FB2022_03, released June 9, 2022

Gene: Dmel\Kr

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General Information
Symbol
Dmel\Kr
Species
D. melanogaster
Name
Kruppel
Annotation Symbol
CG3340
Feature Type
protein_coding_gene
FlyBase ID
FBgn0001325
Gene Model Status
Current
Stock Availability
567 publicly available
Gene Summary
Krueppel is a gap class segmentation protein. It is involved in the segmentation of the embryo and in the differentiation of the Malpighian tubules. (UniProt, P07247)
Contribute a Gene Snapshot for this gene.
All Summaries
Also Known As

If, Dm-Kr

Key Links
Genomic Location
Cytogenetic map
60F5-60F5
Sequence location
Recombination map
2-107
RefSeq locus
NT_033778 REGION:25226611..25231394
Sequence
Get Decorated FASTA
Genomic Maps
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
NCBI
UCSC
Ensembl
PopFly
Function
GO Summary Ribbons
Gene Ontology (GO) Annotations (20 terms)
Molecular Function (4 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
enables DNA-binding transcription factor activity, RNA polymerase II-specific
inferred from direct assay
(Shimell et al., 2000, Zuo et al., 1991)
enables DNA-binding transcription factor activity
inferred from direct assay
(Li et al., 2008)
enables sequence-specific DNA binding
inferred from direct assay
(Nibu et al., 2003, Zuo et al., 1991, Treisman and Desplan, 1989)
Terms Based on Predictions or Assertions (2 terms)
CV Term
Evidence
References
enables DNA-binding transcription factor activity
inferred from biological aspect of ancestor with PANTHER:PTN002811104
(assigned by GO_Central )
(Gaudet et al., 2011)
enables RNA polymerase II cis-regulatory region sequence-specific DNA binding
inferred from biological aspect of ancestor with PANTHER:PTN002811104
(assigned by GO_Central )
(Gaudet et al., 2011)
Biological Process (15 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
involved_in axon guidance
inferred from mutant phenotype
(Abrell and Jackle, 2001)
involved_in compound eye development
inferred from mutant phenotype
(Anderson et al., 2006, Baker et al., 1992)
involved_in heterochromatin assembly
inferred from mutant phenotype
(Matyash et al., 2009)
involved_in Malpighian tubule bud morphogenesis
inferred from genetic interaction with FLYBASE:ct; FB:FBgn0004198
(Hatton-Ellis et al., 2007)
involved_in negative regulation of transcription by RNA polymerase II
inferred from direct assay
(Shimell et al., 2000, Zuo et al., 1991)
involved_in negative regulation of transcription, DNA-templated
inferred from direct assay
(Nibu et al., 2003)
involved_in neuroblast fate determination
inferred from expression pattern
(Isshiki et al., 2001)
inferred from mutant phenotype
(Isshiki et al., 2001)
involved_in regulation of hemocyte proliferation
inferred from mutant phenotype
(Sinenko et al., 2010)
involved_in regulation of neural precursor cell proliferation
inferred from mutant phenotype
(Bahrampour et al., 2017)
involved_in regulation of neurogenesis
inferred from mutant phenotype
(Bahrampour et al., 2017)
Terms Based on Predictions or Assertions (5 terms)
CV Term
Evidence
References
involved_in muscle organ development
traceable author statement
(Dworak and Sink, 2002)
involved_in regulation of development, heterochronic
traceable author statement
(Banerjee and Slack, 2002, Pasquinelli and Ruvkun, 2002)
involved_in regulation of transcription by RNA polymerase II
inferred from biological aspect of ancestor with PANTHER:PTN002811104
(assigned by GO_Central )
(Gaudet et al., 2011)
involved_in trunk segmentation
traceable author statement
(Peel, 2004)
involved_in zygotic determination of anterior/posterior axis, embryo
traceable author statement
(Peel, 2004)
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Sharma and Nirenberg, 2007, Gursky et al., 2001, Lamka and Lipshitz, 1999, Small et al., 1991, Hoch et al., 1990, Gaul et al., 1987)
Terms Based on Predictions or Assertions (0 terms)
Gene Group (FlyBase)
Protein Family (UniProt)
Belongs to the krueppel C2H2-type zinc-finger protein family. (P07247)
Protein Signatures (InterPro)
Summaries
Gene Group (FlyBase)
C2H2 ZINC FINGER TRANSCRIPTION FACTORS -
Zinc finger C2H2 transcription factors are sequence-specific DNA binding proteins that regulate transcription. They possess DNA-binding domains that are formed from repeated Cys2His2 zinc finger motifs. (Adapted from PMID:1835093, FBrf0220103 and FBrf0155739).
Protein Function (UniProtKB)
Krueppel is a gap class segmentation protein. It is involved in the segmentation of the embryo and in the differentiation of the Malpighian tubules.
(UniProt, P07247)
Phenotypic Description (Red Book; Lindsley and Zimm 1992)
Kr: Kruppel
The wild-type allele of Kruppel controls the development of the thoracic and abdominal segments of Drosophilia; homozygous mutants show a gap in the larval pattern in these regions (Nusslein-Volhard and Wieschaus, 1980; Knipple, et al., 1987). Kr/+ adult sometimes has thoracic malformation; a leg or a wing may be absent; penetrance low. Heterozygous larvae show small defects in denticle bands of thorax and abdomen as do heterozygous deficiencies for Kr; penetrance about 80%. Homozygotes are embryonic lethal. Kr1 homozygotes exhibit shortened germ band of but three to four segments with three to four tracheal pits; visible beginning at 7h of embryogenesis; head and gnathal segments apparently normal. At later stages only three to four abdominal and thoracic segments clearly visible; normal telson and segments 8 and 7 followed by enlarged sixth, a rudimentary fifth and apparently mirror image sixth segment. Weak and intermediate mutant alleles lack the mirror-image duplications (Gaul and Jackle, 1987, Trends Genet. 3: 127-30). Ventral chain of ganglia disconnected; tracheal system defective; Malpighian tubules missing; salivary glands normal. Homozygotes for hypomorphic alleles display more nearly complete segmentation. Homozygous M+ Kr clones develop normally in all parts of adult cuticle of M/+ flies. Metamorphic potential of Kr/Kr embryos cultured in female abdomens restricted in that wing-disc-derived structures not observed. Germline clones of homozygous Kr cells capable of normal oogenesis; no maternal effect of Kr+ observed. Requirement for Kr+ function apparently restricted to early embryogenesis. Kr affects ftz producing abnormal intensity and spacing of ftz stripes in thorax and anterior abdomen (Carroll and Scott, 1986, Cell 45: 113-26).
If: Irregular facets
In heterozygote, eye area about one-half normal; narrow and pointed ventrally; facets irregular and often missing across middle of eyes, sometimes fused or absent in ventral portion. In homozygote, eyes are narrow slits with smooth glossy surface. In the eye disk of late third instar larvae, fairly large number of cell clusters in irregular arrangement, especially in ventral half of disk (Renfranz and Benzer, 1989). Viability and fertility good. RK1.
Summary (Interactive Fly)

transcription factor - zinc finger transcriptional repressor - gap gene required later for proper generation of neural sublineages - enriched in the larval eye and controls photoreceptor differentiation by promoting Rh5 and Rh6 expression

Gene Model and Products
Number of Transcripts
2
Number of Unique Polypeptides
1

Please see the JBrowse view of Dmel\Kr for information on other features

To submit a correction to a gene model please use the Contact FlyBase form

Protein Domains (via Pfam)
Isoform displayed:
Pfam protein domains
InterPro name
classification
start
end
Protein Domains (via SMART)
Isoform displayed:
SMART protein domains
InterPro name
classification
start
end
Structure New Section
Protein 3D structure   (Predicted by AlphaFold)   (AlphaFold entry P07247)

If you don't see the viewer to the right, refresh your browser.
Model Confidence:
  • Very high (pLDDT > 90)
  • Confident (90 > pLDDT > 70)
  • Low (70 > pLDDT > 50)
  • Very low (pLDDT < 50)

AlphaFold produces a per-residue confidence score (pLDDT) between 0 and 100. Some regions with low pLDDT may be unstructured in isolation.

Comments on Gene Model

Low-frequency RNA-Seq exon junction(s) not annotated.

Gene model reviewed during 5.46

Tissue-specific extension of 3' UTRs observed during later stages (FBrf0218523, FBrf0219848); all variants may not be annotated

Sequence Ontology: Class of Gene
nuclear_gene
protein_coding_gene
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
Kr-RA
FBtr0072449
NM_079143
2536
502
Kr-RB
FBtr0333162
NM_001274252
4411
502
Additional Transcript Data and Comments
Reported size (kB)

2.5 (northern blot)

(Jiang et al., 1986, Preiss et al., 1985)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
UniProt
RefSeq ID
GenBank
Kr-PA
FBpp0072351
54.7
502
7.47
P07247
NP_523867
AAF47321
Kr-PB
FBpp0305365
54.7
502
7.47
P07247
NP_001261181
AGB93711
Polypeptides with Identical Sequences

The group(s) of polypeptides indicated below share identical sequence to each other.

502 aa isoforms: Kr-PA, Kr-PB
Additional Polypeptide Data and Comments
Reported size (kDa)

402, 256 (aa)

(Sauer and Jackle, 1993)
Comments
External Data
Crossreferences
InterPro - A database of protein families, domains and functional sites
Linkouts
Sequences Consistent with the Gene Model
Mapped Features

Click to get a list of regulatory features (enhancers, TFBS, etc.) and gene disruptions (point mutations, indels, etc.) within or overlapping Dmel\Kr using the Feature Mapper tool.

External Data
Crossreferences
Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
Linkouts
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that the Fly Cell Atlas project found the gene expressed in that cell type. Darker colors mean that more cells of that cell type express the gene:
 low
high 
Colorless tiles indicate that there is no scRNAseq data for the gene in that cell type.
Colored tiles in ribbon indicate that expression data (RNA and/or protein) has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
Colored tiles in the ribbon indicate the average RNA expression level of the gene at the indicated stages:
 low
high 
as determined by RNA-seq (RPKM) using whole organism samples modENCODE, Brown et al., 2014. For complete stage-specific expression data, view the modENCODE Development RNA-Seq section under High-Throughput Expression below.
Transcript Expression
No Assay Recorded
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 14
embryonic/larval muscle system
(Gaul et al., 1987)
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
blastoderm stage
organism | 40-60% egg length

Comment: 39-57% egg length

(Hulskamp et al., 1990)
organism | medial
(Gavis and Lehmann, 1992)
embryonic stage 4
organism | 40-60% egg length
(Boring et al., 1993)
embryonic stage 4 -- 6
organism | gap expression pattern
(Fisher et al., 2012, Hoch et al., 1990)
organism | 40-60% egg length
(Hoch et al., 1990)
anlage in statu nascendi
(Fisher et al., 2012)
central brain anlage in statu nascendi
(Fisher et al., 2012)
dorsal ectoderm anlage
(Fisher et al., 2012)
hindgut anlage in statu nascendi
(Fisher et al., 2012)
posterior ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
ventral ectoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)
visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
antennal anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
dorsal head epidermis anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)
embryonic cycle 10 -- embryonic cycle 13
organism | 40-60% egg length
(Ali-Murthy and Kornberg, 2016)
embryonic stage 5
organism | 40-60% egg length
(Jacob et al., 1991)
organism | 90% egg length

Comment: faint second band at 88% egg length

(Jacob et al., 1991)
organism | gap expression pattern
(Capovilla et al., 1992)
embryonic stage 5 -- 6
organism | 80% egg length
(Hoch et al., 1990)
organism | 0-10% egg length
(Hoch et al., 1990)
embryonic stage 7 -- 8
antennal anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
central brain anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal head epidermis anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
visual anlage

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)
dorsal ectoderm
(Fisher et al., 2012)
hindgut anlage
(Fisher et al., 2012)
organism | gap expression pattern
(Fisher et al., 2012)
posterior endoderm
(Fisher et al., 2012)
ventral ectoderm
(Fisher et al., 2012)
ventral nerve cord primordium

Comment: reported as ventral nerve cord anlage

(Fisher et al., 2012)
embryonic stage 9
embryonic ventral midline neuroblast
(Jacob et al., 1991)
embryonic stage 9 -- 10
central brain primordium
(Fisher et al., 2012)
Malpighian tubule primordium

Comment: reported as Malpighian tubule anlage

(Fisher et al., 2012)
stomatogastric nervous system primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 9 -- 12
midline primordium | subset
(Kearney et al., 2004)
embryonic stage 10
Malpighian tubule primordium
(Hatton-Ellis et al., 2007)
embryonic/larval specialized Malpighian tubule cell
(Hatton-Ellis et al., 2007)
embryonic stage 11
epidermis | striped
(Jacob et al., 1991)
embryonic stage 11 -- 12
Bolwig organ primordium

Comment: reported as larval eye primordium

(Fisher et al., 2012)
central brain primordium
(Fisher et al., 2012)
Malpighian tubule primordium
(Fisher et al., 2012)
stomatogastric nervous system primordium
(Fisher et al., 2012)
ventral nerve cord primordium
(Fisher et al., 2012)
embryonic stage 12 -- 17
Bolwig organ
(Sheng et al., 1997)
embryonic stage 13 -- 16
Bolwig organ
(Fisher et al., 2012)
dorsal sensory complex proper primordium

Comment: reported as dorsal/lateral sensory complexes

(Fisher et al., 2012)
embryonic brain
(Fisher et al., 2012)
embryonic head sensory system
(Fisher et al., 2012)
embryonic/larval nervous system
(Fisher et al., 2012)
sensory nervous system primordium
(Fisher et al., 2012)
larval ventral nerve cord
(Fisher et al., 2012)
ventral sensory complex primordium
(Fisher et al., 2012)
larval midline neuron
(Kearney et al., 2004)
embryonic stage 14
embryonic brain
(Gaul et al., 1987)
embryonic/larval nervous system
(Gaul et al., 1987)
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 4 -- 10
(Preiss et al., 1985)
Additional Descriptive Data

Nascent Kr transcripts can be detected as early as cycle 7, but do not become numerous enough to be visible on whole-embryo imaging until cycle 10.

(Ali-Murthy and Kornberg, 2016)

Kr is detected in 4-6 Bolwig's organ (BO) precursor cells in stage 12 embryos. In stage 16 embryos, Kr is detected in 12 BO cells.

(Sheng et al., 1997)

The Kr transcript is expressed in the early embryo in a central stripe spanning from 40% to 60% egg length.

(Boring et al., 1993)

No Kr protein is detected in nosbcd.3UTR embryos.

(Gavis and Lehmann, 1992)

Kr transcript is first expressed in early blastoderm stage embryos in a broad central stripe. Late blastoderm stage embryos have an additional anterior stripe (80% egg length) and posterior stripe (90-100% egg length) of Kr transcript expression. By early gastrulation, the anterior stripe resolves into two stripes, and the central band into four stripes of differing intensities.

(Hoch et al., 1990)

Kr is expressed in the middle of the blastoderm embryo from approximately 39-57% egg length.

(Hulskamp et al., 1990)

In germ band retracted embryos, Kr transcripts are detected in the embryonic brain, as well as the precursor lateral muscle cells, significant levels are also detected in developing neural tissue.

(Gaul et al., 1987)

The primary Kr transcript is most abundant in 2-5 hour embryos, but transcripts are detected through pupal stages at lower levels.

(Jiang et al., 1986)

A 2.5kb Kr transcript is detected at high levels in 2-5 hour embryos.

(Preiss et al., 1985)
Marker for
 
Malpighian tubule primordium
(Hatton-Ellis et al., 2007)
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
photoreceptor precursor cell
(Mishra et al., 2016)
embryonic stage 4
organism | 40-60% egg length
(Gaul et al., 1987)
embryonic stage 4 -- 5
organism | gap expression pattern
• nucleus
(Small et al., 1991)
embryonic stage 4 -- 6
organism | gap expression pattern
• nucleus
(Hoch et al., 1990)
organism | 40-60% egg length
• nucleus
(Hoch et al., 1990)
embryonic stage 5
organism | gap expression pattern
(Hoch and Jackle, 1998, Langeland et al., 1994, Langeland and Carroll, 1993, Stanojevic et al., 1989)
posterior ectoderm derivative
(Hoch and Jackle, 1998)
organism | 40-60% egg length
(Hoch et al., 1991)
embryonic stage 5 -- 6
organism | 80% egg length
• nucleus
(Hoch et al., 1990)
embryonic stage 6
organism | 0-10% egg length
• nucleus
(Hoch et al., 1990)
embryonic stage 8
Malpighian tubule primordium
(Hoch and Jackle, 1998)
embryonic stage 8 -- 11
neuroblast NB5-6 of thorax
(Karlsson et al., 2010)
neuroblast NB5-6 of abdomen
(Karlsson et al., 2010)
embryonic cycle 14
organism | 40-60% egg length
(Janssens et al., 2013)
embryonic cycle 14A
organism | gap expression pattern
• nucleus
(Gursky et al., 2001)
embryonic stage 9
ganglion mother cell
(Karlsson et al., 2010)
embryonic stage 9 -- 12
Malpighian tubule primordium
• nucleus
(Hoch et al., 1990)
stomodeal invagination
• nucleus
(Hoch et al., 1990)
embryonic stage 9 -- 14
amnioserosa
• nucleus
(Hoch et al., 1990)
embryonic stage 9 -- 17
presumptive embryonic/larval central nervous system
• nucleus
(Hoch et al., 1990)
embryonic stage 10
neuroblast NB3-3
(Tsuji et al., 2008)
early stomodeal invagination
(Gonzalez-Gaitan and Jackle, 2000)
embryonic stage 10 -- 12
somatic muscle primordium
(Ruiz Gomez and Bate, 1997)
embryonic stage 10 -- 17
embryonic Malpighian tubule
(Ainsworth et al., 2000)
embryonic stage 11
amnioserosa
• nucleus
(Lamka and Lipshitz, 1999)
muscle founder cell of muscle cell of mesothoracic lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle founder cell of muscle cell of metathoracic lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle founder cell of muscle cell of A1-7 lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle founder cell of muscle cell of mesothoracic dorsal oblique muscle 4
(Enriquez et al., 2012)
muscle founder cell of muscle cell of metathoracic dorsal oblique muscle 4
(Enriquez et al., 2012)
embryonic stage 11 -- 12
Malpighian tubule tip cell primordium
(Hoch and Jackle, 1998)
embryonic stage 12 -- 13
embryonic myoblast
(Yarnitzky et al., 1998)
embryonic stage 12 -- 14
somatic muscle primordium
• nucleus
(Hoch et al., 1990)
embryonic stage 12 -- 17
photoreceptor cell of Bolwig organ
(Sprecher et al., 2007)
embryonic stage 13
embryonic primordium of adult muscle precursor | restricted
(Chen and Olson, 2001)
embryonic Malpighian tubule tip cell
(Ainsworth et al., 2000)
embryonic stage 13 -- 16
amnioserosa
(Stronach and Perrimon, 2001)
embryonic stage 14 -- 15
muscle cell of A1-7 lateral transverse muscle 2 | precursor
(Jagla et al., 1999)
muscle cell of mesothoracic lateral transverse muscle 2 | precursor
(Jagla et al., 1999)
muscle cell of metathoracic lateral transverse muscle 2 | precursor
(Jagla et al., 1999)
muscle cell of A1-7 lateral transverse muscle 4 | precursor
(Jagla et al., 1999)
muscle cell of mesothoracic lateral transverse muscle 4 | precursor
(Jagla et al., 1999)
muscle cell of metathoracic lateral transverse muscle 4 | precursor
(Jagla et al., 1999)
muscle cell of lateral longitudinal muscle 1 | precursor
(Jagla et al., 1999)
muscle cell of ventral acute muscle 2 | precursor
(Jagla et al., 1999)
embryonic stage 15
abdominal lateral adult muscle precursor cell
(Figeac et al., 2010)
muscle cell of mesothoracic lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle cell of metathoracic lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle cell of A1-7 lateral longitudinal muscle 1
(Enriquez et al., 2012)
muscle cell of lateral transverse muscle 4
(Enriquez et al., 2012)
embryonic stage 16
U3 neuron
(Touma et al., 2012)
embryonic stage 16 -- 17
Bolwig organ
• nucleus
(Hoch et al., 1990)
embryonic/larval brain | restricted
• nucleus
(Hoch et al., 1990)
larval ventral nerve cord | restricted
• nucleus
(Hoch et al., 1990)
embryonic stage 17
EL neuron | subset
(Tsuji et al., 2008)
GW neuron
(Karcavich and Doe, 2005)
EW1 neuron
(Karcavich and Doe, 2005)
EW2 neuron
(Karcavich and Doe, 2005)
larval stage
photoreceptor cell of Bolwig organ
(Sprecher et al., 2007)
third instar larval stage
photoreceptor neuron
(Mishra et al., 2016)
Additional Descriptive Data

Kr is expressed in the larval photoreceptors but is absent from the photoreceptors of adult ocelli. It is expressed in all larval photoreceptor precursors throughout embryogenesis. Thereafter, it is maintained in both photoreceptor subtypes (Rh5-expressing and Rh6-expressing) during larval stages.

(Mishra et al., 2016)

Kr is differentially expressed in the LAMPs (lateral adult muscle precursor cells). It is expressed strongly in the anterior LAMP but not in the posterior LAMP.

(Figeac et al., 2010)

Kr is expressed in Bolwig organ precursors beginning in embryonic stage 12 and continuing through the larval stages.

(Sprecher et al., 2007)

Protein is detected in a subset of myoblasts, the founder cells, in stage 13 embryos. As additional cells are recruited to the developing myotubes expression is observed in these clusters corresponding to fusing myotubes.

(Chen and Olson, 2001)

The Malpighian tubule cells are alocated and evert from the embryonic hindgut during extended germband stage. From this stage on protein is detected in the tubule cells.

(Ainsworth et al., 2000)

In blastoderm embryos, the Kr protein expression domain extends from the posterior border of eve stripe 2 to the anterior border of eve stripe 5.

(Small et al., 1991)

In blastoderm stageembryos, Kr protein is localized to the dorsal and ventral periphery, in aband ranging from 10-12 nuclei dorsally and 14-16 nuclei ventrally (40-60% egglength). Staining is highest in the central domain, and fades in a gradedfashion towards the edges of this "Kr domain".

(Gaul et al., 1987)
Marker for
 
Bolwig organ primordium
(Sprecher et al., 2007)
U3 neuron
(Touma et al., 2012)
amnioserosa
(Stronach and Perrimon, 2001)
embryonic myoblast
(Yarnitzky et al., 1998)
Subcellular Localization
CV Term
Evidence
References
located_in nucleus
inferred from direct assay
(Sharma and Nirenberg, 2007, Gursky et al., 2001, Lamka and Lipshitz, 1999, Small et al., 1991, Hoch et al., 1990, Gaul et al., 1987)
Expression Deduced from Reporters
Reporter: P{1BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length

Comment: reference stages ~42-55% egg length

(Hoch et al., 1991)
Reporter: P{1BSKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length

Comment: reference stages ~42-55% egg length

(Hoch et al., 1991)
Reporter: P{4BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length
(Hoch et al., 1991)
Reporter: P{4BSKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length

Comment: reference stages ~45-53% egg length

(Hoch et al., 1991)
Reporter: P{8BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length

Comment: Reference states slightly posterior to native Kr

(Hoch et al., 1991)
Reporter: P{BΔNc0.7HZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 5
organism | 40-60% egg length
(Hoch et al., 1991)
Reporter: P{dPN5.4KrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 4 -- 5
organism | gap expression pattern
(Hoch et al., 1990)
organism | 40-60% egg length
(Hoch et al., 1990)
Reporter: P{GAL4-Kr.C}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
ectoderm
(Golembo et al., 1996)
parasegment 4 -- parasegment 9
(Schweitzer et al., 1995)
embryonic stage 6 -- 10
parasegment 5 -- parasegment 9
(Castelli-Gair et al., 1994)
embryonic stage 9 -- 12
amnioserosa
(Casso et al., 2000)
posterior ectoderm
(Casso et al., 2000)
embryonic stage 13 -- 15
amnioserosa
(Narasimha and Brown, 2004)
embryonic stage 16 -- 17
embryonic/larval plasmatocyte
(Casso et al., 2000)
Bolwig organ
(Casso et al., 2000)
embryonic stage 17
embryonic/larval posterior spiracle
(Casso et al., 2000)
first instar larval stage
Bolwig organ
(Casso et al., 2000)
embryonic/larval posterior spiracle
(Casso et al., 2000)
second instar larval stage
embryonic/larval salivary gland
(Casso et al., 2000)
third instar larval stage
embryonic/larval fat body
(Casso et al., 2000)
prepupal stage
embryonic/larval fat body
(Casso et al., 2000)
pharate adult stage
pupal abdomen
(Casso et al., 2000)
adult stage
adult abdomen | restricted
(Casso et al., 2000)
adult stage | male
testis pigment cell
(DeFalco et al., 2008)
embryonic stage 17
gonadal sheath
(DeFalco et al., 2008)
Reporter: P{GAL4-Kr.zen.8}
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage 9 -- 17
ectoderm
(Frasch, 1995)
Reporter: P{GawB}AG1
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
Bolwig organ
(Tang and Sun, 2002)
third instar larval stage
ventral thoracic disc | restricted
(Tang and Sun, 2002)
eye-antennal disc | restricted
(Tang and Sun, 2002)
oogenesis
follicle cell
(Tang and Sun, 2002)
spermatogenesis
testis sheath
(Tang and Sun, 2002)
Reporter: P{GawB}sl(2)AM1AM1
Stage
Tissue/Position (including subcellular localization)
Reference
embryonic stage
Bolwig organ
(Tang and Sun, 2002)
third instar larval stage
ventral thoracic disc | restricted
(Tang and Sun, 2002)
eye-antennal disc | restricted
(Tang and Sun, 2002)
oogenesis
follicle cell
(Tang and Sun, 2002)
spermatogenesis
testis sheath
(Tang and Sun, 2002)
Reporter: P{Kr-GAL4.5.1}
Stage
Tissue/Position (including subcellular localization)
Reference
third instar larval stage
photoreceptor cell of Bolwig organ
(Yamanaka et al., 2013)
embryonic/larval optic lobe
(Yamanaka et al., 2013)
High-Throughput Expression Data
Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\Kr in GBrowse 2
RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region
Bulk Downloads
RNA-Seq RPKM values for all genes
Reference
See Gelbart and Emmert, 2013 for analysis details and data files for all genes.
Developmental Proteome: Life Cycle
Developmental Proteome: Embryogenesis
External Data and Images
Linkouts
BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
Flygut - An atlas of the Drosophila adult midgut
Images
FlyExpress - Embryonic expression images (BDGP data)
  • Stages(s) 4-6
  • Stages(s) 7-8
  • Stages(s) 9-10
  • Stages(s) 11-12
  • Stages(s) 13-16
FlyBase Wiki
Image Based Resources
Alleles, Insertions, Transgenic Constructs, and Aberrations
Classical and Insertion Alleles ( 39 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 51 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Kr
Transgenic constructs containing regulatory region of Kr
Aberrations (Deficiencies and Duplications) ( 15 )
Alleles Representing Disease-Implicated Variants
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
lethal
lethal (with Df(2R)Kr10)
lethal | recessive
lethal - all die during embryonic stage | gap | recessive
lethal - all die during embryonic stage | recessive
viable, with Dcr-2UAS.cDa, Scer\GAL4elav.PLu
viable, with Scer\GAL4Mef2.PR
viable, with Scer\GAL4pnr-MD237
viable, with Scer\GAL4Pxn.PS
viable, with Scer\GAL4tin.CΔ4
Other Phenotypes
Allele
abnormal neuroanatomy, with Scer\GAL4sca-4512
abnormal neuroanatomy | heat sensitive
abnormal neuroanatomy | heat sensitive, with Scer\GAL4pros.PMG
decreased occurrence of cell division | embryonic stage 13, with Kr1
decreased occurrence of cell division | embryonic stage 13, with KrmCD
decreased occurrence of cell division | embryonic stage 14, with Kr1
decreased occurrence of cell division | embryonic stage 14, with KrmCD
increased occurrence of cell division | embryonic stage 16, with Scer\GAL4elav-C155
melanotic mass phenotype, with Scer\GAL4Pxn.PS
some die during embryonic stage | recessive
visible
visible, with Scer\GAL4Bx-MS1096
visible, with Scer\GAL4ey.PH
visible, with Scer\GAL4hs.2sev
visible | dominant
Phenotype manifest in
Allele
A1-7 lateral transverse muscle cell, with Scer\GAL4how-24B
abdomen
abdominal 1 ventral denticle belt
abdominal 2 ventral denticle belt
abdominal 3 ventral denticle belt
abdominal 4 ventral denticle belt
abdominal segment 6
abdominal segment 7
adult cuticle | ectopic
adult mesothoracic segment
embryonic/first instar larval cuticle
embryonic/larval anterior Malpighian tubule
embryonic/larval hemocyte | third instar larval stage, with Scer\GAL4Pxn.PS
embryonic/larval hindgut | embryonic stage
embryonic/larval Malpighian tubule
embryonic/larval midgut | embryonic stage | posterior
embryonic/larval posterior Malpighian tubule
embryonic/larval ureter
embryonic abdomen | anterior
embryonic abdominal segment
embryonic abdominal segment 1
embryonic abdominal segment 2
embryonic abdominal segment 3
embryonic abdominal segment 4
embryonic abdominal segment 5
embryonic ganglion mother cell | embryonic stage 14, with Kr1
embryonic ganglion mother cell | embryonic stage 14, with KrmCD
embryonic ganglion mother cell | embryonic stage 16, with Scer\GAL4elav-C155
embryonic Malpighian tubule
embryonic mesothoracic segment
embryonic metathoracic segment
embryonic midgut constriction
embryonic neuroblast | embryonic stage 13, with Kr1
embryonic neuroblast | embryonic stage 13, with KrmCD
embryonic neuroblast | embryonic stage 14, with Kr1
embryonic neuroblast | embryonic stage 14, with KrmCD
embryonic neuroblast | embryonic stage 16, with Scer\GAL4elav-C155
embryonic prothoracic segment
embryonic segment
EW1 neuron, with Kr1
EW1 neuron, with KrmCD
EW2 neuron, with Kr1
EW2 neuron, with KrmCD
EW2 neuron | ectopic, with Scer\GAL4en-e16E
EW3 neuron, with Kr1
EW3 neuron, with KrmCD
eye
eye, with Scer\GAL4ey.PH
eye, with Scer\GAL4hs.2sev
eye disc
ganglion mother cell GMC1-1a, with Kr1
ganglion mother cell GMC1-1a, with KrmCD
glial cell
lamellocyte | third instar larval stage, with Scer\GAL4Pxn.PS
larval abdomen | anterior | embryonic stage
larval abdominal segment
larval abdominal segmental nerve, with Scer\GAL4ftz.ng
larval longitudinal connective
larval mesothoracic segment
larval thorax
larval thorax | embryonic stage
Malpighian tubule
Malpighian tubule primordium
Malpighian tubule primordium, with Scer\GAL4ptc-559.1
Malpighian tubule tip cell
Malpighian tubule tip cell, with Scer\GAL4da.G32
mesoderm, with Kr1
mesoderm, with KrmCD
mesoderm, with Scer\GAL4how-24B, Scer\GAL4twi.PB
mesothoracic segment
metathoracic segment
mitotic domain 1 | extended germ band stage
muscle cell of A1-7 dorsal acute muscle 1, with Kr1
muscle cell of A1-7 dorsal acute muscle 1, with KrmCD
muscle cell of A1-7 lateral longitudinal muscle 1, with Kr1
muscle cell of A1-7 lateral longitudinal muscle 1, with KrmCD
muscle cell of A1-7 lateral transverse muscle 2, with Kr1
muscle cell of A1-7 lateral transverse muscle 2, with KrmCD
muscle cell of A1-7 lateral transverse muscle 4, with Kr1
muscle cell of A1-7 lateral transverse muscle 4, with KrmCD
muscle cell of A1-7 ventral oblique muscle 2, with Kr1
muscle cell of A1-7 ventral oblique muscle 2, with KrmCD
muscle cell of abdominal 2 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of abdominal 3 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of abdominal 4 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of abdominal 5 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of abdominal 6 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of abdominal 7 ventral acute muscle 1, with Scer\GAL4how-24B, Scer\GAL4twi.PB
muscle cell of ventral acute muscle 2, with Kr1
muscle cell of ventral acute muscle 2, with KrmCD
muscle cell of ventral oblique muscle 5, with Kr1
muscle cell of ventral oblique muscle 5, with KrmCD
neuroblast NB7-1, with Kr1
neuroblast NB7-1, with KrmCD
neuroblast NB7-3, with Scer\GAL4en-e16E
ommatidium
ommatidium, with Scer\GAL4hs.2sev
retina | ventral, with Scer\GAL4ey.PH
rhabdomere
RP neuron, with Scer\GAL4ftz.ng
RP neuron, with Scer\GAL4how-24B
serotonergic neuron
symmetrical commissure
thorax
U3 neuron | ectopic, with Scer\GAL4en-e16E
U3 neuron | ectopic, with Scer\GAL4sca-4512
U3 neuron | ectopic | heat sensitive
U3 neuron | ectopic | heat sensitive, with Scer\GAL4pros.PMG
wing, with Scer\GAL4Bx-MS1096
Orthologs
Downloads
Download All DIOPT Orthologs
Human Orthologs (via DIOPT v8.0)
Homo sapiens (Human) (9)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Hsap\BCL6
3 of 15
Yes
Yes
0  
Hsap\BCL6B
1 of 15
No
Yes
Hsap\E4F1
1 of 15
No
Yes
Hsap\SNAI1
1 of 15
No
No
Hsap\ZFP64
1 of 15
No
No
Hsap\ZNF131
1 of 15
No
Yes
Hsap\ZNF662
1 of 15
No
No
Hsap\ZNF692
1 of 15
No
Yes
Hsap\ZNF707
1 of 15
No
No
Model Organism Orthologs (via DIOPT v8.0)
Mus musculus (laboratory mouse) (8)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Source
Compara
eggNOG
Hieranoid
Homologene
Inparanoid
Isobase
OMA
OrthoDB
OrthoFinder
OrthoInspector
orthoMCL
Panther
Phylome
RoundUp
TreeFam
ZFIN
Alignment
Complementation?
Transgene?
Mmus\Bcl6
3 of 15
Yes
Yes
Mmus\Bcl6b
2 of 15
No
Yes
Mmus\AI854703
1 of 15
No
Yes
Mmus\E4f1
1 of 15
No
Yes
Mmus\Sp5
1 of 15
No
No
Mmus\Zfp410
1 of 15
No
Yes
Mmus\Zfp652
1 of 15
No
No
Mmus\Zfp711
1 of 15
No
Yes
Rattus norvegicus (Norway rat) (7)
Rnor\Bcl6
2 of 13
Yes
Yes
Rnor\E4f1
1 of 13
No
Yes
Rnor\Gfi1b
1 of 13
No
No
Rnor\RGD1311188
1 of 13
No
Yes
Rnor\Snai1
1 of 13
No
No
Rnor\Sp5
1 of 13
No
No
Rnor\Zfp652
1 of 13
No
No
Xenopus tropicalis (Western clawed frog) (1)
Xtro\bcl6
3 of 12
Yes
Yes
Danio rerio (Zebrafish) (9)
Drer\bcl6aa
2 of 15
Yes
Yes
Drer\bcl6ab
2 of 15
Yes
Yes
Drer\LOC103909987
1 of 15
No
Yes
Drer\si:ch211-284e13.5
1 of 15
No
Yes
Drer\si:dkeyp-69b9.6
1 of 15
No
No
Drer\zgc:66443
1 of 15
No
Yes
Drer\zgc:113886
1 of 15
No
No
Drer\znf367
1 of 15
No
Yes
Drer\znf865
1 of 15
No
No
Caenorhabditis elegans (Nematode, roundworm) (3)
Cele\B0310.2
1 of 15
Yes
Yes
Cele\lsl-1
1 of 15
Yes
Yes
Cele\ztf-6
1 of 15
Yes
Yes
Arabidopsis thaliana (thale-cress) (1)
Atha\AT3G29340
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (5)
Scer\AZF1
1 of 15
Yes
No
Scer\CMR3
1 of 15
Yes
No
Scer\STP3
1 of 15
Yes
Yes
Scer\STP4
1 of 15
Yes
Yes
Scer\YPR015C
1 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Other Organism Orthologs (via OrthoDB)
Paralogs
Downloads
Download All DIOPT Paralogs
Paralogs (via DIOPT v8.0)
Drosophila melanogaster (Fruit fly) (36)
Gene Symbol
Score
Source
Compara
eggNOG
Homologene
Inparanoid
Isobase
OrthoDB
OrthoFinder
orthoMCL
Panther
RoundUp
Alignment
CG14655
2 of 10
Clamp
2 of 10
sens
2 of 10
Aef1
1 of 10
Boh1
1 of 10
Cf2
1 of 10
CG2129
1 of 10
CG3407
1 of 10
CG6654
1 of 10
CG8319
1 of 10
CG8944
1 of 10
CG10147
1 of 10
CG10631
1 of 10
CG10959
1 of 10
CG11695
1 of 10
CG11696
1 of 10
CG11902
1 of 10
CG12236
1 of 10
CG12391
1 of 10
CG17359
1 of 10
CG17568
1 of 10
CG17612
1 of 10
CG18262
1 of 10
CG31388
1 of 10
CG31441
1 of 10
CTCF
1 of 10
hkb
1 of 10
Kr-h1
1 of 10
odd
1 of 10
Paris
1 of 10
pita
1 of 10
sna
1 of 10
su(Hw)
1 of 10
sug
1 of 10
topi
1 of 10
Zif
1 of 10
Human Disease Associations
FlyBase Human Disease Model Reports
    Disease Model Summary Ribbon
    Disease Ontology (DO) Annotations
    Models Based on Experimental Evidence ( 0 )
    Allele
    Disease
    Evidence
    References
    Potential Models Based on Orthology ( 0 )
    Human Ortholog
    Disease
    Evidence
    References
    Modifiers Based on Experimental Evidence ( 1 )
    Allele
    Disease
    Interaction
    References
    Kr2
    ameliorates  acute myeloid leukemia
    modeled by Hsap\RUNX1::Hsap\RUNX1T1UAS.cSa
    (Sinenko et al., 2010)
    Disease Associations of Human Orthologs (via DIOPT v8.0 and OMIM)
    Note that ortholog calls supported by only 1 or 2 algorithms (DIOPT score < 3) are not shown.
    Homo sapiens (Human)
    Gene name
    Score
    OMIM
    OMIM Phenotype
    DO term
    Complementation?
    Transgene?
    BCL6; BCL6 transcription repressor
    3 of 15
    109565
         
        Functional Complementation Data
        Functional complementation data is computed by FlyBase using a combination of the orthology data obtained from DIOPT and OrthoDB and the allele-level genetic interaction data curated from the literature.
        Interactions
        Summary of Physical Interactions
        Summary of Genetic Interactions
        esyN Network Diagram
        esyN Network Key:
        Suppression
        Enhancement
        View in Interactions Browser
        Please look at the allele data for full details of the genetic interactions
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Kr
        suppressible
        bun
        (Abrell et al., 2000)
        Kr
        suppressible
        caps
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        enhanceable
        caps
        (Abrell and Jackle, 2001)
        Kr
        enhanceable
        CCT8
        (Abrell et al., 2000)
        Kr
        enhanceable
        chinmo
        (Abrell et al., 2000)
        Kr
        enhanceable
        crol
        (Abrell et al., 2000)
        Kr
        enhanceable
        dpp
        (Abrell et al., 2000)
        Kr
        suppressible
        Eip75B
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        suppressible
        emc
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        enhanceable
        ex
        (Abrell et al., 2000)
        Kr
        enhanceable
        Fer2LCH
        (Carrera et al., 1998)
        Kr
        enhanceable
        GstS1
        (Abrell et al., 2000)
        Kr|svp
        suppressible
        hb
        (Kanai et al., 2005)
        Kr
        suppressible
        kra
        (Carrera et al., 1998)
        Kr
        enhanceable
        Kr-h1
        (Abrell et al., 2000)
        Kr
        enhanceable
        l(2)04548
        (Abrell et al., 2000)
        Kr
        enhanceable
        l(2)08492
        (Abrell et al., 2000)
        Kr
        suppressible
        l(3)05967
        (Carrera et al., 1998)
        Kr
        suppressible
        l(3)j2C3
        (Carrera et al., 1998)
        Kr
        suppressible
        l(3)neo3
        (Carrera et al., 1998)
        Kr
        suppressible
        l(3)neo39
        (Carrera et al., 1998)
        Kr
        suppressible
        l(3)neo47
        (Carrera et al., 1998)
        Kr
        enhanceable
        Mi-2
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        enhanceable
        osa
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        enhanceable
        piwi
        (Gangaraju et al., 2011)
        Kr
        suppressible
        Psa
        (Carrera et al., 1998)
        Kr
        enhanceable
        Rab6
        (Abrell et al., 2000)
        Kr
        suppressible
        rept
        (Carrera et al., 1998)
        Kr
        enhanceable
        S
        (Abrell et al., 2000)
        Kr
        suppressible
        Sap130
        (Carrera et al., 1998)
        Kr
        suppressible
        Sc2
        (Abrell et al., 2000, Carrera et al., 1998)
        Kr
        enhanceable
        Sema2a
        (Abrell et al., 2000)
        Kr
        enhanceable
        srp
        (Uddin et al., 2011)
        Kr
        enhanceable
        Stip1
        (Gangaraju et al., 2011)
        Kr
        suppressible
        sty
        (Abrell et al., 2000)
        Kr
        enhanceable
        sty
        (Abrell et al., 2000)
        Kr
        suppressible
        Vha55
        (Abrell et al., 2000, Carrera et al., 1998)
        Starting gene(s)
        Interaction type
        Interacting gene(s)
        Reference
        Acsl
        enhanceable
        Kr
        (Zhang et al., 2011)
        Bx
        enhanceable
        Kr
        (Bronstein et al., 2010)
        caps
        enhanceable
        Kr
        (Abrell and Jackle, 2001)
        fru
        enhanceable
        Kr
        (Goto et al., 2011)
        hb
        suppressible
        Kr
        (Wu et al., 2001)
        nej
        enhanceable
        Kr
        (Anderson et al., 2005)
        srp
        enhanceable
        Kr
        (Uddin et al., 2011)
        svp
        enhanceable
        Kr
        (Kanai et al., 2005)
        External Data
        Linkouts
        BioGRID - A database of protein and genetic interactions.
        DroID - A comprehensive database of gene and protein interactions.
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Pathways
        Signaling Pathways (FlyBase)
        Metabolic Pathways
        External Data
        Linkouts
        Genomic Location and Detailed Mapping Data
        Chromosome (arm)
        2R
        Recombination map
        2-107
        Cytogenetic map
        60F5-60F5
        Sequence location
        FlyBase Computed Cytological Location
        Cytogenetic map
        Evidence for location
        60F5-60F5
        Limits computationally determined from genome sequence between P{EP}CG2790EP412&P{EP}CG3776EP835 and P{EP}zipEP856&P{PZ}l(2)1048103263
        Experimentally Determined Cytological Location
        Cytogenetic map
        Notes
        References
        60F1-60F5
        (determined by in situ hybridisation)
        (Spradling et al., 1999)
        60F1-60F5
        (BDGP Project Members, 1994-1999)
        60F3-60F3
        (determined by in situ hybridisation)
        (Knipple et al., 1985)
        Experimentally Determined Recombination Data
        Location

        2-107

        (FlyBase, 2016)

        2-111.8

        (Comeron et al., 2012)

        2-107.6

        (Tearle and Nusslein-Volhard, 1987, Ives, 1967)
        Left of (cM)
        Right of (cM)
        (Ives, 1967)
        (Ives, 1967)
        Notes

        Mapped using KrIf-1.

        (Ives, 1967)
        Stocks and Reagents
        Stocks (567)
        Genomic Clones (9)
         

        Please Note FlyBase no longer curates genomic clone accessions so this list may not be complete

        cDNA Clones (72)
         

        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
        Drosophila Genomics Resource Center cDNA clones

        For each fully sequenced cDNA the DGRC maintains various forms of the cDNA (e.g tagged or untagged) in several different host vectors for subsequent cloning and expression in Drosophila and Drosophila cell lines.

        cDNA Clones, End Sequenced (ESTs)
        BDGP DGC clones
        Other clones
        RNAi and Array Information
        Linkouts
        DRSC - Results frm RNAi screens
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        Antibody Information
        Laboratory Generated Antibodies
         

        polyclonal

        (Kumar et al., 2015, Dubuis et al., 2013, Li et al., 2008, Gaul et al., 1987)
        Commercially Available Antibodies
         
        Other Information
        Relationship to Other Genes
        Source for database identify of

        Source for identity of: Kr CG3340

        (Zhong, 1999.11)
        Source for database merge of
        Additional comments
        Other Comments

        Haploinsufficient locus (not associated with strong haplolethality or haplosterility).

        (Cook et al., 2012)

        DNA-protein interactions: genome-wide binding profile assayed for Kr protein in 0-12 hr embryos; see mE1_TFBS_Kr collection report.

        (Negre et al., 2011)

        The Kr product negatively regulates dac expression in the embryonic head.

        (Anderson et al., 2006)

        Kr interacts with caps in establishing the proper axonal pathway of SNb including the RP5 axons.

        (Abrell and Jackle, 2001)

        hb and Kr control early born temporal identity in neuroblast cell lineages.

        (Isshiki et al., 2001)

        Kr is activated by peb in the amnioserosa.

        (Lamka and Lipshitz, 1999)

        CtBP mediates transcriptional repression by the kni, Kr and sna products in the Drosophila embryo.

        (Nibu et al., 1998)

        Kr is found to have three specific regions within the coding sequence that are highly conserved during Drosophila speciation.

        (Hanna-Rose et al., 1997)

        KrT95D is a putative Kr target gene. Expression patterns of KrT95D and Kr suggest that Kr activity is necessary to activate KrT95D expression in the VO5 muscle precursors, consistent with recent results indicating that Kr activity is necessary for specification of a subset of muscles and their proper innervation during embryogenesis.

        (Hartmann and Jäckle, 1997)

        Kr activity is required to maintain ko expression in specific muscles in the embryo.

        (Hartmann et al., 1997)

        Kr is not required in the mesoderm for the segregation of a normal pattern of muscle founder cells, or to initiate patterns of gene expression in these founders. It is required for the maintenance of normal patterns of gene expression in the precursors that the founders form, and for the acquisition of proper muscle identity during embryogenesis. Gain and loss of Kr expression in sibling founder cells is sufficient to switch the cells, and the muscle they give rise to, between alternative cell fates.

        (Ruiz-Gomez et al., 1997)

        The "If" mutation is probably allelic to Kr.

        (Baker, 1996.3.7)

        The Kr and sna proteins function as short range repressors, which can mediate either quenching or direct repression of a transcription complex, depending on the location of repressor sites. Local quenching and dominant repression require close linkage of the repressor with either upstream activators or the transcription complex.

        (Gray and Levine, 1996)

        Kr is expressed in neural precursor cells, neurons and glial cells at different stages of neurogenesis and Kr mutants develop aberrant peripheral and central nervous systems. Phenotypic analysis of rescued embryos (by a Kr minigene) indicates that Kr expression in the nervous system is functionally required for establishing particular neural and glial fates.

        (Romani et al., 1996)

        The CD2 939bp cis acting control element in the Kr regulatory region has been analysed using Ecol\lacZ reporter gene constructs.

        (Carrera and Jäckle, 1995)

        Kr monomer (the activator) interacts with TfIIB, Kr dimers do not (FBrf0082559). Kr dimers engage in protein-protein interaction with TfIIEβ (FBrf0082559). Kr monomer-dimer transition alters Krs ability to interact with the general transcription machinery (FBrf0082559).

        (Roberts and Green, 1995)

        Transcription factors hb and kni can associate with Kr in vitro and they interact functionally with Kr-dependent target gene expression mediated by a single Kr-binding site close to an heterologous promoter in Schneider cells.

        (Sauer and Jäckle, 1995)

        Kr-dependent control of transcription involves functional interactions with components of the basal RNA polymerase II transcription machinery.

        (Sauer et al., 1995)

        The anterior boundaries of h stripes 5 and 6 are set by Kr.

        (Langeland et al., 1994)

        The region of Kr important for transcriptional repression has been fine mapped and is defined by a minimal 31 amino acid motif rich in Ala and Gln residues.

        (Licht et al., 1994)

        Comparisons of early development to that in other insects have revealed conservation of some aspects of development, as well as differences that may explain variations in early patterning events.

        (Patel, 1994)

        hb is autonomously capable of activating the target gene Kr at low concentrations and repressing it at high concentrations.

        (Schulz and Tautz, 1994)

        Ectopic ttk expression has no effect on expression of Kr and hb.

        (Brown and Wu, 1993)

        Expression of prd depends on activation by gap gene hb, Kr, kni and gt products. Primary pair rule gene products act primarily in subsequent modulation rather than activation of prd stripes. Factors activating prd expression in the pair rule mode interact with those activating it along the dorso-ventral axis.

        (Gutjahr et al., 1993)

        The role of Kr in the regulation of run mRNA expression in the early embryo has been investigated.

        (Klingler and Gergen, 1993)

        Examination of conserved tracts in the D.virilis and D.melanogaster h gene promoters identified potential ftz-f1, Kr and gt binding sites. Kr and gt products establish the anterior and posterior borders of h stripe 5, respectively, through spatial repression.

        (Langeland and Carroll, 1993)

        The Kr gene product selectively represses transcription activated by a glutamine rich activator but not by an acidic activator. This indicates that the Kr protein quenches transcription due to specific protein-protein interactions between Kr protein and part of the transcriptional apparatus mediating stimulation by a glutamine-rich activator.

        (Licht et al., 1993)

        Phenotypic rescue of Kr mutants by P-element mediated transformation of a Kr minigene demonstrates that the Kr gene product is required for nervous system development and its activity is driven by the neuro-specific enhancer elements which are lacking in the transgenic embryos.

        (Romani et al., 1993)

        Kr gene product is able to form homodimers through sequences located within the C terminus, termed C64KR sequences: when fused to separated functional parts of the yeast transcription factor GAL4 they reconstituted a functional transcriptional activator on dimerization in vivo. Results of in vitro experiments suggest that Kr monomer product is a transcriptional activator, whereas at higher concentrations it forms a homodimer that acts as a repressor acting on the same target sites as recognised by the activator.

        (Sauer and Jackle, 1993)

        Mutations affect eye morphology.

        (Baker et al., 1992)

        In vitro footprinting of bacterially expressed kni and tll protein found one strong kni binding site and seven tll binding sites in a 730 bp fragment of the Kr promoter. Binding studies demonstrate that each of the proteins can bind but their binding is mutually exclusive. Ecol\lacZ reporter gene constructs carrying 16 bp fragment of the overlapping kni and bcd protein binding sites demonstrated that expression mediated by the 16 bp element is dependent on bcd activity, kni represses expression by competitive binding.

        (Hoch et al., 1992)

        DNaseI footprinting analysis reveals core histones His2A, His2B, His3 and His4 (but not His1) bind to the Kr minimal enhancer element in a periodic manner.

        (Kerrigan and Kadonaga, 1992)

        Kr acts downstream of ct in the Malpighian tubule regulatory pathway. Kr activity is required for cad expression in the tubules.

        (Liu and Jack, 1992)

        Kr activity required for neurons to differentiate into Bolwig organs and for the fasciculation of the Bolwig nerve.

        (Schmucker et al., 1992)

        The gene products of bcd, hb, Kr and gt all bind within the 480bp region that is necessary and sufficient for the expression of eve stripe 2. Forming the posterior border of the stripe involves a delicate balance between Kr repressor and bcd activator.

        (Small et al., 1992)

        Orthologs present in M.domestica, S.coprophila, P.cinerea, A.mellifera, T.castaneum, E.plebejus, A.salina, P.phalangoides, C.salei and L.forficatus.

        (Sommer et al., 1992)

        Sequence alignments of orthologous fragments of hb, Kr and sna from a variety of arthropods and other phyla show that amino acid differences are not normally correlated with evolutionary distance between respective species. Amino acids directly involved in DNA binding are the most conserved, and binding specificity of a hb finger from different species is not changed.

        (Sommer et al., 1992)

        A plasmid containing a minimal Kr promoter has been used in in vitro transcription assays to study TfIIB activity.

        (Wampler and Kadonaga, 1992)

        Kr has a direct negative effect on gt expression.

        (Eldon and Pirrotta, 1991)

        Ecol\lacZ reporter gene constructs carrying Kr promoter deletions have identified a 142bp core sequence (the Kr730 element) within the cis-acting control element (CD1) which mediates gene expression in a central region of the embryo in response to hb and bcd activities.

        (Hoch et al., 1991)

        The expression pattern of a number of Kr-Ecol\lacZ fusion constructs has been studied, to identify regulatory elements controlling Kr expression.

        (Jacob et al., 1991)

        Transcriptional analysis of Kr deletion constructs has identified a 44bp fragment from -31bp to +13bp with significant promoter activity.

        (Kerrigan et al., 1991)

        Kr is a strong repressor of gt in the embryo.

        (Kraut and Levine, 1991)

        It has been shown that Kr and gt are expressed in complementary, non-overlapping sets of cells in the early embryo. Interactions between the two genes have been studied.

        (Kraut and Levine, 1991)

        Mutations in zygotic gap gene Kr interact with RpII140wimp.

        (Parkhurst and Ish-Horowicz, 1991)

        Mutant Kr embryos do not alter the expression of the Ubx bx region enhancer element, BRE.

        (Qian et al., 1991)

        Low amounts of Kr expression lead to transcriptional activation, whereas high amounts result in repression. Distinct portions of Kr protein, other than the DNA-binding domain, are required for gene activation and repression, suggesting that Kr itself can act as a concentration- dependent positive and negative regulator of transcription.

        (Sauer and Jackle, 1991)

        Kr protein directly regulates the expression of eve stripe 2 expression by DNA binding to the stripe 2 promoter element.

        (Small et al., 1991)

        Kr- embryos show an expansion of the posterior eve stripe 2 border.

        (Stanojevic et al., 1991)

        Zygotically active locus involved in the terminal developmental program in the embryo.

        (Strecker et al., 1991)

        Kr is a transcriptional repressor.

        (Zuo et al., 1991)

        Kr mutants exhibit deletions of the thorax and anterior abdomen.

        (Gaul and Jäckle, 1990)

        The regulatory relationships of Kr with respect to its role in the development of the Malpighian tubules differs from its interactions with the segmentation process.

        (Gaul and Weigel, 1990)

        Ecol\lacZ reporter gene has been used to examine the function of the cis regulatory elements of the Kr gene.

        (Hoch et al., 1990)

        Ubx, Kr and eve expression are altered in fs(1)h mutant embryos. Defects in the segmental organisation in fs(1)h-deficient progeny are mediated primarily, but not exclusively, through a restriction in the Kr expression domain.

        (Huang and Dawid, 1990)

        The effect of hb protein concentration on the expression of kni and Kr in the embryo has been studied.

        (Hulskamp et al., 1990)

        Characterisation of the crude RNA polymerase II transcription system using transcription initiation of the Kr promoter.

        (Kadonaga, 1990)

        The transcriptional repression function of Kr maps to an alanine-rich amino terminal region of the protein (between amino acids 26 to 110).

        (Licht et al., 1990)

        The expression patterns of Kr and kni demonstrate the proteins form overlapping concentration gradients that generate the periodic pair-rule expression pattern.

        (Pankratz et al., 1990)

        Kr exhibits a homeotic function in addition to its role as a segmentation gene and is involved in separating hindgut and Malpighian tubule cells and in the elongation process as well.

        (Harbecke and Janning, 1989)

        An investigation of the role of gap genes in expression from Ubx and Antp promoters in the blastoderm embryo reveals that a unique combination of gap genes and pair rule genes is required for their initial activation.

        (Irish et al., 1989)

        Mutations in Kr alter gt expression in the posterior of the embryo.

        (Mohler et al., 1989)

        Expression of kni-Ecol\lacZ regulatory fusion constructs in mutant embryos shows that kni expression is enhanced by Kr activity.

        (Pankratz et al., 1989)

        The on/off periodicity of the pair-rule gene eve involves the interaction of the hb and Kr proteins with defined eve promoter elements.

        (Stanojevic et al., 1989)

        Genetic analysis demonstrates that the effect of the gap gene product Kr on homeotic gene expression in the visceral mesoderm is indirect and mediated by the genes that establish parasegment borders, eve and ftz.

        (Tremml and Bienz, 1989)

        Kr activity is required for the establishment of the Antp T3 domain. Kr is involved in restricting Abd-B products within the A8--A9 domain.

        (Harding and Levine, 1988)

        Mutant embryos exhibit a slight increase in the number of Dfd expressing cells in ventral and lateral positions.

        (Jack et al., 1988)

        Involved in functions related to that of tll.

        (Strecker et al., 1988)

        The wild-type allele of Kruppel controls the development of the thoracic and abdominal segments of Drosophila; homozygous mutants show a gap in the larval pattern in these regions (Nusslein-Volhard and Wieschaus, 1980; Knipple et al., 1987). Kr/+ adult sometimes has thoracic malformation; a leg or a wing may be absent; penetrance low. Heterozygous larvae show small defects in denticle bands of thorax and abdomen as do heterozygous deficiencies for Kr; penetrance about 80%. Homozygotes are embryonic lethal. Kr1 homozygotes exhibit shortened germ band of but three to four segments with three to four tracheal pits; visible beginning at 7h of embryogenesis; head and gnathal segments apparently normal. At later stages only three to four abdominal and thoracic segments clearly visible; normal telson and segments 8 and 7 followed by enlarged sixth, a rudimentary fifth and apparently mirror image sixth segment. Weak and intermediate mutant alleles lack the mirror-image duplications (Gaul and Jackle, 1987b). Ventral chain of ganglia disconnected; tracheal system defective; Malpighian tubules missing; salivary glands normal. Homozygotes for hypomorphic alleles display more nearly complete segmentation. Homozygous Minute+ Kr clones develop normally in all parts of adult cuticle of M/+ flies. Metamorphic potential of Kr/Kr embryos cultured in female abdomens restricted in that wing-disc-derived structures not observed. Germline clones of homozygous Kr cells capable of normal oogenesis; no maternal effect of Kr+ observed. Requirement for Kr+ function apparently restricted to early embryogenesis. Kr affects ftz producing abnormal intensity and spacing of ftz stripes in thorax and anterior abdomen (Carroll and Scott, 1986).

        Origin and Etymology
        Discoverer

        Graber.

        Etymology
        Identification
        External Crossreferences and Linkouts ( 45 )
        Sequence Crossreferences
        NCBI Gene - Gene integrates information from a wide range of species. A record may include nomenclature, Reference Sequences (RefSeqs), maps, pathways, variations, phenotypes, and links to genome-, phenotype-, and locus-specific resources worldwide.
        GenBank Nucleotide - A collection of sequences from several sources, including GenBank, RefSeq, TPA, and PDB.
        GenBank Protein - A collection of sequences from several sources, including translations from annotated coding regions in GenBank, RefSeq and TPA, as well as records from SwissProt, PIR, PRF, and PDB.
        RefSeq - A comprehensive, integrated, non-redundant, well-annotated set of reference sequences including genomic, transcript, and protein.
        UniProt/GCRP - The gene-centric reference proteome (GCRP) provides a 1:1 mapping between genes and UniProt accessions in which a single 'canonical' isoform represents the product(s) of each protein-coding gene.
        UniProt/Swiss-Prot - Manually annotated and reviewed records of protein sequence and functional information
        UniProt/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information
        Other crossreferences
        AlphaFold DB - AlphaFold provides open access to protein structure predictions for the human proteome and other key proteins of interest, to accelerate scientific research.
        BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
        Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
        DRscDB - A single-cell RNA-seq resource for data mining and data comparison across species
        EMBL-EBI Single Cell Expression Atlas - Single cell expression across species
        Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
        FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
        FlyAtlas2 - A Drosophila melanogaster expression atlas with RNA-Seq, miRNA-Seq and sex-specific data
        Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns
        Flygut - An atlas of the Drosophila adult midgut
        GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents
        iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)
        InterPro - A database of protein families, domains and functional sites
        KEGG Genes - Molecular building blocks of life in the genomic space.
        MARRVEL_MODEL - MARRVEL (model organism gene)
        modMine - A data warehouse for the modENCODE project
        SignaLink - A signaling pathway resource with multi-layered regulatory networks.
        Linkouts
        ApoDroso - Functional genomic database for photoreceptor development, survival and function
        BioGRID - A database of protein and genetic interactions.
        DroID - A comprehensive database of gene and protein interactions.
        DRSC - Results frm RNAi screens
        FlyCyc Genes - Genes from a BioCyc PGDB for Dmel
        FlyMine - An integrated database for Drosophila genomics
        Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
        MIST (genetic) - An integrated Molecular Interaction Database
        MIST (protein-protein) - An integrated Molecular Interaction Database
        Synonyms and Secondary IDs (16)
        Reported As
        Symbol Synonym
        CG3340
        (Bronstein et al., 2010, Ni et al., 2010.12.1, Keleman et al., 2009.8.5, Liu et al., 2009, Mohammad et al., 2009, Perkins et al., 2009, Kreiman, 2004, Zhang et al., 2002)
        Dm-Kr
        (Diaz-Cuadros et al., 2021, Brent et al., 2007)
        If
        (Sollars and Ruden, 2000, Sollars and Ruden, 1998, Bhadra et al., 1997, Vosshall and Young, 1995)
        KR
        (Dresch et al., 2016, Pettie et al., 2016, Peng et al., 2015, Drewell et al., 2014, Duque et al., 2014, Cheng et al., 2013, McKay and Lieb, 2013, Paris et al., 2013, Stringham et al., 2013, Fisher et al., 2012, Bradley et al., 2010, Lusk and Eisen, 2010, modENCODE Consortium et al., 2010, Ho et al., 2009, MacArthur et al., 2009, Li et al., 2008)
        Kr
        (Hafer et al., 2022, Singh et al., 2022, Banisch et al., 2021, Bargiela and Artero, 2021, Duk et al., 2021, Fukaya, 2021, Han et al., 2021, Irizarry and Stathopoulos, 2021, Kvon et al., 2021, Ma et al., 2021, Malin and Desplan, 2021, Nässel, 2021, Parra-Peralbo et al., 2021, Rossi et al., 2021, Sood et al., 2021, Tkačik and Gregor, 2021, Zmojdzian and Jagla, 2021, Cohen et al., 2020, Díaz-de-la-Peña et al., 2020, Huang et al., 2020, Jaeger and Verd, 2020, Lammers et al., 2020, Liu et al., 2020, Lopes et al., 2020, Maurange, 2020, Myasnikova and Spirov, 2020, Poovathumkadavil and Jagla, 2020, Soluri et al., 2020, Yaghmaeian Salmani and Thor, 2020, Yang et al., 2020, Barr et al., 2019, Bozek et al., 2019, Brunet Avalos et al., 2019, Kulik et al., 2019, Oberst et al., 2019, Ramon-Cañellas et al., 2019, Sapar and Han, 2019, Scholes et al., 2019, Shokri et al., 2019, Verd et al., 2019, Zhornikova et al., 2019, Basu et al., 2018, Bischof et al., 2018, Bothma et al., 2018, Drechsler et al., 2018, Haines and Eisen, 2018, Mortensen et al., 2018, Myasnikova and Spirov, 2018, Myasnikova and Spirov, 2018, Bataillé et al., 2017, Chertkova et al., 2017, Doe, 2017, Goyal et al., 2017, Gursky et al., 2017, Huang et al., 2017, Karaiskos et al., 2017, Reichert, 2017, Stratmann and Thor, 2017, Transgenic RNAi Project members, 2017-, Werner et al., 2017, Ali-Murthy and Kornberg, 2016, Altenhein et al., 2016, El-Sherif and Levine, 2016, Fukaya et al., 2016, Hoermann et al., 2016, Ma et al., 2016, Mishra et al., 2016, Pinto-Teixeira et al., 2016, Schwartz et al., 2016, Stratmann et al., 2016, Vincent et al., 2016, Yasugi and Nishimura, 2016, Baëza et al., 2015, Briscoe and Small, 2015, Dequéant et al., 2015, Duque and Sinha, 2015, Gula and Samsonov, 2015, Kim et al., 2015, Kozlov et al., 2015, Kumar et al., 2015, Liu and Ma, 2015, Nadimpalli et al., 2015, Sawala et al., 2015, Schertel et al., 2015, Tikhonov et al., 2015, Tkačik et al., 2015, Ugrankar et al., 2015, Villaverde et al., 2015, Boyle et al., 2014, Ciglar et al., 2014, Dobi et al., 2014, Ferrero et al., 2014, Harumoto et al., 2014, Jiang and Singh, 2014, Krotov et al., 2014, Mannervik, 2014, Marjoram et al., 2014, Martinez et al., 2014, Myasnikova and Kozlov, 2014, Palsson et al., 2014, Slattery et al., 2014, Victorsen and White, 2014.5.13, Aleksic et al., 2013, Becker et al., 2013, Combs and Eisen, 2013, Denholm, 2013, Hartmann et al., 2013, Kim et al., 2013, Little et al., 2013, Liu and Ma, 2013, McKay and Lieb, 2013, Mishra et al., 2013, Samee and Sinha, 2013, Saunders et al., 2013, Spirov and Holloway, 2013, Sung et al., 2013, Surkova et al., 2013, Umulis and Othmer, 2013, Webber et al., 2013, Amrute-Nayak and Bullock, 2012, Aswani et al., 2012, Boukhatmi et al., 2012, Brody et al., 2012, Chen et al., 2012, Cook et al., 2012, Crombach et al., 2012, Dottermusch-Heidel et al., 2012, Enriquez et al., 2012, He et al., 2012, Homem and Knoblich, 2012, Hou et al., 2012, Hurtado et al., 2012, Jaeger et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kao et al., 2012, Kim et al., 2012, Kozlov et al., 2012, Kvon et al., 2012, Liang et al., 2012, Nikulova et al., 2012, Patel et al., 2012, Perry et al., 2012, Tamari and Barkai, 2012, Touma et al., 2012, Ueda et al., 2012, Ajuria et al., 2011, Carrasco-Rando et al., 2011, Dobi et al., 2011, Fowlkes et al., 2011, Gangaraju et al., 2011, Goto et al., 2011, Gursky et al., 2011, Haralalka et al., 2011, Harrison et al., 2011, Juarez et al., 2011, Kaplan et al., 2011, Kim et al., 2011, Kuzin et al., 2011, Li et al., 2011, Nègre et al., 2011, Nien et al., 2011, Papatsenko and Levine, 2011, Perry et al., 2011, Pruteanu-Malinici et al., 2011, Tsurumi et al., 2011, Uddin et al., 2011, Vorwald-Denholtz and De Robertis, 2011, Zhang et al., 2011, Aerts et al., 2010, Aswani et al., 2010, Bauer et al., 2010, Bronstein et al., 2010, Figeac et al., 2010, He et al., 2010, Kao and Lee, 2010, Karlsson et al., 2010, Kitajima et al., 2010, Lennox and Stronach, 2010, Mace et al., 2010, Morton de Lachapelle and Bergmann, 2010, Müller et al., 2010, Nakajima et al., 2010, Porcher and Dostatni, 2010, Rauzi et al., 2010, Sawa, 2010, Sinenko et al., 2010, Sousa-Nunes et al., 2010, Stofanko et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Tran et al., 2010, Ashyraliyev et al., 2009, Bhuin and Roy, 2009, Butler et al., 2009, Campbell et al., 2009, Fomekong-Nanfack et al., 2009, Fomekong-Nanfack et al., 2009, Galindo et al., 2009, Guruharsha et al., 2009, He et al., 2009, He et al., 2009, Kozlov et al., 2009, Liu et al., 2009, Liu et al., 2009, Manu et al., 2009, Manu et al., 2009, Marco et al., 2009, Matyash et al., 2009, Meyer et al., 2009, Ochoa-Espinosa et al., 2009, Papatsenko et al., 2009, Pentek et al., 2009, Pisarev et al., 2009, Sellin et al., 2009, Tchuraev and Galimzyanov, 2009, Twombly et al., 2009, Weber et al., 2009, Zúñiga et al., 2009, Ashyraliyev et al., 2008, Beckett et al., 2008, Bornemann et al., 2008, Crocker and Erives, 2008, DeFalco et al., 2008, Fowlkes et al., 2008, Fujimoto et al., 2008, Geisbrecht et al., 2008, Gregor et al., 2008, Hare et al., 2008, Jennings et al., 2008, Joza et al., 2008, Miles et al., 2008, Noyes et al., 2008, Papatsenko and Levine, 2008, Sanders et al., 2008, Segal et al., 2008, Stofanko et al., 2008, Surkova et al., 2008, Surkova et al., 2008, Surkova et al., 2008, Tran and Doe, 2008, Tsuji et al., 2008, Yu and Small, 2008, Aerts et al., 2007, Beckett and Baylies, 2007, Bergmann et al., 2007, Estrada et al., 2007, Estrada et al., 2007, Haecker et al., 2007, Hatton-Ellis et al., 2007, Jaeger et al., 2007, Johnson et al., 2007, Junion et al., 2007, Kim et al., 2007, Komiyama and Luo, 2007, Link et al., 2007, Lott et al., 2007, Orian et al., 2007, Singh et al., 2007, Sprecher et al., 2007, Surkova et al., 2007, Tran and Doe, 2007, Wang et al., 2007, Wei et al., 2007, Xing et al., 2007, Anderson et al., 2006, Azevedo et al., 2006, Beckett and Baylies, 2006, Braendle and Flatt, 2006, Bullock et al., 2006, Grosskortenhaus et al., 2006, Jaeger and Reinitz, 2006, Janssens et al., 2006, Keranen et al., 2006, Luengo Hendriks et al., 2006, Perkins et al., 2006, Sandmann et al., 2006, Fujioka et al., 2005, Grosskortenhaus et al., 2005, Ishihara et al., 2005, Kanai et al., 2005, Karcavich and Doe, 2005, Odenwald, 2005, Peel et al., 2005, Stathopoulos and Levine, 2005, Demakov et al., 2004, Grad et al., 2004, Gurunathan et al., 2004, Kreiman, 2004, Clyde et al., 2003, Daulny et al., 2003, Nibu et al., 2003, Zeremski et al., 2003, Gim et al., 2001, Gursky et al., 2001, Jagla et al., 1999)
        Kru
        (Nuzhdin et al., 2010)
        Kruppel
        (Ciglar et al., 2014, Kim et al., 2013, Fakhouri et al., 2010, Pollard et al., 2007)
        kr
        (Li and Hidalgo, 2020, Mahmud et al., 2020, Crossman et al., 2018, Ghodsi et al., 2017, Cicin-Sain et al., 2015, Kozlov et al., 2015, Guilgur et al., 2014, Knowles and Biggin, 2013, Li and Gilmour, 2013, Markstein, 2013, McKay and Lieb, 2013, Losada-Pérez et al., 2010, Yassin et al., 2010, Kim et al., 2009, Shelton et al., 2009, Sandmann et al., 2007, Choksi et al., 2006, Lifanov et al., 2003, Spirov et al., 2001)
        Name Synonyms
        Irregular facets
        KRUPPEL
        (Pettie et al., 2016, Ho et al., 2009)
        Krpple
        (Sollars and Ruden, 2000)
        Krueppel
        (Mace et al., 2010)
        Kruppel
        (Loganathan et al., 2021, Meng and Heckscher, 2021, Chipman, 2020, Gheisari et al., 2020, Linde-Medina, 2020, Lee and Chen, 2019, Ramon-Cañellas et al., 2019, Zubair et al., 2019, Stratmann and Thor, 2017, Fukaya et al., 2016, Stratmann et al., 2016, Wieschaus and Nüsslein-Volhard, 2016, Boija and Mannervik, 2015, Cicin-Sain et al., 2015, Dequéant et al., 2015, Kozlov et al., 2015, Kumar et al., 2015, Peng et al., 2015, Rebeiz et al., 2015, Sawala et al., 2015, Shimaji et al., 2015, Tkačik et al., 2015, Vlisidou and Wood, 2015, Li et al., 2014, Pichaud, 2014, Slattery et al., 2014, Villar et al., 2014, Becker et al., 2013, McKay and Lieb, 2013, Spirov and Holloway, 2013, Surkova et al., 2013, Enriquez et al., 2012, Fisher et al., 2012, Homem and Knoblich, 2012, Hurtado et al., 2012, Jaeger et al., 2012, Kvon et al., 2012, Liang et al., 2012, Nikulova et al., 2012, Perry et al., 2012, Bieler et al., 2011, Davis et al., 2011, Dobi et al., 2011, Gursky et al., 2011, Kuzin et al., 2011, Lorbeck et al., 2011, Nègre et al., 2011, Papatsenko and Levine, 2011, Perry et al., 2011, Vorwald-Denholtz and De Robertis, 2011, Zhang et al., 2011, Borok et al., 2010, Bulchand et al., 2010, Figeac et al., 2010, Lennox and Stronach, 2010, Losada-Pérez et al., 2010, Lusk and Eisen, 2010, Morton de Lachapelle and Bergmann, 2010, Rauzi et al., 2010, Schreader et al., 2010, Sinenko et al., 2010, Sousa-Nunes et al., 2010, Zhang and Moret, 2010, Bertet et al., 2009, Bhuin and Roy, 2009, Butler et al., 2009, Campbell et al., 2009, Iovino et al., 2009, Kozlov et al., 2009, Liu et al., 2009, MacArthur et al., 2009, Manu et al., 2009, Manu et al., 2009, Ochoa-Espinosa et al., 2009, Pisarev et al., 2009, Twombly et al., 2009, Weber et al., 2009, Zamparo and Perkins, 2009, Arnosti et al., 2008, Beckett et al., 2008, Bornemann et al., 2008, Bosveld et al., 2008, DeFalco et al., 2008, Fujimoto et al., 2008, Hare et al., 2008, Ishihara and Shibata, 2008, Jennings et al., 2008, Lott et al., 2008, Miles et al., 2008, Sanders et al., 2008, Stofanko et al., 2008, Surkova et al., 2008, Yavatkar et al., 2008, Yu and Small, 2008, Arnosti et al., 2007, Estrada et al., 2007, Estrada et al., 2007, Haecker et al., 2007, Hatton-Ellis et al., 2007, Jaeger et al., 2007, Kim et al., 2007, Link et al., 2007, Lott et al., 2007, Newfeld and Johnson, 2007, Schafer et al., 2007, Sharma and Nirenberg, 2007, Singh et al., 2007, Wang et al., 2007, Xing et al., 2007, Anderson et al., 2006, Beckett and Baylies, 2006, Braendle and Flatt, 2006, Doe, 2006, Hallikas et al., 2006, Jaeger and Reinitz, 2006, Janssens et al., 2006, Jennings et al., 2006, Keranen et al., 2006, Ochoa-Espinosa and Small, 2006, Staudt et al., 2006, Veitia, 2006, Yucel and Small, 2006, Yucel and Small, 2006, Karcavich and Doe, 2005, Kulkarni and Arnosti, 2005, Odenwald, 2005, Shav-Tal and Singer, 2005, Chew et al., 2004, Demakov et al., 2004, MacArthur and Brookfield, 2004, Nibu et al., 2003, Zeremski et al., 2003, Gim et al., 2001, Gursky et al., 2001, Wilkie et al., 2001, Gonzalez-Gaitan and Jackle, 2000, Jagla et al., 1999)
        Kuppel
        (Sanders et al., 2008)
        Lruppel
        (Wong et al., 2008)
        krueppel
        (Kreiman, 2004)
        kruppel
        (Chetverina et al., 2021, Saunders and Howard, 2009, Joza et al., 2008, De Renzis et al., 2007, King-Jones and Thummel, 2005, Ambros, 2003, Galis et al., 2002)
        Secondary FlyBase IDs
        • FBgn0001251
        • FBgn0015732
        Datasets (2)
        Study focus (2)
        Experimental Role
        Project
        Project Type
        Title
        • bait_protein
        BDTNP_TFBS
        ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.
        • bait_protein
        modENCODE_regulation_TFs
        Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
        Study result (0)
        Result
        Result Type
        Title
        References (1,071)