The gene huckebein is referred to in FlyBase by the symbol Dmel\hkb (CG9768, FBgn0261434). It is a protein_coding_gene from Drosophila melanogaster. There is experimental evidence that it has the molecular function: transcription factor binding; sequence-specific DNA binding. There is experimental evidence that it is involved in the biological process: central nervous system development; tube morphogenesis; glial cell differentiation; lateral inhibition; cellular membrane organization; negative regulation of transcription from RNA polymerase II promoter; germ cell migration; endoderm development; salivary gland morphogenesis. 28 alleles are reported. The phenotypes of these alleles are annotated with: organ system; organ system subdivision; multicellular structure; portion of tissue; larval head segment; non-connected developing system; embryonic foregut; extended germ band embryo; acellular anatomical structure; integumentary specialisation. It has one annotated transcript and one annotated polypeptide. Protein features are: Zinc finger C2H2-type/integrase DNA-binding domain; Zinc finger, C2H2; Zinc finger, C2H2-like. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of extremely low expression. Peak expression observed within 00-12 hour embryonic stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval central nervous system. Comments on Affy2 ProbeSet: ProbeSet 1638586_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of hkb. Gene sequence location is 3R:170737..172368.
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FB2013_03
FB2013_02
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Expression of the terminal gap gene hkb is first detected in the anterior and posterior tips of the syncytial blastoderm. During gastrulation, hkb expression is shifted to a more ventral position so as to abut the invaginating ventral furrow. Later in gastrulation, the polar expression of hkb disappears. Instead, hkb transcript is present in the salivary gland placodes and in a metameric pattern in the developing CNS. hkb is expressed in the CNS through the rest of embryonic development. In hkb mutant embryos, the mesodermal and ectodermal primordia expand at the expense of the endoderm, indicating that hkb is required for endoderm development. When hkb is ectopically expressed in embryos, the expression of the central gap gene gt is reduced, and invagination of the ventral furrow is disrupted.
hkb transcript is first detected in the salivary gland primordia in the posterior-most cells and then subsequently in dorsal-posterior cells that are the first to invaginate. There is also concomitant expression in a small group of cells in the dorsal-anterior region. Expression is higher in the dorsal aspect of the primordia but there is low level expression in the ventral region of the placode. hkb transcripts decreased and disappeared as invagination proceeded.
hkb transcript disappears from the procephalic ectoderm by stage 11 and is restricted to neuroblasts derived from these regions. However, a lacZ reporter displays a longer perdurance in these regions. Expression in procephalic neuroblasts stage 9-11: tritocerebrum - d6, d8, v1; deuterocerebrum - d1-10, d12, d13, v1-v8; protocerebrum - cd2, cv7
Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: larval central nervous system.
[download data (TSV)]
Guide to FlyAtlas expression level colors
No expression (0 - 9.999)
Low expression (10 - 99.999)
Moderate expression (100 - 499.999)
High level expression (500 - 999.999)
Very high expression (>999.999)
Linear, scaled to maximum expression level
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
Linear, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
High
Linear, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
High
Very high
Linear, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
Very high
log, scaled to maximum expression level
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
log, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
High
log, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
High
Very high
log, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
114.85
Larval Midgut
1.7
Larval Hindgut
2
Larval Malpighian Tubules
4.5
Larval Fat Body
6.4
Larval Salivary Gland
6.2
Larval Trachea
3.475
Larval Carcass
3.175
Adult Head
2.8
Adult Eye
3.55
Adult Brain
6.5
Adult Thoracic-Abdominal Ganglion
61
Adult Crop
3.8
Adult Midgut
0.9
Adult Hindgut
1.9
Adult Malpighian Tubules
1.6
Adult Fat Body
5.7
Adult Salivary Gland
12.5
Adult Heart
2.25
Adult VirginFemale Spermatheca
7.2
Adult InseminatedFemale Spermatheca
6.2
Adult Ovary
1.4
Adult Testis
9.5
Adult Male Accessory Gland
6.8
Adult Carcass
5.5
Expression Level Scale
None
Low
Moderate
High
Very high
Heatmap
Tissue
Expression Level
Larval Central Nervous System
Larval Midgut
Larval Hindgut
Larval Malpighian Tubules
Larval Fat Body
Larval Salivary Gland
Larval Trachea
Larval Carcass
Adult Head
Adult Eye
Adult Brain
Adult Thoracic-Abdominal Ganglion
Adult Crop
Adult Midgut
Adult Hindgut
Adult Malpighian Tubules
Adult Fat Body
Adult Salivary Gland
Adult Heart
Adult VirginFemale Spermatheca
Adult InseminatedFemale Spermatheca
Adult Ovary
Adult Testis
Adult Male Accessory Gland
Adult Carcass
FlyAtlas Organ/Tissue Expression, larval vs. adult
Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of extremely low expression. Peak expression observed within 00-12 hour embryonic stages.
[download data (TSV)]
Please Note FlyBase no
longer curates genomic clone accessions so this list
may not be complete
cDNA Clones ( 51 )
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.
Mutants are isolated in an EMS mutagenesis screen to identify zygotic mutations affecting germ cell migration at discrete points during embryogenesis: mutants exhibit gut development defects.
cad acts in hindgut development through fog, fkh and wg, but does not play a role in activating tll, hkb, byn and bowl which are also required for proper hindgut development.
Repression of oc expression at the anterior terminus of the blastoderm is mediated by hkb and requires input from all three maternal morphogens that specify embryonic head development.
DiI has been used to trace the entire NB4-2 lineage in wild type and hkb mutant embryos. The number, position or type of neurons in the lineage are not affected, but hkb specifies aspects of CNS precursor identity required for motoneuron axon pathfinding.
hkb is necessary for endoderm development and its activity defines the spatial limits within the blastoderm embryo in which the germ layers are established.
twi, sna, hkb and tll gene products define the positions of the primordia of the germ layers and thereby the regions in which the blastoderm epithelium will invaginate.
Mesodermal fate is determined where sna and twi but not hkb are expressed. Anteriorly, hkb together with sna determines endodermal fate, and hkb together with twi and sna are required for foregut development. hkb is a good candidate for factor X and factor Y, necessary for setting the anterior and posterior borders of the ventral furrow.
An artificial bcd responder gene composed of three bcd consensus binding sites driving Ecol\lacZ is activated by bcd and repressed by tor. This repression does not require tll or hkb.
The activation and spatial limitation of tll and hkb expression in the posterior region of the embryo is critically dependent on tor activity. The spatial limitation of hkb and tll expression is not regulated by the "central gap genes" which are essential for the establishment of segmentation in the trunk of the embryo, and also does not involve mutual interactions between hkb and tll.
Shu and Li, 2013, Theor. Biol. Med. Model. 10(1): 11
A statistical thin-tail test of predicting regulatory regions in the Drosophila genome. [FBrf0221021]
Staller et al., 2013, Genetics 193(1): 51--61
Depleting Gene Activities in Early Drosophila Embryos with the "Maternal-Gal4-shRNA" System. [FBrf0220341]
Andrioli et al., 2012, Dev. Biol. 361(1): 177--185
Huckebein is part of a combinatorial repression code in the anterior blastoderm. [FBrf0216872]
Aswani et al., 2012, Methods Cell Biol. 110: 243--261
Nonparametric variable selection and modeling for spatial and temporal regulatory networks. [FBrf0217971]
Crombach et al., 2012, PLoS Comput. Biol. 8(7): e1002589
Efficient reverse-engineering of a developmental gene regulatory network. [FBrf0218873]
Kvon et al., 2012, Genes Dev. 26(9): 908--913
HOT regions function as patterned developmental enhancers and have a distinct cis-regulatory signature. [FBrf0218184]
Popkova et al., 2012, PLoS Genet. 8(12): e1003159
Polycomb controls gliogenesis by regulating the transient expression of the gcm/glide fate determinant. [FBrf0220515]
Shu and Li, 2012, Comput. Biol. Med. 42(9): 935--941
A statistical fat-tail test of predicting regulatory regions in the Drosophila genome. [FBrf0219304]
Turki-Judeh and Courey, 2012, PLoS ONE 7(2): e30610
The unconserved groucho central region is essential for viability and modulates target gene specificity. [FBrf0217447]
Ajuria et al., 2011, Development 138(5): 915--924
Capicua DNA-binding sites are general response elements for RTK signaling in Drosophila. [FBrf0212975]
Bieler et al., 2011, Biophys. J. 101(2): 287--296
Whole-embryo modeling of early segmentation in Drosophila identifies robust and fragile expression domains. [FBrf0214381]
Dunipace et al., 2011, Development 138(18): 4075--4084
Complex interactions between cis-regulatory modules in native conformation are critical for Drosophila snail expression. [FBrf0214809]
Fowlkes et al., 2011, PLoS Genet. 7(10): e1002346
A conserved developmental patterning network produces quantitatively different output in multiple species of Drosophila. [FBrf0216666]
Gabilondo et al., 2011, Mech. Dev. 128(3-4): 208--221
A targeted genetic screen identifies crucial players in the specification of the Drosophila abdominal Capaergic neurons. [FBrf0213290]
Garcia and Stathopoulos, 2011, PLoS ONE 6(12): e29172
Lateral gene expression in Drosophila early embryos is supported by grainyhead-mediated activation and tiers of dorsally-localized repression. [FBrf0217082]
Grillo et al., 2011, Genetics 187(2): 513--521
Control of Germline torso Expression by the BTB/POZ Domain Protein Pipsqueak Is Required for Embryonic Terminal Patterning in Drosophila. [FBrf0213010]
Helman et al., 2011, Curr. Biol. 21(13): 1102--1110
Phosphorylation of Groucho Mediates RTK Feedback Inhibition and Prolonged Pathway Target Gene Expression. [FBrf0214255]
Kaplan et al., 2011, PLoS Genet. 7(2): e1001290
Quantitative Models of the Mechanisms That Control Genome-Wide Patterns of Transcription Factor Binding during Early Drosophila Development. [FBrf0213025]
Kim et al., 2011, Dev. Cell 20(6): 880--887
Gene regulation by MAPK substrate competition. [FBrf0213872]
Kim et al., 2011, Mol. Syst. Biol. 7: 467
Substrate-dependent control of MAPK phosphorylation in vivo. [FBrf0212870]
Kuzin et al., 2011, Mech. Dev. 128(3-4): 165--177
Functional analysis of conserved sequences within a temporally restricted neural precursor cell enhancer. [FBrf0213297]
Li et al., 2011, Genome Biol. 12(4): R34
The role of chromatin accessibility in directing the widespread, overlapping patterns of Drosophila transcription factor binding. [FBrf0216471]
Nègre et al., 2011, Nature 471(7339): 527--531
A cis-regulatory map of the Drosophila genome. [FBrf0213303]
Perry et al., 2011, Proc. Natl. Acad. Sci. U.S.A. 108(33): 13570--13575
Multiple enhancers ensure precision of gap gene-expression patterns in the Drosophila embryo. [FBrf0214713]
Seong et al., 2011, Biogerontology 12(2): 93--107
Genome-wide analysis of low-dose irradiated male Drosophila melanogaster with extended longevity. [FBrf0213307]
Singh et al., 2011, Dev. Biol. 352(1): 104--115
The Bin3 RNA methyltransferase is required for repression of caudal translation in the Drosophila embryo. [FBrf0213172]
Tio et al., 2011, PLoS ONE 6(11): e26879
Asymmetric cell division and notch signaling specify dopaminergic neurons in Drosophila. [FBrf0216651]
Uddin et al., 2011, Int. J. Dev. Biol. 55(10-11-12): 945--952
The drumstick gene acts cell-non-autonomously and triggers specification of the small intestine in the Drosophila hindgut. [FBrf0217187]
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