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General Information
Symbol
Dmel\Kr
Species
D. melanogaster
Name
Kruppel
Annotation Symbol
CG3340
Feature Type
FlyBase ID
FBgn0001325
Gene Model Status
Stock Availability
Gene Snapshot
In progress.Contributions welcome.
Also Known As
If
Key Links
Genomic Location
Cytogenetic map
Sequence location
2R:25,226,611..25,231,394 [+]
Recombination map
2-107
Sequence
Other Genome Views
The following external sites may use different assemblies or annotations than FlyBase.
Function
GO Summary Ribbons
Protein Family (UniProt)
Belongs to the krueppel C2H2-type zinc-finger protein family. (P07247)
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 GBrowse view of Dmel\Kr or 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
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
Transcript Data
Annotated Transcripts
Name
FlyBase ID
RefSeq ID
Length (nt)
Assoc. CDS (aa)
FBtr0072449
2536
502
FBtr0333162
4411
502
Additional Transcript Data and Comments
Reported size (kB)
Comments
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name
FlyBase ID
Predicted MW (kDa)
Length (aa)
Theoretical pI
RefSeq ID
GenBank
FBpp0072351
54.7
502
7.47
FBpp0305365
54.7
502
7.47
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)
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
Gene Ontology (21 terms)
Molecular Function (3 terms)
Terms Based on Experimental Evidence (3 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Biological Process (17 terms)
Terms Based on Experimental Evidence (10 terms)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (8 terms)
CV Term
Evidence
References
Cellular Component (1 term)
Terms Based on Experimental Evidence (1 term)
CV Term
Evidence
References
Terms Based on Predictions or Assertions (0 terms)
Expression Data
Expression Summary Ribbons
Colored tiles in ribbon indicate that expression data has been curated by FlyBase for that anatomical location. Colorless tiles indicate that there is no curated data for that location.
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
in situ
Stage
Tissue/Position (including subcellular localization)
Reference
northern blot
Stage
Tissue/Position (including subcellular localization)
Reference
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.
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.
The Kr transcript is expressed in the early embryo in a central stripe spanning from 40% to 60% egg length.
No Kr protein is detected in nosbcd.3UTR embryos.
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.
Kr is expressed in the middle of the blastoderm embryo from approximately 39-57% egg length.
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.
The primary Kr transcript is most abundant in 2-5 hour embryos, but transcripts are detected through pupal stages at lower levels.
A 2.5kb Kr transcript is detected at high levels in 2-5 hour embryos.
Marker for
 
Subcellular Localization
CV Term
Polypeptide Expression
immunolocalization
Stage
Tissue/Position (including subcellular localization)
Reference
Additional Descriptive Data
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.
Kr is expressed in Bolwig organ precursors beginning in embryonic stage 12 and continuing through the larval stages.
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.
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.
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.
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".
Marker for
Subcellular Localization
CV Term
Evidence
References
Expression Deduced from Reporters
Reporter: P{1BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | 40-60% egg length

Comment: reference stages ~42-55% egg length

Reporter: P{1BSKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | 40-60% egg length

Comment: reference stages ~42-55% egg length

Reporter: P{4BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{4BSKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | 40-60% egg length

Comment: reference stages ~45-53% egg length

Reporter: P{8BKrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
organism | 40-60% egg length

Comment: Reference states slightly posterior to native Kr

Reporter: P{BΔNc0.7HZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{dPN5.4KrZ}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4-Kr.C}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GAL4-Kr.zen.8}
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}AG1
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{GawB}sl(2)AM1AM1
Stage
Tissue/Position (including subcellular localization)
Reference
Reporter: P{Kr-GAL4.5.1}
Stage
Tissue/Position (including subcellular localization)
Reference
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
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
FLIGHT - Cell culture data for RNAi and other high-throughput technologies
FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
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
Alleles, Insertions, and Transgenic Constructs
Classical and Insertion Alleles ( 38 )
For All Classical and Insertion Alleles Show
 
Other relevant insertions
Transgenic Constructs ( 47 )
For All Alleles Carried on Transgenic Constructs Show
Transgenic constructs containing/affecting coding region of Kr
Transgenic constructs containing regulatory region of Kr
Deletions and Duplications ( 15 )
Phenotypes
For more details about a specific phenotype click on the relevant allele symbol.
Lethality
Allele
Other Phenotypes
Allele
Phenotype manifest in
Allele
Orthologs
Human Orthologs (via DIOPT v7.1)
Homo sapiens (Human) (10)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
1 of 15
Yes
Yes
1 of 15
Yes
Yes
 
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
Yes
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Model Organism Orthologs (via DIOPT v7.1)
Mus musculus (laboratory mouse) (7)
Species\Gene Symbol
Score
Best Score
Best Reverse Score
Alignment
Complementation?
Transgene?
2 of 15
Yes
Yes
1 of 15
No
Yes
Mmus\Osr1
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
No
1 of 15
No
Yes
Rattus norvegicus (Norway rat) (7)
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
No
1 of 13
Yes
Yes
1 of 13
Yes
Yes
Xenopus tropicalis (Western clawed frog) (3)
1 of 12
Yes
Yes
1 of 12
Yes
No
1 of 12
Yes
No
Danio rerio (Zebrafish) (7)
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
No
1 of 15
Yes
Yes
1 of 15
Yes
No
Caenorhabditis elegans (Nematode, roundworm) (2)
1 of 15
Yes
Yes
1 of 15
Yes
No
Arabidopsis thaliana (thale-cress) (1)
1 of 9
Yes
No
Saccharomyces cerevisiae (Brewer's yeast) (2)
1 of 15
Yes
Yes
1 of 15
Yes
Yes
Schizosaccharomyces pombe (Fission yeast) (0)
No records found.
Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG091908BU )
Organism
Common Name
Gene
AAA Syntenic Ortholog
Multiple Dmel Genes in this Orthologous Group
Drosophila melanogaster
fruit fly
Drosophila suzukii
Spotted wing Drosophila
Drosophila simulans
Drosophila sechellia
Drosophila erecta
Drosophila yakuba
Drosophila ananassae
Drosophila pseudoobscura pseudoobscura
Drosophila persimilis
Drosophila willistoni
Drosophila virilis
Drosophila mojavensis
Drosophila grimshawi
Orthologs in non-Drosophila Dipterans (via OrthoDB v9.1) ( EOG09150BN5 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Musca domestica
House fly
Glossina morsitans
Tsetse fly
Lucilia cuprina
Australian sheep blowfly
Mayetiola destructor
Hessian fly
Mayetiola destructor
Hessian fly
Aedes aegypti
Yellow fever mosquito
Anopheles darlingi
American malaria mosquito
Anopheles gambiae
Malaria mosquito
Culex quinquefasciatus
Southern house mosquito
Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W0D4C )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Bombyx mori
Silkmoth
Danaus plexippus
Monarch butterfly
Heliconius melpomene
Postman butterfly
Apis florea
Little honeybee
Apis mellifera
Western honey bee
Bombus impatiens
Common eastern bumble bee
Bombus terrestris
Buff-tailed bumblebee
Linepithema humile
Argentine ant
Megachile rotundata
Alfalfa leafcutting bee
Nasonia vitripennis
Parasitic wasp
Dendroctonus ponderosae
Mountain pine beetle
Tribolium castaneum
Red flour beetle
Pediculus humanus
Human body louse
Rhodnius prolixus
Kissing bug
Cimex lectularius
Bed bug
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Acyrthosiphon pisum
Pea aphid
Zootermopsis nevadensis
Nevada dampwood termite
Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X0D24 )
Organism
Common Name
Gene
Multiple Dmel Genes in this Orthologous Group
Strigamia maritima
European centipede
Ixodes scapularis
Black-legged tick
Stegodyphus mimosarum
African social velvet spider
Tetranychus urticae
Two-spotted spider mite
Tetranychus urticae
Two-spotted spider mite
Daphnia pulex
Water flea
Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )
No non-Arthropod Metazoa orthologies identified
Paralogs
Paralogs (via DIOPT v7.1)
Drosophila melanogaster (Fruit fly) (45)
2 of 10
2 of 10
2 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
1 of 10
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
    Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
     
    Disease Associations of Human Orthologs (via DIOPT v7.1 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?
    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

    Please look at the allele data for full details of the genetic interactions
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    Starting gene(s)
    Interaction type
    Interacting gene(s)
    Reference
    suppressible
    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.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Pathways
    Gene Group - Pathway Membership (FlyBase)
    External Data
    Linkouts
    SignaLink - A signaling pathway resource with multi-layered regulatory networks.
    Genomic Location and Detailed Mapping Data
    Chromosome (arm)
    2R
    Recombination map
    2-107
    Cytogenetic map
    Sequence location
    2R:25,226,611..25,231,394 [+]
    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)
    60F3-60F3
    (determined by in situ hybridisation)
    Experimentally Determined Recombination Data
    Left of (cM)
    Right of (cM)
    Notes
    Mapped using KrIf-1.
    Stocks and Reagents
    Stocks (516)
    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 sequences
    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
    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
    Commercially Available Antibodies
     
    Other Information
    Relationship to Other Genes
    Source for database identify of
    Source for identity of: Kr CG3340
    Source for database merge of
    Additional comments
    Other Comments
    Haploinsufficient locus (not associated with strong haplolethality or haplosterility).
    DNA-protein interactions: genome-wide binding profile assayed for Kr protein in 0-12 hr embryos; see mE1_TFBS_Kr collection report.
    The Kr product negatively regulates dac expression in the embryonic head.
    Kr interacts with caps in establishing the proper axonal pathway of SNb including the RP5 axons.
    hb and Kr control early born temporal identity in neuroblast cell lineages.
    Kr is activated by peb in the amnioserosa.
    CtBP mediates transcriptional repression by the kni, Kr and sna products in the Drosophila embryo.
    Kr is found to have three specific regions within the coding sequence that are highly conserved during Drosophila speciation.
    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.
    Kr activity is required to maintain ko expression in specific muscles in the embryo.
    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.
    The "If" mutation is probably allelic to Kr.
    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.
    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.
    The CD2 939bp cis acting control element in the Kr regulatory region has been analysed using Ecol\lacZ reporter gene constructs.
    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).
    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.
    Kr-dependent control of transcription involves functional interactions with components of the basal RNA polymerase II transcription machinery.
    The anterior boundaries of h stripes 5 and 6 are set by Kr.
    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.
    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.
    hb is autonomously capable of activating the target gene Kr at low concentrations and repressing it at high concentrations.
    Ectopic ttk expression has no effect on expression of Kr and hb.
    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.
    The role of Kr in the regulation of run mRNA expression in the early embryo has been investigated.
    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.
    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.
    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.
    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.
    Mutations affect eye morphology.
    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.
    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.
    Kr acts downstream of ct in the Malpighian tubule regulatory pathway. Kr activity is required for cad expression in the tubules.
    Kr activity required for neurons to differentiate into Bolwig organs and for the fasciculation of the Bolwig nerve.
    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.
    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.
    A plasmid containing a minimal Kr promoter has been used in in vitro transcription assays to study TfIIB activity.
    Kr has a direct negative effect on gt expression.
    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.
    The expression pattern of a number of Kr-Ecol\lacZ fusion constructs has been studied, to identify regulatory elements controlling Kr expression.
    Transcriptional analysis of Kr deletion constructs has identified a 44bp fragment from -31bp to +13bp with significant promoter activity.
    Kr is a strong repressor of gt in the embryo.
    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.
    Mutations in zygotic gap gene Kr interact with RpII140wimp.
    Mutant Kr embryos do not alter the expression of the Ubx bx region enhancer element, BRE.
    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.
    Kr protein directly regulates the expression of eve stripe 2 expression by DNA binding to the stripe 2 promoter element.
    Kr- embryos show an expansion of the posterior eve stripe 2 border.
    Zygotically active locus involved in the terminal developmental program in the embryo.
    Kr is a transcriptional repressor.
    Kr mutants exhibit deletions of the thorax and anterior abdomen.
    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.
    Ecol\lacZ reporter gene has been used to examine the function of the cis regulatory elements of the Kr gene.
    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.
    The effect of hb protein concentration on the expression of kni and Kr in the embryo has been studied.
    Characterisation of the crude RNA polymerase II transcription system using transcription initiation of the Kr promoter.
    The transcriptional repression function of Kr maps to an alanine-rich amino terminal region of the protein (between amino acids 26 to 110).
    The expression patterns of Kr and kni demonstrate the proteins form overlapping concentration gradients that generate the periodic pair-rule expression pattern.
    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.
    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.
    Mutations in Kr alter gt expression in the posterior of the embryo.
    Expression of kni-Ecol\lacZ regulatory fusion constructs in mutant embryos shows that kni expression is enhanced by Kr activity.
    The on/off periodicity of the pair-rule gene eve involves the interaction of the hb and Kr proteins with defined eve promoter elements.
    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.
    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.
    Mutant embryos exhibit a slight increase in the number of Dfd expressing cells in ventral and lateral positions.
    Involved in functions related to that of tll.
    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 ( 39 )
    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/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
    BDGP expression data - Patterns of gene expression in Drosophila embryogenesis
    Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones
    Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.
    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.
    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
    FLIGHT - Cell culture data for RNAi and other high-throughput technologies
    FlyAtlas - Adult expression by tissue, using Affymetrix Dros2 array
    FlyMine - An integrated database for Drosophila genomics
    Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution
    InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.
    MIST (genetic) - An integrated Molecular Interaction Database
    MIST (protein-protein) - An integrated Molecular Interaction Database
    Synonyms and Secondary IDs (16)
    Reported As
    Symbol Synonym
    Kr
    (Bozek et al., 2019, Kulik et al., 2019, Ramon-Cañellas et al., 2019, Scholes et al., 2019, Shokri et al., 2019, Zhornikova et al., 2019, Basu et al., 2018, Bischof 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, 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, 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, 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, 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, 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)
    Name Synonyms
    Irregular facets
    Kruppel
    (Ramon-Cañellas 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)
    krueppel
    Secondary FlyBase IDs
    • FBgn0001251
    • FBgn0015732
    Datasets (2)
    Study focus (2)
    Experimental Role
    Project
    Project Type
    Title
    • bait_protein
    ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.
    • bait_protein
    Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.
    References (1,005)