A Database of Drosophila Genes & Genomes

FB2013_03, released May 7th, 2013
 

Gene Dmel\per

General Information
SymbolDmel\perSpeciesD. melanogaster
NameperiodAnnotation symbolCG2647
Feature typeprotein_coding_geneFlyBase IDFBgn0003068
Gene Model StatusCurrent Stock availability 8 publicly available
Also Known Asdper, Clk, EG:155E2.4
Genomic Location
Chromosome (arm)XRecombination map1-
Cytogenetic map3B1-3B2Sequence locationX:2,579,613..2,586,813 [+]

Genomic Maps

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Automatically generated summary

See sections below for more information
The gene period is referred to in FlyBase by the symbol Dmel\per (CG2647, FBgn0003068). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: protein binding; transcription factor binding; protein heterodimerization activity. There is experimental evidence for 16 unique biological process terms, many of which group under: biological regulation; rhythmic process; response to stimulus; multi-organism reproductive process; response to stress; mating; single-organism developmental process; multicellular organismal development; learning or memory; determination of adult lifespan; locomotory behavior; age-dependent response to oxidative stress. 232 alleles are reported. The phenotypes of these alleles are annotated with: trichogen cell; indirect flight muscle motor neuron MN5; Malpighian tubule; mesothoracic tergum; posterior adult hindgut. It has 2 annotated transcripts and 2 annotated polypeptides. Protein features are: PAC motif; PAS domain; PAS fold. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderate expression to a trough of extremely low expression. Peak expression observed during late pupal stages, in stages of adults of both sexes. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, adult central nervous system. Comments on Affy2 ProbeSet: ProbeSet 1638452_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of per. Gene sequence location is X:2579613..2586813.

User Contributed Data
External Summaries
hide Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage
per: period (J.C. Hall; M. Young)
The per gene is essential for biological clock functions and determines the period length of circadian and ultradian rhythms. The per mutants are characterized by aberrant rhythms involving eclosion and locomotor activity (Konopka and Benzer, 1971) and may change the rhythmic component of the male courtship song (Crossley, 1988; Ewing, 1988; Kyriacou and Hall, 1980, 1986, 1988). These mutants also affect the rhythm of the larval heartbeat (Dowse, Ringo, and Kyriacou; Livingstone, 1981, Neurosci. Abstr. 7: 351), the level of tyrosine decarboxylase [Livingstone and Tempel, 1983, Nature (London) 303: 67-70], and fluctuations in membrane potentials in larval salivary glands (Weitzel and Rensing, 1981, J. Comp. Physiol. 143: 229-35), modulate intercellular junctional communication (Bargiello et al., 1987), and alter the location of neural secretory cells in the brain (Konopka and Wells, 1980, J. Neurobiol. 11: 411-15). In wild-type flies the period length is about 24 hr. In general, increases in per+ dosage lead to shortened circadian rhythms and decreases lead to lengthened circadian rhythms (Baylies et al., 1987; Cote and Brody, 1986; Hamblen et al., 1986; Smith and Konopka, 1981, 1982; Young et al., 1985). Females heterozygous for per+ and a deletion of the locus or a per0 allele show longer-than-normal periods. per flies can be classified on the basis of their circadian rhythms as: (1) Cryptic period mutants (per0, per-) which have a 10-15 hr (ultradian) period and appear arrhythmic except in special algorhythmic tests (Dowse et al., 1987); (2) Long period mutants (perL), 29 hr; (3) Long-period variable mutants (perLvar), which in homozygotes or heterozygotes are arrhythmic but in combination with certain partial deletions of the per locus result in a 30-34 hr period. (Konopka, 1987); (4) Short period mutants (pers), 19 hr; (5) Short period variable mutants (persvar), some flies having a 20 hr period and the others a normal 24 hr period for locomotor activity. In temperature-change experiments on pers and perL1, the locomotor activity periods were found to be nearer to 24 hr at low temperatures, but to diverge further from normal upon heating (Konopka, Pittendrigh, and Orr; Hamblen, Ewer, and Hall). perL2 shows lengthening of the periods at high temperatures. The mutant types affecting circadian rhythms (per0, perL, and pers) may cause similar kinds of changes in the rhythmic fluctuations in courtship song interpulse intervals (IPIs) of the male (Crossley, 1988; Ewing, 1988; Kyriacou and Hall, 1980, 1986, 1988). per0 mutants show nonrhythmic variations in the interval between pulses of wing vibration. Neural studies show that transplantation of pers brains into per01 adult hosts causes some of the hosts to be "rescued"; i.e. to show short-period circadian rhythms for locomotor activity (Handler and Konopka, 1979). Octopamine synthesis occurs at subnormal rates in per01 brains, with a corresponding decrease in the enzyme tyrosine decarboxylase (Livingstone and Tempel, 1983); less severe decrements in tyrosine decarboxylase are found in pers and perL1 flies. Physiological studies show that per mutations can affect the level of gap junctional communication among cells in a tissue (Bargiello et al., 1987). In salivary glands the per0 and perL1 mutations cause a lowering of the level of junctional communication, while pers gives a level of communication higher than wild type. Because electrical synapses are composed of gap junctions, per may influence circadian behavioral rhythms through altered conductances at the synapse (Bargiello et al., 1987). Mosaic analysis of pers mutants indicates that the gene influences the brain with respect to aberrant locomotor rhythms (Konopka, Wells, and Lee, 1983, Mol. Gen. Genet. 190: 284-88); per01 and per02 (and, to a lesser degree, pers) are said to cause anomalous photonegative behavior in light-response tests (Palmer, Kendrick, and Hotchkiss, 1985, Ann. N.Y. Acad. Sci., pp 323-24), but in general are not defective in visual responses (phototaxis tests, optomotor behavior, and electroretinogram) according to Dushay and Hall.
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Description
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FB2013_03
FB2013_02
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hide Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map
Evidence for location
3B1-3B2  
Limits computationally determined from genome sequence between P{EP}EP1362 and P{EP}dncEP1395  
Experimentally Determined Cytological Location
Cytogenetic map
Notes
References
3B1-3B1  
(determined by in situ hybridisation)  
3B1-3B2  
(determined by in situ hybridisation)  
3B1-3B2  
(determined by in situ hybridisation)  
3B1-3B2  
(determined by in situ hybridisation)  
3B1-3B1  
(determined by in situ hybridisation)  
3B1-3B2  
(determined by in situ hybridisation)  
Experimentally Determined Recombination Data
Location
Left of (cM)
Right of (cM)
Notes
1-1.2
 
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Please see the GBrowse view of Dmel\per for information on other features GBrowse View Help
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detailed view FBtr0340601 FBtr0332311 FBtr0070477 FBtr0070504 FBtr0070502 FBpp0070455 FBpp0304590 FBpp0070480 FBti0037293 FBti0036763 FBti0071605 FBti0007403 FBti0029753 FBti0110212 FBti0029836 FBti0107707 FBti0007495 FBti0103671 FBti0109436 FBti0033945 FBti0110263 FBti0020932 FBti0071597
Comments on Gene Model
Gene model reviewed during 5.45
hide Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Associated CDS (aa)
FBtr0070477
  4527
  1218
FBtr0332311
  4441
  1218
Additional Transcript Data & Comments
Reported size (kB)
Comments
External Data
Crossreferences
hide Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank protein
per-PA  
FBpp0070455  
127.4  
1218  
7.17  
per-PB  
FBpp0304590  
127.4  
1218  
7.17  
Additional Polypeptide Data & Comments
Reported size (kDa)
Comments
External Data
Linkouts
Crossreferences
InterPro domains - A database of protein families, domains, and functional sites
PAS domain (IPR000014)
PAS fold (IPR013767)
PAC motif (IPR001610)
hide Sequences Consistent with the Gene Model
DDBJ /
EMBL /
GenBank
DNA sequence
Protein sequence
Name
 
 
UniProtKB/Swiss-Prot
UniProtKB/TrEMBL
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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
regulatory region
comment=Region confers robust expression and circadian cycling of mRNA in transgenic flies.
evidence=experimental
regulatory region
comment=18bp E-box-containing region found to be sufficient for Clk-mediated activation of per transcription.
evidence=experimental
regulatory region
regulatory region
hide External Data
Linkouts
Crossreferences
EPD - Eukarytoic Promoter Database, an annotated collection of POL II promoters
hide Expression Data
hideTranscript Expression
expression microarray
Stage
Tissue/Position (including subcellular localization)
Reference
Comment:per mRNA levels do not significantly differ between ZT4 and ZT16.
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
RT-PCR
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
There is a dramatic rise in transcript level between Zeitgeber time (ZT)6 and 10. The level remains high for 8 hours, then drops precipitously by ZT22, suggesting that per mRNA cycling is transcriptionally regulated and that per mRNAs have short half-lives.
per transcripts show circadian cycling with a pattern that is antiphase to that of Clk transcripts. Peak values of per transcripts occur in the early night.
per transcripts are expressed in photoreceptor cells in the eye. They are also expressed in a wide region between the optic lobe and the central brain which includes the lateral neurons. They are expressed in the same pattern as dco transcripts.
The levels of per transcript oscillate throughout the day in a circadian fashion, similarly to tim but in contrast to inc. Expression of per is highest between ZT10 to ZT20.
per transcript levels in adult flies are not affected by exposure to light.
RT-PCR analysis indicates that the levels of the per transcript cycle when in light:dark conditions, in anti-phase with the levels of the Clk transcript. per transcript levels increase gradually from ZT02 to ZT14, and decrease after that until ZT20.
Marker for
Subcellular Localization
CV Term
hide Polypeptide Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Comment:midline and lateral cells
Comment:At the border between adjacent neuropils
Comment:DNa are found 15-30um anterior of DN1.
western blot
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
per protein in the adult is expressed in the s-LNv Pdf positive, l-LNv, LNd and DN neurons. On the third instar larval stage it is expressed in the larval s-LNv Pdf neurons.
Observed at ZT00, 06, 12 and 18, per protein shows oscillation in the LP neurons and DN1a neurons under a 12:12 LD cycle. per protein is found in the nucleus at ZT23. Expression of per is variable in intensity in the DN1p neurons, the DN2 neurons and DN3 neurons at ZT23.
per protein starts being expressed in the embryonic brain at stage 12, extending gradually to 130 neurons at stage 16; a hundred of those also express Scer\GAL4per.PK. per-positive neurons are found throughout the brain but with more cells located caudally. Around 20 cells co-express per and tim. These cells are located in the lateral protocerebrum, close to where the lateral and dorsomedial larval clock neurons are found. Expression of per in the ventral nerve cord starts at stage 12 in 2-3 neurons per segment close to the midline. The number of labeled cells increases until stage 16, when per is detected in 8 midline cells and 11 pairs of lateral cells per segment. Co-expression with en is found in 3 VUM interneurons and one UMI interneuron, four other midline neurons and two lateral cells per hemisegment.
per protein shows robust circadian cycling in the pacemaker neuron of wildtype flies.
Expression levels of per cycle in Malpighian tubules in a circadian pattern.
Period staining is strongest at ZT0-1, but is absent at ZT12. At peak expression, the Period protein is predominantly nuclear. tim protein colocalizes with the per protein.
At 1 hour before lights on in an LD cycle, per is predominantly nuclear localized, close to the nuclear membrane. per protein levels are highest 3hrs before lights on and lowest 12hrs later.
per protein shows strong oscillations under LD conditions, with a peak in the night and a gradual decrease at lights on. per protein is not restricted to the cytoplasm, but migrates to and from the nucleus.
The levels of per protein oscillate throughout the day in a circadian fashion, in contrast to inc. Expression of per is highest between ZT0 to ZT4 and ZT16 to ZT20.
A detailed examination of the distribution of per protein-expressing cells within the adult brain was made. Expression was detected in the ocelli, the photoreceptor cells, and throughout the central brain and the optic ganglia and the detailed locations of expressing cells were described. The most prominent immunoreactive cells are lateral cells in the cortical area between the inner margin of the medulla and the central brain neuropil which may correspond to the medullary tangential neurons. Double staining with an antibody to elav which recognizes only neurons was used to determine the nature of the per protein-expressing cells. The per protein-expressing cells in the ocelli and eyes, the lateral cells, and the dorsal-most cortex cells were shown to be neurons. The cells located at the margins of the cortex and neuropil in the optic lobes and the central brain, the cells within the lamina and central brain neuropil, and the cells in the inner chiasm are not neurons and are thought to be glia.
per is lost from within the nucleus of the l-LNvs by ZT10. It is found to be evenly distributed between the nucleus and the cytoplasm at ZT16, four hours after lights-off. It becomes predominantly nuclear by ZT19 and remains so for the rest of the dark period. In the s-LNv neurons, per is restricted to the cytoplasm until ZT16, but is seen in both the cytoplasm and nucleus by ZT18, becoming predominantly nuclear by ZT20.
Marker for
Subcellular Localization (GO Cellular Component)
CV term
References
hide Expression Deduced from Reporters
Reporter: P{GAL4-per.BS}
Stage
Tissue/Position (including subcellular localization)
Reference
Comment:Variably labelled between samples
Comment:Variably labelled between samples
Comment:Variably labelled between samples
Comment:Cells surrounding the esophageal foramen
Comment:faint expression
Comment:faint expression
Comment:strong expression
Comment:faint expression
Comment:medium expression
hide High-Throughput Expression Data
Associated Tools
Reference
See Gelbart and Emmert, 2010.10.13 for analysis details and data files for all genes.
hide FlyAtlas Anatomy Microarray

FlyAtlas Anatomical Expression Data

(FlyAtlas-RNA.adult)

(FlyAtlas-RNA.larva)


   Styles
   Scales

Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult head, adult eye, adult central nervous system.
[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
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
Linear, scaled to Moderate expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
 High 
Linear, scaled to High level expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
 High 
 Very high 
Linear, scaled to Very high expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 Very high 
log, scaled to maximum expression level
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
log, scaled to Moderate expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
 High 
log, scaled to High level expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
Expression Level Scale
 None 
 Low 
 Moderate 
 High 
 Very high 
log, scaled to Very high expression
Tissue   Expression Level
Larval Central Nervous System
 
 30.8
Larval Midgut
 
 19.6
Larval Hindgut
 
 3.9
Larval Malpighian Tubules
 
 7.2
Larval Fat Body
 
 18.4
Larval Salivary Gland
 
 12.8
Larval Trachea
 
 19.2
Larval Carcass
 
 7.025
Adult Head
 
 159.2
Adult Eye
 
 184.45
Adult Brain
 
 112.9
Adult Thoracic-Abdominal Ganglion
 
 110
Adult Crop
 
 44.6
Adult Midgut
 
 38.7
Adult Hindgut
 
 48
Adult Malpighian Tubules
 
 71.1
Adult Fat Body
 
 38.2
Adult Salivary Gland
 
 69.4
Adult Heart
 
 29.575
Adult VirginFemale Spermatheca
 
 30.9
Adult InseminatedFemale Spermatheca
 
 27.6
Adult Ovary
 
 20.2
Adult Testis
 
 11.4
Adult Male Accessory Gland
 
 26.4
Adult Carcass
 
 54.4
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
Larval Expression Level Tissue Adult Expression Level
 
NA 
Head
 
 159.2
 
NA 
Eye
 
 184.45
 
NA 
Brain
 
 112.9
 
30.8 
Central Nervous System
 
 NA
 
NA 
Thoracic-Abdominal Ganglion
 
 110
 
NA 
Crop
 
 44.6
 
no informative data 
Midgut
 
 38.7
 
3.9 
Hindgut
 
 48
 
7.2 
Malpighian Tubules
 
 no informative data
 
no informative data 
Fat Body
 
 38.2
 
no informative data 
Salivary Gland
 
 69.4
 
NA 
Heart
 
 29.575
 
19.2 
Trachea
 
 NA
 
NA 
VirginFemale Spermatheca
 
 30.9
 
NA 
InseminatedFemale Spermatheca
 
 no informative data
 
NA 
Ovary
 
 20.2
 
NA 
Testis
 
 no informative data
 
NA 
Male Accessory Gland
 
 no informative data
 
7.025 
Carcass
 
 54.4

FlyAtlas Anatomical Expression Data (Chintapalli et al., 2007)
hide modENCODE Anatomy RNA-Seq

modENCODE Tissue Expression Data

(modENCODE_mRNA-Seq_tissues)


   Styles
   Scales


[download data (TSV)]

Guide to modENCODE expression level colors
 
No/Extremely low expression (0 - 0)
 
Very low expression (1 - 3)
 
Low expression (4 - 10)
 
Moderate expression (11 - 25)
 
Moderately high expression (26 - 50)
 
High expression (51 - 100)
 
Very high expression (101 - 1000)
 
Extremely high expression (>1000)

Linear, scaled to maximum expression level
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
Linear, scaled to Moderate expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 (34)
head, mated 4-day male
 (41)
head, mated 20-day male
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
Linear, scaled to High expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
Linear, scaled to Extremely high expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Extremely high 
log, scaled to maximum expression level
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
log, scaled to Moderate expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 (34)
head, mated 4-day male
 (41)
head, mated 20-day male
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
log, scaled to High expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
log, scaled to Extremely high expression
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 1
central nervous system, larvae L3
 
 3
central nervous system, pupae P8
 
 8
head, virgin 1-day female
 
 15
head, virgin 4-day female
 
 26
head, virgin 20-day female
 
 15
head, mated 1-day female
 
 22
head, mated 4-day female
 
 21
head, mated 20-day female
 
 11
head, mated 1-day male
 
 34
head, mated 4-day male
 
 41
head, mated 20-day male
 
 26
salivary gland, larvae L3 wandering
 
 0
salivary gland, white prepupae
 
 0
digestive system, larvae L3 wandering
 
 0
digestive system, 1-day adult
 
 2
digestive system, 4-day adult
 
 3
digestive system, 20-day adult
 
 4
fat body, larvae L3 wandering
 
 1
fat body, white prepupae
 
 0
fat body, pupae P8
 
 5
carcass, larvae L3 wandering
 
 2
carcass, 1-day adult
 
 3
carcass, 4-day adult
 
 3
carcass, 20-day adult
 
 4
ovary, virgin 4-day female
 
 1
ovary, mated 4-day female
 
 2
testis, mated 4-day male
 
 1
accessory gland, mated 4-day male
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
 Extremely high 
Heatmap
Tissue   Expression Level
imaginal disc, larvae L3 wandering
 
 
central nervous system, larvae L3
 
 
central nervous system, pupae P8
 
 
head, virgin 1-day female
 
 
head, virgin 4-day female
 
 
head, virgin 20-day female
 
 
head, mated 1-day female
 
 
head, mated 4-day female
 
 
head, mated 20-day female
 
 
head, mated 1-day male
 
 
head, mated 4-day male
 
 
head, mated 20-day male
 
 
salivary gland, larvae L3 wandering
 
 
salivary gland, white prepupae
 
 
digestive system, larvae L3 wandering
 
 
digestive system, 1-day adult
 
 
digestive system, 4-day adult
 
 
digestive system, 20-day adult
 
 
fat body, larvae L3 wandering
 
 
fat body, white prepupae
 
 
fat body, pupae P8
 
 
carcass, larvae L3 wandering
 
 
carcass, 1-day adult
 
 
carcass, 4-day adult
 
 
carcass, 20-day adult
 
 
ovary, virgin 4-day female
 
 
ovary, mated 4-day female
 
 
testis, mated 4-day male
 
 
accessory gland, mated 4-day male
 
 

hide modENCODE Development RNA-Seq

modENCODE Temporal Expression Data

(modENCODE_mRNA-Seq_U)


   Styles
   Scales

Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderate expression to a trough of extremely low expression. Peak expression observed during late pupal stages, in stages of adults of both sexes.
[download data (TSV)]

Guide to modENCODE expression level colors
 
No/Extremely low expression (0 - 0)
 
Very low expression (1 - 3)
 
Low expression (4 - 10)
 
Moderate expression (11 - 25)
 
Moderately high expression (26 - 50)
 
High expression (51 - 100)
 
Very high expression (101 - 1000)
 
Extremely high expression (>1000)

Linear, scaled to maximum expression level
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
Linear, scaled to Moderate expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
Linear, scaled to High expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
Linear, scaled to Extremely high expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Extremely high 
log, scaled to maximum expression level
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
log, scaled to Moderate expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
log, scaled to High expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
log, scaled to Extremely high expression
Developmental Stage   Expression Level
embryo 00-02hr
 
 0
embryo 02-04hr
 
 0
embryo 04-06hr
 
 1
embryo 06-08hr
 
 1
embryo 08-10hr
 
 0
embryo 10-12hr
 
 1
embryo 12-14hr
 
 1
embryo 14-16hr
 
 2
embryo 16-18hr
 
 1
embryo 18-20hr
 
 1
embryo 20-22hr
 
 1
embryo 22-24hr
 
 1
larva L1
 
 1
larva L2
 
 0
larva L3 12hr old
 
 0
larva L3 puffstage 1-2
 
 0
larva L3 puffstage 3-6
 
 0
larva L3 puffstage 7-9
 
 1
white prepupae new
 
 1
white prepupae 12hr
 
 1
white prepupae 24hr
 
 2
pupae 2d postWPP
 
 4
pupae 3d postWPP
 
 3
pupae 4d postWPP
 
 6
adult male 01day
 
 8
adult male 05day
 
 9
adult male 30day
 
 10
adult female 01day
 
 7
adult female 05day
 
 2
adult female 30day
 
 4
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
 Extremely high 
Heatmap
Developmental Stage   Expression Level
embryo 00-02hr
 
 
embryo 02-04hr
 
 
embryo 04-06hr
 
 
embryo 06-08hr
 
 
embryo 08-10hr
 
 
embryo 10-12hr
 
 
embryo 12-14hr
 
 
embryo 14-16hr
 
 
embryo 16-18hr
 
 
embryo 18-20hr
 
 
embryo 20-22hr
 
 
embryo 22-24hr
 
 
larva L1
 
 
larva L2
 
 
larva L3 12hr old
 
 
larva L3 puffstage 1-2
 
 
larva L3 puffstage 3-6
 
 
larva L3 puffstage 7-9
 
 
white prepupae new
 
 
white prepupae 12hr
 
 
white prepupae 24hr
 
 
pupae 2d postWPP
 
 
pupae 3d postWPP
 
 
pupae 4d postWPP
 
 
adult male 01day
 
 
adult male 05day
 
 
adult male 30day
 
 
adult female 01day
 
 
adult female 05day
 
 
adult female 30day
 
 

modENCODE Temporal Expression Data (Graveley et al., 2011)
hide modENCODE Cell Lines RNA-Seq

modENCODE Cell Line Expression Data

(modENCODE_mRNA-Seq_cell.A)

(modENCODE_mRNA-Seq_cell.B)


   Styles
   Scales


[download data (TSV)]

Guide to modENCODE expression level colors
 
No/Extremely low expression (0 - 0)
 
Very low expression (1 - 3)
 
Low expression (4 - 10)
 
Moderate expression (11 - 25)
 
Moderately high expression (26 - 50)
 
High expression (51 - 100)
 
Very high expression (101 - 1000)
 
Extremely high expression (>1000)

Linear, scaled to maximum expression level
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
Linear, scaled to Moderate expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 (47)
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
Linear, scaled to High expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
Linear, scaled to Extremely high expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Extremely high 
log, scaled to maximum expression level
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
log, scaled to Moderate expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 (47)
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
log, scaled to High expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
log, scaled to Extremely high expression
Cell Line   Expression Level
Schneider line 2 S2R+
 
 0
Schneider line 2 Sg4
 
 0
embryonic 1182-4H
 
 1
embryonic GM2
 
 1
embryonic Kc167
 
 0
embryonic S1
 
 6
embryonic S3
 
 1
leg disc CME L1
 
 0
wing disc CME-W2
 
 0
wing disc ML-DmD8
 
 0
wing disc ML-DmD9
 
 1
wing disc ML-DmD16-c3
 
 2
wing disc ML-DmD21
 
 0
wing disc ML-DmD32
 
 0
haltere disc ML-DmD17-c3
 
 0
eye-antennal disc ML-DmD11
 
 0
antennal disc ML-DmD20-c5
 
 0
mixed discs ML-DmD4-c1
 
 2
CNS ML-DmBG1-c1
 
 8
CNS ML-DmBG2-c2
 
 7
tumorous blood cells mbn2
 
 0
ovary fGS/OSS
 
 47
ovary OSC
 
 1
ovary OSS
 
 2
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
 Extremely high 
Heatmap
Cell Line   Expression Level
Schneider line 2 S2R+
 
 
Schneider line 2 Sg4
 
 
embryonic 1182-4H
 
 
embryonic GM2
 
 
embryonic Kc167
 
 
embryonic S1
 
 
embryonic S3
 
 
leg disc CME L1
 
 
wing disc CME-W2
 
 
wing disc ML-DmD8
 
 
wing disc ML-DmD9
 
 
wing disc ML-DmD16-c3
 
 
wing disc ML-DmD21
 
 
wing disc ML-DmD32
 
 
haltere disc ML-DmD17-c3
 
 
eye-antennal disc ML-DmD11
 
 
antennal disc ML-DmD20-c5
 
 
mixed discs ML-DmD4-c1
 
 
CNS ML-DmBG1-c1
 
 
CNS ML-DmBG2-c2
 
 
tumorous blood cells mbn2
 
 
ovary fGS/OSS
 
 
ovary OSC
 
 
ovary OSS
 
 

hide modENCODE Treatments RNA-Seq

modENCODE Treatment Expression Data

(modENCODE_mRNA-Seq_treatments)


   Styles
   Scales


[download data (TSV)]

Guide to modENCODE expression level colors
 
No/Extremely low expression (0 - 0)
 
Very low expression (1 - 3)
 
Low expression (4 - 10)
 
Moderate expression (11 - 25)
 
Moderately high expression (26 - 50)
 
High expression (51 - 100)
 
Very high expression (101 - 1000)
 
Extremely high expression (>1000)

Linear, scaled to maximum expression level
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
Linear, scaled to Moderate expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
Linear, scaled to High expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
Linear, scaled to Extremely high expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Extremely high 
log, scaled to maximum expression level
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
log, scaled to Moderate expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
log, scaled to High expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
log, scaled to Extremely high expression
Treatment   Expression Level
extended cold, 4-day adult
 
 10
cold shock, 4-day adult
 
 5
heat shock, 4-day adult
 
 2
Cadmium 50 mM 6 hrs, larvae L3
 
 0
Cadmium 50 mM 12 hrs, larvae L3
 
 0
Cadmium 50 mM 48 hrs, 4-day adult
 
 3
Cadmium 100 mM 48 hrs, 4-day adult
 
 10
Copper 0.5 mM 12 hrs, larvae L3
 
 0
Copper 15 mM 48 hrs, 4-day adult
 
 3
Zinc 5 mM 12 hrs, larvae L3
 
 1
Zinc 4.5 mM 48 hrs, 4-day adult
 
 5
Ethanol 2.5% 3 hrs, larvae L3
 
 7
Ethanol 5% 3 hrs, larvae L3
 
 0
Ethanol 10% 3 hrs, larvae L3
 
 0
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 0
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 5
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 5
Paraquat 5 mM 48 hrs, 4-day adult
 
 6
Paraquat 10 mM 48 hrs, 4-day adult
 
 9
Rotenone 2 μg 12 hrs, larvae L3
 
 0
Rotenone 8 μg 12 hrs, larvae L3
 
 0
Expression Level Scale
 Very low 
 Low 
 Moderate 
 Moderately high 
 High 
 Very high 
 Extremely high 
Heatmap
Treatment   Expression Level
extended cold, 4-day adult
 
 
cold shock, 4-day adult
 
 
heat shock, 4-day adult
 
 
Cadmium 50 mM 6 hrs, larvae L3
 
 
Cadmium 50 mM 12 hrs, larvae L3
 
 
Cadmium 50 mM 48 hrs, 4-day adult
 
 
Cadmium 100 mM 48 hrs, 4-day adult
 
 
Copper 0.5 mM 12 hrs, larvae L3
 
 
Copper 15 mM 48 hrs, 4-day adult
 
 
Zinc 5 mM 12 hrs, larvae L3
 
 
Zinc 4.5 mM 48 hrs, 4-day adult
 
 
Ethanol 2.5% 3 hrs, larvae L3
 
 
Ethanol 5% 3 hrs, larvae L3
 
 
Ethanol 10% 3 hrs, larvae L3
 
 
Caffeine 1.5 mg/ml 4 hrs, larvae L3
 
 
Caffeine 2.5 mg/ml 48 hrs, 4-day adult
 
 
Caffeine 25 mg/ml 48 hrs, 4-day adult
 
 
Paraquat 5 mM 48 hrs, 4-day adult
 
 
Paraquat 10 mM 48 hrs, 4-day adult
 
 
Rotenone 2 μg 12 hrs, larvae L3
 
 
Rotenone 8 μg 12 hrs, larvae L3
 
 

hide Expression Clusters
A cluster of genes with similar mRNA expression dynamics across development.
hide External Data & Images
Linkouts
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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hide Summary of Allele Phenotypes
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Allele
hide Classical Alleles ( 55 )
For All Classical Alleles Show

Allele of perClassMutagenStocksKnown lesion
perJC43
2 --
perKG005461 --
per01amorphic allele - genetic evidence, loss of function allele0 Yes
per020 Yes
per030 Yes
per040 Yes
perClk0 Yes
perE02
0 Yes
perE11
0 Yes
perE12
0 Yes
perE15
0 Yes
perE30
0 Yes
perE33
0 Yes
perK1
0 --
perK2
0 --
perL1hypomorphic allele - genetic evidence0 Yes
perL20 Yes
perLI1
0 --
perLI2
0 --
perLvar0 --
perNJ1
0 --
perNJ2
0 --
perNW01
0 Yes
perNW03
0 Yes
perNW05
0 Yes
perNW06
0 Yes
perNW09
0 Yes
perNW12
0 Yes
perRD01
0 Yes
perRD02
0 Yes
perRD03
0 Yes
perRD05
0 Yes
perRD07
0 Yes
perRD08
0 Yes
perRD13
0 Yes
perRD15
0 Yes
perRD16
0 Yes
perRD20
0 Yes
perRD21
0 Yes
perRD23
0 Yes
perRD25
0 Yes
pershypermorphic allele - genetic evidence0 Yes
perSLIH
0 Yes
persvar0 --
perT0 Yes
perThr-Gly-140 Yes
perThr-Gly-150 Yes
perThr-Gly-170 Yes
perThr-Gly-200 Yes
perThr-Gly-210 Yes
perThr-Gly-23a0 Yes
perThr-Gly-23b0 Yes
perThr-Gly-240 Yes
perVT1.10 Yes
perVT1.20 Yes
hide Alleles Carried on Transgenic Constructs ( 177 )
For All Alleles Carried on Transgenic Constructs Show

Allele of perClassMutagenStocksKnown lesion
perGD12602 Yes
perHMS020451 Yes
perJF012261 Yes
perJF014521 Yes
per+mCA0 Yes
per+mCB0 Yes
per+mCC0 Yes
per+mCG0 Yes
per+t12.30 Yes
per+t13.20 Yes
per+t14.60 Yes
per+t7.10 Yes
per+t7.20 Yes
per+t8.00 Yes
per+tCa0 Yes
per01.tYa0 Yes
per1-1224.Act5C.T:SV5\V50 Yes
per1-23.T:Ecol\lacZ0 Yes
per1-450.Act5C.T:SV5\V5,T:SV40\nls20 Yes
per1-450.Act5C.T:SV5\V50 Yes
per1-450.M73-7.Act5C.T:SV5\V50 Yes
per1-516.Act5C.T:SV5\V5,T:SV40\nls20 Yes
per1-516.Act5C.T:SV5\V50 Yes
per1-95.T:Ecol\lacZ0 Yes
per1.T:Ecol\lacZ0 Yes
per447-1224.Act5C.T:SV5\V50 Yes
per513-1224.835-7.Act5C.T:SV5\V5,T:SV40\nls20 Yes
per513-1224.Act5C.T:SV5\V50 Yes
per513-1224.M788-91.Act5C.T:SV5\V50 Yes
per513-1224.M822-3.Act5C.T:SV5\V5,T:SV40\nls20 Yes
per513-1224.M822-3.Act5C.T:SV5\V50 Yes
per513-1224.M835-7.Act5C.T:SV5\V50 Yes
per513-813.Act5C.T:SV5\V50 Yes
per513-840.Act5C.T:SV5\V50 Yes
per513-892.Act5C.T:SV5\V50 Yes
per586-984.Act5C.T:SV5\V50 Yes
per764-1224.Act5C.T:SV5\V50 Yes
per8.0.T:Ecol\lacZ0 Yes
per8.0.T:Ppyr\LUC0 Yes
per807-1224.Act5C.T:SV5\V50 Yes
per829.T:Ecol\lacZ0 Yes
per843-1224.Act5C.T:SV5\V50 Yes
per8:80 Yes
per9.00 Yes
per95.gl.T:Ecol\lacZ0 Yes
per95.T:Ecol\lacZ0 Yes
perA0 Yes
perAct5C.cNa.T:SV5\V50 Yes
perAct5C.PD0 Yes
perAct5C.T:Avic\GFP-ECFP0 Yes
perAct5C.T:Ivir\HA10 Yes
perAct5C.T:SV5\V5,T:Zzzz\His60 Yes
perAct5C.T:SV5\V50 Yes
perB0 Yes
perBG.T:Ecol\lacZ0 Yes
perBG.T:Ppyr\LUC0 Yes
perC.T:Hsap\MYC0 Yes
perC.T:Ivir\HA10 Yes
perCG.T:Ecol\lacZ0 Yes
perClk.t13.20 Yes
perCRS.hs0 Yes
perCRS.P\T0 Yes
perCt-as.Scer\UAS0 Yes
perCt-s.Scer\UAS0 Yes
perD588F0 Yes
perD588V0 Yes
perD600E,S604G0 Yes
perD600V0 Yes
perdel0 Yes
perdsRNA.Ct.Scer\UAS0 Yes
perdsRNA.PAS.Scer\UAS0 Yes
perdsRNA.Scer\UAS0 Yes
perDyak\per.3'0 Yes
perE575G0 Yes
perE579A0 Yes
perE586G,D588A,M591L0 Yes
perE586V,V590D0 Yes
perE0 Yes
perF0 Yes
perG249F.Act5C.T:Hsap\MYC0 Yes
perG593C0 Yes
perG593S0 Yes
perG0 Yes
pergl.PV0 Yes
perH0 Yes
perhs.PEa0 Yes
perhs.PEb0 Yes
perhs.PH0 Yes
perhs.PN0 Yes
perhs.PS0 Yes
perhs.T:Avic\GFP-ECFP0 Yes
perKKKKNKG74-80PKKKRKV0 Yes
perL.hs.T:Avic\GFP-ECFP0 Yes
perL1.tBa0 Yes
perL415R.Act5C.T:Hsap\MYC0 Yes
perL574M0 Yes
perLN0 Yes
perLR0 Yes
perM.T:Hsap\MYC0 Yes
perM2M10 Yes
perM560D.Act5C.T:Hsap\MYC0 Yes
perN.T:Ivir\HA10 Yes
perNIG.2647R0 Yes
perninaE.PZ0 Yes
perninaE.T:Ivir\HA10 Yes
perP577R,H578Q0 Yes
perP597A0 Yes
perP597T,H599Q,Y601F0 Yes
perRKKKK73-76LNNNN0 Yes
pers.hs0 Yes
pers.tBa0 Yes
perS149-151-153A.tG0 Yes
perS149A.tG0 Yes
perS151-153D.tG0 Yes
perS151A.tG0 Yes
perS153.tG0 Yes
perS47A.Scer\UAS0 Yes
perS585A.T:Ivir\HA1.T:Zzzz\His60 Yes
perS585A0 Yes
perS589A.T:Ivir\HA1.T:Zzzz\His60 Yes
perS589D0 Yes
perS589E0 Yes
perS589G0 Yes
perS589L0 Yes
perS589T0 Yes
perS589Y0 Yes
perS596A.T:Ivir\HA1.T:Zzzz\His60 Yes
perS59A.tG0 Yes
perScer\UAS.cBa0 Yes
perScer\UAS.cKa0 Yes
perScer\UAS.cYa0 Yes
perSG.T:Ecol\lacZ0 Yes
perSL.T:Ivir\HA1,T:Zzzz\His60 Yes
perSMSTS1202-1206VLDR0 Yes
perT583A.S596A.T:Ivir\HA1.T:Zzzz\His60 Yes
perT:Ivir\HA1,T:Zzzz\His60 Yes
pertA0 Yes
pertB'0 Yes
pertG0 Yes
pertim.Aa0 Yes
pertThr-Gly-17a0 Yes
pertThr-Gly-10 Yes
pertThr-Gly-20a0 Yes
pertVT1.1-SNP3.SNP40 Yes
pertVT1.1-SNP30 Yes
pertVT1.1-SNP40 Yes
pertVT1.10 Yes
pertVT1.20 Yes
perV243D.Act5C.T:Hsap\MYC0 Yes
perW482A.Act5C.T:Hsap\MYC0 Yes
perXLG.T:Ecol\lacZ0 Yes
perXLG.T:Ppyr\LUC0 Yes
perαTub84B.3'UTR0 Yes
perΔ(Thr-Gly)0 Yes
perΔ1-95.hs0 Yes
perΔ118-293.hs0 Yes
perΔ118-530.hs0 Yes
perΔ240-293.hs0 Yes
perΔ365-454.hs0 Yes
perΔ452-512.hs0 Yes
perΔ452-569.hs0 Yes
perΔ515-569.hs0 Yes
perΔ524-685.hs0 Yes
perΔ540-569.hs0 Yes
perΔ585-601.hs0 Yes
perΔ598-1224.hs0 Yes
perΔ65-93,A94R,S95A0 Yes
perΔ830-1224.hs0 Yes
perΔ95-530.hs0 Yes
perΔ96-529.T:Ecol\lacZ0 Yes
perΔC20 Yes
perΔCBD.T:Ivir\HA1,T:Zzzz\His60 Yes
perΔcld0 Yes
perΔNLS.T:Ecol\lacZ0 Yes
perΔS0 Yes
perΔTG13.20 Yes
perΔTG8.00 Yes
hide Aneuploid Aberrations
Disrupted in
Duplicated in
Not disrupted in
Partially disrupted in
Partially duplicated in
Not duplicated in
hide Transgenic Constructs & Insertions
Transgenic Constructs
reporter construct
NameExpression Data
P{8.0-βgal}No
P{CATper.BS}No
P{CATper.H1}No
P{CATper.HS}No
P{CATper.R1}No
P{CATper.Spe1}No
P{CG-βgal}No
P{CperNlZ}No
P{CγB}No
P{glass-PER1-95β-gal}No
P{lacZper.P}No
P{NγB}No
P{PER1-23lacZ}No
P{PER1-95lacZ}No
P{PER1-95β-gal}No
P{PER1-636ΔNLS66-79lacZ}No
P{per21e-lacZ}No
P{per21-lacZ}No
P{per69x3-lacZ}No
P{per-hs43-lacZ.-341-156}No
P{per-hs43-lacZ.-449-603}No
P{per-hs43-lacZ.-467-325}No
P{per-hs43-lacZ.-505-449}No
P{per-hs43-lacZ.-563-494}No
P{per-hs43-lacZ.-603-449}No
P{per-hs43-lacZ.-603-550}No
P{per-hs43-lacZ.-1313-34}No
P{per-lacZ.829}No
P{per-lacZ.BG}No
P{per-lacZ.CRS}No
P{per-lacZ.NG}No
P{per-lacZ.SG}No
P{per-P\T-lacZ.-175-34}No
P{per-P\T-lacZ.-341-34}No
P{per-P\T-lacZ.-341-156}No
P{per-P\T-lacZ.-467-34}No
P{per-P\T-lacZ.-467-325}No
P{per-P\T-lacZ.-603-34}No
P{per-P\T-lacZ.-603-156}No
P{per-P\T-lacZ.-603-449}No
P{per-P\T-lacZ.-1313-34}No
P{per-P\T-lacZ.-1313-587}No
P{PER-SGΔ96-529}No
P{SγB}No
P{XLG}No
UAS construct
NameExpression Data
P{GD1260}NA
P{NIG.2647R}NA
P{TRiP.HMS02045}NA
P{TRiP.JF01226}NA
P{TRiP.JF01452}NA
P{UAS-per.B}NA
P{UAS-per.Ct-as}NA
P{UAS-per.Ct-IR}NA
P{UAS-per.Ct-s}NA
P{UAS-per.IR}NA
P{UAS-per.K}NA
P{UAS-per.PAS-IR}NA
P{UAS-per.S47A}NA
P{UAS-per.Y}NA
heat-shock construct
NameExpression Data
P{hsp70-per7.2}NA
P{hsp-per.E}NA
P{perCRS.hs}NA
P{perhs.PH}NA
GAL4 construct
NameExpression Data
P{GAL4-per.BS}No
characterization construct
NameExpression Data
P{13.2(perAsp589)}NA
P{13.2(perGlu589)}NA
P{13.2(perGly589)}NA
P{13.2(perThr589)}NA
P{13.2(perTyr589)}NA
P{BG-luc}NA
P{CB}NA
P{CP1}NA
P{glass-PERc}NA
P{MM1}NA
P{mps2}NA
P{mps3}NA
P{mps4}NA
P{mps5}NA
P{ninaE-per.-180}NA
P{per5.8}NA
P{per7.3}NA
P{per8.0(PA)}NA
P{per8.0}NA
P{per8.0ΔTG}NA
P{per9.0}NA
P{per9.8}NA
P{per12.3}NA
P{per13.2.cp}NA
P{per13.2.HA10His}NA
P{per13.2.VT1.1}NA
P{per13.2.VT1.1-SNP3.SNP4}NA
P{per13.2.VT1.1-SNP3}NA
P{per13.2.VT1.1-SNP4}NA
P{per13.2.VT1.2}NA
P{per13.2.W8}NA
P{per13.2(HA/C)}NA
P{per13.2(HA/N)}NA
P{per13.2(myc/C)}NA
P{per13.2(myc/M)}NA
P{per13.2}NA
P{per13.2C}NA
P{per13.2m-TGs}NA
P{per13.2s-TGm}NA
P{per13.2ΔCBD}NA
P{per13.2ΔTG}NA
P{per14.6}NA
P{per69x3-luc}NA
P{per.8:8}NA
P{per.8.0-luc}NA
P{per.dyp3'}NA
P{per.M2M1}NA
P{per.S59A}NA
P{per.S149A}NA
P{per.S151-153D}NA
P{per.S151A}NA
P{per.S153A}NA
P{per.S585A.HA.His}NA
P{per.S589A.HA.His}NA
P{per.S596A.HA.His}NA
P{per.T583A.S596A.HA.His}NA
P{per.XLG-luc}NA
P{per(Thr-Gly)1}NA
P{per(Thr-Gly)17a}NA
P{per(Thr-Gly)20a}NA
P{per+tCa}NA
P{perCRS.P\T}NA
P{perD588F}NA
P{perD588V}NA
P{perD600E,S604G}NA
P{perD600V}NA
P{perdel}NA
P{perE575G}NA
P{perE579A}NA
P{perE586G,D588A,M591L}NA
P{perE586V,V590D}NA
P{perG593C}NA
P{perG593S}NA
P{perKKKKNKG74-80PKKKRKV}NA
P{perL574M}NA
P{perl}NA
P{perLN}NA
P{perLR}NA
P{perP577R,H578Q}NA
P{perP597A}NA
P{perP597T,H599Q,Y601F}NA
P{perRKKKK73-76LNNNN}NA
P{perS585A}NA
P{perS589L}NA
P{pers}NA
P{perSMSTS1202-1206VLDR}NA
P{pertA}NA
P{pertB'}NA
P{pertG}NA
P{perΔ65-93,A94R,S95A}NA
P{perΔC2}NA
P{perαTub84B.3'UTR}NA
P{perA}NA
P{per-Aper\per.Ap}NA
P{perB}NA
P{perC7.2}NA
P{perCA}NA
P{perCC}NA
P{perCG}NA
P{perD}NA
P{perE}NA
P{perF}NA
P{perG}NA
P{perH}NA
P{per-luc.NOG}NA
P{perSL.HA10His}NA
P{perΔ(Thr-Gly)}NA
P{perΔcld}NA
P{perΔS.g}NA
P{rh-per}NA
P{tim-per.A}NA
Insertions
Type of insertions
Name
Expression data
insertion of mobile activating element
miscellaneous insertions
hide Gene Ontology: Function, Process & Cellular Component ( 33 unique terms )
hide Terms Based on Experimental Evidence ( 21 terms )
Molecular Function
CV term
References
inferred from physical interaction with UniProtKB:O77059
(assigned by UniProtKB)
inferred from physical interaction with Clk
inferred from physical interaction with Clk
Biological Process
CV term
References
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
inferred from mutant phenotype
Cellular Component
CV term
References
hide Terms Based on Predictions or Assertions ( 19 terms )
Molecular Function
CV term
References
non-traceable author statement
traceable author statement
non-traceable author statement
Biological Process
CV term
References
non-traceable author statement
traceable author statement
non-traceable author statement
non-traceable author statement
traceable author statement
non-traceable author statement
traceable author statement
traceable author statement
traceable author statement
Cellular Component
CV term
References
non-traceable author statement
traceable author statement
hide Sequence Ontology: Class of Gene
hide Interactions & Pathways
hide Summary of Physical Interactions
protein-protein
Interacting group
Assay
References
protein kinase assay, western blot
pull down, anti tag western blot
hide Summary of Genetic Interactions
Interacts with
Please look at the allele data for full details of the genetic interactions
per allele
Gene
References
hide External Data
Linkouts
BioGRID - A database of protein and genetic interactions
DroID - A comprehensive database of gene and protein interactions.
InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets.
hide Orthologs
hide OrthoDB Orthologs (30) - based on analysis using Dmel annotation version 5.41
OrthoDB Ortholog Groups
Drosophila inclusive ortholog search
Dipteran inclusive ortholog search
Insect inclusive ortholog search
Arthropod inclusive ortholog search
Metazoa inclusive ortholog search
hideOrthologs in Drosophila Species (EOG6Z0BRD)
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly 
 
 
Drosophila erecta
 
Y
 
Drosophila yakuba
 
 
 
Drosophila pseudoobscura pseudoobscura
 
Y
 
Drosophila willistoni
 
Y
 
Drosophila virilis
 
Y
 
Drosophila mojavensis
 
Y
 
Drosophila grimshawi
 
Y
 
hideOrthologs in non-Drosophila Dipterans (EOG6SJ5P6)
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Aedes aegypti
Yellow fever mosquito 
 
Anopheles gambiae
Malaria mosquito 
 
Culex quinquefasciatus
Southern house mosquito 
 
hideOrthologs in non-Dipteran Insects (EOG6D51SZ)
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Apis mellifera
Western honey bee 
Amel\GB52077
 
Apis mellifera
Western honey bee 
Amel\GB52078
 
Nasonia vitripennis
Parasitic wasp 
Nvit\Nasvi2EG013801
 
Acromyrmex echinatior
Panamanian leafcutter ant 
Aech\AECH12493
 
Atta cephalotes
Leafcutter ant 
Acep\ACEP14492
 
Camponotus floridanus
Florida carpenter ant 
Cflo\CFLO16807
 
Harpegnathos saltator
Jerdons jumping ant 
Hsal\HSAL28493
 
Linepithema humile
Argentine ant 
Lhum\LH14371
 
Pogonomyrmex barbatus
Red harvester ant 
Pbar\PB27286
 
Solenopsis invicta
Red fire ant 
Sinv\SINV16729
 
Acyrthosiphon pisum
Pea aphid 
 
Bombyx mori
Silkmoth 
 
Bombyx mori
Silkmoth 
 
Pediculus humanus
Human body louse 
 
Tribolium castaneum
Red flour beetle 
 
hideOrthologs in non-Insect Arthropods (EOG647D8H)
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Daphnia pulex
Water flea 
 
Ixodes scapularis
Deer tick 
 
hideOrthologs in non-Arthropod Metazoa (EOG6PZX0V)
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Caenorhabditis elegans
Nematode 
 
Danio rerio
Zebrafish 
 
hide Human Orthologs (0)
Gene
OMIM
HGNC
hideAAA Orthologs (9) based on analysis using Dmel annotation version 4.3
Organism
Gene
Drosophila sechellia
Drosophila erecta
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
hide Stocks & Reagents
hide Stocks Listed in FlyBase ( 8 )
Bloomington
Kyoto
VDRC
hide Genomic Clones ( 2 )
Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete
hide cDNA Clones ( 32 )
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
Other clones
cDNA Clones, End Sequenced (ESTs)
BDGP DGC clones
Other clones
hide RNAi & Array Information
Linkouts
DRSC - Results from RNAi screens.
GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents
hide Antibody Information
hide Other Information
hide Discoverer
Konopka.
 
hide Etymology
hide Identification
hide Relationship to Other Genes
Source for database identity of
Source for identity of: per CG2647
Source for database merge of
Additional comments
hide Other Comments
DNA-protein interactions: genome-wide binding profile (ChIP-chip) assayed for per protein in adult heads; see GEO_GSE32613 (http://www.ncbi.nlm.nih.gov/geo/query/acc.cgi?acc=GSE32613).
Gene expression is increased in response to the presence of two copies of Scer\GAL4hs.PB.
per is required for entrainment of the circadian clock by temperature.
per is necessary for nuclear localization of tim.
Nuclear transport is essential for the inhibitory action of per.
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.
Identified as one of 10 highest fold cycling genes as assessed by expression analysis using high density oligonucleotide arrays with probe generated from adult heads harvested over six time points over the course of a day.
per has a role in circadian mating rhythm regulation.
Maintenance of the per-tim feedback loop is not an absolute requirement for behavioral rhythms, but cyclic expression of the two proteins is essential.
per dependent molecular oscillators may have a role in the modulation of amine receptor responsiveness.
"Gene order: Overall orientation not stated: per+ CG2650- CG2658- Csat-" was stated as revision.
Excretory organs exhibit autonomous per and tim cycling.
Light and post-transcriptional regulation play major roles in defining the temporal properties of the per and tim protein curves. The lag between mRNA and protein accumulation is unecessary for the feedback regulation of per and tim protein on per and tim transcription.
Sensitization to repeated cocaine exposures, a phenomenon also seen in humans and animal models and associated with enhanced drug craving, is eliminated in flies mutant for per, dco, Clk, and cyc but not tim.
per, along with other circadian genes Clk, cyc, and dco (but not tim) have roles in regulating cocaine sensitization and may function as regulators of Tdc.
Mutants in per genes effect CrebB-17A activity, suggesting that the two genes participate in the same regulatory feedback loop.
The cry gene product blocks the function of the per/tim heterodimeric complex in a light-dependent fashion.
Clk mRNA cycling is regulated by per-tim-mediated release of Clk- and cyc-dependent repression.
Mutations in per abolish circadian rhythms in olfactory responses.
Daily cycles in the association of per and tim proteins with the Clk-cyc complex may contribute to rhythmic expression of per and tim.
A thermosensitive splicing event in the 3' UTR of the per mRNA plays an important role in how the circadian clock adapts to seasonally cold days. Enhanced splicing at cold temperatures advances the steady state phases of the per mRNA and protein cycles, contributing to the preferential day time activity of flies on cold days. There is a temperature-dependent switch in the molecular logic governing cycles in per mRNA levels.
timrit lengthens circadian period in a temperature-dependent manner through suppression of per protein cycling and nuclear localization.
tim increases per mRNA levels through a post-transcriptional mechanism.
per controls courtship song cycles.
Photoreceptors R1-R6 contain an autonomous circadian oscillator that can function without per mRNA cycling.
per mRNA cycling is not required for per protein cycling.
Transcription of per and tim, inhibitors if Clk, is induced by Clk.
Study of per and Dpse\per fusions reveals striking phenotypic differences between transgenic flies, these difference support the idea of an intragenic coevolution between the repeat and flanking regions of the two genes.
The gene product of the dco locus regulates per protein accumulation. The function of dco may be to reduce the stability and thus the level of accumulation of monomeric per proteins, promoting the delay between per/tim transcription and the function of the per/tim protein complex, which is essential for molecular rhythmicity.
Mutations at the per locus do not influence mean courtship duration.
Heat pulses at all times in a daily cycle elicited dramatic and rapid decreases in the levels of per and tim proteins, the proteins can be independently degraded by heat pulses. These two modalities produce markedly different long term effects on the circadian time-keeping mechanism. Heat-induced phase delays in behavioural rhythms are accompanied by long-term delays in the per and tim biochemical oscillations. Heat pulses in the late night elicit transient and rapid increases in the speed of the per-tim cycles. The timing of per and tim mRNA cycles is perturbed by heat pulses in a manner consistent with the direction and magnitude of the behavioural phase shift.
per coding region contains posttrancriptional regulatory information that is important for per protein cycling.
per protein expression in the pupal prothoracic gland has been studied in vivo and in ring glands cultured in vitro.
A circadian transcriptional enhancer within a 69bp DNA fragment is identified in per upstream sequences. The enhancer drives high-amplitude mRNA cycling under light-dark-cycling or constant-dark conditions and this activity is per protein-dependent. An E-box sequence within this fragment is necessary for high-level expression, but not for rhythmic expression, indicating per mediates circadian transcription through other sequences in this fragment.
Analysis of the rhythmic expression of a per reporter construct suggests that the Malpighian tubules contain a circadian pacemaker that functions independently of the brain.
The products of per and tim are detectable in a limited number of neurons in the larval CNS, the expression patterns in several such cells is cyclical.
The temporal regulation of per protein and RNA products is used to evaluate the status of the oscillatory mechanism in Pka-C1 mutants and to determine the site of action on the circadian timing system that is affected by reduced levels of Pka-C1.
per driven Ppyr\LUC reporter gene expression can be used to investigate transcriptional regulation of per and allows study of the molecular rhythmicity of an individual fly.
Quantifying rates of protein sequence divergence within and between species reveals that the Drosophila genome harbors a substantial proportion of genes with a very high divergence rate.
Nuclear run-on assays for fly heads and the in vivo transcription rate of per and tim suggest that there is an important circadian regulation at a post-transcriptional level. Results suggest this additional regulatory mode serves to ensure proper circadian fluctuations of clock gene expression.
The spatial and temporal expression patterns of per-Ecol\lacZ constructs containing differing amounts of the per coding region have been analysed.
A new regulatory element necessary for the correct temporal expression of per is identified by monitoring real-tie per expression in living individual flies carrying two different per-Ppyr\LUC transgenes.
Circadian rhythms are clearly exhibited in constant darkness even in flies reared in constant light and constant darkness as well as flies reared in light-dark cycles, but the freerunning period differs. Results suggest that the circadian clock is assembled without any cyclical photic information and light influences the developing circadian clock to alter the freerunning period.
A Ppyr\LUC reporter construct permits per circadian rhythms to be monitored by bioluminescence over several days in living adults and reveals novel features of per transcription.
Nuclear expression of tim depends on per. The expression of tim, but not per, is rapidly reduced by light, suggesting that tim mediates light-induced resetting of the circadian clock.
Photic stimuli perturb the timing of the per protein and mRNA cycles in a manner consistent with the direction and magnitude of the phase shift. The tim protein interacts with per in vivo, and the association is rapidly decreased by light. The disruption of the per-tim complex in the cytoplasm is accompanied by a delay in per phosphorylation and nuclear entry and disruption in the nucleus by an advance in per phosphorylation and disappearance.
Light inhibits the level and phosphorylation status of per and tim proteins, this then delays the negative feedback circuit (in which per or the per-tim complex participates) and extends the RNA profiles. This light-mediated delay compensates for the accelerated RNA increase in per mutant strains and restores rhythms to wild-type like periodicities.
The tim and per proteins physically interact and the timing of their association and nuclear localization promotes cycles of per and tim transcription through an autoregulatory feedback loop. tim couples the per-tim pacemaker to the environment: the tim gene product is rapidly degraded on exposure to light.
Studies of per mRNA cycling and locomotor activity rhythms under different light/dark cycling regimes indicate that the per feedback loop uses lights-off as a phase reference point and suggest that per mRNA cycling is not regulated via simple negative feedback from the per protein.
The complete sequence of the Thr-Gly region is examined and reveals that the region has evolved largely by the action of deletions/duplications and point mutations. Polymorphic sites upstream and downstream of the Thr-Gly region are also examined.
The identification of a period-altering tim allele provides further evidence that tim is a major component of the clock, and the allele-specific interactions with per provide evidence that the per/tim heterodimer is a unit of circadian function.
tim and per accumulate in the cytoplasm when independently expressed in S2 cells and move to the nucleus when coexpressed. Domains of per and tim have been identified that block nuclear localisation of the monomeric proteins. In vitro protein interaction studies indicate that the sequence inhibiting the nuclear accumulation of per forms a binding site for tim. Results indicate a mechanism for controlled nuclear localisation in which suppression of cytoplasmic localisation is accomplished by direct interaction of per and tim.
per and tim are in a large protein complex, they are heterodimeric partners.
The conformation of (Thr-Gly)n peptides (the conserved (Thr-Gly)n repeat in the per gene may have an important role in the temperature compensation of the circadian clock) has been analysed.
Key pacemaker neurons of the brain were examined to determine the changes of subcellular distribution of per with the time of day, per accumulates in the cytoplasm for several hours before entering the nucleus during a narrow time window. Long-period mutations cause a delay in the timing of nuclear translocation and a further delay at elevated temperature. This data indicates that regulation of per nuclear entry is critical for circadian oscillations by providing a necessary temporal delay between per synthesis and its effect on transcription.
The interaction between the tim and per products determines the timing of per nuclear entry and therefore the duration of part of the circadian cycle.
Identification: as a protein that interacts specifically with per protein in a yeast two-hybrid assay.
The entire arborisation pattern of per-containing pacemaker cells is revealed by immunostaining with crustacean pigment-dispersing hormone (PDH) antiserum. The arborisations of these neurons are appropriate for the modulation of the activity of many neurons and they might interact with per containing glial cells.
Temperature compensation of circadian period may be due in part to temperature-independent PER activity, which is based on competition between inter- and intramolecular interactions with similar temperature coefficients.
The period protein from the silkmoth A.pernyi can mediate clock functions in D.melanogaster.
tim and per interact and both are required for production of circadian rhythms. Absence of tim sequence similarity to the PER dimerization motif (PAS) indicates that direct interaction between the per and tim products would require a heterotypic protein association.
Mutations of tim lead to loss of circadian rhythms. tim suppresses circadian oscillations of per abundance and phosphorylation in light/dark cycles, depresses levels of per and blocks nuclear localisation of a per reporter gene due to a primary affect on per expression at the post-transcriptional level. Constant light has no effect on per in tim flies.
A linear relationship between temperature compensation and per Thr-Gly repeat length has been demonstrated. The major natural variants differ by units of (Thr-Gly)3, and rarer variants whose lengths fall out of phase with this pattern show more erratic temperature compensation, providing a correlation between behaviour and protein structure. Interspecific comparisons reveal a co-evolutionary process between the Thr-Gly region and flanking regions.
The tim RNA oscillations are dependent on the presence of per and tim proteins. The cyclic expression of tim appears to dictate the timing of per protein accumulation and nuclear localization, suggesting that tim promotes circadian rhythms of per and tim transcription by restricting per RNA and protein accumulation to separate times of day.
DNA binding coimmunoprecipitation assays studying the interaction of human Arnt with other PAS proteins demonstrates human Arnt forms heterodimers with per and sim, by means of the PAS domain, in a cooperative way.
Expression of per under the control of the gl promoter confers both behavioral and molecular rhythmicity. Expression in a few central brain cells producing gl and per product are capable of generating biological rhythms.
Ecol\lacZ reporter gene constructs demonstrate that per contains multipartite regulatory information for dosage compensation within the first large intron and also within the 3' half of the locus.
Ectopic expression of per supports the contention that the per gene product is a clock component.
The per protein undergoes daily oscillations in apparent molecular mass and abundance. The mobility changes are exclusively due to multiple phosphorylation events, phosphorylation is an important determinant in the clock time keeping mechanism.
Analysis of per mRNA cycling suggests that circadian oscillators are present in head and body tissues in which per protein is nuclear, and that these oscillators behave differently.
A hypothesis has been suggested that a controlled chaotic attractor may provide the central oscillator responsible for the generation of circadian and ultradian rhythms. this can be tested by seeking chaotic dynamics in systems where the controls have been disrupted.
The "Thr-Gly" region of the per gene has been compared with the corresponding region of the per genes of a number of Dipteran species.
The process of reinitiation of a 24 hour rhythmicity in individual flies reared in constant darkness (DD) is studied.
D.melanogaster females do not discriminate between males carrying D.melanogaster or D.simulans per genes, indicating that the per locus may only make a small contribution to total premating isolation between the two species.
per gene does not influence an observable locomotor behavioural phenotype in the larval stage of development.
per gene contains sequences that can inhibit per protein nuclear localisation in the absence of tim protein.
Autoregulation of per transcription is a direct, intracellular event and each per expressing cell may contain an autonomous clock, of which the per autoregulation loop is a component.
Sequences homologous to per demonstrated in the genome of the mole rat, Spalax ehrenbergi.
The per gene contains an amino acid motif of approximately 270 residues, termed PAS, whose function is unknown. PAS is also present in sim gene product, and in the two subunits of the mammalian dioxin receptor. Coprecipitation experiments showed that PAS functions in vitro as a novel protein dimerization motif which can mediate interactions between different members of the PAS protein family. Missense mutations in the PAS domain, including the original perL1, decrease the dimerization efficiency. In vivo experiments using transformants with tagged per coding regions suggest that dimerization also occurs in vivo.
Protein-protein interactions mediated through the PAS domain may be a crucial aspect of per biochemical function.
A 1.9 kb region of per has been compared in D.melanogaster, D.simulans, D.sechellia and D.mauritiana, and reveals a complex history. D.simulans appears to be a parent species to D.sechellia and D.mauritiana, but the order of appearance of the two species remains unclear. Whereas D.simulans and D.mauritiana share a large number of polymorphisms, D.sechellia shows very little variation.
Validity of experiments that led to the conclusions that strong differences in intercellular coupling distinguished per genotypes have been called into question. A reinvestigation of dye coupling concluded that no consistent differences in intercellular coupling in salivary glands can be attributed to mutations at the per lous (Flint, Rosbash and Hall, and Spray and Siwicki, unpublished information).
Mutations altering the structure of an approximately 20 amino acid segment, surrounding the location of the pers mutation, confer short per phenotypes. Loss or lowered function may dramatically increase measured level of protein activity.
Mutants exhibit defective courtship song.
Effects of per mutant alleles on visual pigment, sensitivity and rhabdomere size in 12hr light/12hr dark cycles was measured.
Minisatellite region encoding thr-gly repeat is polymorphic in length in natural populations. Geographical analysis of this polymorphism cloned by PCR reveals a robust clinal pattern observed along a north-south axis: the higher the latitude the more likely a population is to have thr-gly20 and less likely to have thr-gly17. Length polymorphism may be maintained by thermal selection, because thr-gly region has been shown to provide thermostability to the circadian phenotype.
A 13.2kb construct cloned from Clk-bearing flies will rescue arrhythmicity of per mutants. Transformant flies have shorter than normal rhythms demonstrating that the Clk mutation is within the 13.2b per fragment.
Internally marked mosaics determined that the pacemaker location is in the brain but not exclusively in the eyes, optic lobe or the ocelli. Although the pacemaker may be paired, the function of one of them is sufficient for rhythmicity. Glial expression is sufficient for some, albeit weak, rhythmicity.
Courtship song rhythms and locomotor activity rhythms assayed in D.melanogaster carrying Dsim\per or D.melanogaster/D.simulans per gene fusions. In all cases the circadian periodicities were slightly longer than for wild type, suggesting that the 13.2kb per fragment used to make the transgenic constructs is slightly inadequate at performing wild type per function.
Phases of the evening peaks of activity under LD conditions are correspondingly earlier than normal for the short-period mutants and later than normal for long cycle durations. The morning peaks, however, move minimally under the influence of a given per variant.
Fluctuations in per mRNA are primarily controlled by fluctuations in per transcription: per mRNA has a relatively short half life, and sequences sufficient to drive per mRNA cycling are present in 1.3kb of 5' flanking sequences.
per protein is found predominantly in the nuclei in adult Drosophila.
Mutagenesis of the per gene product reveals that the only the presence of a serine at residue 589 gives a 24h period but mutation of position 589 gives short period mutants. These short period mutants give rise to pacemakers with elements that fail to respond to the negative feedback loops of circadian oscillators.
The threonine-glycine encoding repeat region of the per gene shows high levels of length polymorphism in natural populations of D.melanogaster.
Associative learning of per mutants is assayed using the classical conditioning procedure of Tully (FBrf0043081). Results lend little support to the possibility that a short-cycle oscillation plays an integral role in learning processes; neither do they indicate any general effect on learning attributable to an abnormally lengthened circadian rhythm.
Chimeric per gene constructs from D.melanogaster and D.simulans have been used to map the genetic control of their courtship song rhythm difference to a small segment of the amino acid encoding information within per.
Ectopic expression of per demonstrates that per gene action during preimaginal stages is neither necessary or sufficient for locomotor activity rhythms to be expressed in the adult. per gene function appears to be necessary for pacemaker functioning itself. By analysing the singing of D.simulans and D.melanogaster reciprocally hybrid males the genetic etiology of the song rhythm difference is due to 1-4 amino acid replacements that have occurred over evolutionary time.
per mRNA levels in the fly head undergo circadian fluctuations during both 12 hour light/12 hour dark cycles and constant darkness.
The per gene product is thought to be involved in the regulation of the cell cycle as per mutant development time differs from wild type. A role of per in timers required for conditioning is suggested as mutations can slow down the biological timers. Transformation experiments involving the Thr-Gly encoding region of per (Yu, Nature 326:765) suggest that the Thr-Gly residues may play a role in determining song cycles, per has 17 to 23 Thr-Gly pairs and sings with 55 second song cycles.
The rhythmic components of male courtship songs have been spectral analysed.
Fluctuations in per protein expression in the visual system and central brain of adult flies at different times of the day have been studied.
per has an indirect influence on the levels of CG2650 transcript via its own regulation of eclosion rhythm.
Sequence analysis of frequency (frq) of N.crassa identifies a common element between frq and per suggesting a common element in the clock mechanisms of these two organisms.
Aberrant intervals between song pulses were observed in per mutant songs.
Comparison of the predicted protein sequence of the Thr-Gly repeat region of different D.melanogaster strains reveals a high degree of polymorphism and evolutionary plasticity. Deletion derivatives of the per gene lacking all the perfect Thr-Gly repeats indicates that the Thr-Gly region may have an important function in the courtship song phenotype.
The lesions associated with per01 and pers have been molecularly mapped.
per has been cloned and sequenced.
Germ line transformation demonstrates that a 7.1kb fragment including the per transcript can restore rhythmicity of eclosion and activity to per mutants.
per01 is complemented by l(1)3A and l(1)3B mutants nearby (Young and Judd, 1978; Smith and Konopka, 1981). The mutant per01 of D.melanogaster can be rescued (i.e. made to show rhythmic behavior) by transformation with a hybrid gene carrying the coding region of the D.pseudoobscura.pseudoobscura per gene (Peterson et al., 1988).
 
The per gene is essential for biological clock functions and determines the period length of circadian and ultradian rhythms. The per mutants are characterized by aberrant rhythms involving eclosion and locomotor activity (Konopka and Benzer, 1971) and may change the rhythmic component of the male courtship song (Crossley, 1988; Ewing, 1988; Kyriacou and Hall, 1980, 1986, 1988). These mutants also affect the rhythm of the larval heartbeat (Dowse, Ringo and Kyriacou) (Livingstone, 1981), the level of tyrosine decarboxylase (Livingstone and Tempel, 1983) and fluctuations in membrane potentials in larval salivary glands (Weitzel and Rensing, 1981), modulate intercellular junctional communication (Bargiello et al., 1987), and alter the location of neural secretory cells in the brain (Konopka and Wells, 1980). In wild-type flies the period length is about 24 hr. In general, increases in per+ dosage lead to shortened circadian rhythms and decreases lead to lengthened circadian rhythms (Baylies et al., 1987; Cote and Brody, 1986; Hamblen et al., 1986; Smith and Konopka, 1981; Smith and Konopka, 1982; Young et al., 1985). Females heterozygous for per+ and a deletion of the locus or a per01 allele show longer-than-normal periods. per flies can be classified on the basis of their circadian rhythms as: (1) Cryptic period mutants (per01, per-) which have a 10-15 hr (ultradian) period and appear arrhythmic except in special algorhythmic tests (Dowse, Hall and Ringo, 1987); (2) Long period mutants (perL), 29 hr; (3) Long-period variable mutants (perLvar), which in homozygotes or heterozygotes are arrhythmic but in combination with certain partial deletions of the per locus result in a 30-34 hr period (Konopka, 1987); (4) Short period mutants (pers), 19 hr; (5) Short period variable mutants (persvar), some flies having a 20 hr period and the others a normal 24 hr period for locomotor activity. In temperature-change experiments on pers and perL1, the locomotor activity periods were found to be nearer to 24 hr at low temperatures, but to diverge further from normal upon heating (Konopka, Pittendrigh, and Orr, 1989; Hamblen, Ewer and Hall). perL2 shows lengthening of the periods at high temperatures. The mutant types affecting circadian rhythms (per01, perL1 and pers) may cause similar kinds of changes in the rhythmic fluctuations in courtship song interpulse intervals (IPIs) of the male (Crossley, 1988; Ewing, 1988; Kyriacou and Hall, 1980; Kyriacou and Hall, 1986; Kyriacou and Hall, 1988). per01 mutants show nonrhythmic variations in the interval between pulses of wing vibration. Neural studies show that transplantation of pers brains into per01 adult hosts causes some of the hosts to be 'rescued'; i.e. to show short-period circadian rhythms for locomotor activity (Handler and Konopka, 1979). Octopamine synthesis occurs at subnormal rates in per01 brains, with a corresponding decrease in the enzyme tyrosine decarboxylase (Livingstone and Tempel, 1983); less severe decrements in tyrosine decarboxylase are found in pers and perL1 flies. Physiological studies that claimed that per mutations can affect the level of gap junctional communication among cells in a tissue (Bargiello et al., 1987). In salivary glands and that the per01 and perL1 mutations cause a lowering of the level of junctional communication have been withdrawn (Saez, Young, Baylies, Gasic, Bargiello and Spray, 1992). Mosaic analysis of pers mutants indicates that the gene influences the brain with respect to aberrant locomotor rhythms (Konopka, Wells and Lee, 1983); per01 and per02 (and, to a lesser degree, pers) are said to cause anomalous photonegative behavior in light-response tests (Palmer, Kendrick, and Hotchkiss, 1985), but in general are not defective in visual responses (phototaxis tests, optomotor behavior and electroretinogram) according to Dushay and Hall.
 
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Sequence Crossreferences
RefSeq (Transcripts)
RefSeq (Proteins)
Entrez Gene - A searchable database of RefSeq genes.
Other Crossreferences
EPD - Eukarytoic Promoter Database, an annotated collection of POL II promoters
InterPro domains - A database of protein families, domains, and functional sites
PAS domain (IPR000014)
PAS fold (IPR013767)
PAC motif (IPR001610)
Linkouts
BioGRID - A database of protein and genetic interactions
DroID - A comprehensive database of gene and protein interactions.
DRSC - Results from RNAi screens.
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyMine - Integrated genomics database for Drosophila, Anopheles, and C.elegans
FlyReactome - A curated knowledgebase of Drosophila melanogaster pathways
GenomeRNAi - GenomeRNAi – A database for cell-based and in vivo RNAi phenotypes and reagents
Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
InterologFinder Protein-protein interactions (PPI) from both known and predicted PPI data sets.
modMine - Data generated by the modENCODE project.
REDfly - A database of transcriptional regulatory elements.
hide Synonyms & Secondary IDs ( 18 )
Reported As
Symbol Synonym
clk-6
 
per
(Fujii et al., 2007, Wijnen et al., 2006, Akashi et al., 2006, Merrow, 2006, Krupp et al., 2007, Wijnen et al., 2007, Fang et al., 2007, Hart and Warrick, 2007, Kashi and King, 2006, Reppert, 2006, Dunlap, 2006, Ko et al., 2006, Wheeler et al., 2006, Kadener et al., 2006, Koh et al., 2006, Koh et al., 2006, Bartolome and Charlesworth, 2006, Chen et al., 2012, Megighian et al., 2001, Mazzoni et al., 2005, Lear et al., 2005, Shigeyoshi et al., 2002, Pezzulo et al., 2012, Kurata et al., 2007, Geiger-Thornsberry and Mackay, 2004, Mehnert et al., 2007, Hall, 2002, Yoshii et al., 2004, Choi et al., 2005, Fujii and Amrein, 2010, Van Gelder, 2006, Cyran et al., 2005, Lin et al., 2005, Klarsfeld et al., 2004, Stoleru et al., 2007, Sawyer et al., 2006, Konopka et al., 2007, Preuss et al., 2004, Majercak et al., 2004, Hung et al., 2007, Lim et al., 2007, Fang et al., 2007, Matsumoto et al., 2007, Yu and Hardin, 2006, Vosshall, 2007, Dunlap, 2008, Kaneko et al., 2000, Liu and Lehmann, 2008, Driver, 2000, Dolezelova et al., 2007, Xie et al., 2010, Ito et al., 2011, Boothroyd et al., 2007, Nawathean et al., 2007, Lim et al., 2007, Darlington et al., 2000, Lyons et al., 2000, Muskus et al., 2007, van der Linde et al., 2010, Kilman and Allada, 2009, Beaver et al., 2010, Smith et al., 2008, Kadener et al., 2007, Mehnert and Cantera, 2011, Kolaczkowski et al., 2011, Linde and Lyons, 2011, Lone et al., 2011, Hung et al., 2009, Taylor and Hardin, 2008, Kempinger et al., 2009, Lu et al., 2008, DiTacchio et al., 2011, Pereanu et al., 2010, Kaiser and Cobb, 2008, Tomaiuolo et al., 2008, Lone and Sharma, 2011, Abruzzi et al., 2011, Itoh et al., 2011, Ko et al., 2007, Yoshii et al., 2007, Gunawan and Doyle, 2007, Rush et al., 2006, Chen et al., 2006, Kaushik et al., 2007, McDermott and Kliman, 2008, Picot et al., 2007, Murad et al., 2007, Suh and Jackson, 2007, Kadener et al., 2008, Fernandez et al., 2008, Ito et al., 2008, Lee and Edery, 2008, Kivimäe et al., 2008, Diangelo et al., 2011, Tanoue et al., 2008, Krishnan et al., 2008, Krupp et al., 2008, Chiu et al., 2008, Busza et al., 2007, Cao and Nitabach, 2008, Meissner et al., 2008, Lear et al., 2005, Fujii et al., 2008, Akten et al., 2009, Mohammad et al., 2009, Yu et al., 2009, Paranjpe et al., 2005, Weber et al., 2009, Lyons and Roman, 2009, Landskron et al., 2009, Kilman et al., 2009, Donlea et al., 2009, Kotwica et al., 2009, Hung et al., 2009, Hall, 2003, Górska-Andrzejak et al., 2009, Noor and Kliman, 2003, Rogers et al., 2004, Kuo et al., 2010, Anaka et al., 2008, Rieger et al., 2009, Lear et al., 2009, Larsen et al., 2009, Tanenhaus et al., 2012, Wright, 1987, Dierick and Greenspan, 2006, Johard et al., 2009, Yoshii et al., 2008, Shafer et al., 2006, Blanchardon et al., 2001, T et al., 2008, Helfrich-Forster et al., 2007, Sehadova et al., 2009, Cusumano et al., 2009, Ni et al., 2009, Zheng et al., 2009, Hodge and Stanewsky, 2008, Currie et al., 2009, Shahidullah et al., 2009, Levine et al., 2002, Levine et al., 2002, Jaramillo et al., 2004, Yang et al., 2008, Houl et al., 2008, Krishnan et al., 2009, Krishnan et al., 2008, Chatterjee et al., 2010, Morozova et al., 2006, Fernández-Ayala et al., 2010, Menet et al., 2010, Barth et al., 2010, Yoshii et al., 2005, Zhang et al., 2010, Li et al., 2010, Sekine et al., 2008, Richier et al., 2008, Benito et al., 2008, Zheng et al., 2008, Wang et al., 2008, Bagheri et al., 2008, Zhang et al., 2010, Kula-Eversole et al., 2010, Blanchard et al., 2010, Nagoshi et al., 2010, Kadener et al., 2009, Codd et al., 2007, Smolen et al., 2004, Li and Lang, 2008, Fathallah-Shaykh et al., 2009, Lim et al., 2011, Ng et al., 2011, Saez et al., 2011, Thimgan et al., 2010, Syed et al., 2011, Gleason, 2005, Oliveira et al., 2011, Chiu et al., 2011, Fathallah-Shaykh, 2010, Lamaze et al., 2011, Ogawa et al., 2008, Miura et al., 2008, Ruiz et al., 2010, Powell et al., 2003, Zhou et al., 2005, Beaver et al., 2003, Hendricks et al., 2003, Stanewsky et al., 2002, Ivanchenko et al., 2001, Petri and Stengl, 2001, Stavropoulos and Young, 2011, Hara et al., 2011, Lone and Sharma, 2011, Mukherjee et al., 2012, Kaasik et al., 2013, Ling et al., 2012, Mizrak et al., 2012, Rakshit et al., 2012, Zheng et al., 2007, Japanese National Institute of Genetics, 2012.5.21, Shaik et al., 2008, Krishnan et al., 2012, Bywalez et al., 2012, Vanin et al., 2012, Grima et al., 2012, De et al., 2012, Kaneko et al., 2012, Ruben et al., 2012)
Name Synonym
Clock
 
clock-6
 
period
(Turner et al., 2008, Kliman, 1993.2.25, Kliman, 1993.2.25, Kliman, 1993.2.25, Kliman, 1993.2.25, Kliman, 1993.2.25, Kliman, 1993.2.25, Krupp et al., 2007, Wijnen et al., 2007, Fang et al., 2007, Wheeler et al., 2006, Megighian et al., 2001, Mazzoni et al., 2005, Lear et al., 2005, Wu and Silverman, 2007, Shigeyoshi et al., 2002, Geiger-Thornsberry and Mackay, 2004, Mehnert et al., 2007, Hall, 2002, Nitabach et al., 2005, Van Gelder, 2006, Korol et al., 2006, Cyran et al., 2005, Sawyer et al., 2006, Reaume and Sokolowski, 2006, Konopka et al., 2007, Sherazee et al., 2008, Preuss et al., 2004, Majercak et al., 2004, Lim et al., 2007, Yu and Hardin, 2006, Vosshall, 2007, Dunlap, 2008, Kaneko et al., 2000, Driver, 2000, Dolezelova et al., 2007, Pereanu and Hartenstein, 2006, Boothroyd et al., 2007, Nawathean et al., 2007, Lim et al., 2007, Darlington et al., 2000, Lyons et al., 2000, Mehnert and Cantera, 2011, Lone et al., 2011, Lu et al., 2008, Pereanu et al., 2010, Ruiz et al., 2010, Kaiser and Cobb, 2008, Tomaiuolo et al., 2008, Kaushik et al., 2007, Kadener et al., 2008, Lee and Edery, 2008, Krupp et al., 2008, Busza et al., 2007, Fan et al., 2009, Akten et al., 2009, Yu et al., 2009, Paranjpe et al., 2005, Weber et al., 2009, Yoshii et al., 2009, Lyons and Roman, 2009, Landskron et al., 2009, Kilman et al., 2009, Laayouni et al., 2007, Donlea et al., 2009, Hall, 2003, Górska-Andrzejak et al., 2009, Rogers et al., 2004, Kuo et al., 2010, Picot et al., 2009, Rieger et al., 2009, Larsen et al., 2009, Wright, 1987, Yoshii et al., 2008, Blanchardon et al., 2001, T et al., 2008, Sehadova et al., 2009, Zheng et al., 2009, Levine et al., 2002, Levine et al., 2002, Jaramillo et al., 2004, Houl et al., 2008, Krishnan et al., 2009, Krishnan et al., 2008, Morozova et al., 2006, Menet et al., 2010, Barth et al., 2010, Li et al., 2010, Bagheri et al., 2008, Blanchard et al., 2010, Saez et al., 2011, Syed et al., 2011, Gleason, 2005, Oliveira et al., 2011, Lamaze et al., 2011, Zhou et al., 2005, Beaver et al., 2003, Hendricks et al., 2003, Stanewsky et al., 2002, Ivanchenko et al., 2001, Petri and Stengl, 2001, Cirelli et al., 2005, Hara et al., 2011, Lone and Sharma, 2011, Rakshit et al., 2012, Krishnan et al., 2012, Bywalez et al., 2012, De et al., 2012, Kaneko et al., 2012)
period clock protein
Secondary FlyBase IDs
  • FBgn0000321
  • FBgn0020918
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hide Recent research papers ( 59 )
Gentile et al., 2013, Curr. Biol. 23(3): 185--195
Cryptochrome Antagonizes Synchronization of Drosophila's Circadian Clock to Temperature Cycles. [FBrf0220748]
Górska-Andrzejak et al., 2013, Dev. Neurobiol. 73(1): 14--26
Circadian expression of the presynaptic active zone protein bruchpilot in the lamina of Drosophila melanogaster. [FBrf0220165]
Kaasik et al., 2013, Cell Metab. 17(2): 291--302
Glucose Sensor O-GlcNAcylation Coordinates with Phosphorylation to Regulate Circadian Clock. [FBrf0220792]
Beaver et al., 2012, PLoS ONE 7(11): e50454
Circadian Regulation of Glutathione Levels and Biosynthesis in Drosophila melanogaster. [FBrf0220135]
Bywalez et al., 2012, Chronobiol. Int. 29(4): 395--407
The Dual-Oscillator System of Drosophila melanogaster Under Natural-Like Temperature Cycles. [FBrf0218098]
Chen et al., 2012, Science 335(6069): 678--685
Visualizing long-term memory formation in two neurons of the Drosophila brain. [FBrf0217453]
Collins et al., 2012, Neuron 74(4): 706--718
Balance of Activity between LN(v)s and Glutamatergic Dorsal Clock Neurons Promotes Robust Circadian Rhythms in Drosophila. [FBrf0218423]
De et al., 2012, J. Biol. Rhythms 27(4): 280--286
Adult Emergence Rhythm of Fruit Flies Drosophila melanogaster under Seminatural Conditions. [FBrf0219091]
Fontana and Crews, 2012, Dev. Biol. 372(1): 131--142
Transcriptome analysis of Drosophila CNS midline cells reveals diverse peptidergic properties and a role for castor in neuronal differentiation. [FBrf0219766]
Grima et al., 2012, PLoS Biol. 10(8): e1001367
CULLIN-3 Controls TIMELESS Oscillations in the Drosophila Circadian Clock. [FBrf0219137]
Hughes et al., 2012, Genome Res. 22(7): 1266--1281
Deep sequencing the circadian and diurnal transcriptome of Drosophila brain. [FBrf0218741]
Jepson et al., 2012, PLoS Genet. 8(4): e1002671
dyschronic, a Drosophila Homolog of a Deaf-Blindness Gene, Regulates Circadian Output and Slowpoke Channels. [FBrf0218124]
Kaneko et al., 2012, Curr. Biol. 22(19): 1851--1857
Circadian rhythm of temperature preference and its neural control in Drosophila. [FBrf0219631]
Kim et al., 2012, Nat. Neurosci. 15(6): 876--883
Contribution of visual and circadian neural circuits to memory for prolonged mating induced by rivals. [FBrf0219349]
Krishnan et al., 2012, Neurobiol. Disease 45(3): 1129--1135
Loss of circadian clock accelerates aging in neurodegeneration-prone mutants. [FBrf0217432]
Lagisz et al., 2012, Heredity 108(6): 602--608
Two distinct genomic regions, harbouring the period and fruitless genes, affect male courtship song in Drosophila montana. [FBrf0218390]
Linford et al., 2012, PLoS Genet. 8(5): e1002668
Re-Patterning Sleep Architecture in Drosophila through Gustatory Perception and Nutritional Quality. [FBrf0218228]
Ling et al., 2012, J. Neurosci. 32(47): 16959--16970
KAYAK-α Modulates Circadian Transcriptional Feedback Loops in Drosophila Pacemaker Neurons. [FBrf0219997]
Low et al., 2012, PLoS ONE 7(11): e49536
Natural Variation in the Drosophila melanogaster Clock Gene Period Modulates Splicing of Its 3'-Terminal Intron and Mid-Day Siesta. [FBrf0219973]
Luo and Sehgal, 2012, Cell 148(4): 765--779
Regulation of Circadian Behavioral Output via a MicroRNA-JAK/STAT Circuit. [FBrf0217497]
Luo et al., 2012, Aging Cell 11(3): 428--438
Old flies have a robust central oscillator but weaker behavioral rhythms that can be improved by genetic and environmental manipulations. [FBrf0218380]
Maeda and Kurata, 2012, PLoS ONE 7(2): e30489
A symmetric dual feedback system provides a robust and entrainable oscillator. [FBrf0218819]
Menegazzi et al., 2012, J. Biol. Rhythms 27(6): 433--442
Laboratory versus Nature: The Two Sides of the Drosophila Circadian Clock. [FBrf0220145]
Mizrak et al., 2012, Curr. Biol. 22(20): 1871--1880
Electrical activity can impose time of day on the circadian transcriptome of pacemaker neurons. [FBrf0219786]
Mukherjee et al., 2012, J. Exp. Biol. 215(17): 2960--2968
A model based on oscillatory threshold and build-up of a developmental substance explains gating of adult emergence in Drosophila melanogaster. [FBrf0219138]
Pezzulo et al., 2012, PLoS ONE 7(8): e43777
Expression of Human Paraoxonase 1 Decreases Superoxide Levels and Alters Bacterial Colonization in the Gut of Drosophila melanogaster. [FBrf0219363]
Rakshit et al., 2012, Chronobiol. Int. 29(1): 5--14
Effects of aging on the molecular circadian oscillations in Drosophila. [FBrf0217133]
Risau-Gusman and Gleiser, 2012, J. Theor. Biol. 307: 53--61
Modelling the effect of phosphorylation on the circadian clock of Drosophila. [FBrf0218753]
Ruben et al., 2012, J. Biol. Rhythms 27(5): 353--364
A mechanism for circadian control of pacemaker neuron excitability. [FBrf0219602]
Tanenhaus et al., 2012, PLoS ONE 7(10): e45130
In Vivo Circadian Oscillation of dCREB2 and NF-κB Activity in the Drosophila Nervous System. [FBrf0219734]
Vanin et al., 2012, Nature 484(7394): 371--375
Unexpected features of Drosophila circadian behavioural rhythms under natural conditions. [FBrf0218149]
Vesala et al., 2012, Insect Mol. Biol. 21(1): 107--118
Cold tolerance and cold-induced modulation of gene expression in two Drosophila virilis group species with different distributions. [FBrf0217233]
Abruzzi et al., 2011, Genes Dev. 25(22): 2374--2386
Drosophila CLOCK target gene characterization: implications for circadian tissue-specific gene expression. [FBrf0216798]
Chen et al., 2011, Curr. Biol. 21(9): 719--729
QUASIMODO, a Novel GPI-Anchored Zona Pellucida Protein Involved in Light Input to the Drosophila Circadian Clock. [FBrf0213611]
Chiu et al., 2011, Cell 145(3): 357--370
NEMO/NLK Phosphorylates PERIOD to Initiate a Time-Delay Phosphorylation Circuit that Sets Circadian Clock Speed. [FBrf0213577]
Depetris-Chauvin et al., 2011, Curr. Biol. 21(21): 1783--1793
Adult-specific electrical silencing of pacemaker neurons uncouples molecular clock from circadian outputs. [FBrf0216659]
Diangelo et al., 2011, PLoS ONE 6(5): e19921
The central clock neurons regulate lipid storage in Drosophila. [FBrf0213791]
DiTacchio et al., 2011, Science 333(6051): 1881--1885
Histone lysine demethylase JARID1a activates CLOCK-BMAL1 and influences the circadian clock. [FBrf0216307]
Goda et al., 2011, PLoS Genet. 7(7): e1002167
Adult Circadian Behavior in Drosophila Requires Developmental Expression of cycle, But Not period. [FBrf0214302]
Hara et al., 2011, J. Neurosci. 31(27): 9982--9990
Post-Translational Regulation and Nuclear Entry of TIMELESS and PERIOD Are Affected in New timeless Mutant. [FBrf0214253]
Ito et al., 2011, J. Biol. Rhythms 26(1): 14--23
Temperature Entrainment of the Circadian Cuticle Deposition Rhythm in Drosophila melanogaster. [FBrf0212848]
Itoh et al., 2011, Genes Cells 16(12): 1159--1167
Membrane-bound transporter controls the circadian transcription of clock genes in Drosophila. [FBrf0216813]
King et al., 2011, J. Mol. Biol. 413(3): 561--572
Structure of an enclosed dimer formed by the Drosophila period protein. [FBrf0216421]
Kolaczkowski et al., 2011, Genetics 187(1): 245--260
Genomic Differentiation Between Temperate and Tropical Australian Populations of Drosophila melanogaster. [FBrf0212762]
Lamaze et al., 2011, EMBO Rep. 12(6): 549--557
The E3 ubiquitin ligase CTRIP controls CLOCK levels and PERIOD oscillations in Drosophila. [FBrf0213851]
Lan and Mezić, 2011, BMC Syst. Biol. 5: 37
On the architecture of cell regulation networks. [FBrf0214351]
Lim et al., 2011, Nature 470(7334): 399--403
The novel gene twenty-four defines a critical translational step in the Drosophila clock. [FBrf0213059]
Linde and Lyons, 2011, Chronobiol. Int. 28(5): 397--406
Circadian modulation of acute alcohol sensitivity but not acute tolerance in Drosophila. [FBrf0214064]
Lone et al., 2011, Chronobiol. Int. 28(6): 497--508
Cyclic Presence and Absence of Conspecifics Alters Circadian Clock Phase But Does Not Entrain the Locomotor Activity Rhythm of the Fruit Fly Drosophila melanogaster. [FBrf0214519]
Lone and Sharma, 2011, J. Exp. Biol. 214(22): 3742--3750
Social synchronization of circadian locomotor activity rhythm in the fruit fly Drosophila melanogaster. [FBrf0216518]
Lone and Sharma, 2011, PLoS ONE 6(12): e28336
Circadian Consequence of Socio-Sexual Interactions in Fruit Flies Drosophila melanogaster. [FBrf0217074]
Mehnert and Cantera, 2011, Cell Tissue Res. 344(3): 381--389
Circadian rhythms in the morphology of neurons in Drosophila. [FBrf0213790]
Ng et al., 2011, Curr. Biol. 21(8): 625--634
Glial cells physiologically modulate clock neurons and circadian behavior in a calcium-dependent manner. [FBrf0213518]
Oliveira et al., 2011, BMC Evol. Biol. 11: 179
Variations on a theme: diversification of cuticular hydrocarbons in a clade of cactophilic Drosophila. [FBrf0214778]
Saez et al., 2011, Genetics 188(3): 591--600
A key temporal delay in the circadian cycle of Drosophila is mediated by a nuclear localization signal in the timeless protein. [FBrf0214282]
Stavropoulos and Young, 2011, Neuron 72(6): 964--976
insomniac and Cullin-3 Regulate Sleep and Wakefulness in Drosophila. [FBrf0217043]
Syed et al., 2011, J. Biol. Chem. 286(31): 27654--27662
Kinetics of Doubletime Kinase-dependent Degradation of the Drosophila Period Protein. [FBrf0214559]
Tio et al., 2011, PLoS ONE 6(11): e26879
Asymmetric cell division and notch signaling specify dopaminergic neurons in Drosophila. [FBrf0216651]
Yu et al., 2011, Curr. Biol. 21(9): 756--761
NEMO Kinase Contributes to Core Period Determination by Slowing the Pace of the Drosophila Circadian Oscillator. [FBrf0213626]
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