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Gene Dmel\grh

General Information
Symbol	Dmel\grh	Species	D. melanogaster
Name	grainy head	Annotation symbol	CG42311
Feature type	protein_coding_gene	FlyBase ID	FBgn0259211
Gene Model Status	Current	Stock availability	15 publicly available
Also Known As	NTF-1, Elf-1, Gh, Elf1, CG5058, l(2)06850, Ntf
Genomic Location
Chromosome (arm)	2R	Recombination map	2-86
Cytogenetic map	54E10-54F1	Sequence location	2R:13,690,294..13,729,584 [+]
Genomic Maps Select View:Gene Models/EvidenceExpression/RegulationMegaViewGene Disruptions, Stocks and Reagents modENCODE GBrowse	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene grainy head is referred to in FlyBase by the symbol Dmel\grh (CG42311, FBgn0259211). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: sequence-specific DNA binding transcription factor activity; sequence-specific DNA binding; protein homodimerization activity; DNA binding. There is experimental evidence for 13 unique biological process terms, many of which group under: biological regulation; open tracheal system development; system development; cellular component organization or biogenesis; chitin-based cuticle development; cuticle development; multicellular organismal development; sensory organ development; regulation of cell proliferation; membrane organization; positive regulation of cellular biosynthetic process; regulation of developmental process; regulation of anatomical structure size; regulation of wound healing; establishment of planar polarity; epithelial cell morphogenesis. 76 alleles are reported. The phenotypes of these alleles are annotated with: late extended germ band embryo; organ system; tracheal branch primordium; non-connected developing system; acellular anatomical structure; larval abdominal segment; cephalopharyngeal skeleton; adult segment; organ system subdivision; dorsal trunk primordium; dorsal branch primordium; adherens junction; neuroblast NB5-6 of thorax. It has 8 annotated transcripts and 8 annotated polypeptides. Protein features are: CP2 transcription factor. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of very low expression. Peak expression observed at stages throughout embryogenesis, at stages throughout the pupal period. Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression in all larval and adult organs/tissues ranges from low to undetected. Expression at high levels in the following post-embryonic organs or tissues: larval carcass. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval central nervous system, adult crop, larval trachea, adult carcass. Gene sequence location is 2R:13690294..13729584.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	Ntf: Neurogenic element binding transcription factor Gene encodes at least three isoforms of a protein that binds to upstream sequences of Ddc, en, ftz, and Ubx. The proteins share epitopes as recognized by two monoclonal antibodies. Expression first detected in 4-to-8 hour embryos, peaking from 8 to 12 hours and declining from 12 to 16 hours; NTF protein detected in nuclei of ectodermal derivatives; seen in all epidermal cells and changing subsets of neurons in the developing central nervous system.
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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FB2011_10	Alleles grh[PanA.Scer\UAS] grh[2A.Scer\UAS] grh[2E.Scer\UAS] grh[Scer\UAS.cKa] Controlled Vocabulary Terms chitin-based cuticle development regulation of wound healing
FB2012_01	Controlled Vocabulary Terms positive regulation of transcription from RNA polymerase II promoter protein homodimerization activity References Benito-Sipos et al., 2011, Development 138(24): 5311--5320 Jennings et al., 1999, Molec. Cell. Biol. 19(7): 4600--4610 Sequence features BKN31269 HFA07625 HFA05680 HFA07624 BKN29135 HFA05683 Transgenic Constructs P{UAS-grh.PanA} P{UAS-grh.K} P{UAS-grh.2A} P{UAS-grh.2E}
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 54E10-54F1   Limits computationally determined from genome sequence between P{EP}subEP616 and P{PZ}grh06850&P{lacW}olf186-Fk11505
Experimentally Determined Cytological Location
Cytogenetic map Notes References 54F1-54F2   (determined by in situ hybridisation)   (Spradling et al., 1999) 54F1-54F2     (BDGP Project Members, 1994-1999) 54F-54F   (determined by in situ hybridisation)   (Bray and Kafatos, 1991) 54F1-54F2   (determined by in situ hybridisation)   (Dynlacht et al., 1989) 54F-54F   (determined by in situ hybridisation)   (Bray et al., 1989)
Experimentally Determined Recombination Data
Location	2-86 (Nusslein-Volhard et al., 1984, Tearle and Nusslein-Volhard, 1987)
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\grh for information on other features To submit a correction to a gene model please use the Contact FlyBase form
Comments on Gene Model
	Due to an oversight in the implementation of gene annotation merges, the symbols for some of the transcripts and polypeptides of this gene were changed to non-unique designations in r5.9. Although there were no conflicts within a given release, the same symbols were used for different isoforms in different releases. The affected gene product symbols have now been changed to symbols not used previously in the database; see individual transcript reports for details. Gene merge based on EST/cDNA data
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) grh-RH FBtr0300538  NM_057495   4753   1032 grh-RI FBtr0299705  NM_057494   4846   1063 grh-RJ FBtr0299706  NM_057497   5656   1333 grh-RK FBtr0299707  NM_166250   3437   784 grh-RL FBtr0300539  NM_057496   6029   1302 grh-RM FBtr0300540  NM_166253   1677   155 grh-RN FBtr0300541  NM_166252   3187   729 grh-RO FBtr0306625     3094   698
Additional Transcript Data & Comments
Reported size (kB)	10.6, 7.4 (northern blot) (Dynlacht et al., 1989)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein grh-PH   FBpp0289765   113.0   1032   6.16     NP_476843   AAM68468 grh-PI   FBpp0288983   116.1   1063   6.19     NP_476842   AAF57782 grh-PJ   FBpp0288984   143.9   1333   6.04     NP_476845   AAM68467 grh-PK   FBpp0288985   87.6   784   6.41     NP_725723   AAM68470 grh-PL   FBpp0289766   140.9   1302   6.01     NP_476844   AAF57784 grh-PM   FBpp0289767   17.7   155   5.86     NP_725726   AAF57783 grh-PN   FBpp0289768   80.4   729   6.51     NP_725725   AAM68472 grh-PO   FBpp0297580   77.4   698   6.48
Additional Polypeptide Data & Comments
Reported size (kDa)	1333, 1303, 1063, 1032 (aa) (Uv et al., 1997) 1063 (aa); 116 (kD predicted) (Bray et al., 1989) 1032 (aa) (Attardi and Tjian, 1993) 120, 83 (aa); 140 (kD observed) (Dynlacht et al., 1989)
Comments	The DNA-binding region of the 1063aa grh protein maps to a 263 amino acid region between residues 603 and 865. Data suggests that grh protein functions as a dimer, both when bound to DNA and free in solution. The major dimerization is in the carboxyl region and is dependent on residues between 949 and 1063. Dimerization of grh protein stabilizes the interaction with DNA. (Uv et al., 1994) grh protein was purified from embyronic nuclear extracts. (Bray et al., 1989) Binds to element I of Ddc promoter. (Bray et al., 1989) The 1032aa form of grh protein binds DNA as a dimer and forms dimers in solution. The binding and dimerization domains map to a 400aa region encompassing the HLH homology region and extending beyond it. grh is a potent transcriptional activator in a cotransfection assay in Schneider cells. The activation domain maps to an isoleucine-rich domain of 56 amino acids between residues 173-228. The domain is active both in Schneider cells and in embryo extracts. No evidence was found for protein interaction between grh protein and Taf4 protein. (Bray et al., 1989) grh protein was purified from embryonic extracts. Three prominent polypeptides of 140, 120, and 83kD were observed. All of these bind to promoter elements within the Ddc, Ubx, and ftz genes. The purified protein was shown to act as binding site-dependent transcription factor in vitro. (Dynlacht et al., 1989)
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • CP2 transcription factor (IPR007604)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AE013599 AAF57782   AE013599 AAF57783   AE013599 AAF57784   AE013599 AAM68467   AE013599 AAM68468   AE013599 AAM68470   AE013599 AAM68472   AC004328   AC004640   AI134262   AI295313   AI296916   AI515542   AI519280   AL035312 CAA22954   AW943443   AW944140   AY052066 AAK93490   AY061451 AAL28999   AY231723   BG632642   BI164023   BI228296   BI232917   BI234840   BI372231   BI481731   BI486656   BI568011   BI579463   BI580402   BI611032   BI617868   BI630412   BI641593   BT001414 AAN71169   BT003182 AAO24937   BT011460   BT029431   CO186688   CO188339   CO191068   CO263171   CO267254   CO268547   CO269248   CO270203   CO270213   CO275630   CO279202   CO281118   CO282754   CO283857   CO287470   CO287531   CO287926   CO288575   CO292515   CO304794   CO312625   CO315519   CO319158   CO329352   CO344422   CO345085   CZ466147   CZ466182   CZ466194   CZ467185   CZ471255   CZ481719   CZ481720   CZ486189   EC059664   EC070890   EC076175   EC079786   EC084564   EC084577   EC198742   EC199262   EC204097   EC207493   EC208178   EC212452   EC213734   EC214542   EC215196   EC218934   EC221061   EC222089   EC222534   EC227761   EC237013   EC240773   EC241979   EC242136   EC242804   EC250342   EC252582   EL876070   EL884760   EL884888   G00456   X15657 CAA33692
UniProtKB/Swiss-Prot	P13002
UniProtKB/TrEMBL	Q95RD9 A1ZB05
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
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
in situ Stage Tissue/Position (including subcellular localization) Reference embryonic stage   neuroblast   (Uv et al., 1997) embryonic brain   (Dynlacht et al., 1989) epidermis   (Dynlacht et al., 1989) ventral nerve cord   (Dynlacht et al., 1989) embryonic cycle 13   organism | dorsal & ubiquitous   (Huang et al., 1995) Comment:stronger on dorsal than ventral side embryonic stage 5   organism | dorsal & lateral   (Huang et al., 1995) Comment:absent from a ventral midline band 15-20 cells wide larval stage   neuroblast   (Uv et al., 1997)   (Uv et al., 1997) oogenesis stage S6   oocyte   • perinuclear region of cytoplasm (Huang et al., 1995) nurse cell   • cytoplasm (Huang et al., 1995) northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage     (Dynlacht et al., 1989)
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
No Assay Recorded Stage Tissue/Position (including subcellular localization) Reference third instar larval stage   wing disc   (Santaren et al., 1993) immunolocalization Stage Tissue/Position (including subcellular localization) Reference embryonic stage   embryonic/larval foregut   (Bray et al., 1989) embryonic/larval hindgut   (Bray et al., 1989) embryonic stage 11 -- 17   ventral nerve cord | segmentally repeated   (Bray et al., 1989) epidermis   (Bray et al., 1989) embryonic stage 13 -- 17   neuroblast NB3-3 of embryonic abdomen   (Tsuji et al., 2008) embryonic stage 13 -- second instar larval stage   neuroblast NB3-3 of thorax   (Tsuji et al., 2008) larval stage   neuroblast   (Uv et al., 1997) embryonic/larval optic lobe   (Uv et al., 1997) embryonic/larval anterior spiracle   (Uv et al., 1997) imaginal disc   (Uv et al., 1997) epidermis   (Uv et al., 1997) embryonic/larval foregut   (Uv et al., 1997) embryonic/larval trachea   (Uv et al., 1997) embryonic/larval hindgut   (Uv et al., 1997) embryonic/larval proventriculus   (Uv et al., 1997)
Additional Descriptive Data
	grh protein is first detected at stage 11 in the epidermis and CNS. grh protein expression remains fairly constant in the epidermis but changes with developmental time in the CNS. Early on, grh protein is detected in one or two medial nuclei per segmental neuromere. Later, it is detected in more lateral nuclei in a pattern that changes as development proceeds. At stage 15, expression is observed in around 30 cells per neuromere in a segmentally repeated pattern. By stage 17, the number of nuclei expressing the protein has decreased particularly in the abdominal regions of the ventral nerve cord. grh protein is also detected in small regions of the foregut and the hindgut. grh protein and Ddc protein are not co-expressed in the same cells in late embryos. Co-expression was observed in primary cultures of neuroblasts. (Bray et al., 1989) grh protein expression persists in epidermal cells and in parts of the foregut and hindgut in the larval period. It is also detected in the trachea, anterior spiracles, proventriculus, and in cells that secrete the head skeleton. grh protein is expressed uniformly in the imaginal discs and in the optic lobes. (Uv et al., 1997)
Marker for
Subcellular Localization
CV Term	nucleus (Bray et al., 1989) nucleus (Bray et al., 1989)
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. modENCODE Temporal Expression Data for FBgn0259211    Styles Linear Logarithmic Heatmap    Scales max expr for FBgn0259211 Very low expression bin max 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 moderately high expression to a trough of very low expression. Peak expression observed at stages throughout embryogenesis, at stages throughout the pupal period. [download data (TSV)] Guide to modENCODE expression level colors   No expression (0 - 0)   Extremely low expression (1 - 10)   Very low expression (11 - 100)   Low expression (101 - 400)   Moderate expression (401 - 1400)   Moderately high expression (1401 - 4000)   High expression (4001 - 10000)   Very high expression (10001 - 100000)   Extremely high expression (100001 - 2000000) Linear, scaled to maximum FBgn0259211 expression level Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr  (1034) embryo 04-06hr  (1507) embryo 06-08hr  (2725) embryo 08-10hr  (2705) embryo 10-12hr  (1950) embryo 12-14hr  (1717) embryo 14-16hr  (1544) embryo 16-18hr  (1018) embryo 18-20hr  (1440) embryo 20-22hr  (808) embryo 22-24hr  (775) larva L1  (623) larva L2  (481) larva L3 12hr old  (625) larva L3 puffstage 1-2  (371) larva L3 puffstage 3-6  (931) larva L3 puffstage 7-9  (936) white prepupae new  (1114) white prepupae 12hr  (1216) white prepupae 24hr  (1803) pupae 2d postWPP  (2175) pupae 3d postWPP  (779) pupae 4d postWPP  (914) adult male 01day  (426) adult male 05day  (266) adult male 30day  (274) adult female 01day  (289) adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low  Linear, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr  1507 embryo 06-08hr  (2725) embryo 08-10hr  (2705) embryo 10-12hr  (1950) embryo 12-14hr  (1717) embryo 14-16hr  (1544) embryo 16-18hr    1018 embryo 18-20hr  1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr  (1803) pupae 2d postWPP  (2175) pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  Linear, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  Linear, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high   Extremely high  log, scaled to maximum FBgn0259211 expression level Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High  log, scaled to Very low expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr  (1034) embryo 04-06hr  (1507) embryo 06-08hr  (2725) embryo 08-10hr  (2705) embryo 10-12hr  (1950) embryo 12-14hr  (1717) embryo 14-16hr  (1544) embryo 16-18hr  (1018) embryo 18-20hr  (1440) embryo 20-22hr  (808) embryo 22-24hr  (775) larva L1  (623) larva L2  (481) larva L3 12hr old  (625) larva L3 puffstage 1-2  (371) larva L3 puffstage 3-6  (931) larva L3 puffstage 7-9  (936) white prepupae new  (1114) white prepupae 12hr  (1216) white prepupae 24hr  (1803) pupae 2d postWPP  (2175) pupae 3d postWPP  (779) pupae 4d postWPP  (914) adult male 01day  (426) adult male 05day  (266) adult male 30day  (274) adult female 01day  (289) adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low  log, scaled to Moderate expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr  1507 embryo 06-08hr  (2725) embryo 08-10hr  (2705) embryo 10-12hr  1950 embryo 12-14hr  1717 embryo 14-16hr  1544 embryo 16-18hr    1018 embryo 18-20hr  1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr  1803 pupae 2d postWPP  2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high  log, scaled to High expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   Very low   Low   Moderate   Moderately high   High   Very high  log, scaled to Extremely high expression Developmental Stage   Expression Level embryo 00-02hr    51 embryo 02-04hr    1034 embryo 04-06hr    1507 embryo 06-08hr    2725 embryo 08-10hr    2705 embryo 10-12hr    1950 embryo 12-14hr    1717 embryo 14-16hr    1544 embryo 16-18hr    1018 embryo 18-20hr    1440 embryo 20-22hr    808 embryo 22-24hr    775 larva L1    623 larva L2    481 larva L3 12hr old    625 larva L3 puffstage 1-2    371 larva L3 puffstage 3-6    931 larva L3 puffstage 7-9    936 white prepupae new    1114 white prepupae 12hr    1216 white prepupae 24hr    1803 pupae 2d postWPP    2175 pupae 3d postWPP    779 pupae 4d postWPP    914 adult male 01day    426 adult male 05day    266 adult male 30day    274 adult female 01day    289 adult female 05day    65 adult female 30day    53 Expression Level Scale  None   Extremely low   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     FlyAtlas Anatomical Expression Data for FBgn0259211    Styles Linear Logarithmic Heatmap Back-to-back    Scales max expr for FBgn0259211 Moderate expression bin max High level expression bin max Very high expression bin max Summary of FlyAtlas Anatomical Expression Data: Two or more Affy2 ProbeSets identify exons of this gene. This is a summary of the tissue expression peaks exhibited in at least one of these ProbeSets. Expression in all larval and adult organs/tissues ranges from low to undetected. Expression at high levels in the following post-embryonic organs or tissues: larval carcass. Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, larval central nervous system, adult crop, larval trachea, adult carcass. [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 (1000 - 25000) Heatmap Tissue   Expression Level Larval Central Nervous System no informative data Larval Midgut     Larval Hindgut no informative data Larval Malpighian Tubules     Larval Fat Body     Larval Salivary Gland     Larval Trachea no informative data Larval Carcass no informative data Adult Head no informative data Adult Eye no informative data Adult Brain     Adult Thoracic-Abdominal Ganglion     Adult Crop no informative data Adult Midgut     Adult Hindgut no informative data Adult Malpighian Tubules     Adult Fat Body no informative data Adult Salivary Gland     Adult Heart no informative data Adult VirginFemale Spermatheca     Adult InseminatedFemale Spermatheca     Adult Ovary     Adult Testis no informative data Adult Male Accessory Gland     Adult Carcass no informative data modENCODE Temporal Expression Data (Graveley et al., 2011) FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
Expression Clusters
	A cluster of genes with similar mRNA expression dynamics across development. (The modENCODE Consortium, 2010) mE2_34_mRNA_expression_cluster_11
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0259211 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG30111-RA
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal (with Df(2R)Pcl7B) grh370 lethal | embryonic stage grhB28 grhB32 grhB35 grhB37 lethal | embryonic stage, with Scer\GAL4Act5C.PU grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | embryonic stage, with Scer\GAL4arm.PS grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | embryonic stage, with Scer\GAL4da.G32 grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | embryonic stage | recessive grh300 grh370 grh436 grhIM lethal | larval stage, with Scer\GAL4Act5C.PU grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | larval stage, with Scer\GAL4arm.PS grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | larval stage, with Scer\GAL4da.G32 grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa lethal | polyphasic grhNΔ447.hs lethal | polyphasic | heat sensitive grhhs.PA grhNΔ447.hs lethal | recessive grh370 grh380 grh06850 grhrH247 grhs2140 semi-lethal | embryonic stage | maternal effect, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 grhRH.Scer\UAS.T:Zzzz\FLAG semi-lethal | pupal stage, with Scer\GAL4pnr-MD237 grhGD9996 semi-lethal | second instar larval stage (with Df(2R)Pcl7B) grh370 viable, with Scer\GAL4e22c grh2A.Scer\UAS grh2E.Scer\UAS grhPanA.Scer\UAS grhScer\UAS.cKa Other Phenotypes Allele cell death defective (with Df(2R)Pcl7B) grh370 increased cell death | somatic clone grhB37 mitotic cell cycle defective (with Df(2R)Pcl7B) grh370 mitotic cell cycle defective | embryonic stage | maternal effect, with Scer\GAL4mat.αTub67C.T:Hsim\VP16 grhRH.Scer\UAS.T:Zzzz\FLAG mitotic cell cycle defective | somatic clone grhB37 neuroanatomy defective (with Df(2R)Pcl7B) grhIM planar polarity defective | dominant grhFK2131 planar polarity defective | somatic clone grhIM grhs2140 visible, with Scer\GAL4pnr-MD237 grhGD9996 visible | dominant grhFK2131 wound healing defective grhIM wound healing defective | embryonic stage 17 | recessive grhIM Phenotype manifest in Allele abdominal neuroblast | supernumerary (with Df(2R)Pcl7B) grh370 abdominal neuroblast ventro-medial cluster (with Df(2R)Pcl7B) grh370 abdominal neuroblast ventro-medial cluster | larval stage (with Df(2R)Pcl7B) grh370 abdominal ventro-lateral cluster neuroblast (with Df(2R)Pcl7B) grh370 adherens junction grhunspecified amnioserosa, with Scer\GAL4332.3 grhScer\UAS.cHa amnioserosa, with Scer\GAL4c381 grhScer\UAS.cHa cephalopharyngeal skeleton grhIM cephalopharyngeal skeleton | embryonic stage grh300 grh370 grh436 grhIM chaeta | somatic clone grhIM chaeta | somatic clone | supernumerary grhIM chaeta | supernumerary, with Scer\GAL4pnr-MD237 grhGD9996 cuticle | heat sensitive grhNΔ447.hs denticle belt grhB37 grhIM dorsal branch primordium, with Scer\GAL4btl.PS grhScer\UAS.cHa dorsal closure embryo | heat sensitive grhhs.PA dorsal neurohemal organ (with Df(2R)Pcl7B) grhIM dorsal trunk primordium grhunspecified dorsal trunk primordium, with Scer\GAL4btl.PS grhScer\UAS.cHa dorsal trunk primordium (with Df(2R)Pcl7B) grhs2140 embryonic/first instar larval cuticle grhB37 grhIM embryonic/first instar larval cuticle | dorsal | heat sensitive grhhs.PA embryonic/larval cuticle grhIM embryonic/larval epidermis | embryonic stage 17 grhIM embryonic dorsal epidermis, with Scer\GAL4332.3 grhScer\UAS.cHa embryonic dorsal epidermis, with Scer\GAL4c381 grhScer\UAS.cHa eye | somatic clone grhIM eye photoreceptor cell | somatic clone grhIM ganglionic branch primordium (with Df(2R)Pcl7B) grhs2140 larval thorax & neuroblast (with Df(2R)Pcl7B) grh370 larval thorax & neuroblast | somatic clone grhB37 lateral trunk primordium, with Scer\GAL4btl.PS grhScer\UAS.cHa lateral trunk primordium (with Df(2R)Pcl7B) grhs2140 mouth hooks grhB37 neuroblast NB5-6 of thorax (with Df(2R)Pcl7B) grhIM ommatidium | somatic clone grhIM grhs2140 presumptive embryonic/larval tracheal system grhB37 presumptive embryonic/larval tracheal system, with Scer\GAL4bs-23.26 grhScer\UAS.cHa presumptive embryonic/larval tracheal system, with Scer\GAL4btl.PS grhScer\UAS.cHa presumptive embryonic/larval tracheal system (with Df(2R)Pcl7B) grhs2140 thoracic neuroblast | larval stage (with Df(2R)Pcl7B) grh370 thoracic neuron | somatic clone grhB37 transverse connective primordium (with Df(2R)Pcl7B) grhs2140 triple row grhFK2131 visceral branch primordium, with Scer\GAL4btl.PS grhScer\UAS.cHa wing | somatic clone grhIM wing hair grhFK2131 wing hair | cell non-autonomous | somatic clone grhIM wing hair | cell non-autonomous | somatic clone | supernumerary grhIM wing hair | somatic clone grhIM wing hair | somatic clone | supernumerary grhIM wing margin bristle | somatic clone grhIM wing vein | ectopic grhFK2131 wing vein | ectopic | somatic clone grhIM
Classical Alleles ( 23 )
For All Classical Alleles Show
Allele of grh Class Mutagen Stocks Known lesion grhIM amorphic allele - genetic evidence ethyl methanesulfonate 2 -- grhs2140 P-element activity 2 Yes grh06850 P-element activity 1 Yes grh5-HA-1720 1 -- grhc00006 piggyBac transposase 1 -- grhc01458 piggyBac transposase 1 -- grhc07119 piggyBac transposase 1 -- grhf03615 piggyBac transposase 1 -- grh300 ethyl methanesulfonate 0 -- grh370 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 Yes grh380 hypomorphic allele - genetic evidence ethyl methanesulfonate 0 -- grh436 ethyl methanesulfonate 0 -- grhB28 ethyl methanesulfonate 0 -- grhB32 ethyl methanesulfonate 0 -- grhB35 ethyl methanesulfonate 0 -- grhB37 loss of function allele ethyl methanesulfonate 0 Yes grhc00076 piggyBac transposase 0 -- grhc00094 piggyBac transposase 0 -- grhd00702 0 -- grhe03957 piggyBac transposase 0 -- grhFK2131 0 -- grhrH247 P-element activity 0 -- grhunspecified 0 --
Alleles Carried on Transgenic Constructs ( 53 )
For All Alleles Carried on Transgenic Constructs Show
Allele of grh Class Mutagen Stocks Known lesion grhGD9996 in vitro construct - RNAi in vitro construct - regulatory fusion 2 Yes grhJF02305 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes grhKK109135 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes grhKK111269 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes grh2A.Scer\UAS in vitro construct - amino acid replacement 0 Yes grh2E.Scer\UAS in vitro construct - amino acid replacement 0 Yes grhAct5C.PA in vitro construct - regulatory fusion 0 Yes grhCΔ437.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhCΔ707.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhCΔ748.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhCΔ92.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhCΔ930.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhGD4690 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes grhhs.PA in vitro construct - regulatory fusion 0 Yes grhhs.PB in vitro construct - regulatory fusion 0 Yes grhN.Act5C in vitro construct - regulatory fusion 0 Yes grhNΔ114.Act5C in vitro construct - deletion 0 Yes grhNΔ197.Act5C in vitro construct - deletion 0 Yes grhNΔ447.Act5C in vitro construct - deletion 0 Yes grhNΔ447.hs in vitro construct - deletion 0 Yes grhNΔ495.Act5C in vitro construct - deletion 0 Yes grhNΔ619.Act5C in vitro construct - deletion 0 Yes grhNΔ632.Act5C in vitro construct - deletion 0 Yes grhO.Act5C in vitro construct - regulatory fusion 0 Yes grhPanA.Scer\UAS in vitro construct - amino acid replacement 0 Yes grhPCR1.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR11.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR12.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR2.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR4.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR5.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR6.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR7.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR8.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhPCR9.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhRH.Scer\UAS.T:Zzzz\FLAG in vitro construct - regulatory fusion in vitro construct - coding region fusion 0 Yes grhScer\UAS.cBa in vitro construct - regulatory fusion 0 Yes grhScer\UAS.cHa in vitro construct - regulatory fusion 0 Yes grhScer\UAS.cKa in vitro construct - regulatory fusion 0 Yes grhScer\UAS.cUa in vitro construct - regulatory fusion 0 Yes grhsdm1.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhsdm2.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhsdm3.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhsdm4.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhsdm6.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhsdm7.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhsdm8.T:Scer\GAL4 in vitro construct - amino acid replacement in vitro construct - site directed mutagenesis 0 Yes grhT:Scer\GAL4 in vitro construct - coding region fusion in vitro construct - deletion 0 Yes grhΔ114-143.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhΔ114-172.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhΔ114-226.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhΔ176-226.T:Scer\GAL4 in vitro construct - deletion 0 Yes grhΔ204-226.T:Scer\GAL4 in vitro construct - deletion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(2R)02B10W-08 (Mohr and Gelbart, 2002) Df(2R)14H10W-31 (Mohr and Gelbart, 2002) Df(2R)14H10W-33 (Mohr and Gelbart, 2002) Df(2R)14H10W-34 (Mohr and Gelbart, 2002) Df(2R)14H10W-35 (Mohr and Gelbart, 2002) Df(2R)PC4 (Nusslein-Volhard et al., 1984, Tearle and Nusslein-Volhard, 1987, Tearle, 1996.5.14) Df(2R)Pcl7B (Spradling et al., 1999, BDGP Project Members, 1994-1999, Tearle, 1996.5.14, Almeida and Bray, 2005) Df(2R)PclXM82 (Tearle, 1996.5.14) Df(2R)RM2-1 (Spradling et al., 1999, BDGP Project Members, 1994-1999)
Not disrupted in	Df(2R)02B10W-06 (Mohr and Gelbart, 2002) Df(2R)02B10W-07 (Mohr and Gelbart, 2002) Df(2R)14H10W-23 (Mohr and Gelbart, 2002) Df(2R)14H10W-24 (Mohr and Gelbart, 2002) Df(2R)14H10W-25 (Mohr and Gelbart, 2002) Df(2R)14H10W-26 (Mohr and Gelbart, 2002) Df(2R)14H10W-27 (Mohr and Gelbart, 2002) Df(2R)14H10W-28 (Mohr and Gelbart, 2002) Df(2R)14H10W-29 (Mohr and Gelbart, 2002) Df(2R)14H10W-30 (Mohr and Gelbart, 2002) Df(2R)Pcl11B (Tearle, 1996.5.14)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD4690} NA P{KK109135} NA P{KK111269} NA P{TRiP.JF02305} NA P{UAS-grh.2A} NA P{UAS-grh.2E} NA P{UAS-grh.B} NA P{UAS-grh.H} NA P{UAS-grh.K} NA P{UAS-grh.O} NA P{UAS-grh.PanA} NA P{UAS-grh.RH.FLAG} NA reporter construct P{grh-lacZ.NB} No heat-shock construct P{hs-grh.A} NA P{hs-grh.B} NA P{hs-grh.NΔ447} NA
Insertions
	Type of insertions Name Expression data insertion of enhancer trap P{lacW}grhs2140 No P{lacW}l(2)SH0521SH0521 No P{PZ}grh06850 No P{PZ}grhrH247 No miscellaneous insertions P{lacO.hsp26-SIP1.hsp70-w}54F1 NA P{RS5}grh5-HA-1720 NA PBac{PB}grhc00006 NA PBac{PB}grhc00076 NA PBac{PB}grhc00094 NA PBac{PB}grhc01458 NA PBac{PB}grhc07119 NA PBac{RB}grhe03957 NA PBac{WH}grhf03615 NA insertion of mobile activating element P{XP}grhd00702 NA
Gene Ontology: Function, Process & Cellular Component ( 24 unique terms )
Terms Based on Experimental Evidence ( 18 terms )
Molecular Function
CV term Evidence References
DNA binding inferred from direct assay (Uv et al., 1994) protein homodimerization activity inferred from physical interaction with grh (Harrison et al., 2010) sequence-specific DNA binding inferred from direct assay (Harrison et al., 2010) sequence-specific DNA binding transcription factor activity inferred from direct assay (Liaw et al., 1995)
Biological Process
CV term Evidence References
chitin-based cuticle development inferred from mutant phenotype (Kim and McGinnis, 2011) chitin-based embryonic cuticle biosynthetic process inferred from mutant phenotype (Ostrowski et al., 2002) epithelial cell morphogenesis inferred from mutant phenotype (Narasimha et al., 2008) nervous system development inferred from mutant phenotype (Cenci and Gould, 2005) open tracheal system development inferred from expression pattern (Hemphala et al., 2003) plasma membrane organization inferred from mutant phenotype (Hemphala et al., 2003) positive regulation of transcription from RNA polymerase II promoter inferred from expression pattern (Jennings et al., 1999) R3/R4 cell fate commitment inferred from mutant phenotype (Gambis et al., 2011) regulation of cell proliferation inferred from mutant phenotype (Cenci and Gould, 2005) regulation of cell shape inferred from mutant phenotype (Hemphala et al., 2003) regulation of tube length, open tracheal system inferred from mutant phenotype (Hemphala et al., 2003) regulation of wound healing inferred from mutant phenotype (Kim and McGinnis, 2011) tracheal outgrowth, open tracheal system inferred from mutant phenotype (Hemphala et al., 2003)
Cellular Component
CV term Evidence References
nucleus inferred from direct assay (Liaw et al., 1995, Harrison et al., 2010)
Terms Based on Predictions or Assertions ( 8 terms )
Molecular Function ( 0 terms)
Biological Process
CV term Evidence References
cellular membrane organization traceable author statement (Sudarsan, 2003) ganglion mother cell fate determination traceable author statement (Skeath and Thor, 2003) open tracheal system development traceable author statement (Sudarsan, 2003, Ghabrial et al., 2003) primary branching, open tracheal system traceable author statement (Uv et al., 2003) regulation of transcription from RNA polymerase II promoter non-traceable author statement (Swiss-Prot Project Members, 1990.1.1) regulation of tube architecture, open tracheal system traceable author statement (Uv et al., 2003, Ghabrial et al., 2003) ventral cord development non-traceable author statement (Skeath and Thor, 2003)
Cellular Component
CV term Evidence References
nucleus non-traceable author statement (Swiss-Prot Project Members, 1990.1.1)
Sequence Ontology: Class of Gene
	nuclear_gene   protein_coding_gene
Interactions & Pathways
Summary of Physical Interactions
Protein-protein
Interacting group Assay References
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions grh allele Gene References grhFK2131 fz (Lee and Adler, 2004) stan (Lee and Adler, 2004) Vang (Lee and Adler, 2004) grhs2140 bnl (Hemphala et al., 2003)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 62723 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0259211 DroID - A comprehensive database of gene and protein interactions. • FBgn0259211 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37038
Orthologs
Genome-wide drosophilid orthologs	Dana\GF11679 Dere\GG21827 Dgri\GH22029 Dmoj\GI19118 Dsim\GD11321 Dvir\GJ22249 Dwil\GK22231 Dyak\GE11904
Curated drosophilid orthologs	Dpse\GA15658   Dsec\grh (Gleason and Ritchie, 2004) Dsim\grh (Gleason and Ritchie, 2004) Dvir\grh (Uv et al., 1997) Dyak\grh
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AGAP005564-PA Anopheles gambiae (mosquito) XP_394553.3 Apis mellifera (honey bee) 106909 Branchiostoma floridae (lancelet) CE34614 Caenorhabditis elegans (worm) CPIJ004513 Culex pipens (mosquito) ENSCPOP00000005075 Cavia porcellus (guinea pig) ENSGACP00000004851 Gymnopilus aculeatus (mushroom) ENSGALP00000006748 Gallus gallus (chicken) 157385 Lottia gigantea (owl limpet) XP_001601451.1 Nasonia vitripennis (wasp) ENSORLP00000003636 Oryzias latipes (japanese killifish) PHUM009404-PA Pediculus humanus (human body louse) XP_972054.1 Tribolium castaneum (red flour beetle) ENSTNIP00000003891 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000004868 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000008467 Xenopus tropicalis (frog) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB14883 NV11321 LH18591 ISCW008081 FBgn0067967 FBgn0224217 FBgn0209362 FBgn0183071 FBgn0176705 JGI_V11_64332 FBgn0246433 FBgn0154830 FBgn0141857 FBgn0129490 FBgn0114006 FBgn0088718 CPIJ004513 BGIBMGA003479 GB13030 AGAP005564 AAEL000577 GLEAN_04589
Stocks & Reagents
Stocks Listed in FlyBase ( 15 )
Bloomington	10460 w1118; P{lacW}grhs2140/CyO 28820 y1 v1; P{TRiP.JF02305}attP2 12325 P{PZ}grh06850 cn1/CyO; ry506
Harvard	- PBac{PB}grhc00006 - PBac{PB}grhc01458 - PBac{PB}grhc07119 - PBac{WH}grhf03615
Kyoto	125365 w1118; P{RS5}grh5-HA-1720
VDRC	v101428 P{KK109135}VIE-260B v106879 P{KK111269}VIE-260B v33678 w1118; P{GD9996}v33678 v33680 w1118; P{GD9996}v33680
	More stocks available...
Genomic Clones ( 1 )
	BACR20E20 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 77 )
	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	GH11672 IP16310 LD38807 LP11035 RE30607 RE74590
Other clones	DPiM_GH11672
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD47416 LP08909 RE03580 RE25778 RE29067 RE59142 RE64331 RE70111 RE73494 RH17116 RH39342 RH48332 RH59450 SD24995
Other clones	15244 18867 37041 37276 38249 39055 45610 71832 101293 103536 104996 123407 158851 162840 175115 190355 214575.38 CK01.3A.H11 CK01.24D.E9 CK01.82D.E5 CK01.84B.D11 CK01.84D.D11 CK01.90D.D7 CK01.104C.H6 CK01.108C.C3 CK01.108C.G3 CK01.123C.E12 EK022214 EK039248 EK050622 EK065934 EK066119 EK067314 EK068914 EK087663 EK091907 EK093952 EK097910 EK097926 EK118112 EK133754 EK149242 EK156016 EK161140 EK164614 EK200348 EK229530 EK246234 EK261730 EK276423 EK292651 EP19239 EP21259 IP05936 IP16010 IP16110
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0259211 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37038 246463
Antibody Information
	monoclonal (Bray et al., 1989, Bray et al., 1989, Bray et al., 1989) polyclonal (Baumgardt et al., 2009, Harrison et al., 2010, Kim and McGinnis, 2011)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: CG30111 BcDNA:LD38807 (FlyBase, 1996-) Source for merge of: grh CG30111 (FlyBase Genome Annotators, 2008) Source for merge of: grh l(2)06850 (Misra, 2000.11.18)
Additional comments	Annotations CG5058 and CG30111 merged as CG42311 in release 5.9 of the genome annotation. (FlyBase Genome Annotators, 2008) Source for merge of CG30111 BcDNA:LD38807 was a shared cDNA (date:030728). (FlyBase, 1996-)
Other Comments
	In the thorax, grh prolongs neural proliferation by maintaining a mitotically active neuroblast. In the abdomen, grh is a terminal neuroblast factor that restricts postembryonic lineage size by promoting the proapoptotic subfunction of abd-A. (Cenci and Gould, 2005) New annotation (CG30111) in release 3 of the genome annotation. (FlyBase Genome Annotators, 2002-2003) The structure and function of grh mRNAs has been examined. (Uv et al., 1997) grh mutations do not show any dominant interaction with Ubx mutants, or affect expression of Ubx when also mutant for z. (Laney and Biggin, 1996) Variation of a microsatellite within the grh locus has been studied in North American populations of D.melanogaster. (Goldstein and Clark, 1995) Binding sites for the grh transcription factor contribute to the ventral repression of dpp. (Huang et al., 1995) The grh and Trl factors bind to the 11bp tor response element in the tll promoter, raising the possibility that they are involved in the regulation of tll expression. (Liaw et al., 1995) The interaction of grh with its cognate DNA has been studied to characterise the functional domains. Data suggests that domains within grh may interact to regulate the activity of grh protein. (Uv et al., 1994) In vivo and in vitro assays have been used to map the DNA binding domains, the dimerization domain and transcriptional activation domain of the grh gene product. (Attardi and Tjian, 1993) Identified in 2D gels of CMW W2 wing imaginal disc cell proteins. (Santaren et al., 1993) Ubx promoter-Ecol\lacZ reporter gene constructs have been used to study the role of grh in the regulation of Ubx transcription. (Laney and Biggin, 1992) Gene encodes at least three isoforms of a protein that binds to upstream sequences of Ddc, en, ftz and Ubx. The proteins share epitopes as recognized by two monoclonal antibodies. Expression first detected in 4-to-8 hour embryos, peaking from 8 to 12 hours and declining from 12 to 16 hours; NTF protein detected in nuclei of ectodermal derivatives; seen in all epidermal cells and changing subsets of neurons in the developing central nervous system.
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_057494 NM_057495 NM_057496 NM_057497 NM_166250 NM_166252 NM_166253 RefSeq (Proteins) • NP_476842 NP_476843 NP_476844 NP_476845 NP_725723 NP_725725 NP_725726 DDBJ / EMBL / GenBank • AAF57782 AAF57783 AAF57784 AAK93490 AAL28999 AAM68467 AAM68468 AAM68470 AAM68472 AAN71169 AAO24937 AC004328 AC004640 AI134262 AI296916 AI519280 AL035312 AW943443 AW944140 AY052066 AY061451 BG632642 BI234840 BI580402 BT001414 BT003182 CAA22954 CAA33692 CZ466147 CZ466182 CZ466194 CZ467185 CZ471255 CZ481719 CZ481720 CZ486189 G00456 X15657 Entrez Gene - A searchable database of RefSeq genes. • 37038 UniProtKB/Swiss-Prot • P13002 UniProtKB/TrEMBL • A1ZB05 Q95RD9
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • CP2 transcription factor (IPR007604)
Linkouts
	BioGRID - A database of protein and genetic interactions • 62723 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0259211 DroID - A comprehensive database of gene and protein interactions. • FBgn0259211 DRSC - Results from RNAi screens. • FBgn0259211 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0259211 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG30111-RA FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0259211 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 37038 246463 InParanoid A subset of ortholog calls from InParanoid. • AGAP005564-PA Anopheles gambiae (mosquito) XP_394553.3 Apis mellifera (honey bee) 106909 Branchiostoma floridae (lancelet) CE34614 Caenorhabditis elegans (worm) CPIJ004513 Culex pipens (mosquito) ENSCPOP00000005075 Cavia porcellus (guinea pig) ENSGACP00000004851 Gymnopilus aculeatus (mushroom) ENSGALP00000006748 Gallus gallus (chicken) 157385 Lottia gigantea (owl limpet) XP_001601451.1 Nasonia vitripennis (wasp) ENSORLP00000003636 Oryzias latipes (japanese killifish) PHUM009404-PA Pediculus humanus (human body louse) XP_972054.1 Tribolium castaneum (red flour beetle) ENSTNIP00000003891 Tetraodon nigroviridis (green spotted pufferfish) ENSTRUP00000004868 Takifugu rubripes (pufferfish Takifugu) ENSXETP00000008467 Xenopus tropicalis (frog) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /gene/grnhead.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 37038 modMine - Data generated by the modENCODE project. • FBgn0259211 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • PB14883 NV11321 LH18591 ISCW008081 FBgn0067967 FBgn0224217 FBgn0209362 FBgn0183071 FBgn0176705 JGI_V11_64332 FBgn0246433 FBgn0154830 FBgn0141857 FBgn0129490 FBgn0114006 FBgn0088718 CPIJ004513 BGIBMGA003479 GB13030 AGAP005564 AAEL000577 GLEAN_04589 REDfly - A database of transcriptional regulatory elements. • FBgn0259211
Synonyms & Secondary IDs ( 35 )
Reported As
Symbol Synonym	BcDNA:LD38807   CG5058 (Whitfield, 2002.7.26, Carrera et al., 2008, Keleman et al., 2009.8.5) CG30111 (Keleman et al., 2009.8.5) CG42311 (FlyBase Genome Annotators, 2008, Perkins et al., 2009) DREB (Valentine et al., 1998) EG:191D12.1 (Benos, 1999.2.9, ) Elf1 (Tootle and Rebay, 2005, Li and Perrimon, 1997, ) Elf-1 (Gravot et al., 2004, Cobb et al., 2000, Capy et al., 2000, Michalakis and Veuille, 1996, Moussian and Uv, 2005) elf1 (Goldstein and Clark, 1995) Elf-1/NTF-1 (Read and Manley, 1992) elf1(CAG)7 (Flores and Engels, 1999) Gh (Skeath and Thor, 2003, Brody and Odenwald, 2000, Brody and Odenwald, 1999, Brody and Odnewald, 1999) grh (Aerts et al., 2007, Baumgardt et al., 2009, Luschnig et al., 2006, Krattinger et al., 2007, Mace et al., 2005, Almeida and Bray, 2005, Cenci and Gould, 2005, Moussian and Uv, 2005, Odenwald, 2005, Walker et al., 2006, Lovegrove et al., 2006, Nishimura et al., 2007, Mace et al., 2005, Muller and Kassis, 2006, Tuckfield et al., 2002, Blastyak et al., 2006, Maurange et al., 2008, Choksi et al., 2006, Kucherenko et al., 2008, Lichtneckert et al., 2008, Xylourgidis et al., 2006, Narasimha et al., 2008, Steinel and Whitington, 2009, Pearson et al., 2009, Carrera et al., 2008, Dworkin et al., 2009, Maurel-Zaffran et al., 2010, Karlsson et al., 2010, Losada-Pérez et al., 2010, Harrison et al., 2010, Boyle et al., 2010, Gambis et al., 2011, Shaik et al., 2011) Grh (Wang et al., 2007, Juarez et al., 2007, Bello et al., 2006, Bello et al., 2007, Lichtneckert et al., 2007, Nagel et al., 2005, Ringrose and Paro, 2007, Tsuji et al., 2008, Wang et al., 2009) GRH (Tuckfield et al., 2002, Hauenschild et al., 2008, Colonques et al., 2011, Kim and McGinnis, 2011) hsk   l(2)06850 (Spradling et al., 1999, Nicholls and Gelbart, 1998, BDGP Project Members, 1994-1999, ) l(2)IM45   l(2)s2140   Ntf (Lindsley and Zimm, 1992) NTF (Moussian and Uv, 2005) NTF1 (Duncan, 2002) NTF-1 (Valentine et al., 1998, Shirra and Hansen, 1998, Treisman, 1996, Attardi et al., 1993, Attardi and Tjian, 1993, Laney and Biggin, 1992, Muller and Kassis, 2006, Chang et al., 2007) Ntf1   ntf-1 (Muller and Kassis, 2006) NTF-1/Elf-1 (Huang et al., 1995)
Name Synonym	CG30111   CG42311 (FlyBase Genome Annotators, 2008) decapentaplegic repression element binding protein (Valentine et al., 1998) Element I binding activity   grainyhead (Baumgardt et al., 2009, Edenfeld et al., 2002, Uv et al., 1994, Walker et al., 2006, Tuckfield et al., 2002, Blastyak et al., 2006, Karlsson et al., 2010, Harrison et al., 2010, Boyle et al., 2010) Grainyhead (Jones, 2005, Bray and Furriols, 2001, Brody and Odenwald, 2000, Maurange and Gould, 2006, Schuettengruber, 2006, Odenwald, 2005, Bello et al., 2007, Muller and Kassis, 2006, Nagel et al., 2005, Maurange et al., 2008, Tsuji et al., 2008, Xylourgidis et al., 2006, Losada-Pérez et al., 2010) grainy head (Mace et al., 2005, Moussian and Uv, 2005, Choksi et al., 2006, Lichtneckert et al., 2008, Narasimha et al., 2008, Truman et al., 2010) Grainy head (Juarez et al., 2007, Bello et al., 2006, Juarez et al., 2008, Ringrose and Paro, 2007, Wang et al., 2009, Kim and McGinnis, 2011) head skeleton   Neurogenic element binding transcription factor (Lindsley and Zimm, 1992) neuronal transcription factor 1 (Muller and Kassis, 2006)
Secondary FlyBase IDs
	FBgn0000572 FBgn0001136 FBgn0004586 FBgn0010628 FBgn0010683 FBgn0013964 FBgn0014968 FBgn0026798 FBgn0047195 FBgn0050111
References ( 164 )
Generate a list of	All references relating to this gene ( 164 )
List References by type	Research paper  ( 93 ) Supplementary material  ( 6 ) Review  ( 27 ) Personal communication to FlyBase  ( 9 ) Abstract  ( 14 ) FlyBase analysis  ( 5 ) DNA/RNA sequence record  ( 1 ) Computer file  ( 4 ) Book  ( 1 ) Stock list  ( 1 ) Protein sequence record  ( 1 ) Curated genome annotation  ( 1 ) Automatic genome annotation  ( 1 )
Recent research papers ( 12 )
	Benito-Sipos et al., 2011, Development 138(24): 5311--5320 Seven up acts as a temporal factor during two different stages of neuroblast 5-6 development. [FBrf0216799] Colonques et al., 2011, PLoS ONE 6(4): e19342 A Transient Expression of Prospero Promotes Cell Cycle Exit of Drosophila Postembryonic Neurons through the Regulation of Dacapo. [FBrf0213608] Gabilondo et al., 2011, Mech. Dev. 128(3-4): 208--221 A targeted genetic screen identifies crucial players in the specification of the Drosophila abdominal Capaergic neurons. [FBrf0213290] Gambis et al., 2011, Dev. Biol. 351(1): 128--134 Two-color in vivo imaging of photoreceptor apoptosis and development in Drosophila. [FBrf0213000] Kim and McGinnis, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(2): 650--655 Phosphorylation of Grainy head by ERK is essential for wound-dependent regeneration but not for development of an epidermal barrier. [FBrf0214191] Shaik et al., 2011, Glycobiology 21(6): 743--756 The Alg5 ortholog Wollknäuel is essential for correct epidermal differentiation during Drosophila late embryogenesis. [FBrf0214472] Boyle et al., 2010, Dev. Biol. 346(1): 68--79 Division of labor: Subsets of dorsal-appendage-forming cells control the shape of the entire tube. [FBrf0211761] Harrison et al., 2010, Dev. Biol. 345(2): 248--255 Grainyhead and Zelda compete for binding to the promoters of the earliest-expressed Drosophila genes. [FBrf0211626] Karlsson et al., 2010, PLoS Biol. 8(5): e1000368 Segment-specific neuronal subtype specification by the integration of anteroposterior and temporal cues. [FBrf0210822] Losada-Pérez et al., 2010, Mech. Dev. 127(9-12): 458--471 Lineage-unrelated neurons generated in different temporal windows and expressing different combinatorial codes can converge in the activation of the same terminal differentiation gene. [FBrf0212045] Maurel-Zaffran et al., 2010, Dev. Biol. 343(1-2): 18--27 Reiterative use of signalling pathways controls multiple cellular events during Drosophila posterior spiracle organogenesis. [FBrf0211024] Truman et al., 2010, Development 137(1): 53--61 Role of Notch signaling in establishing the hemilineages of secondary neurons in Drosophila melanogaster. [FBrf0209606] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...