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Gene Dmel\Hsp70Aa

General Information
Symbol	Dmel\Hsp70Aa	Species	D. melanogaster
Name	Heat-shock-protein-70Aa	Annotation symbol	CG31366
Feature type	protein_coding_gene	FlyBase ID	FBgn0013275
Created / Updated	2003-12-01/2003-12-01
Genomic Location
Chromosome (arm)	3R	Recombination map
Cytogenetic map	87A2-87A2	Sequence location	3R:7,779,885..7,782,514 [-]
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 Heat-shock-protein-70Aa is referred to in FlyBase by the symbol Hsp70Aa (CG31366, FBgn0013275). It has the cytological map location 87A2. Its sequence location is 3R:7779885..7782514. Its molecular function is described as ATP binding. It is involved in the biological processes: heat shock-mediated polytene chromosome puffing; response to heat. 5 alleles are reported. No phenotypic data is available. It has one annotated transcript and one annotated polypeptide.
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	Hsp70 The structural genes that code for the 70,000 dalton heat-shock protein (HSP70), the most abundant of the heat-shock proteins. HSP70 returns to preshock levels more rapidly than other heat-shock proteins following return to 25 (DiDomenico, Bugaisky, and Lindquist, 1982, Proc. Nat. Acad. Sci. USA 79: 6181-85). The protein becomes concentrated in nuclei during heat shock; disperses to cytoplasm during recovery; returns to nucleus upon further heat shock (Velazquez and Lindquist, 1984, Cell 36: 655-62). Appears not to be expressed in the testis in response to heat-shock stimulation (Bonner, Parks, Parker-Thornberg, Mortin, and Pelham, 1984, Cell 37: 979-91). Deletion of either the 87A7 or the 87C1 sequences does not eliminate the HSP70 heat-shock response; simultaneous deletion of both sequences does eliminate the HSP70 heat-shock response (Ish-Horowitz et al., 1979).
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 87A2-87A2   Limits computationally determined from genome sequence between P{EP}Lk6EP886 and P{PZ}svp07842
Experimentally Determined Cytological Location
Cytogenetic map Notes References 87A7-87A7 (determined by in situ hybridisation)   (Ranz et al., 1999) 87A-87A (determined by in situ hybridisation)   (Segarra et al., 1996) 87A-87A (determined by in situ hybridisation)   (Papaceit and Juan, 1993) 87A7-87A7 (determined by in situ hybridisation)   (Goldschmidt-Clermont, 1980) 87A7-87A7 (determined by in situ hybridisation)   (Ish-Horowicz et al., 1979) 87A-87A (determined by in situ hybridisation)   (Schedl et al., 1978)
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References In direction of increasing cytology: Hsp70Aa- Hsp70Ab+ (Ish-Horowicz and Pinchin, 1980) Gene Order (overall orientation not stated) References Overall orientation not stated: Hsp70Aa- Hsp70Ab+ (Bettencourt et al., 1999)
Gene Model & Products
Please see the GBrowse view of Dmel\Hsp70Aa for information on other features
Comments on Gene Model
	this is Hsp70Aa; CG18743 (on + strand) is Hsp70Ab. (see Leigh Brown and Ish-Horowicz, 1981, Nature 290:677-682; Bettencourt and Feder 2001, Mol. Biol. Evol. 18:1272-1282; Bettencourt and Feder, 2002, J. Mol. Evol. 54:569-586).
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) Hsp70Aa-RA FBtr0082512  NM_169441   2630   642
Additional Transcript Data & Comments
Reported size (kB)	2.55 (northern blot) (Holmgren et al., 1979)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein Hsp70Aa-PA   FBpp0081986   70.2   642   5.49     NP_731651   AAN13535
Additional Polypeptide Data & Comments
Reported size (kD)	70 (kD) (Holmgren et al., 1979)
Comments
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0013275
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Heat shock protein Hsp70 (IPR001023) Heat shock protein 70 (IPR013126) TRANSFAC - Eukaryotic transcription factors, their genomic binding sites, and DNA-binding profiles • R00751 R00752 R00754 R00755
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA802879   AE003693 AAN13535   AF295933 AAG26887   AF295934 AAG26888   AF295935 AAG26889   AF295936 AAG26890   AF295937 AAG26891   AF295938 AAG26892   AF350452 AAK30209   AF350453 AAK30210   AF350454 AAK30211   AF350455 AAK30212   AF350456 AAK30213   AF350457 AAK30214   AF350458 AAK30215   AF350459 AAK30216   AI110149   AY118457 AAM49826   BE977138   BG633852   BT011058   BT021339 AAX33487   CK659005   CK661022   CL526167   EC054270   EC054622   EC054721   EC062495   EC073968   EC198499   EC213927   EC234458   EC265405   K01292 AAA28639   U03466   X63732
UniProtKB/Swiss-Prot	P82910
UniProtKB/TrEMBL	Q5BI85
Maps to
Does NOT map to
Identified with	AT29796 LP23554 (BDGP Project Members, 2000-) D1362 (BDGP Project Members, 1994-1999)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 9100
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
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference
Marker for
Subcellular Localization
CV Term
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0013275
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele viable Hsp70AaEY09953 Sterility Allele fertile Hsp70AaEY09953 Phenotype manifest in Allele
Classical Alleles ( 3 )
For All Classical Alleles Show
Allele of Hsp70Aa Class Mutagen Stocks Known lesion Hsp70AaDG16104 P-element activity 1 -- Hsp70AaEY09953 P-element activity 1 -- Hsp70AaSze-13 spontaneous 0 Yes
Alleles Carried on Transgenic Constructs ( 2 )
For All Alleles Carried on Transgenic Constructs Show
Allele of Hsp70Aa Class Mutagen Stocks Known lesion Hsp70AacKa in vitro construct | genomic fragment 0 Yes Hsp70Aarand in vitro construct | genomic fragment 0 Yes
Aneuploid Aberrations
Useful deficiency	Df(3R)E229
Useful duplication	Dp(3;3)M86D+2
Disrupted in	Df(3R)E229 (Gausz et al., 1981) Df(3R)E79 (Gausz et al., 1981) Df(3R)Exel6162 (Ryder, 2004.4.26) Df(3R)Exel6163 (Ryder, 2004.4.26) Df(3R)Hsp70A (Gong and Golic, 2004, Gong and Golic, 2005.6.2) Df(3R)P-29 (Yang and Ballinger, 1994) Df(3R)T-45 (Gausz et al., 1981) Df(3R)T-47 (Gausz et al., 1981) Df(3R)kar-H5 (Gausz et al., 1981) Df(3R)kar1W (Gausz et al., 1981)
Not disrupted in	Df(3R)T-55 (Gausz et al., 1981, Paro et al., 1983) Df(3R)kar-D1 (Gausz et al., 1981, Paro et al., 1983, Yang and Ballinger, 1994) Df(3R)kar-H10 (Gausz et al., 1981) Df(3R)kar-Sz12 (Gausz et al., 1981) Df(3R)kar3J (Gausz et al., 1981) Df(3R)kar3Q (Gausz et al., 1981) Df(3R)ry615 (Gausz et al., 1981)
Transgenic Constructs & Insertions
Transgenic Constructs
	Type of construct Name Expression data characterization construct P{r-w.cKb} NA P{scs-w.cKb} NA
Insertions
	Type of insertions Name Expression data miscellaneous insertions P{wHy}Hsp70AaDG16104 NA insertion of mobile activating element P{EPgy2}105Ig NA P{EPgy2}111IIa NA P{EPgy2}121II NA P{EPgy2}123Ia NA P{EPgy2}134Ib NA P{EPgy2}163Ia NA P{EPgy2}169Ig NA P{EPgy2}177I NA P{EPgy2}179I NA P{EPgy2}184I NA P{EPgy2}186Ib NA P{EPgy2}1IIa NA P{EPgy2}246I NA P{EPgy2}253Ia NA P{EPgy2}253Ib NA P{EPgy2}258II NA P{EPgy2}284Ia NA P{EPgy2}288Ia NA P{EPgy2}288Ib NA P{EPgy2}288Ic NA P{EPgy2}288Id NA P{EPgy2}288Ie NA P{EPgy2}288Ik NA P{EPgy2}295II NA P{EPgy2}30IIb-Aa NA P{EPgy2}31IIa NA P{EPgy2}369I NA P{EPgy2}48Ia NA P{EPgy2}5IId NA P{EPgy2}60IIa NA P{EPgy2}61Ib NA P{EPgy2}86I NA P{EPgy2}Hsp70AaEY09953 NA P{EPgy2}Hsp70AbEY01148 NA P{EPgy2}α138IIa NA P{EPgy2}α148IIa NA P{EPgy2}α196Ia NA P{EPgy2}α198I-Aa NA P{EPgy2}α21Ib NA P{EPgy2}α250Ia NA P{EPgy2}α33Ib NA P{EPgy2}α9Ib NA
Related Comments
	Please look at the allele reports for the complete phenotype data One of five structural genes (in two clusters, Hsp70A and Hsp70B) that code for the 70,000 dalton heat-shock protein (HSP70), the most abundant of the heat-shock proteins. Hsp70A usually includes two HSP70 encoding genes (Hsp70Aa (proximal), Hsp70Ab (distal)) (Holmgren et al., 1979) with slightly different restriction maps (Artavanis-Tsakonas et al., 1978). HSP70 returns to preshock levels more rapidly than other heat-shock proteins following return to 25oC (DiDomenico, Bugaisky and Lindquist, 1982). The protein becomes concentrated in nuclei during heat shock; disperses to cytoplasm during recovery; returns to nucleus upon further heat shock (Velazquez and Lindquist, 1984). Appears not to be expressed in the testis in response to heat-shock stimulation (Bonner et al., 1984). Deletion of either Hsp70A or Hsp70B does not eliminate the HSP70 heat-shock response; simultaneous deletion of both does (Ish-Horowicz et al., 1979).   The effect of aging on the expression of the "Hsp70" genes (Hsp70Aa, Hsp70Ab, Hsp70Ba, Hsp70Bb and Hsp70Bc) has been studied. (Niedzwiecki et al., 1991) The Hsp70A promoter is unsuitable for use in fusion gene constructs for long term expression studies where repeated heat shocks are required as the amount of RNA is substantially reduced. (Hunt et al., 1992) Probes from D.melanogaster used in chromosome in situ hybridisation to study response to heat shock in D.guanche, D.madeirensis and D.subobscura. Results suggest that the 18C, 94A, 89A and 27A loci of the three obscura group species are homologous to the D.melanogaster loci Hsp83, Hsp70A, Hsp68 and the small Hsp group Hsp22, Hsp23, Hsp26 and Hsp27 respectively. (Molto et al., 1992) Psoralen cross-linking studies reveal that the Hsp70A and 18S rRNA DNA is wound with a significant level of superhelical tension irrespective of state of transcriptional activation. Conversely DNA flanking Hsp70A shows substantially less tension, indicating stable, torsionally stressed topological domains. Thus the relaxing activity of topoisomerases is not ubiquitous throughout the nucleus but is tightly regulated. (Jupe et al., 1993) Identified in 2D gels of CMW W2 wing imaginal disc cell proteins. (Santaren et al., 1993) Members of the hsc70 gene family (heat shock cognate genes) that reside within the same intracellular compartment in different organisms share greater amino acid identity than hsc70 proteins from the same organism but different organelles. This pattern of conservation indicates specialisation of hsc70 function. (Rubin et al., 1993) RNA polymerase distribution on uninduced Hsp70 genes has been compared with the distribution of RNA polymerase on DRB (5,6,-Dichloro-1-β-D-ribofuranosylbenzimidazole)-inhibited induced Hsp70 genes. (Giardina and Lis, 1993) Heat shock alters the distribution of Top2 in the 87A7 heat shock locus. scs and scs' are targets for Top2 localisation. A return to normal temperatures relocalises Top2. (Udvardy and Schedl, 1993) Introduction of Trl protein during or after nucleosome assembly in vitro results in disruption of nucleosome structure at the Hsp70Aa promoter. Hsp70Aa promoter deletion analysis reveals that disruption of a preassembled nucleosome requires three or four GA/CT elements, nucleosome displacement requires at least two GA/CT elements. The disruption is characterised by hypersensitivity to DNase I digestion and a realignment of adjacent nucleosomes. Disruption is facilitated by the presence of hydrolysable ATP. (Tsukiyama et al., 1994) Activation of Hsp70Aa is a multistep process. RNA polymerase II is engaged, release of the polymerase and start of transcription requires the binding of heat shock transcription factors (HSF), whose binding is in turn conditional on binding of Trl. (van Holde, 1994) The Hsp70Aa and Hsp70Ab loci are flanked by special chromatin structures, scs and scs'. Each structure is defined by a pair of nuclease hypersensitivity sites bordering a nuclease resistant core of 250-350bp in length. Both scs and scs' have properties that suggest they may correspond to the boundaries of the 87A7 chromomere. scs and scs' are capable of establishing a domain of independent gene activity: w gene flanked by scs and scs' inserted into the genome by P element mediated transformation is isolated against both positive and negative position effects at most insertion sites. scs and scs' also have enhancer blocking activities: when inserted between a Yp1 enhancer and a heat shock promoter-Ecol\lacZ fusion little or no Ecol\lacZ expression is detected even though the promoter is fully heat shock inducible. (Vazquez et al., 1993) The transcription of heat shock proteins, except Hsp83, is independent of the p200 subunit of initiation factor eIF-4F, eIF-4G. (Zapata et al., 1994) Synthesis of heat shock proteins is inhibited by both short-chain fatty acids and their corresponding alcohols, compounds which have no observable effect on histone acetylation. (Munks and Turner, 1994) UV cross linking technique has been used to study the in vivo distribution of Trl protein on Hsp70 and Hsp26. Prior to heat shock Trl protein is associated with the promoter regions of the uninduced Hsp70 and Hsp26 genes. Upon heat shock induction Trl protein is recruited to their transcription units with its distribution coincident with that of RNA polymerase II. (O'Brien et al., 1995) RNA levels do not increase with age, so the observed increase in protein levels is due to post-transcriptional regulation. Aging-specific expression may be a result of oxidative damage. (Wheeler et al., 1995) Gene contains an RNA polymerase II complex which pauses after synthesis of a short transcript. In vivo ultraviolet crosslinking techniques demonstrate phosphorylation of the carboxy terminal domain (CTD) of the large subunit of RNA polymerase II could either regulate the transition of polymerase from a paused to an elongated complex or be a consequence of the transition from paused to elongated. (O'Brien et al., 1994) Sodium salicylate induces activation of Hsf binding activity in salivary gland cells and Schneider SL2 tissue cells. Puffing of heat shock gene loci occurs in salivary glands but Hsp70 transcription is not induced suggesting puffing and transcription are separable events. (Winegarden et al., 1996) d(GA.TC)n sequences can be found in the promoters of Hsp26 and the Hsp70 genes. In vitro assembly of mononucleosomes into short DNA fragments carrying d(GA.TC)n sequences of different lengths is very efficient. Nucleosome assembly is inhibited strongly when the d(GA.TC)n sequence forms a triple-stranded conformation. Triplex formation requires partial destabilisation of the nucleosome. Results indicate nucleosome assembly and triplex formation are competing processes. (Espinas et al., 1996) Flies with no copies of the Hsp70 genes (Hsp70Aa, Hsp70Ab, Hsp70Ba, Hsp70Bb, Hsp70Bbb and Hsp70Bc) are viable and fertile. (Gong and Golic, 2004) Flies with no copies of the Hsp70 genes (Hsp70Aa, Hsp70Ab, Hsp70Ba, Hsp70Bb, Hsp70Bbb and Hsp70Bc) are unable to survive a severe heat shock. These flies show a lengthened heat-shock response and developmental delay following a non-lethal heat shock. (Gong and Golic, 2006)
Gene Ontology: Function, Process & Cellular Component ( 3 )
Molecular Function
CV term Evidence References
ATP binding inferred from electronic annotation with InterPro:IPR001023 (FlyBase Curators et al., 2004-)
Biological Process
CV term Evidence References
heat shock-mediated polytene chromosome puffing inferred from mutant phenotype (Gong and Golic, 2006)
response to heat inferred from mutant phenotype (Gong and Golic, 2006) non-traceable author statement (Swiss-Prot Project Members, 2000.10.1)
Cellular Component
CV term Evidence References
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 Hsp70Aa allele Gene References
External Data
Linkouts	BioGRID - Interaction data, including yeast 2-hybrid and genetic interactions • 71511
Orthologs
Genome-wide drosophilid orthologs
Curated drosophilid orthologs	Damb\Hsp70Aa   Dbio\Hsp70   Dbir\Hsp70   Dbuz\Hsp70Ba   Dbuz\Hsp70Bb   Dbuz\Hsp70Bc   Dcub\Hsp70   Ddip\Hsp70   Dequ\Hsp70   Dere\Hsp70Aa   Dgua\Hsp70A   Dgua\Hsp70B   Dhyd\Hsp70   Dins\Hsp70   Djam\Hsp70   Dkik\Hsp70   Dleb\Hsp70A   Dleb\Hsp70B   Dlum\Hsp70a   Dlum\Hsp70b   Dlum\Hsp70c   Dlum\Hsp70d   Dmad\Hsp70A   Dmad\Hsp70B   Dmir\Hsp70A   Dmir\Hsp70B   Dmoj\Hsp70   Dneb\Hsp70   Dore\Hsp70Aa (Bettencourt, 2000.9.2) Dpau\Hsp70   Dper\Hsp70A   Dper\Hsp70B   Dpse\Hsp70A   Dpse\Hsp70B   Dqua\Hsp70   Drep\Hsp70Ba   Drep\Hsp70Bb   Drep\Hsp70Bc   Dsec\Hsp70Aa   Dseg\Hsp70   Dser\Hsp70   Dsim\Hsp70Aa   Dsub\Hsp70A   Dsub\Hsp70B   Dtei\Hsp70Aa   Dtra\Hsp70   Dtro\Hsp70   Dtrz\Hsp70   Dvir\Hsp70A   Dvir\Hsp70a   Dvir\Hsp70b   Dvir\Hsp70c   Dvir\Hsp70d   Dvir\Hsp70e   Dvir\Hsp70f   Dvir\Hsp70g   Dvul\Hsp70   Dwat\Hsp70   Dwil\Hsp70   Dyak\Hsp70Aa
Linkouts
Functional Complementation between Species
Inter-Species Misexpression Data
Produces phenotype in
Produces NO phenotype in
Stocks & Reagents
Stocks Listed in FlyBase ( 3 )
Bloomington	21515 y1 w67c23; P{wHy}Hsp70AaDG16104 15327 y1 w67c23; P{EPgy2}Hsp70AbEY01148 17624 y1 w67c23; P{EPgy2}Hsp70AaEY09953
Genomic Clones ( 2 )
	BACR03N24 BACR48E12
cDNA Clones ( 10 )
	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	LP05203 LP23554
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones	LP20090 GH09635 LP16393
Other clones	EC054270 EC054721 EC054622 bs60b11 GM06511
RNAi & Array Information
Affy Oligo
Linkouts	DRSC - RNAi screening (Harvard): high-throughput cell culture data and design • FBgn0013275 GenomeRNAi - RNAi phenotypes (Heidelberg): high-throughput cell culture data • FBgn0013275
Antibody Information
Other Information
Discoverer