The gene Heat-shock-protein-70Bc is referred to in FlyBase by the symbol Dmel\Hsp70Bc (CG6489, FBgn0013279). It is a protein_coding_gene from Drosophila melanogaster. Its molecular function is unknown. There is experimental evidence that it is involved in the biological process: response to hypoxia; heat shock-mediated polytene chromosome puffing; response to methotrexate; response to heat. 4 alleles are reported. No phenotypic data is available. It has one annotated transcript and one annotated polypeptide. Protein features are: Heat shock protein 70 family; Heat shock protein 70, conserved site. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed at stages throughout the pupal period. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval salivary gland. Comments on Affy2 ProbeSet: ProbeSet 1632841_x_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of Hsp70Bc. Gene sequence location is 3R:8334798..8337183.
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Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval salivary gland.
[download data (TSV)]
Guide to FlyAtlas expression level colors
No expression (0 - 9.999)
Low expression (10 - 99.999)
Moderate expression (100 - 499.999)
High level expression (500 - 999.999)
Very high expression (>999.999)
Linear, scaled to maximum expression level
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Linear, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
(769.3)
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Linear, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Very high
Linear, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
Very high
log, scaled to maximum expression level
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Very high
log, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
log, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Very high
log, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
16.9
Larval Midgut
7.7
Larval Hindgut
60
Larval Malpighian Tubules
17.2
Larval Fat Body
23
Larval Salivary Gland
157.4
Larval Trachea
10.725
Larval Carcass
20.65
Adult Head
7.5
Adult Eye
2.1
Adult Brain
18.7
Adult Thoracic-Abdominal Ganglion
15.5
Adult Crop
13.5
Adult Midgut
8.1
Adult Hindgut
4.4
Adult Malpighian Tubules
769.3
Adult Fat Body
8.7
Adult Salivary Gland
46.3
Adult Heart
16.025
Adult VirginFemale Spermatheca
2.4
Adult InseminatedFemale Spermatheca
50.3
Adult Ovary
2.5
Adult Testis
7.2
Adult Male Accessory Gland
2.6
Adult Carcass
6.5
Expression Level Scale
None
Low
Moderate
High
Very high
Heatmap
Tissue
Expression Level
Larval Central Nervous System
Larval Midgut
Larval Hindgut
Larval Malpighian Tubules
Larval Fat Body
Larval Salivary Gland
Larval Trachea
Larval Carcass
Adult Head
Adult Eye
Adult Brain
Adult Thoracic-Abdominal Ganglion
Adult Crop
Adult Midgut
Adult Hindgut
Adult Malpighian Tubules
Adult Fat Body
Adult Salivary Gland
Adult Heart
Adult VirginFemale Spermatheca
Adult InseminatedFemale Spermatheca
Adult Ovary
Adult Testis
Adult Male Accessory Gland
Adult Carcass
FlyAtlas Organ/Tissue Expression, larval vs. adult
Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of high expression to a trough of very low expression. Peak expression observed at stages throughout the pupal period.
[download data (TSV)]
Please Note FlyBase no
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cDNA Clones ( 40 )
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.
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.
Some Drosophila strains, including that sequenced in the Drosophila genome project have three, not two, tandemly repeated Hsp70 genes at 87C1 (in addition to a reverse orientation Hsp70 gene, Hsp70Ba, approximately 40kb upstream). The distal-most gene of the three tandem copies is Hsp70Bc. The inner two tandem copies are Hsp70Bb and Hsp70Bbb.
The D.melanogaster strain used for sequencing the genome contains an additional, third tandem copy of an Hsp70 gene at 87C as well as the two tandemly repeated copies (Hsp70Bb and Hsp70Bc) which are found in many strains. This additional copy is named "Hsp70Bbb" (see FBrf0137034 and Bettencourt, 2002.1.16, personal communication to FlyBase).
Variation in copy number of Hsp70Bb and Hsp70Bc demonstrates while extra Hsp70 provides additional protection against the immediate damage from heat stress, abnormally high concentrations can decrease growth, development and survival to adulthood.
There is significant variation among 74 different 2nd chromosome lines and 70 different 3rd chromosome lines in response to heat shock, measured by mRNA accumulation.
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.
Variation in copy number of Hsp70Bb and Hsp70Bc affects Hsp70 protein concentration in whole larvae and pupae, which in turn affects their tolerance of natural thermal stress and, potentially, their fitness.
Chromosome homologies of Muller's element D (J chromosome in the Paleartic species and XR chromosome arm in Nearctic species) and of element E (O chromosome in the Paleartic species and 2 chromosome in Nearctic species) have been confirmed by single copy probes in the species of the obscura group and in D.melanogaster.
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.
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.
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.
Heat shock does not block the polyadenylation of Hsp70Bc RNA, instead poly(A)-deficient transcripts arise from the rapid removal of poly(A) from previously polyadenylated RNAs. Removal of poly(A) is regulated according to the severity of the heat shock and precedes degradation, suggesting that it predisposes the transcript to turnover. Deadenylation appears to play an important role in regulating Hsp70Bc expression.
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.
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.
The TFIID complex interacts with the promoter of Hsp70Bc making contacts at the TATA element, initiator, +18 and +28 regions. The complex recognises a sequence within the context of the promoter region that corresponds to an initiator consensus sequence.
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.
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.
Nascent chain nuclear run-on assays in KC161 cells reveal different responses to heat shock for different genes. Transcription of His1 is severely inhibited under mild heat shocks, of Act5C decreases proportionally with increasing temperature while that of the core histone genes or the heat shock cognates is repressed only under extreme heat shock. In unshocked cells Hsp83 is moderately transcribed while transcription from the other heat shock genes is undetectable. Engaged but paused RNA molecules are found at the various Hsp70 and Hsp26 genes but not at the other heat shock genes. Increased transcription of the heat shock genes is observed within 1-2 mins of heat shock and maximal rates were reached within 2-5 minutes. Rates of transcription vary over a 20-fold range. Hsrω is transcribed at a very high rate under non-heat shock conditions, and its response to elevated temperatures is different from that of the protein coding heat shock genes.
Expression of transgenes carrying Hsp70Bb and Hsp70Bc demonstrate that at early embryonic stages the accumulation of Hsp70 is a rate-limiting factor in the acquisition of thermotolerance. Genetic manipulation can improve stress tolerance.
Hsp70 promoter driven expression in heat induced and non-heat induced conditions is more efficient in D.melanogaster embryos than in L.cuprina embryos.
The interaction of Top1 and Top2 gene products with transcriptionally active and inactive Hsp70B has been compared. Topoisomerase I binding sites are found in the transcribed portions of the Hsp70B gene, and only when Hsp70B is transcriptionally active. Topoisomerase II binding to Hsp70B sequences occurs on both transcriptionally active and inactive chromatin. An unusual type of topoisomerase II binding site is associated with the 5' ends of inactive Hsp70 gene, suggesting that this enzyme may be associated with repression of gene transcription.
initiates the heat-shock response. A restricted subset of genes, the Hsp genes, is activated and the majority of transcription and translation is shut down. However, mitochondrial- and histone-gene activities persist (Spradling, Pardue and Penman, 1977). This response follows a pulse of 36oC to 40oC; treatments above 40oC inhibit all activity and lead to death; treatments of 30oC-35oC induce heat-shock-protein synthesis without repressing normal protein synthesis (Tissieres, Mitchell and Tracy, 1974). Similar response inducible by other stressful treatments. The response may be elicited at all stages of the life cycle and in cultured cells. Stage specific phenocopies result from heat shocking early stages of Drosophila development (Mitchell and Petersen, 1982). In polytene cells existing puffs regress and a novel group quickly appears at 33B, 63C, 64F, 67B, 70A, 87A, 87C, 93D, 95D (Ashburner, 1970; Tissieres, Mitchell and Tracy, 1974). Activation of transcription of Hsp genes apparently involves the sequential binding of two or more protein factors in vicinity of TATA box (Wu, 1984). Binding sites for these proteins are multiple short upstream sequence elements called HSEs or heat shock consensus elements (Pelham, 1982; Xiao and Lis, 1988). Polymerase II dissociates from most chromosome regions and accumulates at the new puff sites (Bonner and Kerby, 1982). 3H-uridine incorporation ceases at its usual positions and commences at new puff sites. Preexisting polysomes disaggregate and within a few minutes a new population of polysomes appears containing newly transcribed mRNA; this RNA hybridizes to some of the heat-shock puffs. The effects of heat shock may be abrogated to some degree by pretreatment with a pulse of a slightly lower temperature (Mitchell, Moller, Petersen and Lipps-Sarmiento, 1979; Peterson and Mitchell, 1981). For reviews of the heat-shock response see Ashburner and Bonner (1978).
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. Hsp70B usually includes three HSP70 encoding genes (Hsp70Ba (proximal), Hsp70Bb (middle), Hsp70Bc (distal)) (Holmgren et al., 1979) with slightly different restriction maps (Artavanis-Tsakonas, Steward, Gehring, Mirault, Goldschmidt-Clermont, Moran and Tissieres, 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, Parks, Parker-Thornberg, Mortin and Pelham, 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).
Sørensen et al., 2013, J. Exp. Biol. 216(5): 809--814
Cellular damage as induced by high temperature is dependent on rate of temperature change - investigating consequences of ramping rates on molecular and organismal phenotypes in Drosophila melanogaster. [FBrf0220807]
Calabria et al., 2012, J. Evol. Biol. 25(4): 691--700
Hsp70 protein levels and thermotolerance in Drosophila subobscura: a reassessment of the thermal co-adaptation hypothesis. [FBrf0217786]
Coléno-Costes et al., 2012, PLoS Genet. 8(10): e1003006
New partners in regulation of gene expression: the enhancer of trithorax and polycomb corto interacts with methylated ribosomal protein l12 via its chromodomain. [FBrf0219702]
Defays and Bertoli, 2012, Alcohol 46(8): 737--745
Quantitative trait loci for response to ethanol in an intercontinental set of recombinant inbred lines of Drosophila melanogaster. [FBrf0220080]
Engel et al., 2012, Toxicol. In Vitro 26(3): 485--492
The effects of methylmercury on Notch signaling during embryonic neural development in Drosophila melanogaster. [FBrf0217650]
Fredriksson et al., 2012, Aging Cell 11(4): 634--643
Effects of aging and reproduction on protein quality control in soma and gametes of Drosophila melanogaster. [FBrf0218923]
Fuda et al., 2012, Mol. Cell. Biol. 32(17): 3428--3437
Fcp1 Dephosphorylation of the RNA Polymerase II C-Terminal Domain Is Required for Efficient Transcription of Heat Shock Genes. [FBrf0219151]
Herranz et al., 2012, BMC Genomics 13: 52
Microgravity simulation by diamagnetic levitation: effects of a strong gradient magnetic field on the transcriptional profile of Drosophila melanogaster. [FBrf0217750]
Mathieu et al., 2012, Nucleic Acids Res. 40(11): 4879--4891
Recruitment of the ATP-dependent chromatin remodeler dMi-2 to the transcribed region of active heat shock genes. [FBrf0219222]
Paré et al., 2012, PLoS ONE 7(5): e36254
The functions of grainy head-like proteins in animals and fungi and the evolution of apical extracellular barriers. [FBrf0218280]
Schneiderman et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(48): 19721--19726
Nucleosome-depleted chromatin gaps recruit assembly factors for the H3.3 histone variant. [FBrf0220071]
Ardehali et al., 2011, EMBO J. 30(14): 2817--2828
Drosophila Set1 is the major histone H3 lysine 4 trimethyltransferase with role in transcription. [FBrf0214333]
Boeke et al., 2011, PLoS ONE 6(6): e20761
The RNA Helicase Rm62 Cooperates with SU(VAR)3-9 to Re-Silence Active Transcription in Drosophila melanogaster. [FBrf0213941]
Cernilogar et al., 2011, Nature 480(7377): 391--395
Chromatin-associated RNA interference components contribute to transcriptional regulation in Drosophila. [FBrf0216979]
Chan et al., 2011, Hum. Mol. Genet. 20(9): 1738--1750
Expanded polyglutamine domain possesses nuclear export activity which modulates subcellular localization and toxicity of polyQ disease protein via exportin-1. [FBrf0214367]
Ghosh et al., 2011, Mol. Cell. Biol. 31(20): 4232--4243
Negative Elongation Factor Accelerates the Rate at Which Heat Shock Genes Are Shut off by Facilitating Dissociation of Heat Shock Factor. [FBrf0215827]
Morettini et al., 2011, Nucleic Acids Res. 39(8): 3103--3115
The chromodomains of CHD1 are critical for enzymatic activity but less important for chromatin localization. [FBrf0213515]
Murawska et al., 2011, PLoS Genet. 7(7): e1002206
Stress-Induced PARP Activation Mediates Recruitment of Drosophila Mi-2 to Promote Heat Shock Gene Expression. [FBrf0214629]
Takahashi et al., 2011, PLoS ONE 6(4): e17295
Environmental Stress-Dependent Effects of Deletions Encompassing Hsp70Ba on Canalization and Quantitative Trait Asymmetry in Drosophila melanogaster. [FBrf0213649]
Teves and Henikoff, 2011, Genes Dev. 25(22): 2387--2397
Heat shock reduces stalled RNA polymerase II and nucleosome turnover genome-wide. [FBrf0216728]
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