive embryonic salivary gland, with Scer\GAL4twi.PG twi2x.Scer\UAS principal midgut epithelial cell twi3 skeletal muscle of abdomen twiv50 skeletal muscle of abdomen | heat sensitive twiRY50 ventral furrow twi1 twidsRNA.cGa twiunspecified visceral mesoderm twi1 twiRY50 twiey53 twiunspecified visceral mesoderm, with Scer\GAL4twi.PB twiScer\UAS.cBa visceral muscle, with Scer\GAL4twi.PB twiScer\UAS.cBa visceral muscle primordium, with Scer\GAL4twi.PB, twi1 twi2x.Scer\UAS visceral muscle primordium, with Scer\GAL4twi.PB, twi2x.Scer\UAS twi1 wing twiv50 wing, with Scer\GAL4Mef2.PR twiGD1553 wing | heat sensitive twiRY50
Classical Alleles ( 20 )
For All Classical Alleles Show
Allele of twi Class Mutagen Stocks Known lesion twi1 loss of function allele, amorphic allele - genetic evidence, hypomorphic allele - genetic evidence ethyl methanesulfonate 2 -- twi3 ethyl methanesulfonate 2 -- twiey50 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes twiey53 amorphic allele - genetic evidence ethyl methanesulfonate 0 Yes twi2 ethyl methanesulfonate 0 -- twi4 ethyl methanesulfonate 0 -- twi5 ethyl methanesulfonate 0 -- twi6 ethyl methanesulfonate 0 -- twi7 ethyl methanesulfonate 0 -- twi8 ethyl methanesulfonate 0 -- twid50 ethyl methanesulfonate 0 Yes twiEY53R1 0 -- twiey55 ethyl methanesulfonate 0 Yes twiey62 0 -- twiey63 ethyl methanesulfonate 0 -- twiHH07 0 -- twiRY50 hypomorphic allele - genetic evidence X ray 0 Yes twiry51 X ray 0 -- twiunspecified 0 -- twiv50 0 Yes
Alleles Carried on Transgenic Constructs ( 38 )
For All Alleles Carried on Transgenic Constructs Show
Allele of twi Class Mutagen Stocks Known lesion twiGD1553 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes twiHMS01317 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes twiJF02003 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes 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 twicRa in vitro construct 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 twig.PE in vitro construct 0 Yes twiHsp83.bcd.3'UTR in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes twiNIG.2956R in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes twiScer\UAS.cBa in vitro construct - regulatory fusion 0 Yes twiScer\UAS.cUa 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 twiΔ142-346.Act5C in vitro construct - regulatory fusion 0 Yes twiΔN173.Act5C in vitro construct - regulatory fusion 0 Yes twiΔN273.Act5C in vitro construct - regulatory fusion 0 Yes twiΔN346.Act5C in vitro construct - regulatory fusion 0 Yes twiΔN85.Act5C in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)59AD (Knoblich and Lehner, 1993) Df(2R)BSC698 (Christensen et al., 2008.12.28) Df(2R)BSC769 (Christensen et al., 2009.2.28) Df(2R)bw-D31 (Tearle and Nusslein-Volhard, 1987) Df(2R)P (Muller et al., 1999) Df(2R)twi (Leiss et al., 1988, Leptin, 1991, Jaglarz and Howard, 1994, Ip et al., 1994) Df(2R)vir130 (Schutt et al., 1998, Hilfiker et al., 1995) Ts(Y;2Lt)L116 (Muller et al., 1999) Ts(YLt;2Lt)A169 (Muller et al., 1999) Ts(YLt;2Lt)H144 (Muller et al., 1999)
Not disrupted in	Df(2R)bw-D23 (Tearle and Nusslein-Volhard, 1987)
Transgenic Constructs & Insertions
Transgenic Constructs
reporter construct	Name Expression Data P{1.0twi-lacZ} No P{1.2twi-lacZ} No P{2.0twi-lacZ} No P{2.9twi-lacZ} No P{2PE-2xscs-E3} No P{2PE-2xSU-E3} No P{2PE-Fab7-E3} No P{2PE-scs-E3} No P{2PE-scs-Fab7-E3} No P{2PE-scs-MAR-E3} No P{2PE-SF-1b-E3} No P{2PE-SF-1b-Fab7-E3} No P{2PE-SF-1b-scs-E3} No P{2PE-SF-1-SU-E3} No P{2PE-SU-E3} No P{2PE-SU-Fab7-E3} No P{2PE-SU-MAR-E3} No P{2PE-SU-scs-E3} No P{2PE-λ1.4-E3} No P{2XPE-Kr-rhoNEE} No P{2XPE-rhoNEE} No P{2XPE-rhoNEE-Kr} No P{180twi-lacZ} No P{-269twi/lacZ} No P{440twi-lacZ} No P{-648twi/lacZ} No P{800twi-lacZ} No P{-920twi/lacZ} No P{-1829twi/lacZ} No P{-5000twi/lacZ} No P{CA1xDEtwi-lacZ} No P{CA2xDEtwi-lacZ} No P{Ctd(55)U(-55)-lacZ} No P{Ctd(55)U(-130)-lacZ} No P{Ctd(110)U(-55)-lacZ} No P{Ctd(110)U(-130)-lacZ} No P{Ctd(165)U(-55)-lacZ} No P{Ctd(165)U(-130)-lacZ} No P{CtdU(-55)-lacZ} No P{CtdU(-130)-lacZ} No P{DmD/hsp70} No P{lacZtwi.2PEe-Et} No P{lacZzen/twi-VR/2PE} No P{twi/hsp-1} No P{twi/hsp-2} No P{twi/hsp-3} No P{twi/hsp-4} No P{twi/hsp-5} No P{twi/hsp-6} No P{twi/hsp-7} No P{twi/hsp-8} No P{twi/hsp-9} No P{twi-BR-eve-lacZ.PS3} No P{twi-BR-eve-lacZ.PSA3} No P{twi-BR-eve-lacZ.PSS3} No P{twi-CD2} No P{twi-eve-lacZ.PL3} No P{twi-lacZ.2PEe} No P{twi-lacZ.648VAR-zen.VR} No P{twi-lacZ.660VARD4} No P{twi-lacZ.PEe-6Et} No P{twi-lacZ.PEe-11Et} No P{twi-lacZ.PEe-20Et} No P{twi-lacZ.PEe-25Et} No P{twi-lacZ.PEe-86Et} No P{twi-lacZ.PEe} No P{twi-lacZ.PEe-Etprox} No P{twi-lacZ.PEm} No P{twist-lacZ} No P{twi-βgal-0.8a} No P{twi-βgal-0.8t} No P{twi-βgal-1.4a} No P{twi-βgal-1.4t} No P{twi-βgal-3.0a} No P{twi-βgal-3.0t} No P{twi-βgal-5.0a} No P{twi-βgal-5.0t} No P{zen-lacZ.D4} No
UAS construct	Name Expression Data P{GD1553} NA P{NIG.2956R} NA P{TRiP.HMS01317} NA P{TRiP.JF02003} NA P{UAS-twi.B} NA P{UAS-twi.U} NA P{UAS-twi-twi.C} NA
heat-shock construct	Name Expression Data P{Hsp83-twi.bcd.3'UTR} NA
GAL4 construct	Name Expression Data P{GAL4-twi.2xPE} No P{GAL4-twi.B} No P{GAL4-twi.BS} No P{GAL4-twi.G} No P{twi-GAL4.Ga} No P{twi-GAL4.U} No
characterization construct	Name Expression Data P{E3-SF-1b-ftz-distal-PE} NA P{GAL4-twi.Bc} NA P{Ptwig} NA P{twi-CD2*} NA P{twi-lacZ.2PE} NA P{twi-lacZ.2PEe-Eds} NA P{twi-lacZ.2PE-Et} NA P{twi-lacZ.4PE} NA P{twi-lacZ.P} NA P{twi-lacZ.PE} NA P{twi-lacZ.PEe'} NA P{twi-lacZ.PEe-Eds} NA P{twi-lacZ.PEe-Em} NA P{twi-lacZ.PE-Et} NA P{twi-lacZ.PEm-Et.PEe} NA P{twi-NTF} NA P{UAS-twi-da.C} NA P{WT9} NA P{WT10} NA P{WT11} NA P{WT36} NA P{WT40} NA P{WT48} NA P{WT64} NA P{WT71} NA P{WT72} NA P{WT135} NA
Insertions
	Type of insertions Name Expression data
Gene Ontology: Function, Process & Cellular Component ( 26 unique terms )
Terms Based on Experimental Evidence ( 15 terms )
Molecular Function
CV term Evidence References
DNA binding inferred from direct assay (assigned by UniProtKB) (Castanon et al., 2001) protein heterodimerization activity inferred from physical interaction with UniProtKB:P11420 (assigned by UniProtKB) (Castanon et al., 2001) protein homodimerization activity inferred from physical interaction with UniProtKB:P10627 (assigned by UniProtKB) (Castanon et al., 2001) RNA polymerase II distal enhancer sequence-specific DNA binding transcription factor activity inferred from direct assay (Yin et al., 1997) contributes_to RNA polymerase II distal enhancer sequence-specific DNA binding transcription factor activity inferred from genetic interaction with dl (Bhaskar and Courey, 2002)
Biological Process
CV term Evidence References
actomyosin structure organization inferred from mutant phenotype (Martin et al., 2010) Malpighian tubule morphogenesis inferred from mutant phenotype (Jack and Myette, 1999) mesodermal cell fate commitment inferred from mutant phenotype (assigned by UniProtKB) (Castanon et al., 2001) mesoderm development inferred from expression pattern (Furlong et al., 2001) muscle fiber development inferred from mutant phenotype (Lovato et al., 2005) negative regulation of gene expression inferred from direct assay (Anderson et al., 2006) positive regulation of transcription from RNA polymerase II promoter inferred from direct assay (Yin et al., 1997, Postigo et al., 1999) regulation of striated muscle tissue development inferred from mutant phenotype (assigned by UniProtKB) (Castanon et al., 2001) salivary gland morphogenesis inferred from mutant phenotype (Vining et al., 2005)
Cellular Component
CV term Evidence References
cytoplasm inferred from direct assay (Tipping et al., 2010) nucleus inferred from direct assay (Thisse et al., 1988)
Terms Based on Predictions or Assertions ( 16 terms )
Molecular Function
CV term Evidence References
DNA binding non-traceable author statement (Swiss-Prot Project Members, 1989.7.1) sequence-specific DNA binding transcription factor activity inferred from sequence or structural similarity with UniProtKB:Q8WVJ9 (FlyBase, 1992-)
Biological Process
CV term Evidence References
anterior midgut invagination non-traceable author statement (Reuter and Casal, 1994) dorsal/ventral axis specification traceable author statement (Lall and Patel, 2001) ectoderm and mesoderm interaction non-traceable author statement (Stainier, 2002) gastrulation non-traceable author statement (Leptin, 1999) gastrulation involving germ band extension traceable author statement (Stathopoulos and Levine, 2004) heart development traceable author statement (Cripps and Olson, 2002) mesodermal cell fate commitment traceable author statement (Reuter and Casal, 1994, Furlong et al., 2001) mesodermal cell fate determination traceable author statement (Bate et al., 1999) mesodermal cell fate specification traceable author statement (Lall and Patel, 2001, Stathopoulos and Levine, 2002) mesoderm development non-traceable author statement (Swiss-Prot Project Members, 1989.7.1, Taylor, 2002) traceable author statement (Wilson and Leptin, 2000, Cripps and Olson, 2002) positive regulation of transcription from RNA polymerase II promoter non-traceable author statement (Leptin, 1999) regulation of transcription from RNA polymerase II promoter inferred from sequence or structural similarity with UniProtKB:Q8WVJ9 (FlyBase, 1992-) ventral furrow formation traceable author statement (Reuter and Casal, 1994)
Cellular Component
CV term Evidence References
nucleus inferred from sequence or structural similarity with UniProtKB:Q8WVJ9 (FlyBase, 1992-) non-traceable author statement (Swiss-Prot Project Members, 1989.7.1)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
protein-protein
Interacting group Assay References
twi - Nmnat experimental knowledge based, anti tag coimmunoprecipitation, peptide massfingerprinting (Guruharsha et al., 2011, Mintseris et al., 2009.11.23) twi - akirin pull down, autoradiography (Nowak et al., 2012) twi - da pull down, autoradiography (Nowak et al., 2012) twi - mts anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23) twi - βTub85D anti tag coimmunoprecipitation, peptide massfingerprinting (Mintseris et al., 2009.11.23)
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions twi allele Gene References twi1 da (Castanon et al., 2001) fog (Morize et al., 1998) twi3 sna (Reuter and Leptin, 1994, De Velasco et al., 2004) twiScer\UAS.cBa da (Castanon et al., 2001) Ras85D (Halfon et al., 2000) wg (Halfon et al., 2000, Cox and Baylies, 2005) unspecified br (Lovato et al., 2005) Mef2 (Lovato et al., 2005) RpII140 (Parkhurst and Ish-Horowicz, 1991)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 63264 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0003900 DroID - A comprehensive database of gene and protein interactions. • FBgn0003900 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37655 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P10627
Orthologs
OrthoDB Orthologs (47) - based on analysis using Dmel annotation version 5.41
OrthoDB Ortholog Groups
	OrthoDB orthology group searches (Waterhouse et al., 2011, Nucleic Acids Res. 39(Database issue): D283--D288) Drosophila inclusive ortholog search EOG6TXCBW Dipteran inclusive ortholog search EOG6HT91W Insect inclusive ortholog search EOG6RN951 Arthropod inclusive ortholog search EOG634TNM Metazoa inclusive ortholog search EOG664C5K
Orthologs in Drosophila Species (EOG6TXCBW)
Organism Common Name Gene AAA Syntenic Ortholog Multiple Dmel Genes in this Orthologous Group Drosophila melanogaster fruit fly  Dmel\twi     Drosophila simulans   Dsim\twi Y   Drosophila sechellia   Dsec\GM15978 Y   Drosophila erecta   Dere\twi Y   Drosophila yakuba   Dyak\twi Y   Drosophila ananassae   Dana\GF13559 Y   Drosophila pseudoobscura pseudoobscura   Dpse\twi Y   Drosophila persimilis   Dper\GL11229 Y   Drosophila willistoni   Dwil\GK19547 Y   Drosophila virilis   Dvir\twi Y   Drosophila mojavensis   Dmoj\GI21307 Y   Drosophila grimshawi   Dgri\GH20971 Y
Orthologs in non-Drosophila Dipterans (EOG6HT91W)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Aedes aegypti Yellow fever mosquito  Aaeg\AAEL002902   Anopheles gambiae Malaria mosquito  Agam\AGAP011985   Culex quinquefasciatus Southern house mosquito  Cqui\CPIJ012828   Culex quinquefasciatus Southern house mosquito  Cqui\CPIJ012827
Orthologs in non-Dipteran Insects (EOG6RN951)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Apis mellifera Western honey bee  Amel\GB40416   Nasonia vitripennis Parasitic wasp  Nvit\Nasvi2EG000292   Acromyrmex echinatior Panamanian leafcutter ant  Aech\AECH16106   Atta cephalotes Leafcutter ant  Acep\ACEP23430   Camponotus floridanus Florida carpenter ant  Cflo\CFLO22783   Harpegnathos saltator Jerdons jumping ant  Hsal\HSAL22479   Linepithema humile Argentine ant  Lhum\LH16875   Pogonomyrmex barbatus Red harvester ant  Pbar\CG:02546957   Solenopsis invicta Red fire ant  Sinv\SINV20920   Acyrthosiphon pisum Pea aphid  Apim\ACYPI001448   Bombyx mori Silkmoth  Bmor\BGIBMGA008990   Pediculus humanus Human body louse  Phum\PHUM396880   Tribolium castaneum Red flour beetle  Tcas\twi
Orthologs in non-Insect Arthropods (EOG634TNM)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Daphnia pulex Water flea  Dpux\Lefty2   Ixodes scapularis Deer tick  Isca\ISCW009433
Orthologs in non-Arthropod Metazoa (EOG664C5K)
Organism Common Name Gene Multiple Dmel Genes in this Orthologous Group Caenorhabditis elegans Nematode  Cele\hlh-8   Strongylocentrotus purpuratus Purple sea urchin  Spur\SPU_030059gn   Danio rerio Zebrafish  Drer\twist1b   Danio rerio Zebrafish  Drer\twist2   Danio rerio Zebrafish  Drer\twist3   Danio rerio Zebrafish  Drer\twist1a   Xenopus tropicalis Western clawed frog  Xtro\twist2   Xenopus tropicalis Western clawed frog  Xtro\twist1   Gallus gallus Domestic chicken  Ggal\cdermo-1   Gallus gallus Domestic chicken  Ggal\ENSGALG00000006321   Mus musculus House mouse  Mmus\Twist2   Mus musculus House mouse  Mmus\Twist1   Rattus norvegicus Norway rat  Rnor\Twist2   Rattus norvegicus Norway rat  Rnor\Twist1   Homo sapiens Human  Hsap\TWIST2   Homo sapiens Human  Hsap\TWIST1
Human Orthologs (2)
Gene OMIM HGNC Hsap\TWIST2     Hsap\TWIST1
AAA Orthologs (11) based on analysis using Dmel annotation version 4.3
Organism Gene Drosophila simulans Dsim\twi Drosophila sechellia Dsec\GM15978 Drosophila erecta Dere\twi Drosophila yakuba Dyak\twi Drosophila ananassae Dana\GF13559 Drosophila pseudoobscura pseudoobscura Dpse\twi Drosophila persimilis Dper\GL11229 Drosophila willistoni Dwil\GK19547 Drosophila virilis Dvir\twi Drosophila mojavensis Dmoj\GI21307 Drosophila grimshawi Dgri\GH20971
Stocks & Reagents
Stocks Listed in FlyBase ( 8 )
Bloomington	25981 y1 v1; P{TRiP.JF02003}attP2 2381 cn1 twi1 bw1 sp1/CyO 34330 y1 sc* v1; P{TRiP.HMS01317}attP2 3299 cn1 sna18 twi3 bw1 sp1/CyO
Kyoto	107298 cn1 twi1 bw1 sp1/CyO 106965 cn1 sna18 twi3 bw1 sp1/CyO
VDRC	v37091 w1118; P{GD1553}v37091 v37092 w1118; P{GD1553}v37092
Genomic Clones ( 2 )
	BACR08P09 BACR37H13 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 14 )
	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 GBrowse for alignment of the cDNAs and ESTs to the gene model.
cDNA Clones, Fully Sequenced
BDGP DGC clones	AT15089 AT17310
Other clones	DPiM_FI01303 FI01303
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	AT19956
Other clones	145480 165373 EK218706 EK250203 EK252727 EK259702 EK269540 EK281502 RP004548488
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0003900 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37655
Antibody Information
	polyclonal (Maqbool et al., 2006, Thisse et al., 1988, Figeac et al., 2010)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of
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	DNA-protein interactions: genome-wide binding profile assayed for twi protein in 2-3 hr embryos; see BDTNP1_TFBS_twi collection report. (MacArthur et al., 2009) The twi product negatively regulates dac expression in the embryonic head. (Anderson et al., 2006) twi homodimers specify mesoderm and the subsequent allocation of mesodermal cells to the somatic muscle fate. twi, da heterodimers repress genes required for somatic myogenesis. (Castanon et al., 2001) dl can activate transcription from zen and twi promoters, and additional Dsp1 inhibits the zen activation and increases the twi activation. (Brickman et al., 1999) An interplay of dl and twi proteins with Taf4 protein is required for the activation of mesoderm-determining gene expression in the embryo. (Pham et al., 1999) twi and N negatively regulate adult muscle differentiation in Drosophila. (Anant et al., 1998) twi protein directly regulates Mef2 expression in adult somatic muscle precursor cells via a 175bp enhancer located 2245bp upstream of the Mef2 transcriptional start site. Activation of Mef2 transcription via this enhancer by twi protein is essential for normal adult muscle development. (Cripps et al., 1998) twi function is required during the late larval stage for splitting of the larval oblique muscles that form the templates of the adult dorsal longitudinal indirect flight muscles. (Cripps and Olson, 1998) In a sample of 79 genes with multiple introns, 33 showed significant heterogeneity in G+C content among introns of the same gene and significant positive correspondence between the intron and the third codon position G+C content within genes. These results are consistent with selection adding against preferred codons at the start of genes. (Kliman and Eyre-Walker, 1998) The normal location of the ventral furrow in embryos with uniformly expressed fog suggests the existence of a fog-independent pathway determining mesoderm-specific cell behaviours and invagination. Epistasis experiments indicate this pathway requires sna but not twi expression. (Morize et al., 1998) nej is necessary for dl-mediated activation of the twi promoter. (Akimaru et al., 1997) tin is a direct transcriptional target for twi and its own gene product in visceral mesodermal cells, supporting the idea that twi and tin function in the subdivision of the mesoderm during embryogenesis. (Lee et al., 1997) dl and twi proteins synergistically activate transcription in cell culture from a promoter containing binding sites for both factors. A region outside of the conserved bHLH domain of the twi protein is required for the synergy. Protein-protein interaction assays show that dl and twi proteins bind to one another in vitro. (Shirokawa and Courey, 1997) Expression of esg in the neuroectoderm is studied, the expression pattern prefigures that of the ASC genes. Dorsoventral pattern of esg expression in the blastoderm is determined by three independent repressive cues (dpp, sna and twi). (Yagi and Hayashi, 1997) High levels of twi product are required for somatic myogenesis and block the formation of other mesodermal derivatives. Ectopic expression of twi can drive ectodermal cells into myogenesis. After an initial role in gastrulation, twi regulates mesodermal differentiation and propels a specific subset of mesodermal cells into somatic myogenesis. (Baylies and Bate, 1996) Analysis of the motorneuron structure in twi mutant embryos demonstrates post-synaptic target muscles are not required for the outgrowth of presynaptic motor axons into the periphery, nor are they required to induce the expression of genes coding for the components of active zones or for the assembly of active zones themselves. Synthesis of the active zone appears to be an independent function of the presynaptic motorneuron, which is integrated into the development of the neuromuscular synapse. (Prokop et al., 1996) At stage 10 two different groups of mesodermal cells can be observed by twi protein expression: group A have weak twi expression and give rise to internal mesodermal structures (visceral mesoderm) and group A have strong twi expression and give rise to external mesodermal structures (heart and somatic mesoderm). (Dunin-Borkowski et al., 1995) Mef2 is a downstream target of twi. (Taylor et al., 1995) The genomic region of tin contains several twi binding sites suggesting that twi is a direct activator of tin. (Venkatesh et al., 1995) sna is sufficient to induce the formation of an attenuated ventral furrow in the absence of twi+ gene activity. Expression of sna in the absence of twi uncouples ventral furrow formation and mesoderm differentiation, invaginating cells fail to express various mesoderm marker genes. (Ip et al., 1994) twi, sna, hkb and tll gene products define the positions of the primordia of the germ layers and thereby the regions in which the blastoderm epithelium will invaginate. (Leptin, 1994) Lack of Mef2 gene expression in twi mutant embryos suggests that Mef2 lies downstream in the regulatory pathway. (Lilly et al., 1994) The coding region and regulatory regions of twi and Dvir\twi are compared. Within the coding region the basic helix-loop-helix DNA binding and dimerisation domain are highly conserved. A high degree of conservation is seen between the more distal of the two ventral activator elements in the 5' regulatory regions of twi and Dvir\twi. The more proximal ventral activator is missing in the equivalent position in Dvir\twi, instead a region in the second intron resembles the proximal element. Transformation experiments reveal the evolved enhancer elements are similar in structure and functional properties. (Pan et al., 1994) Mesodermal fate is determined where sna and twi but not hkb are expressed. Anteriorly, hkb together with sna determines endodermal fate, and hkb together with twi and sna are required for foregut development. (Reuter and Leptin, 1994) The prepattern of the adult musculature, revealed in persistent twi-expressing cells in the embryo, is not dependent on the embryonic peripheral motor nerves. (Broadie and Bate, 1993) twist-expressing cells associate with the segmental nerves in the thoracic as well as the abdominal segments of the third larval instar. (Fernandes and Vijay Raghavan, 1993) Promoter fusions using elements of the twi, ve, da and sna promoters indicate that low affinity dl-binding sites restrict target gene expression to the presumptive mesoderm, where there are peak levels of dl expression. The twi promoter includes a proximal and a distal element, and the proximal element contains two low affinity dl binding sites. The proximal element alone directs twi expression in the ventralmost 12 to 14 nuclei while the complete promoter with both proximal and distal elements directs twi expression in the ventralmost 18 to 20 cells. Promoters containing the Et (veinlet) or Eds (dl and snail) E boxes display opposite behaviour in da and twi mutants, suggesting they are regulated by different basic helix loop helix proteins. (Jiang and Levine, 1993) The BRE region of Ubx includes binding sites for hb, ftz, tll, en and twi. The binding of their products and the interplay between them is responsible for generating the expression pattern directed by the BRE. (Qian et al., 1993) In mutant embryos lacking the entire mesoderm or failing to differentiate the visceral mesoderm, the anterior and posterior midgut primordia form but do not migrate properly. (Reuter et al., 1993) twi mutants fail to differentiate ventrally derived mesoderm. (Arora and Nusslein-Volhard, 1992) Expression of twi assayed in embryos from females expressing wild type dl and dl-lacZ fusion proteins. Reduction in intensity of twi expression at gastrulation correlates well with degree of dorsalization of embryos, suggesting effect of dl mediated through its downstream target genes. (Govind et al., 1992) A twi binding site has been identified in the neural ectoderm expression region (NEE) of ve promoter: twi activates ve expression. The twi gene is a target gene of dl. (Ip et al., 1992) dl binding site domain exchange experiments, using Ecol\lacZ reporter gene constructs, between the zen and twi promoters demonstrate that dl is intrinsically an activator and that repression requires additional factors present in the distal region of the zen promoter, the VR. (Jiang et al., 1992) Deletion analysis of the twi promoter using Ecol\lacZ reporter gene constructs demonstrates that the repressor/activator activity of the dl binding sites depends on its context, not an intrinsic property. (Pan and Courey, 1992) The zygotically acting DV genes repress ac expression within specific DV domains. (Skeath et al., 1992) twi expression marks embryonic precursors of the cells that make the adult muscles. (Bate et al., 1991) twi-expressing lineages of cells have been ablated in the larva, and the impact on muscle pattern in the adult studied. (Broadie and Bate, 1991) A combination of promoter fusion-P-element transformation assays (1.2kb twi promoter fragment is sufficient to generate normal twi pattern of Ecol\lacZ expression) and in vitro DNA binding assays coupled with site directed mutagenesis (revealing four dl-binding sites in the twi promoter) have been used to establish a link between the dl-binding sites and twi expression in early embryos. The dorsal ventral limits of twi expression depend on the number and affinity of dl binding sites present in the twi promoter. (Jiang et al., 1991) Establishment of the mesoderm neuroectoderm boundary involves the interaction of twi, sna and dl proteins. (Kosman et al., 1991) twi is required for the activation of downstream mesoderm genes. (Leptin, 1991) Deletion analysis of the twi promoter region using Ecol\lacZ reporter gene construct demonstrate that the region extending from -68 to -269 relative to the transcription start site contains an element that is required for ventral activation of twi in the embryo. This element "z" lies between -55 and -205 and it influences overall promoter strength. A second functional element lies between -537 and -269 and constitutes a ventral activator region. In vitro transcription reactions and S1 nuclease analysis demonstrate the presence of z binding sites within 120bp region of twi promoter proximal to the transcription start site. (Pan et al., 1991) Mutations in zygotic dorsal class gene twi interact with RpII140wimp. (Parkhurst and Ish-Horowicz, 1991) twi mutant embryos have a few extra neuroblasts around the midline. (Rao et al., 1991) Lateralized embryos have twi expression at the poles. Dorsalized embryos show twi expression during late stages of embryogenesis. Polar expression of twi requires genes of the terminal group. twi is a primary patterning gene for dorsal mesoderm as expression is unaffected by mutations in zygotic dorsal-ventral genes. (Ray et al., 1991) Zygotically active locus involved in the terminal developmental program in the embryo. (Strecker et al., 1991) Phenotypic rescue and twi-lacZ expression studies demonstrate that 5' sequences -4.0kb up to +6.2kb 3' sequences are required for normal twi expression. Recurrent deletions in the 3kb twi promoter region have found dl responsive elements (DREs). dl acts as a sequence specific trans-activator of the twi promoter. Gel retardation assays have been used to investigate binding of dl protein to synthetic oligonucleotides corresponding to the proximal and distal activator region of the twi promoter. (Thisse et al., 1991) twi and sna have been shown to independently control different aspects of ventral cell behaviour during gastrulation. (Leptin and Grunewald, 1990) Involved in the regulatory hierarchy responsible for the asymmetric distribution and function of zygotic regulatory gene products along the DV axis of early embryos. (Rushlow and Levine, 1990) Molecular analysis of the twi gene and its product reveals that it encodes a nuclear protein expressed in the endoderm and mesoderm anlagen and derived tissues. (Thisse et al., 1988) In homozygous embryos invagination of the ventral presumptive mesodermal cells fails to occur and the resulting embryos are devoid of internal organs. (Simpson, 1983) Mutations in dl, pll, ea, or Tl abolish the expression of twi. At least one dose of dl+ in females is necessary for transcription of twi (Thisse, Stoetzel, El Messal and Perrin-Schmitt, 1987). twi shows extensive identity to a pair of myc-related polypeptides whose dimerized products bind to a sequence in the immunoglobulin kappa chain enhancer; the identical regions have the potential to form two amphipathic helices separated by an intervening loop (Murre, McCaw and Baltimore, 1989).   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 germ-line chimeras. twi/+ heterozygous embryos have delayed ventral furrow formation. The TSP of the twi gene is around gastrulation (Thisse, Stoetzel, El Messal and Perrin-Schmitt, 1987).
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_079092 NM_001038878 RefSeq (Proteins) • NP_523816 NP_001033967 DDBJ / EMBL / GenBank • AAF46941 AAF46941.1 AAL68080 AAM49823 ABC66041 AY075213 AY118454 BF500249 BF501764 CAA31024 CAA32707 S59100 X12506 X14569 Entrez Gene - A searchable database of RefSeq genes. • 37655 UniProtKB/Swiss-Prot • P10627
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Twist-related (IPR015789) Myc-type, basic helix-loop-helix (bHLH) domain (IPR011598) BDGP expression data - Patterns of gene expression in Drosophila embryogenesis • AT15089
Linkouts
	BioGRID - A database of protein and genetic interactions • 63264 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0003900 DroID - A comprehensive database of gene and protein interactions. • FBgn0003900 DRSC - Results from RNAi screens. • FBgn0003900 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0003900 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG2956-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0003900 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37655 Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /torstoll/twist1.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37655 modMine - Data generated by the modENCODE project. • FBgn0003900 REDfly - A database of transcriptional regulatory elements. • FBgn0003900 SignaLink - A protein-protein and protein-miRNA interaction database with multi-layered structure and multiple confidence scores. • P10627
Synonyms & Secondary IDs ( 11 )
Reported As
Symbol Synonym	bHLHb17 (Stevens et al., 2008) CG2956 (Gonzalez-Aguero et al., 2005, Mohit et al., 2006, Perkins et al., 1.15.2009, Stevens et al., 2008, Stevens et al., 2008, Krejcí et al., 2009, Ni et al., 2010.12.1) dip5 (Bhaskar et al., 2000) DTwist (Castanon and Baylies, 2002) EC9 (Thisse et al., 1987) Tw (Fakhouri et al., 2010) twi (Dutta et al., 2005, Prothmann et al., 2006, Akashi et al., 2006, Chen et al., 2007, Lovato et al., 2005, Chen et al., 2006, Mohit et al., 2006, Shapiro and Anderson, 2006, Zeitlinger et al., 2007, Sandmann et al., 2007, Anderson et al., 2006, Stathopoulos and Levine, 2005, Ratnaparkhi et al., 2006, Aerts et al., 2007, Turki-Judeh and Courey, 2012, Steiner et al., 2012, Chen et al., 2000, Nibu et al., 2003, Zhang et al., 2008, Thomas and Cripps, 2008, Seher et al., 2007, Dietzl et al., 2007, Jia and Huan, 2010, Frise et al., 2010, Chung et al., 2011, Wang et al., 2007, Johnson et al., 2011, Wilczyński and Furlong, 2010, Fernandez-Sanchez et al., 2010, Nègre et al., 2011, Li et al., 2011, Kuzin et al., 2011, Christensen et al., 2008.12.28, Astigarraga et al., 2007, Zinzen et al., 2006, Choksi et al., 2006, Beckett and Baylies, 2007, Yan and Lin, 2007, Wong et al., 2008, Li et al., 2008, Caussinus et al., 2008, Hewitt et al., 1999, Liberman and Stathopoulos, 2009, Liu et al., 2009, Crocker et al., 2008, Christensen et al., 2009.2.28, Martin et al., 2009, van Impel et al., 2009, Pouille et al., 2009, Bernard et al., 2009, Haghayeghi et al., 2010, Salzer et al., 2010, Figeac et al., 2010, Bernard et al., 2010, Jia et al., 2002, Ismat et al., 2010, Cherbas et al., 2011, Martin et al., 2010, Garcia et al., 2011, Park et al., 2011, Sandmann et al., 2006, Cunha et al., 2010, Ozdemir et al., 2011, Konikoff et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Aswani et al., 2012, Bonn et al., 2012, Kvon et al., 2012, Mathew et al., 2011, Haskel-Ittah et al., 2012) Twi (Wong and Baylies, 2007, Castanon et al., 2001, Nguyen and Frasch, 2006, Harris and Peifer, 2004, Maqbool et al., 2006, Wang et al., 2005, Philippakis et al., 2006, Chung et al., 2011, Papatsenko et al., 2009, Desprat et al., 2008, Duan et al., 2007, Chandraratna et al., 2007, Sellin et al., 2009, Sawyer et al., 2009, Klingseisen et al., 2009, Zinzen et al., 2009, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Pruteanu-Malinici et al., 2011, Ratnaparkhi et al., 2008, Puri et al., 2008, Mariappa et al., 2011, Aboukhalil and Bulyk, 2012) TWI (Bernard et al., 2006, Toledano-Katchalski et al., 2007, MacArthur et al., 2009, modENCODE Consortium et al., 2010, Yang et al., 2012, Grzegorczyk and Husmeier, 2012) twist (Bhuin and Roy, 2009)
Name Synonym	Twist (Bhaskar et al., 2002, Sivatchenko and Letsou, 2006, Zinzen et al., 2007, Wong and Baylies, 2007, Nowak et al., 2007, Zinzen et al., 2006, Harris and Peifer, 2004, Murray and Saint, 2007, Zeitlinger et al., 2007, Beckett and Baylies, 2006, Jang et al., 2006, Sanny et al., 2006, Schaub et al., 2012, Zhu et al., 2007, de Navascués and Modolell, 2007, Fox and Peifer, 2007, Nowak et al., 2008, Levine et al., 2008, Koelsch and Leptin, 2008, Stark et al., 2007, Cho, 2004, Liotta et al., 2007, Orgogozo et al., 2007, Philippakis et al., 2006, Dutta et al., 2010, Soler and Taylor, 2009, Mellerick and Liu, 2004, Papatsenko et al., 2009, Nègre et al., 2011, Hong et al., 2008, Blanke and Jackle, 2006, McMahon et al., 2008, Sandmann et al., 2006, Desprat et al., 2008, Toledano-Katchalski et al., 2007, Yan and Lin, 2007, Wu and Sato, 2008, Chandraratna et al., 2007, Atreya and Fernandes, 2008, van Impel et al., 2009, Jia et al., 2002, DeLotto, 2001, Klingseisen et al., 2009, Potthoff and Olson, 2007, MacArthur et al., 2009, Caygill and Johnston, 2008, Haghayeghi et al., 2010, Kheradpour et al., 2007, Honkela et al., 2010, Zinzen et al., 2009, Figeac et al., 2010, Bernard et al., 2010, Nowak et al., 2012, Fakhouri et al., 2010, Dougherty et al., 2008, Kvon et al., 2012, Aboukhalil and Bulyk, 2012, Haruta et al., 2010) twist (Prothmann et al., 2006, Leptin, 2005, Mathew et al., 2007, Castanon et al., 2001, Nguyen and Frasch, 2006, Sandmann et al., 2007, Kolsch et al., 2007, Ratnaparkhi et al., 2006, Maqbool et al., 2006, Gullaud et al., 2003, Nibu et al., 2003, Montell, 2006, Bousum et al., 2008, Thomas and Cripps, 2008, Seher et al., 2007, Meinhardt, 2004, Taylor, 2006, Kulkarni and Arnosti, 2005, Frise et al., 2010, Wang et al., 2007, Ishihara and Shibata, 2008, Mathew et al., 2011, Astigarraga et al., 2007, Duan et al., 2007, Zhang et al., 2004, Wong et al., 2008, Li et al., 2008, Caussinus et al., 2008, Peng et al., 2007, Sellin et al., 2009, Liu et al., 2009, Crocker et al., 2008, Martin et al., 2009, Kadam et al., 2009, Atreya and Fernandes, 2008, Hurlbut et al., 2009, Krejcí et al., 2009, Widmann and Dahmann, 2009, Pouille et al., 2009, Smith et al., 2009, Butler et al., 2009, Fernandez-Gonzalez et al., 2009, Sokol et al., 2008, Neely et al., 2010, Zimmermann et al., 2006, Salzer et al., 2010, Gurunathan et al., 2004, Martin et al., 2010, Tipping et al., 2010, Aerts et al., 2010, Sandmann et al., 2006, Cunha et al., 2010, Copley et al., 2007, Ozdemir et al., 2011, Ratnaparkhi et al., 2008, Yang et al., 2012, Chiu et al., 2012, Haskel-Ittah et al., 2012)
Secondary FlyBase IDs
	FBpp0071864 FBtr0071953
References ( 729 )
Generate a list of	All references relating to this gene ( 729 )
List References by type	Research paper  ( 454 ) Supplementary material  ( 7 ) Review  ( 148 ) Personal communication to FlyBase  ( 10 ) Abstract  ( 92 ) FlyBase analysis  ( 1 ) Stock list  ( 2 ) Protein sequence record  ( 1 ) Website  ( 1 ) Letter  ( 1 ) Book  ( 1 ) Curated genome annotation  ( 2 ) Conference report  ( 3 ) Spoof  ( 1 ) Poem  ( 1 ) DNA/RNA sequence record  ( 3 )
Recent research papers ( 40 )
	Rockel et al., 2013, Nucleic Acids Res. 41(4): e52 iSLIM: a comprehensive approach to mapping and characterizing gene regulatory networks. [FBrf0220859] Shu and Li, 2013, Theor. Biol. Med. Model. 10(1): 11 A statistical thin-tail test of predicting regulatory regions in the Drosophila genome. [FBrf0221021] Staller et al., 2013, Genetics 193(1): 51--61 Depleting Gene Activities in Early Drosophila Embryos with the "Maternal-Gal4-shRNA" System. [FBrf0220341] Aboukhalil and Bulyk, 2012, Bioinformatics 28(11): 1446--1454 LOESS correction for length variation in gene set-based genomic sequence analysis. [FBrf0218376] Aswani et al., 2012, Methods Cell Biol. 110: 243--261 Nonparametric variable selection and modeling for spatial and temporal regulatory networks. [FBrf0217971] Bonn et al., 2012, Nat. Protoc. 7(5): 978--994 Cell type-specific chromatin immunoprecipitation from multicellular complex samples using BiTS-ChIP. [FBrf0218169] Chiu et al., 2012, Math. Biosci. 236(2): 132--140 A two-scale mathematical model for DNA transcription. [FBrf0217901] Fisher et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(52): 21330--21335 DNA regions bound at low occupancy by transcription factors do not drive patterned reporter gene expression in Drosophila. [FBrf0220344] Gaertner et al., 2012, Cell Rep. 2(6): 1670--1683 Poised RNA Polymerase II Changes over Developmental Time and Prepares Genes for Future Expression. [FBrf0220312] Gildor et al., 2012, Development 139(21): 4040--4050 Bidirectional Notch activation represses fusion competence in swarming adult Drosophila myoblasts. [FBrf0219658] Grzegorczyk and Husmeier, 2012, Stat. Appl. Genet. Mol. Biol. 11(4): A non-homogeneous dynamic bayesian network with sequentially coupled interaction parameters for applications in systems and synthetic biology. [FBrf0219097] Haskel-Ittah et al., 2012, Cell 150(5): 1016--1028 Self-organized shuttling: generating sharp dorsoventral polarity in the early Drosophila embryo. [FBrf0219335] Holmqvist et al., 2012, PLoS Genet. 8(6): e1002769 Preferential Genome Targeting of the CBP Co-Activator by Rel and Smad Proteins in Early Drosophila melanogaster Embryos. [FBrf0218675] Konikoff et al., 2012, Dev. Dyn. 241(1): 150--160 Comparison of embryonic expression within multigene families using the flyexpress discovery platform reveals more spatial than temporal divergence. [FBrf0216961] Kvon et al., 2012, Genes Dev. 26(9): 908--913 HOT regions function as patterned developmental enhancers and have a distinct cis-regulatory signature. [FBrf0218184] Nowak et al., 2012, PLoS Genet. 8(3): e1002547 Akirin Links Twist-Regulated Transcription with the Brahma Chromatin Remodeling Complex during Embryogenesis. [FBrf0217616] Schaub et al., 2012, Development 139(5): 1001--1012 Org-1, the Drosophila ortholog of Tbx1, is a direct activator of known identity genes during muscle specification. [FBrf0217455] Shu and Li, 2012, Comput. Biol. Med. 42(9): 935--941 A statistical fat-tail test of predicting regulatory regions in the Drosophila genome. [FBrf0219304] Steiner et al., 2012, Genome Res. 22(4): 766--777 Cell-type-specific nuclei purification from whole animals for genome-wide expression and chromatin profiling. [FBrf0217906] Stojnic et al., 2012, PLoS Comput. Biol. 8(11): e1002725 A graphical modelling approach to the dissection of highly correlated transcription factor binding site profiles. [FBrf0219905] Turki-Judeh and Courey, 2012, PLoS ONE 7(2): e30610 The unconserved groucho central region is essential for viability and modulates target gene specificity. [FBrf0217447] Yang et al., 2012, Genome Res. 22(11): 2199--2207 The BEAF-32 insulator coordinates genome organization and function during the evolution of Drosophila species. [FBrf0219877] Yáñez-Cuna et al., 2012, Genome Res. 22(10): 2018--2030 Uncovering cis-regulatory sequence requirements for context-specific transcription factor binding. [FBrf0219649] Cherbas et al., 2011, Genome Res. 21(2): 301--314 The transcriptional diversity of 25 Drosophila cell lines. [FBrf0213077] Chung et al., 2011, Nat. Methods 8(2): 171--176 A microfluidic array for large-scale ordering and orientation of embryos. [FBrf0212871] Chung et al., 2011, Dev. Biol. 360(1): 160--172 Trachealess (Trh) regulates all tracheal genes during Drosophila embryogenesis. [FBrf0216563] Clark et al., 2011, Development 138(13): 2705--2715 Fibroblast growth factor signalling controls successive cell behaviours during mesoderm layer formation in Drosophila. [FBrf0213889] Garcia et al., 2011, BMC Mol. Biol. 12: 32 Functional characterization of the sciarid BhC4-1 core promoter in transgenic Drosophila. [FBrf0214770] Hafer et al., 2011, Genetics 189(3): 907--921 The Drosophila CPEB Protein Orb2 Has a Novel Expression Pattern and Is Important for Asymmetric Cell Division and Nervous System Function. [FBrf0216612] He et al., 2011, Nat. Genet. 43(5): 414--420 High conservation of transcription factor binding and evidence for combinatorial regulation across six Drosophila species. [FBrf0213592] Johnson et al., 2011, Development 138(21): 4627--4638 JAK/Stat signaling regulates heart precursor diversification in Drosophila. [FBrf0216404] Kuzin et al., 2011, Mech. Dev. 128(3-4): 165--177 Functional analysis of conserved sequences within a temporally restricted neural precursor cell enhancer. [FBrf0213297] Li et al., 2011, Genome Biol. 12(4): R34 The role of chromatin accessibility in directing the widespread, overlapping patterns of Drosophila transcription factor binding. [FBrf0216471] Mariappa et al., 2011, Sci. Signal. 4(204): ra89 Protein O-GlcNAcylation Is Required for Fibroblast Growth Factor Signaling in Drosophila. [FBrf0217579] Mathew et al., 2011, Mol. Cell. Biol. 31(24): 4978--4993 Role for traf4 in polarizing adherens junctions as a prerequisite for efficient cell shape changes. [FBrf0216703] Nègre et al., 2011, Nature 471(7339): 527--531 A cis-regulatory map of the Drosophila genome. [FBrf0213303] Ozdemir et al., 2011, Genome Res. 21(4): 566--577 High resolution mapping of Twist to DNA in Drosophila embryos: Efficient functional analysis and evolutionary conservation. [FBrf0213359] Park et al., 2011, PLoS Genet. 7(8): e1002241 Specification of Drosophila corpora cardiaca neuroendocrine cells from mesoderm is regulated by Notch signaling. [FBrf0215231] Pruteanu-Malinici et al., 2011, PLoS Comput. Biol. 7(7): e1002098 Automatic Annotation of Spatial Expression Patterns via Sparse Bayesian Factor Models. [FBrf0214618] Sayal et al., 2011, Fly 5(1): 47--52 Optimization of reporter gene architecture for quantitative measurements of gene expression in the Drosophila embryo. [FBrf0213005] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...