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Gene Dmel\hep

General Information
Symbol	Dmel\hep	Species	D. melanogaster
Name	hemipterous	Annotation symbol	CG4353
Feature type	protein_coding_gene	FlyBase ID	FBgn0010303
Gene Model Status	Current	Stock availability	19 publicly available
Also Known As	JNKK, HEP/MKK7, MKK7, DJNKK, hp
Genomic Location
Chromosome (arm)	X	Recombination map	1-
Cytogenetic map	11D10-11D10	Sequence location	X:12,973,240..12,984,466 [-]
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 hemipterous is referred to in FlyBase by the symbol Dmel\hep (CG4353, FBgn0010303). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; protein kinase activity. There is experimental evidence for 26 unique biological process terms, many of which group under: anatomical structure development; cellular component organization or biogenesis; biological regulation; cellular process involved in reproduction; multicellular organismal reproductive process; cellular response to inorganic substance; chemical homeostasis; post-embryonic organ morphogenesis; response to stress; response to chemical stimulus. 47 alleles are reported. The phenotypes of these alleles are annotated with: organ system subdivision; adult segment; organ system; imaginal precursor; external compound sense organ; adult mesothoracic segment; thoracic segment; non-connected developing system; nervous system; larval thorax. It has 3 annotated transcripts and 3 annotated polypeptides. Protein features are: Protein kinase, ATP binding site; Protein kinase, catalytic domain; Protein kinase-like domain; Serine/threonine-protein kinase, active site; Serine/threonine-protein kinase, catalytic domain; Serine/threonine-protein kinase-like domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of moderate expression. Peak expression observed at stages throughout embryogenesis, in adult female stages. 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 at moderate levels in the following post-embryonic organs or tissues: adult eye, larval/adult central nervous system, adult crop, larval midgut, larval hindgut, larval Malpighian tubules, adult fat body, larval/adult salivary gland, larval trachea, adult female reproductive system, larval carcass. Gene sequence location is X:12973240..12984466.
External Summaries
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	Transgenic Constructs P{NIG.4153R} Alleles hep[NIG.4153R]
FB2012_01	Sequence features BKN51194 BKN42143 BKN42747 HFA19839 BKN51002 BKN51003 BKN51196 HFA20337 References Jezowska et al., 2011, Dev. Biol. 360(1): 143--159 Umehara et al., 2010, Molec. Cell. Biochem. 342(1-2): 223--232 Cobreros-Reguera et al., 2010, EMBO Rep. 11(12): 943--949 Kwon et al., 2010, Biochem. Biophys. Res. Commun. 393(4): 656--661
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 11D10-11D10   Limits computationally determined from genome sequence between P{EP}EP1461 and P{EP}CG1640EP1626
Experimentally Determined Cytological Location
Cytogenetic map Notes References 11D1-11D4   (determined by in situ hybridisation) 11D (determined by in situ hybridisation)   (Peter et al., 2002) 11D-11D   (determined by in situ hybridisation)   (Schaefer et al., 1999.12.2) 11D1-11D4   (determined by in situ hybridisation)   (Schaefer et al., 1999.12.10) 11D-11D     (Noselli, 1994.1.18) 11D1-11D2     (Noselli, 1996.4.28) 11D1-11D2   (determined by in situ hybridisation)   (Glise et al., 1995) 11B12-13F   Determined by complementation mapping, aberrations not specified.   (Perrimon et al., 1989)
Experimentally Determined Recombination Data
Location	1-
Left of (cM)
Right of (cM)
Notes
Gene Model & Products
Please see the GBrowse view of Dmel\hep 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) hep-RA FBtr0073747  NM_167346   4908   1178 hep-RB FBtr0073749  NM_167347   2986   580 hep-RC FBtr0073748  NM_078587   2724   492
Additional Transcript Data & Comments
Reported size (kB)	6.0 (northern blot) (Glise et al., 1995)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kDa) Length (aa) Theoretical pI RefSeq ID GenBank protein hep-PA   FBpp0073578   125.1   1178   10.05     NP_727661   AAN09646 hep-PB   FBpp0073580   61.9   580   10.22     NP_727662   AAG22351 hep-PC   FBpp0073579   53.1   492   9.19     NP_511142   AAF48222
Additional Polypeptide Data & Comments
Reported size (kDa)	487 (aa) (Glise et al., 1995)
Comments
External Data
Linkouts
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Protein kinase, catalytic domain (IPR000719) Serine/threonine-protein kinase, catalytic domain (IPR002290) Serine/threonine-protein kinase, active site (IPR008271) Protein kinase-like domain (IPR011009) Protein kinase, ATP binding site (IPR017441) Serine/threonine-protein kinase-like domain (IPR017442)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA202573   AA942111   AE014298 AAF48222   AE014298 AAG22351   AE014298 AAN09646   AI062601   AI296171   AI515005   AJ426887   AJ426888   AW943860   AY069695 AAL39840   AY071210 AAL48832   BG636753   BG637754   BG637756   BI228126   BI363974   BI364064   BI365530   BI366407   BI367334   BI368106   BI368259   BI376190   BI638067   BI642020   BT044138   CK659618   CK660969   CO154210   CO156435   CO265915   CO271003   CO276812   CO279004   CO283822   CO295582   CO296410   CO298147   CO308665   CO338382   CO341236   CZ485498   EC050995   EC051985   EC054479   EC054992   EC055875   EC072734   EC197439   EC197700   EC198938   EC204096   EC213066   EC215075   EC217374   EC228883   EC235957   EC255180   EC263496   EC263998   EC267360   GH775842   GH807138   GH894015   GH915979   U05240 AAC46944   U93032 AAB63449
UniProtKB/Swiss-Prot	Q23977
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
External Data
Linkouts
Crossreferences
Expression Data
Transcript Expression
northern blot Stage Tissue/Position (including subcellular localization) Reference embryonic stage   organism | ubiquitous   (Glise et al., 1995) embryonic stage -- adult stage     (Glise et al., 1995)
Additional Descriptive Data
	hep transcripts are detected throughout development on northern blots. In situ hybridization shows that they are homogeneously distributed throughout the embryo. (Glise et al., 1995)
Marker for
Subcellular Localization
CV Term
Notes
Polypeptide Expression
Additional Descriptive Data
Marker for
Subcellular Localization
CV Term
Notes
High-Throughput Expression Data
or
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes. [an error occurred while processing this directive] 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_01
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0010303 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4353-RC;CG4353-RA;CG4353-RB
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele cell lethal | partially | somatic clone, with Scer\GAL4αTub84B.PL hepAct.Scer\UAS lethal, with Scer\GAL469B hepAct.Scer\UAS lethal | embryonic stage, with Scer\GAL469B hepAct.Scer\UAS lethal | embryonic stage, with Scer\GAL4e22c hepCA.Scer\UAS lethal | embryonic stage | maternal effect | recessive hep1 lethal | embryonic stage | recessive hep1 lethal | embryonic stage | rescuable maternal effect hep1 lethal | heat sensitive, with Scer\GAL4vg.PU hepCA.Scer\UAS lethal | pharate adult stage hepG0390 lethal | pupal stage hepG0107 hepG0390 hepr75 lethal | pupal stage | recessive hep699 hepr75 lethal | recessive hep7P1 hepG0107 hepr39 hepr45 hepr51 hepr75 heprh1 heprh8 heprh19 heprh31 heprh99 lethal | recessive | rescuable maternal effect hep1 semi-lethal | female hepG0208 semi-lethal | male hepG0208 semi-lethal | recessive hepG0208 viable hep1 viable, with Scer\GAL4pnr-MD237 hepGD1461 viable (with hep1) hep699 viable (with hep699) hep1 viable | female hep1 Other Phenotypes Allele cell shape defective | embryonic stage, with Scer\GAL4arm.PS hepAct.Scer\UAS chemical resistant, with Scer\GAL4arm.PS hepScer\UAS.cBa chemical resistant | RU486 conditional, with Scer\GAL4elav.Switch.PO hepScer\UAS.cBa chemical sensitive hep1 increased cell death hepCA increased cell death | heat sensitive, with Scer\GAL4en-e16E, Scer\GAL80ts.αTub84B hepCA increased cell death | somatic clone, with Scer\GAL4GMR.PF hepCA.Scer\UAS long lived, with Scer\GAL4elav-C155 hepScer\UAS.cBa mitotic cell cycle defective | somatic clone hepr75 neuroanatomy defective hep1 hepr75 neuroanatomy defective, with Scer\GAL4ato.3.6 hepCA.Scer\UAS neuroanatomy defective, with Scer\GAL4P2.4.Pdf hepCA.Scer\UAS neuroanatomy defective | somatic clone hepr39 hepr75 planar polarity defective, with Scer\GAL4hs.2sev hepAct.Scer\UAS hepScer\UAS.cBa small body | adult stage, with Scer\GAL4Ilp2.PR hepAct.Scer\UAS stress response defective hep1 stress response defective, with Scer\GAL4ppl.PP hepScer\UAS.cBa visible hep1 visible, with Scer\GAL4GMR.PF hepCA.Scer\UAS visible, with Scer\GAL4hs.2sev hepAct.Scer\UAS visible, with Scer\GAL4pnr-MD237 hepScer\UAS.cBa visible (with hep1) hep699 hepr39 hepr75 visible (with hep699) hep1 visible (with hepr39) hep1 visible (with hepr75) hep1 visible | dominant hepCA visible | recessive hep1 wild-type hephs.PG hepUbi-p63E.PG wound healing defective hepr75 Phenotype manifest in Allele adult head hepr75 adult head | rescuable maternal effect hep1 adult mesothoracic segment | dorsal | rescuable maternal effect hep1 adult thorax (with hep1) hepr39 hepr75 adult thorax (with hepr39) hep1 adult thorax (with hepr75) hep1 adult thorax & macrochaeta, with Scer\GAL4pnr-MD237 hepScer\UAS.cBa adult thorax | dorsal hep1 adult thorax | dorsal, with Scer\GAL4pnr-MD237 hepScer\UAS.cBa adult thorax | dorsal | maternal effect hep1 amnioserosa hep1 hepr75 axon & dorsal cluster neuron hep1 axon & dorsal cluster neuron, with Scer\GAL4ato.3.6 hepCA.Scer\UAS axon | somatic clone hepr39 hepr75 basement membrane | pupal stage hepr75 cephalopharyngeal skeleton hepr75 cephalopharyngeal skeleton | germline clone hepr75 chaeta (with hep1) hepr39 hepr75 chaeta (with hepr39) hep1 chaeta (with hepr75) hep1 cuticle & abdomen hepr75 cuticle & abdomen | germ-line clone hepr75 cuticle & thorax hepr75 cuticle & thorax | germ-line clone hepr75 cuticle | dorsal | rescuable maternal effect (with hep1) hepr75 cuticle | dorsal | rescuable maternal effect (with hepr75) hep1 dorsal appendage hep1 dorsal appendage | maternal effect | somatic clone hepr75 dorsal closure embryo hep1 hepr75 dorsal closure embryo | germline clone hep7P1 hepG24 hepH6 dorsal mesothoracic disc & cytoskeleton hepr75 egg chamber | anterior | somatic clone hepr75 embryonic/first instar larval cuticle, with Scer\GAL469B hepAct.Scer\UAS hepScer\UAS.cBa embryonic/first instar larval cuticle, with Scer\GAL4arm.PS hepAct.Scer\UAS embryonic/first instar larval cuticle | dorsal hep1 embryonic/first instar larval cuticle | dorsal | rescuable maternal effect hep1 embryonic/larval hemolymph | third instar larval stage, with Scer\GAL4Act5C.PU hepdsRNA.Scer\UAS embryonic dorsal epidermis hep1 hepr75 embryonic epidermis hep1 hepr75 embryonic leading edge cell hepunspecified embryonic leading edge cell, with Scer\GAL4ptc-559.1 hepAct.Scer\UAS hepScer\UAS.cBa embryonic leading edge cell & actin filament hep1 hepr75 embryonic leading edge cell & filopodium hepr75 embryonic leading edge cell | rescuable maternal effect hep1 embryonic leading edge cell | rescuable maternal effect (with hep1) hepr75 embryonic leading edge cell | rescuable maternal effect (with hepr75) hep1 embryonic maxillary segment, with Scer\GAL4arm.PS hepAct.Scer\UAS eye hep1 eye, with Scer\GAL4GMR.PF hepCA.Scer\UAS hepScer\UAS.cGa eye, with Scer\GAL4hs.2sev hepAct.Scer\UAS hepScer\UAS.cBa eye, with Scer\GAL4sev.EP hepAct.Scer\UAS eye | somatic clone, with Scer\GAL4GMR.PF hepCA.Scer\UAS eye-antennal disc hepr75 eye disc, with Scer\GAL4hs.2sev hepAct.Scer\UAS eye photoreceptor cell hepr75 female terminalia (with hep1) hep699 female terminalia (with hep699) hep1 follicle cell | somatic clone hepr75 genital disc, with Scer\GAL4Abd-B-LDN hepScer\UAS.cBa genital disc, with Scer\GAL4ptc-559.1 hepScer\UAS.cBa imaginal disc hepr39 hepr75 interommatidial bristle hepr75 interommatidial bristle, with Scer\GAL4GMR.PF hepScer\UAS.cGa Kenyon cell | somatic clone hepr39 hepr75 lamellocyte, with Scer\GAL4He.PZ hepCA.Scer\UAS lamellocyte, with Scer\GAL4Hml.PG hepCA.Scer\UAS lamina hepr75 leg hep1 leg | pupal stage hepr75 macrochaeta | maternal effect hep1 maxillary palpus (with hep1) hep699 maxillary palpus (with hep699) hep1 melanotic mass, with Scer\GAL4He.PZ hepCA.Scer\UAS melanotic mass, with Scer\GAL4Hml.PG hepCA.Scer\UAS mesothoracic tergum hep1 micropyle | maternal effect | somatic clone hepr75 neuron hepr75 ommatidial precursor cluster, with Scer\GAL4hs.2sev hepAct.Scer\UAS ommatidium, with Scer\GAL4hs.2sev hepScer\UAS.cBa peripodial epithelium & dorsal mesothoracic disc & prepupa hepr75 peripodial stalk & dorsal mesothoracic disc & prepupa hepr75 pigment cell hepr75 scutellum, with Scer\GAL4pnr-MD237 hepScer\UAS.cBa thorax hep1 thorax, with Scer\GAL4pnr-MD237 hepdsRNA.Scer\UAS ventral adult lateral neuron & axon, with Scer\GAL4P2.4.Pdf hepCA.Scer\UAS ventral thoracic disc hepr75 wing hep1 hepCA wing, with Scer\GAL4vg.PU hepCA.Scer\UAS wing | pupal stage hepr75 wing | rescuable maternal effect hep1 wing disc hep1 hepCA hepr39 hepr75 wing disc | prepupal stage hepr75 wing margin, with Scer\GAL4vg.PU hepCA.Scer\UAS wing vein, with Scer\GAL4vg.PU hepCA.Scer\UAS wing vein L5, with Scer\GAL4CY2 hepCA.Scer\UAS
Classical Alleles ( 29 )
For All Classical Alleles Show
Allele of hep Class Mutagen Stocks Known lesion hepc02950 piggyBac transposase 2 -- hepG0107 P-element activity 2 -- hepf02941 piggyBac transposase 1 -- hepG0208 P-element activity 1 -- hepGG01725 P-element activity 1 -- hepr75 loss of function allele, hypomorphic allele - genetic evidence PM hybrid dysgenesis 1 Yes hep1 hypomorphic allele - genetic evidence PM hybrid dysgenesis P-element activity 0 Yes hep699 0 -- hep75 0 -- hep7P1 0 -- hepCA P-element activity 0 Yes hepG0390 P-element activity 0 -- hepG24 P-element activity 0 Yes hepGG01622 0 -- hepH49 P-element activity 0 Yes hepH6 P-element activity 0 Yes hepH81 P-element activity 0 Yes hepM56 P-element activity 0 Yes hepr39 PM hybrid dysgenesis 0 Yes hepr45 PM hybrid dysgenesis 0 Yes hepr51 PM hybrid dysgenesis 0 Yes heprh19 0 -- heprh1 0 -- heprh31 0 -- heprh8 0 -- heprh99 0 -- hepunspecified 0 -- hepv39 0 -- hepv75 0 --
Alleles Carried on Transgenic Constructs ( 18 )
For All Alleles Carried on Transgenic Constructs Show
Allele of hep Class Mutagen Stocks Known lesion hepGD1461 in vitro construct - RNAi in vitro construct - regulatory fusion 3 Yes hepAct.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 2 Yes hepCA.Scer\UAS in vitro construct - regulatory fusion in vitro construct - amino acid replacement 2 Yes hepScer\UAS.cBa in vitro construct - regulatory fusion 2 Yes hepGL00089 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hepJF03137 in vitro construct - RNAi in vitro construct - regulatory fusion 1 Yes hepdsRNA.cBa in vitro construct - RNAi 0 Yes hepdsRNA.cCa in vitro construct - RNAi 0 Yes hepdsRNA.cIa in vitro construct - RNAi 0 Yes hepdsRNA.cSa in vitro construct - RNAi 0 Yes hepdsRNA.Scer\UAS in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes hepGD8726 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes hephs.PG in vitro construct - regulatory fusion 0 Yes hepKK116295 in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes hepNIG.2190R in vitro construct - RNAi in vitro construct - regulatory fusion 0 Yes hepNIG.4153R in vitro construct - regulatory fusion in vitro construct - RNAi 0 Yes hepScer\UAS.cGa in vitro construct - regulatory fusion 0 Yes hepUbi-p63E.PG in vitro construct - regulatory fusion 0 Yes
Aneuploid Aberrations
Disrupted in	Df(1)G24 (Suzanne et al., 1999, Paricio et al., 1999) Df(1)H6 (Suzanne et al., 1999, Paricio et al., 1999, Weber et al., 2000) Df(1)H49 (Suzanne et al., 1999) Df(1)H81 (Suzanne et al., 1999) Df(1)M56 (Suzanne et al., 1999) Df(1)N12 (Glise et al., 1995)
Duplicated in	Dp(1;3)DC264 (Popodi et al., 2010-)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD1461} NA P{GD8726} NA P{KK116295} NA P{TRiP.GL00089} NA P{TRiP.JF03137} NA P{UAS-Hep.Act} NA P{UAS-hep.B} NA P{UAS-hep.CA} NA P{UAS-hep.G} NA P{UAS-hep.IR} NA heat-shock construct P{HShep} NA characterization construct P{NIG.2190R} NA P{NIG.4153R} NA P{UBhep} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{?SDFL}hepr45 NA P{?SDFL}hepr51 NA P{SDFL}hep1 NA P{UAS-hep.CA}CA NA PBac{PB}hepc02950 NA PBac{WH}hepf02941 NA insertion of enhancer trap P{lacW}hepG0107 No P{lacW}hepG0208 No P{lacW}hepG0390 No insertion of mobile activating element P{Mae-UAS.6.11}hepGG01622 NA P{Mae-UAS.6.11}hepGG01725 NA
Gene Ontology: Function, Process & Cellular Component ( 40 unique terms )
Terms Based on Experimental Evidence ( 29 terms )
Molecular Function
CV term Evidence References
protein binding inferred from physical interaction with Cka (Chen et al., 2002) protein kinase activity inferred from direct assay (Glise et al., 1995)
Biological Process
CV term Evidence References
actin filament bundle assembly inferred from mutant phenotype (Kaltschmidt et al., 2002) axon extension inferred from mutant phenotype (Srahna et al., 2006) basement membrane disassembly inferred from mutant phenotype (Srivastava et al., 2007) border follicle cell migration inferred from genetic interaction with Pvr (Mathieu et al., 2007) carbohydrate homeostasis inferred from mutant phenotype (Hull-Thompson et al., 2009) cellular response to arsenic-containing substance inferred from mutant phenotype (Ryabinina et al., 2006) cellular response to cadmium ion inferred from mutant phenotype (Ryabinina et al., 2006) chorion-containing eggshell pattern formation inferred from mutant phenotype (Suzanne et al., 2001) determination of adult lifespan inferred from mutant phenotype (Wang et al., 2003) dorsal appendage formation inferred from mutant phenotype (Suzanne et al., 2001) dorsal closure inferred from mutant phenotype (Kaltschmidt et al., 2002) dorsal closure, spreading of leading edge cells inferred from mutant phenotype (Glise et al., 1995) filopodium assembly inferred from mutant phenotype (Kaltschmidt et al., 2002) imaginal disc eversion inferred from mutant phenotype (Pastor-Pareja et al., 2004, Srivastava et al., 2007) imaginal disc fusion inferred from mutant phenotype (Pastor-Pareja et al., 2004) lamellipodium assembly inferred from mutant phenotype (Kaltschmidt et al., 2002) melanization defense response inferred from mutant phenotype (Bidla et al., 2007) micropyle formation inferred from mutant phenotype (Suzanne et al., 2001) neuron development inferred from mutant phenotype (Rallis et al., 2010) ovarian follicle cell development inferred from mutant phenotype (Jordan et al., 2006) positive regulation of JNK cascade inferred from mutant phenotype (Ryabinina et al., 2006) positive regulation of JUN kinase activity inferred from mutant phenotype (Uhlirova et al., 2005) regulation of hemocyte differentiation inferred from mutant phenotype (Zettervall et al., 2004) response to heat inferred from mutant phenotype (Karpac et al., 2009) triglyceride homeostasis inferred from mutant phenotype (Hull-Thompson et al., 2009) wound healing inferred from mutant phenotype (Bosch et al., 2005)
Cellular Component
CV term Evidence References
axon inferred from direct assay (Rallis et al., 2010)
Terms Based on Predictions or Assertions ( 13 terms )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR002290, InterPro:IPR017441 (FlyBase Curators et al., 2004-) JUN kinase kinase activity inferred from sequence or structural similarity (Glise et al., 1995) traceable author statement (Stronach and Perrimon, 2002, Noselli and Agnes, 1999) MAP kinase kinase activity inferred from sequence or structural similarity with NCBI_gi:4826946 (FlyBase, 1992-) non-traceable author statement (Swiss-Prot Project Members, 1997.2.1) traceable author statement (Martin-Blanco, 2000) protein serine/threonine kinase activity inferred from electronic annotation with InterPro:IPR008271 (FlyBase Curators et al., 2004-) receptor signaling protein serine/threonine kinase activity non-traceable author statement (Morrison et al., 2000)
Biological Process
CV term Evidence References
dorsal closure non-traceable author statement (Swiss-Prot Project Members, 1997.2.1, Van Aelst and Symons, 2002) traceable author statement (Riesgo-Escovar et al., 1996, Noselli and Agnes, 1999) establishment of planar polarity non-traceable author statement (Mlodzik, 2002) imaginal disc fusion, thorax closure traceable author statement (Gibson and Schubiger, 2001) JNK cascade traceable author statement (Riesgo-Escovar et al., 1996, Noselli and Agnes, 1999, Harden, 2002) MAPKKK cascade non-traceable author statement (Martin-Blanco, 2000) ovarian follicle cell development traceable author statement (Dobens and Raftery, 2000) protein phosphorylation non-traceable author statement (Morrison et al., 2000) traceable author statement (Noselli and Agnes, 1999) regulation of transcription from RNA polymerase II promoter non-traceable author statement (Swiss-Prot Project Members, 1997.2.1)
Cellular Component ( 0 terms)
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 hep allele Gene References hep1 arm (Kaltschmidt et al., 2002) by (Lee et al., 2003) cno (Takahashi et al., 1998) dsh (Kaltschmidt et al., 2002) egr (Igaki et al., 2002) Jra (Hou et al., 1997) msn (Su et al., 1998) puc (Martin-Blanco et al., 1998, Zeitlinger and Bohmann, 1999, Agnes et al., 1999, Weber et al., 2000) pyd (Takahashi et al., 1998) Rac1 (Fanto et al., 2000, Stronach and Perrimon, 2002) Rala (Sawamoto et al., 1999) rpr (Kuranaga et al., 2002) Tak1 (Takatsu et al., 2000) wg (Kaltschmidt et al., 2002) hepCA.Scer\UAS Cka (Chen et al., 2002) hepCA bi (Adachi-Yamada et al., 1999) Cka (Chen et al., 2002) dpp (Adachi-Yamada et al., 1999) tkv (Adachi-Yamada et al., 1999) hepr39 puc (Agnes et al., 1999) hepr75 dsh (Boutros et al., 1998, Paricio et al., 1999) lace (Adachi-Yamada et al., 1999) msn (Su et al., 1998, Paricio et al., 1999) puc (Agnes et al., 1999, Bosch et al., 2005) Pvr (Ishimaru et al., 2004) Rac1 (Fanto et al., 2000) Rala (Sawamoto et al., 1999) Src42A (Tateno et al., 2000) Tak1 (Mihaly et al., 2001) tkv (Adachi-Yamada et al., 1999) hepScer\UAS.cBa arm (McEwen et al., 2000) by (Lee et al., 2003) dsh (Boutros et al., 1998) unspecified bsk (Cha et al., 2005, Geuking et al., 2005) btl (Srahna et al., 2006) chm (Miotto et al., 2006) dsh (Srahna et al., 2006) egr (Polaski et al., 2006) flw (Kirchner et al., 2007) grim (Mueller et al., 2005) H (Mueller et al., 2005) Jra (Miotto et al., 2006) kay (Miotto et al., 2006) L (Singh et al., 2006) N (Zecchini et al., 1999) p53 (Mueller et al., 2005) park (Cha et al., 2005) Pvf1 (Mathieu et al., 2007) Pvr (Mathieu et al., 2007) Rac1 (Srahna et al., 2006) Ras85D (Uhlirova and Bohmann, 2006) Rpd3 (Miotto et al., 2006) rpr (Mueller et al., 2005) Traf4 (Cha et al., 2003, Kuranaga et al., 2002) W (Mueller et al., 2005)
External Data
Linkouts	BioGRID - A database of protein and genetic interactions • 58642 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0010303 DroID - A comprehensive database of gene and protein interactions. • FBgn0010303 InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 32256
Orthologs
Genome-wide drosophilid orthologs	Dana\GF20874 Dere\GG19526 Dgri\GH24549 Dmoj\GI16263 Dper\GL16492 Dpse\GA22606 Dsec\GM11518 Dsim\GD15909 Dvir\GJ16904 Dwil\GK25249 Dyak\GE16181
Curated drosophilid orthologs	Dpse\GA18128
Linkouts	InParanoid A subset of ortholog calls from InParanoid. • AAEL003359-PA Aedes aegypti (mosquito) XP_396834.1 Apis mellifera (honey bee) XP_001944527.1 Acyrthosiphon pisum (pea aphid) 83417 Branchiostoma floridae (lancelet) BGIBMGA007646-PA Bombyx mori (silk worm) 191434 Hypsipyla robusta (cedar shoot borer) LmjF21.0853 Leishmania major (virus) XP_001604642.1 Nasonia vitripennis (wasp) PHUM005282-PA Pediculus humanus (human body louse) XP_001182749.1 Strongylocentrotus purpuratus (purple sea urchin) OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0233723 FBgn0204083 FBgn0166461 FBgn0244009 FBgn0154096 FBgn0139012 FBgn0132005 FBgn0111728 FBgn0097880
Stocks & Reagents
Stocks Listed in FlyBase ( 19 )
Bloomington	9308 w*; P{UAS-hep.B}2 9306 w*; P{UAS-Hep.Act}2 6761 w* hepr75/FM7c 6406 y1 w1118; P{UAS-hep.CA}4 9307 P{UAS-hep.B}1, w*/FM6, w* 9305 P{UAS-Hep.Act}1, w*/FM6, w*
Harvard	- PBac{PB}hepc02950 - PBac{WH}hepf02941
Kyoto	108771 y1 w1118; P{UAS-hep.CA}4
VDRC	v2968 w1118 P{GD1461}v2968 v47507 w1118; P{GD1461}v47507 v47509 w1118; P{GD1461}v47509
	More stocks available...
Genomic Clones ( 1 )
	BACR06C04 Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
cDNA Clones ( 58 )
	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	LD46661 RE25570
Other clones	DPiM_FI03256 FI03256
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	GH01858 LD02655 LP10004 LP17496 LP19975 RE48977 RE49105 RE50877 RE51937 RE53076 RE54021 RE54196 RE61253 SD14533 SD19143 SD22516
Other clones	905 19814 33709 77133 80688 156000 162827 214341.5 523258 527928 571344 CK01.6C.F10 CK01.7D.B10 CK01.133C.F2 EC050995 EC051985 EC054479 EC054992 EC055875 EK101353 EK122122 EK132761 EK144442 EK164525 EK171605 EK210751 EK213563 EK259044 EN03722 EN08049 EN16653 EP12105 LD02714 LD26151 RP003033601 RP003165185 RP003241961 RP003625883
RNAi & Array Information
Linkouts	DRSC - Results from RNAi screens. • FBgn0010303 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 32256
Antibody Information
	polyclonal (Rallis et al., 2010)
Other Information
Discoverer
Etymology
Identification
Relationship to Other Genes
Source for database identity of
Source for database merge of	Source for merge of: hep CG2190 (FlyBase Genome Annotators, 2002-2003) Source for merge of: hep l(1)G0107 l(1)G0208 (Peter et al., 2002)
Additional comments	Annotations CG4353 and CG2190 merged as CG4353 in release 3 of the genome annotation. (FlyBase Genome Annotators, 2002-2003)
Other Comments
	dsRNA made from templates generated with primers directed against this gene tested in RNAi screen for effects on Kc167 and S2R+ cell morphology. (Kiger et al., 2003) dsRNA made from templates generated with primers directed against this gene is tested in an RNAi screen for effects on actin-based lamella formation. (Rogers et al., 2003) hep is necessary for actin-cable assembly and actin-based cell process formation in leading edge cells during dorsal closure. (Kaltschmidt et al., 2002) Shows particularly robust cycling of transcription in adult heads, as assessed by expression analysis using high density oligonucleotide arrays with probe generated during three 12-point time course experiments over the course of 6 days. (Claridge-Chang et al., 2001) hep is essential for the correct morphogenesis of the dorsal appendage and micropyle. (Suzanne et al., 2001) hep controls puc expression but not dpp expression in imaginal discs. (Agnes et al., 1999) hep and the JNK pathway are required for correct morphogenesis of the imaginal discs. (Agnes et al., 1999) Candidate gene for posterior lobe area quantitative trait locus. (Macdonald and Goldstein, 1999) Show no or weak ommatidial precursor polarity phenotypes in imaginal tissues. (Paricio et al., 1999) The JNK pathway (which includes hep) is required for thorax closure during metamorphosis. (Zeitlinger and Bohmann, 1999) hep is required for normal dpp expression in the leading edge during dorsal closure. puc is both a repressor and a target of hep function in the leading edge. The combined and antagonistic functions of hep and puc maintain appropriate levels of puc and dpp activities in migrating epithelia during dorsal closure. These results indicate a leading edge cell identity and dorsal closure depend on a balance between bsk activation and puc repression. (Glise and Noselli, 1997) The JNK pathway is conserved between vertebrates and Drosophila and is involved in controlling cell morphogenesis in Drosophila. (Riesgo-Escovar et al., 1996) Data suggests that hep functions in a novel MAPK pathway controlling puc expression and morphogenetic activity of the dorsal epidermis. (Glise et al., 1995)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Transcripts) • NM_078587 NM_167346 NM_167347 RefSeq (Proteins) • NP_511142 NP_727661 NP_727662 DDBJ / EMBL / GenBank • AAB63449 AAC46944 AAF48222 AAG22351 AAL39840 AAL48832 AAN09646 AI515005 AJ426887 AJ426888 AW943860 AY069695 AY071210 BI228126 CZ485498 U05240 U93032 Entrez Gene - A searchable database of RefSeq genes. • 32256 UniProtKB/Swiss-Prot • Q23977
Other Crossreferences
	InterPro domains - A database of protein families, domains, and functional sites • Protein kinase, catalytic domain (IPR000719) Serine/threonine-protein kinase, catalytic domain (IPR002290) Serine/threonine-protein kinase, active site (IPR008271) Protein kinase-like domain (IPR011009) Protein kinase, ATP binding site (IPR017441) Serine/threonine-protein kinase-like domain (IPR017442)
Linkouts
	BioGRID - A database of protein and genetic interactions • 58642 DPiM - DPiM, Drosophila Protein interaction Map • FBgn0010303 DroID - A comprehensive database of gene and protein interactions. • FBgn0010303 DRSC - Results from RNAi screens. • FBgn0010303 FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0010303 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • CG4353-RC;CG4353-RA;CG4353-RB FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans • FBgn0010303 FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways • 92142 GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents • 32256 InParanoid A subset of ortholog calls from InParanoid. • AAEL003359-PA Aedes aegypti (mosquito) XP_396834.1 Apis mellifera (honey bee) XP_001944527.1 Acyrthosiphon pisum (pea aphid) 83417 Branchiostoma floridae (lancelet) BGIBMGA007646-PA Bombyx mori (silk worm) 191434 Hypsipyla robusta (cedar shoot borer) LmjF21.0853 Leishmania major (virus) XP_001604642.1 Nasonia vitripennis (wasp) PHUM005282-PA Pediculus humanus (human body louse) XP_001182749.1 Strongylocentrotus purpuratus (purple sea urchin) Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution • /newgene/hemipter.htm InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets. • 32256 modMine - Data generated by the modENCODE project. • FBgn0010303 OrthoDB (Arthropod subset) The hierarchical catalog of eukaryotic orthologs. • FBgn0233723 FBgn0204083 FBgn0166461 FBgn0244009 FBgn0154096 FBgn0139012 FBgn0132005 FBgn0111728 FBgn0097880
Synonyms & Secondary IDs ( 27 )
Reported As
Symbol Synonym	CG2190 (Claridge-Chang et al., 2001, ) CG4353 (Morrison et al., 2000, Rogers et al., 2003, Perkins et al., 2009, Keleman et al., 2009.8.5, Ni et al., 2011.7.19) DHEP/MKK7 (Chen et al., 2002) DJNKK (Miotto et al., 2006) dJNKK (Bidla et al., 2007) DMKK7 (Ip and Davis, 1998) D-MKK7 (Ryabinina et al., 2006) hem (Chanut-Delalande et al., 2006) HEP (Udomsinprasert et al., 2004, Weitzman, 2000, Sluss and Davis, 1997) hep (Singh et al., 2006, Pastor-Pareja et al., 2004, Walther and Pichaud, 2007, Uhlirova et al., 2005, Murray et al., 2006, Ryoo et al., 2004, Stronach et al., 2007, Mulinari et al., 2006, Blanco et al., 2010, Yoshioka et al., 2008, Karpac et al., 2009, Lennox and Stronach, 2010, Srahna et al., 2006, Cerrato et al., 2006, Kirchner et al., 2007, Jordan et al., 2006, Uhlirova and Bohmann, 2006, Hueber et al., 2007, Polaski et al., 2006, Domingues and Ryoo, 2008, Mathieu et al., 2007, Sato et al., 2006, Fernández et al., 2007, Montrasio et al., 2007, Lee et al., 2007, Peretz et al., 2007, Srivastava et al., 2007, Luo et al., 2007, Xue et al., 2007, Lin et al., 2007, Umemori et al., 2007, Horiuchi et al., 2007, Fiehler and Wolff, 2008, Lee et al., 2005, Bidla et al., 2007, Ayaz et al., 2008, Titen and Golic, 2008, Baril et al., 2009, Pérez-Garijo et al., 2009, Hull-Thompson et al., 2009, Lee et al., 2006, Gesellchen et al., 2005, Buchon et al., 2009, Tokusumi et al., 2009, Igaki et al., 2006, Igaki et al., 2006, Cruz et al., 2009, Hill-Burns and Clark, 2009, Rallis et al., 2010, Popodi et al., 2010-, Guichard et al., 2006, Etter et al., 2005, Tuxworth et al., 2009, Rousset et al., 2010, Gettings et al., 2010, Kim et al., 2010, Djiane and Mlodzik, 2010, Bergantiños et al., 2010, Lee et al., 2009, Portela et al., 2010, Hong et al., 2011, Pastor-Pareja et al., 2008, Sekine et al., 2011, Kwon et al., 2010, Cobreros-Reguera et al., 2010, Umehara et al., 2010) Hep (Cha et al., 2003, Ohayon et al., 2009, Sano et al., 2005, Llense and Martín-Blanco, 2008, Ryabinina et al., 2006, Igaki et al., 2009, Liu et al., 2009, Rodahl et al., 2009, Lee et al., 2006, Wu et al., 2009, Jiang et al., 2009, Glavic et al., 2009, Geuking et al., 2009, Wu et al., 2009, Oliva and Sierralta, 2010, Neisch et al., 2010, Jiang et al., 2011) HEP/MKK7 (Bjorklund et al., 2006, Morrison et al., 2000) Hep/Mkk7 (Geuking et al., 2009) hp (Noselli et al., 1994, Noselli et al., 1993, ) JNKK (Delaney et al., 2005, Stronach and Perrimon, 1999, Homsy et al., 2006, Llense and Martín-Blanco, 2008, Liu et al., 2009, Rodahl et al., 2009, Glavic et al., 2009, Gettings et al., 2010, Jiang et al., 2011) l(1)7P1   l(1)G0107 (Schaefer et al., 1999.12.10, ) l(1)G0208 (Schaefer et al., 1999.12.2, ) MAPKK (Horiuchi et al., 2007) MKK7 (Sathyanarayana et al., 2003, Horiuchi et al., 2007) Mkk7 (Geuking et al., 2009)
Name Synonym	hemipterous (Walther and Pichaud, 2007, Delaney et al., 2006, Yoshioka et al., 2008, Bovolenta et al., 2006, Lennox and Stronach, 2010, Cerrato et al., 2006, Wu et al., 2008, Sato et al., 2006, Fernández et al., 2007, Hashimoto and Yamaguchi, 2006, Homsy et al., 2006, Lee et al., 2007, Bakal et al., 2008, Fiehler and Wolff, 2008, Lee et al., 2005, Bidla et al., 2007, Corl et al., 2009, Buchon et al., 2009, Jiang et al., 2009, Cruz et al., 2009, Hill-Burns and Clark, 2009, Etter et al., 2005, Gettings et al., 2010, Bergantiños et al., 2010, Rhiner et al., 2010, Lee et al., 2011, Portela et al., 2010) Hemipterous (Rogers et al., 2003, Ohayon et al., 2009, Guichard et al., 2008, Horiuchi et al., 2007, Llense and Martín-Blanco, 2008, Ryabinina et al., 2006, Titen and Golic, 2008, Rodahl et al., 2009, Lee et al., 2006, Wu et al., 2009, Tokusumi et al., 2009, Glavic et al., 2009, Rallis et al., 2010, Guichard et al., 2006, Rousset et al., 2010, Oliva and Sierralta, 2010) JNK kinase (Eissenberg, 2006, Rodahl et al., 2009) JNK-Kinase (Glavic et al., 2009) JNK-kinase (Etter et al., 2005) Mitogen-activated protein kinase kinase 7 (Lee et al., 2010)
Secondary FlyBase IDs
	FBgn0013490 FBgn0029076 FBgn0030446 FBgn0040188
References ( 268 )
Generate a list of	All references relating to this gene ( 268 )
List References by type	Research paper  ( 172 ) Supplementary material  ( 12 ) Review  ( 34 ) Personal communication to FlyBase  ( 11 ) Abstract  ( 27 ) FlyBase analysis  ( 3 ) DNA/RNA sequence record  ( 1 ) Protein sequence record  ( 1 ) Stock list  ( 1 ) Computer file  ( 3 ) Automatic genome annotation  ( 1 ) Curated genome annotation  ( 1 )
Recent research papers ( 22 )
	Hong et al., 2011, J. Ethnopharmacol. 134(3): 1028--1032 Neuroprotective effect of SuHeXiang Wan in Drosophila models of Alzheimer's disease. [FBrf0213403] Jezowska et al., 2011, Dev. Biol. 360(1): 143--159 A dual function of Drosophila capping protein on DE-cadherin maintains epithelial integrity and prevents JNK-mediated apoptosis. [FBrf0216680] Jiang et al., 2011, Cell Stem Cell 8(1): 84--95 EGFR/Ras/MAPK Signaling Mediates Adult Midgut Epithelial Homeostasis and Regeneration in Drosophila. [FBrf0212694] Lee et al., 2011, Mol. Cells 31(4): 337--342 The effects of hempseed meal intake and linoleic acid on Drosophila models of neurodegenerative diseases and hypercholesterolemia. [FBrf0213599] Sekine et al., 2011, J. Cell Sci. 124(17): 3006--3016 p38 MAPKs regulate the expression of genes in the dopamine synthesis pathway through phosphorylation of NR4A nuclear receptors. [FBrf0215011] Bergantiños et al., 2010, Development 137(7): 1169--1179 Cell death-induced regeneration in wing imaginal discs requires JNK signalling. [FBrf0210204] Blanco et al., 2010, BMC Dev. Biol. 10: 94 Gene expression following induction of regeneration in Drosophila wing imaginal discs. Expression profile of regenerating wing discs. [FBrf0211812] Cobreros-Reguera et al., 2010, EMBO Rep. 11(12): 943--949 The Ste20 kinase misshapen is essential for the invasive behaviour of ovarian epithelial cells in Drosophila. [FBrf0212408] Djiane and Mlodzik, 2010, PLoS ONE 5(6): e11228 The Drosophila GIPC homologue can modulate myosin based processes and planar cell polarity but is not essential for development. [FBrf0211141] Gettings et al., 2010, PLoS Biol. 8(6): e1000390 JNK Signalling Controls Remodelling of the Segment Boundary through Cell Reprogramming during Drosophila Morphogenesis. [FBrf0211014] Kim et al., 2010, BMC Cell Biol. 11: 9 A genetic screen for modifiers of Drosophila caspase Dcp-1 reveals caspase involvement in autophagy and novel caspase-related genes. [FBrf0209948] Kwon et al., 2010, Biochem. Biophys. Res. Commun. 393(4): 656--661 Nonmuscle myosin II localization is regulated by JNK during Drosophila larval wound healing. [FBrf0210297] Lee et al., 2010, Science 327(5970): 1223--1228 Sestrin as a feedback inhibitor of TOR that prevents age-related pathologies. [FBrf0210161] Lennox and Stronach, 2010, Dev. Dyn. 239(2): 651--664 POSH misexpression induces caspase-dependent cell death in Drosophila. [FBrf0209862] Lesch et al., 2010, Genetics 186(3): 943--957 A Targeted UAS-RNAi Screen in Drosophila Larvae Identifies Wound Closure Genes Regulating Distinct Cellular Processes. [FBrf0212282] Neisch et al., 2010, J. Cell Biol. 189(2): 311--323 Rho1 regulates apoptosis via activation of the JNK signaling pathway at the plasma membrane. [FBrf0210554] Oliva and Sierralta, 2010, Dev. Biol. 344(2): 911--921 Regulation of axonal development by the nuclear protein hindsight (pebbled) in the Drosophila visual system. [FBrf0211428] Portela et al., 2010, Dev. Cell 19(4): 562--573 Drosophila SPARC Is a Self-Protective Signal Expressed by Loser Cells during Cell Competition. [FBrf0212084] Rallis et al., 2010, Dev. Biol. 339(1): 65--77 Signal strength and signal duration define two distinct aspects of JNK-regulated axon stability. [FBrf0209975] Rhiner et al., 2010, Dev. Cell 18(6): 985--998 Flower forms an extracellular code that reveals the fitness of a cell to its neighbors in Drosophila. [FBrf0211273] Rousset et al., 2010, Development 137(13): 2177--2186 The Drosophila serine protease homologue Scarface regulates JNK signalling in a negative-feedback loop during epithelial morphogenesis. [FBrf0211045] Umehara et al., 2010, Molec. Cell. Biochem. 342(1-2): 223--232 Over-expression of transglutaminase in the Drosophila eye imaginal disc induces a rough eye phenotype. [FBrf0211642] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2010 Show reviews ...