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Gene Dmel\lok

General Information
Symbol	Dmel\lok	Species	D. melanogaster
Name	loki	Annotation symbol	CG10895
Feature type	protein_coding_gene	FlyBase ID	FBgn0019686
Created / Updated	2003-12-01/2003-12-01
Genomic Location
Chromosome (arm)	2L	Recombination map	2-
Cytogenetic map	38B2-38B2	Sequence location	2L:20,061,477..20,064,381 [+]
Map ( GBrowse )	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene loki is referred to in FlyBase by the symbol lok (CG10895, FBgn0019686). It has the cytological map location 38B2. Its sequence location is 2L:20061477..20064381. Its molecular function is described as: protein serine/threonine kinase activity; protein kinase activity; ATP binding. It is involved in the biological processes: germ cell development; protein amino acid phosphorylation; DNA damage checkpoint; DNA damage response, signal transduction resulting in induction of apoptosis; response to DNA damage stimulus; cell cycle checkpoint; apoptosis; caspase activation. 10 alleles are reported. No phenotypic data is available. It has 3 annotated transcripts and 3 annotated polypeptides.
External Summaries
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 38B2-38B2   Limits computationally determined from genome sequence between P{lacW}barrk14014 and P{EP}nebEP642
Experimentally Determined Cytological Location
Cytogenetic map Notes References 38B-38B   (Minami, 1997.10.2) 38B-38B   (Larochelle, 1997.1.31) 38B-38B (determined by in situ hybridisation)   (Oishi et al., 1998) 38B-38B (determined by in situ hybridisation)   (Hijal, 1997.4.17)
Experimentally Determined Recombination Data
Location	2- (Larochelle, 1997.1.31)
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References In direction of increasing cytology: barr- lok+ vls- (Butler et al., 2001) Gene Order (overall orientation not stated) References Overall orientation not stated: barr- lok+ (Xu et al., 2001)
Gene Model & Products
Please see the GBrowse view of Dmel\lok for information on other features
Comments on Gene Model
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) lok-RA FBtr0081328  NM_057871   2176   476 lok-RB FBtr0081329  NM_057870   2125   459 lok-RC FBtr0081330  NM_165318   2186   459
Additional Transcript Data & Comments
Reported size (kB)	2.2 (northern blot) (Oishi et al., 1998)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein lok-PA   FBpp0080860   54.3   476   9.54     NP_477219   AAF53867 lok-PB   FBpp0080861   52.3   459   9.39     NP_477218   AAN11062 lok-PC   FBpp0080862   52.3   459   9.39     NP_724241   AAN11063
Additional Polypeptide Data & Comments
Reported size (kD)	476, 459 (aa); 56, 54 (kD predicted) (Oishi et al., 1998)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0019686
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Forkhead-associated (IPR000253) Protein kinase, core (IPR000719) Serine/threonine protein kinase (IPR002290) Serine/threonine protein kinase, active site (IPR008271) SMAD/FHA domain (IPR008984) Protein kinase-like (IPR011009) Protein kinase ATP binding, conserved site (IPR017441) Serine/threonine protein kinase-related (IPR017442)
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA816409   AA949096   AA951171   AB007821 BAA28755   AB007822 BAA28756   AC004364   AC009252   AE003665 AAF53867   AE003665 AAN11062   AE003665 AAN11063   AI532068   AW942781   AY070549 AAL48020   BI164599   BI170187   BI214954   BI239814   BI357201   BI357257   BI578275   BI632532   BI639501   BI639590   BI640595   BP541694   BP556041   BP556042   BP558648   BP558649   CG10895   CK136045   CO299280   CO304724   CO325538   EC051973   EC054478   EC054979   EC056308   EC200985   EC204441   EC226111   EC248698   SD22085   U87984 AAB49642
UniProtKB/Swiss-Prot	O61267
UniProtKB/TrEMBL
Maps to	DS00863 (FlyBase, 1992-, Butler et al., 2001) FBcl0297593 (FlyBase, 1992-)
Does NOT map to
Identified with	LD27857 (BDGP Project Members, 2000-, Butler, 2001) SD22085 (BDGP Project Members, 2000-)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References rescue fragment lok[+tBa] 2L:20,060,954..20,064,555 comment=Genomic fragment extending from 523bp upstream of the transcription start to 72bp downstream of the polyA site. evidence=experimental (Takada et al., 2003, Brodsky, 2003.5.16)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 8166
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  oogenesis stage,adult stage | stage >=S2  nurse cell (Oishi et al., 1998)  embryonic stage | stage 1,2   (Oishi et al., 1998)  embryonic stage | early   (Oishi et al., 1998)  oogenesis stage,adult stage | female  ovary (Oishi et al., 1998)  embryonic stage | stage >=3  pole cell (Oishi et al., 1998)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  embryonic stage | stage 3,4   (Oishi et al., 1998)
Marker for
Subcellular Localization
CV Term	nucleus (Oishi et al., 1998)
Microarray Data
	Developmental timecourse, Costello et al., 2008 (Original data from Arbeitman et al., 2002) AA949096
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0019686 FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array • FBgn0019686 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0019686
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele lethal | embryonic stage | maternal effect lokE51 Other Phenotypes Allele wild-type lokhs.PX cell death decrease lokp6 chemical resistant lokp6 radiation sensitive | recessive lokE51 Phenotype manifest in Allele nucleus & embryo lokE51 spindle & embryo lokE51 mitosis & nuclear chromosome & embryo lokE51
Classical Alleles ( 3 )
For All Classical Alleles Show
Allele of lok Class Mutagen Stocks Known lesion lok30 P-element activity 0 Yes lokE51 Δ2-3 0 Yes lokp6 loss of function P-element activity 0 Yes
Alleles Carried on Transgenic Constructs ( 7 )
For All Alleles Carried on Transgenic Constructs Show
Allele of lok Class Mutagen Stocks Known lesion lok+tBa in vitro construct | genomic fragment 0 Yes lok3'.tMa in vitro construct | genomic fragment 0 Yes lok5'.tMa in vitro construct | genomic fragment 0 Yes lokD303A.Scer\UAS in vitro construct | amino acid replacement 0 Yes lokGD4190 in vitro construct | RNAi in vitro construct | regulatory fusion 2 Yes lokScer\UAS.cPa in vitro construct | regulatory fusion 0 Yes lokhs.PX in vitro construct | regulatory fusion 0 Yes
Aneuploid Aberrations
Useful deficiency	Df(2L)TW150
Disrupted in	Df(2L)E51 (Xu et al., 2001) Df(2L)Exel7077 (Ryder, 2004.4.26)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data UAS construct P{GD4190} NA P{UAS-lok.D303A} NA P{UAS-lok.P} NA characterization construct P{CG10728+tMa} NA P{barr+tMa} NA P{mnk-rescued} NA heat-shock construct P{hs-lok.X} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{}lokp6 NA
Related Comments
	Please look at the allele reports for the complete phenotype data Mutants show defects in maintaining genome stability and are highly sensitive to ionizing radiation. Mutation in lok completely blocks DNA damage-induced apoptosis and partially blocks damage-induced cell cycle arrest. (Xu et al., 2001)
Gene Ontology: Function, Process & Cellular Component ( 13 )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR000719, InterPro:IPR002290, InterPro:IPR017441, InterPro:IPR017442 (FlyBase Curators et al., 2004-)
protein kinase activity inferred from sequence or structural similarity with HGNC_gene:CHEK2; HGNC:16627; OMIM:604373 (Sekelsky et al., 2000)
protein serine/threonine kinase activity non-traceable author statement (Hijal et al., 1997) inferred from sequence or structural similarity (Oishi et al., 1998) non-traceable author statement (Larochelle, 1997.1.31) non-traceable author statement (Morrison et al., 2000) inferred from electronic annotation with InterPro:IPR002290, InterPro:IPR008271 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
DNA damage checkpoint inferred from mutant phenotype (Xu et al., 2001) inferred from mutant phenotype (LaRocque et al., 2007)
DNA damage response, signal transduction resulting in induction of apoptosis inferred from mutant phenotype (Xu et al., 2001)
apoptosis inferred from mutant phenotype (Wichmann et al., 2006)
caspase activation inferred from mutant phenotype (Akdemir et al., 2007)
cell cycle checkpoint traceable author statement (Lee and Orr-Weaver, 2003) inferred from sequence or structural similarity with HGNC_gene:CHEK2; HGNC:16627; OMIM:604373 (Sekelsky et al., 2000)
germ cell development inferred from expression pattern (Oishi et al., 1998)
protein amino acid phosphorylation inferred from sequence or structural similarity (Oishi et al., 1998) non-traceable author statement (Morrison et al., 2000) inferred from sequence or structural similarity with HGNC_gene:CHEK2; HGNC:16627; OMIM:604373 (Sekelsky et al., 2000)
response to DNA damage stimulus inferred from mutant phenotype (Takada et al., 2003)
Cellular Component
CV term Evidence References
cytoplasm non-traceable author statement (Hijal et al., 1997)
nucleus inferred from direct assay (Oishi et al., 1998)
Sequence Ontology: Class of Gene
	protein_coding_gene   nuclear_gene
Interactions & Pathways
Summary of Genetic Interactions
Interacts with	Please look at the allele data for full details of the genetic interactions lok allele Gene References lokD303A.Scer\UAS p53 (Peters et al., 2002) lokE51 stg (Xu and Du, 2003) polo (Xu and Du, 2003) lokp6 wee (Stumpff et al., 2004) spn-D (Abdu et al., 2002) grp (Takada et al., 2003) lokScer\UAS.cPa p53 (Peters et al., 2002) unspecified mus301 (McCaffrey et al., 2006) wee (Stumpff et al., 2005) aub (Klattenhoff et al., 2007) spn-D (Klattenhoff et al., 2007) spn-A (Staeva-Vieira et al., 2003) armi (Klattenhoff et al., 2007)
External Data
Linkouts	BioGRID - Interaction data, including yeast 2-hybrid and genetic interactions • 61259
Orthologs
Genome-wide drosophilid orthologs	Dana\GF14445 Dere\GG21217 Dgri\GH13656 Dmoj\GI17408 Dper\GL26345 Dpse\GA10622 Dsec\GM17387 Dsim\GD24241 Dvir\GJ18340 Dwil\GK18729 Dyak\GE13292
Curated drosophilid orthologs	Dpse\GA10622
Linkouts	InParanoid orthologs - Eukaryotic orthologs • ENSANGP00000019651 Anopheles gambiae (Ensembl) ENSBTAP00000006518 Bos taurus (Ensembl) ENSCAFP00000017544 Canis familiaris (Ensembl) ENSCINP00000010261 Ciona intestinalis (Ensembl) NEWSINFRUP00000140329 Takifugu rubripes (Ensembl) ENSGALP00000009122 Gallus gallus (Ensembl) ENSP00000329178 Homo sapiens (Ensembl) ENSMMUP00000026981 Macaca mulatta (Ensembl) ENSMODP00000010421 Monodelphis domestica (Ensembl) ENSPTRP00000024456 Pan troglodytes (Ensembl) GSTENP00007555001 Tetraodon nigroviridis (Ensembl) ENSXETP00000046701 Xenopus tropicalis (Ensembl) WBGene00011612 Caenorhabditis elegans (Wormbase) ZDB-CDNA-040425-1012 Danio rerio (Zfin) ZDB-GENE-030131-8942 Danio rerio (Zfin) MGI:1355321 Mus musculus (Informatics) YDL101C Saccharomyces cerevisiae (Yeastgenome) SPCC18B5.11c Schizosaccharomyces pombe (Sanger)
Functional Complementation between Species
Inter-Species Misexpression Data
Produces phenotype in
Produces NO phenotype in
Stocks & Reagents
Stocks Listed in FlyBase ( 2 )
VDRC	v44980 w1118; P{GD4190}v44980 v44981 w1118; P{GD4190}v44981
Genomic Clones ( 2 )
	BACR10H13 BACR22E16
cDNA Clones ( 22 )
	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	LD27857
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LD31503 SD22219 RE04305 RE11565 SD03432 RE72126 RE43616 RE21120 SD23521 RE43681 SD26335 LD01443 RE36409
Other clones	EC056308 EC054478 EK175001 EC051973 EK196738 EK245916 SD22085 EC054979
RNAi & Array Information
Affy Oligo
Linkouts	DRSC - RNAi screening (Harvard): high-throughput cell culture data and design • FBgn0019686 GenomeRNAi - RNAi phenotypes (Heidelberg): high-throughput cell culture data • FBgn0019686
Antibody Information
	polyclonal (Oishi et al., 1998)
Other Information
Discoverer
Etymology
Identification
Position Effect Variegation Data
Relationship to Other Genes
Source for database identity of	Source for identity of: lok CG10895 (Burtis, 1999.11)
Source for database merge of
Additional comments	"lok" may be allelic to "vls". (Oishi et al., 1998)
Comments About Role
	lok encodes a serine/threonine protein kinase. (Hijal, 1997.4.17) lok appears to be a transducer of the meiotic checkpoint that is activated by unrepaired double-strand DNA breaks. (Abdu et al., 2002)
Comments About Molecular Function
Other Comments
	Identification and developmental expression of lok are reported, a nuclear protein Ser/Thr kinase. The unique spatio-temporal expression pattern and structural features of lok indicate it is a novel class of nuclear protein kinases. (Oishi et al., 1998) 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) S2 cells are treated with dsRNA made from templates generated with primers directed against this gene to assess effect on proteolysis of dup and geminin genes. (Higa, 2003)
External Crossreferences & Linkouts
Sequence Crossreferences
	RefSeq (Tr