FlyBase Gene Report: Dmel\hkb

General Information

Symbol

Dmel\hkb

Species

D. melanogaster

Name

huckebein

Annotation Symbol

CG9768

Feature Type

protein_coding_gene

FlyBase ID

FBgn0261434

Gene Model Status

Stock Availability

12 publicly available

Gene Snapshot

huckebein (hkb) is expressed in patches within the embryonic neuroectoderm and a subset of neuroblasts and their progeny, where it is required for proper neuronal specification and axon targeting. It is a terminal gap gene mediating the maternal terminal information at the posterior end of the blastoderm embryo. [Date last reviewed: 2019-03-07]

Other Summaries

Alliance
Auto summary
Gene Group
Interactive Fly

Also Known As

gurtelchen, gurt

Key Links

Genomic Location

Cytogenetic map

Sequence location

3R:4,345,015..4,346,646 [-]

Recombination map

3-47.1

RefSeq locus

NT_033777 REGION:4345015..4346646

Sequence

Get Decorated FASTA

Genomic Maps

GBrowse JBrowse

Other Genome Views

The following external sites may use different assemblies or annotations than FlyBase.

Function

GO Summary Ribbons

Gene Ontology (GO) Annotations (20 terms)

Molecular Function (4 terms)

Terms Based on Experimental Evidence (2 terms)

CV Term

Evidence

References

sequence-specific DNA binding

inferred from direct assay

(Brown et al., 2005)

transcription factor binding

inferred from physical interaction with FLYBASE:gro; FB:FBgn0001139

(Goldstein et al., 1999)

Terms Based on Predictions or Assertions (2 terms)

CV Term

Evidence

References

DNA-binding transcription factor activity, RNA polymerase II-specific

inferred from biological aspect of ancestor with PANTHER:PTN004619861

(assigned by GO_Central )

(Gaudet et al., 2011)

transcription regulatory region sequence-specific DNA binding

inferred from biological aspect of ancestor with PANTHER:PTN004619861

(assigned by GO_Central )

(Gaudet et al., 2011)

Biological Process (15 terms)

Terms Based on Experimental Evidence (8 terms)

CV Term

Evidence

References

central nervous system development

inferred from genetic interaction with FLYBASE:eg; FB:FBgn0000560

(Dittrich et al., 1997)

endoderm development

inferred from mutant phenotype

(Wolfstetter et al., 2009)

germ cell migration

inferred from mutant phenotype

(Moore et al., 1998)

glial cell differentiation

inferred from mutant phenotype

(De Iaco et al., 2006)

membrane organization

inferred from mutant phenotype

(Myat and Andrew, 2002)

negative regulation of transcription by RNA polymerase II

inferred from mutant phenotype

(Goldstein et al., 1999, Margolis et al., 1995)

salivary gland morphogenesis

inferred from mutant phenotype

(Myat and Andrew, 2002)

inferred from genetic interaction with FLYBASE:h; FB:FBgn0001168

(Myat and Andrew, 2002)

tube morphogenesis

inferred from mutant phenotype

(Myat and Andrew, 2002)

Terms Based on Predictions or Assertions (8 terms)

CV Term

Evidence

References

cell fate specification

traceable author statement

(Murakami et al., 1999)

ectoderm and mesoderm interaction

non-traceable author statement

(Stainier, 2002)

endoderm formation

traceable author statement

(Stainier, 2002)

germ cell migration

traceable author statement

(Wylie, 1999)

neuroblast fate determination

traceable author statement

(Skeath and Thor, 2003)

positive regulation of transcription by RNA polymerase II

inferred from biological aspect of ancestor with PANTHER:PTN004619861

(assigned by GO_Central )

(Gaudet et al., 2011)

posterior midgut invagination

non-traceable author statement

(Rehorn et al., 1996)

salivary gland development

traceable author statement

(Sudarsan, 2003)

Cellular Component (1 term)

Terms Based on Experimental Evidence (1 term)

CV Term

Evidence

References

inferred from direct assay

(McDonald and Doe, 1997)

Terms Based on Predictions or Assertions (1 term)

CV Term

Evidence

References

inferred from biological aspect of ancestor with PANTHER:PTN004619861

(assigned by GO_Central )

(Gaudet et al., 2011)

Gene Group (FlyBase)

SP1/KLF TRANSCRIPTION FACTORS

C2H2 ZINC FINGER TRANSCRIPTION FACTORS

Protein Family (UniProt)

-

Protein Signatures (InterPro)

Summaries

Gene Group (FlyBase)

SP1/KLF TRANSCRIPTION FACTORS -: The Sp1/KLF (Sp1-like/Kruppel-like factors) family of zinc finger proteins are DNA binding proteins. They can act as activators or repressors and some members bind sequences within Polycomb Response Elements (PREs). (Adapted from FBrf0188187).
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).

Summary (Interactive Fly)

tf - zinc finger - terminal gap gene - limits the extent of mesodermal development - required for proper endoderm formation

Interactive Fly

Gene Model and Products

Number of Transcripts

1

Number of Unique Polypeptides

1

Please see the GBrowse view of Dmel\hkb or the JBrowse view of Dmel\hkb 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

Gene model reviewed during 5.40

Gene model reviewed during 5.47

Sequence Ontology: Class of Gene

(Wolfstetter et al., 2007)

protein_coding_gene

Transcript Data

Annotated Transcripts

Name

FlyBase ID

RefSeq ID

Length (nt)

Assoc. CDS (aa)

FBtr0078951

1567

297

Additional Transcript Data and Comments

Reported size (kB)

1.8 (northern blot)

(Bronner et al., 1994)

Comments

External Data

Crossreferences

Polypeptide Data

Annotated Polypeptides

Name

FlyBase ID

Predicted MW (kDa)

Length (aa)

Theoretical pI

RefSeq ID

GenBank

FBpp0078591

32.6

297

9.34

Polypeptides with Identical Sequences

There is only one protein coding transcript and one polypeptide associated with this gene

Additional Polypeptide Data and Comments

Reported size (kDa)

296 (aa)

(McDonald and Doe, 1997, Bronner et al., 1994)

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\hkb using the Feature Mapper tool.

External Data

Crossreferences

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

hkb_1

Linkouts

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

in situ

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 9 -- 11

tritocerebral neuroblast | subset

(Urbach and Technau, 2003)

protocerebral neuroblast | subset

(Urbach and Technau, 2003)

deutocerebral neuroblast | subset

(Urbach and Technau, 2003)

neuroblast NB2-2

(Chang et al., 2000)

neuroblast NB4-2

(Chang et al., 2000)

embryonic stage 4

organism | 0-20% egg length

Comment: ~0-15% egg length

(Bronner et al., 1994)

organism | 0-10% egg length

Comment: 0-11% egg length

(Gao et al., 1996)

organism | terminal expression pattern

(Gao et al., 1996)

organism | 90-100% egg length

(Gao et al., 1996)

embryonic stage 6 -- 8

ventral neurectoderm | restricted

(Bronner et al., 1994)

presumptive embryonic/larval central nervous system | precursor

(Bronner et al., 1994)

embryonic stage 9 -- 10

procephalic neurogenic region & embryonic intercalary segment | anterior | restricted

(Urbach and Technau, 2003)

procephalic neurogenic region & embryonic ocular segment | restricted

(Urbach and Technau, 2003)

procephalic neurogenic region & embryonic antennal segment | restricted

Comment: stripe of expression

(Urbach and Technau, 2003)

antennal primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

central brain primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

visual primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

dorsal head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

lateral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

ventral head epidermis primordium

Comment: reported as procephalic ectoderm primordium

(Fisher et al., 2012)

salivary gland body primordium

Comment: reported as salivary gland body specific anlage

(Fisher et al., 2012)

ventral nerve cord primordium

(Fisher et al., 2012)

embryonic stage 4 -- 5

organism | gap expression pattern

(Casali and Casanova, 2001)

embryonic stage 10 -- 11

neuroblast NB2-4

(Chang et al., 2000)

neuroblast NB4-4

(Chang et al., 2000)

neuroblast NB5-4

(Chang et al., 2000)

embryonic stage 9

stage 2 neuroblast

(Chang et al., 2000)

embryonic stage 10

stage 3 neuroblast

(Chang et al., 2000)

embryonic stage 11

stage 5 neuroblast

(Chang et al., 2000)

neuroblast NB2-1

(Chang et al., 2000)

neuroblast NB4-3

(Chang et al., 2000)

neuroblast NB5-5

(Chang et al., 2000)

neuroblast NB7-3

(Chang et al., 2000)

embryonic primordium of adult salivary gland | posterior

(Myat and Andrew, 2002)

embryonic primordium of adult salivary gland | anterior

(Myat and Andrew, 2002)

embryonic stage 1 -- 3

organism | anterior

(Fisher et al., 2012)

organism | posterior

(Fisher et al., 2012)

embryonic stage 4 -- 6

anlage in statu nascendi

(Fisher et al., 2012)

anterior endoderm anlage

Comment: anlage in statu nascendi

(Fisher et al., 2012)

central brain anlage in statu nascendi

(Fisher et al., 2012)

foregut anlage in statu nascendi

(Fisher et al., 2012)

posterior ectoderm anlage

Comment: 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)

visual anlage in statu nascendi

Comment: reported as procephalic ectoderm anlage in statu nascendi

(Fisher et al., 2012)

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)

Comment: reported as procephalic ectoderm anlage

(Fisher et al., 2012)

anterior endoderm anlage

(Fisher et al., 2012)

(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 11 -- 12

central brain primordium

(Fisher et al., 2012)

embryonic central brain glial cell

(Fisher et al., 2012)

salivary gland body primordium

(Fisher et al., 2012)

ventral nerve cord primordium

(Fisher et al., 2012)

embryonic stage 13 -- 16

embryonic brain

(Fisher et al., 2012)

embryonic central brain glial cell

(Fisher et al., 2012)

embryonic central brain neuron

(Fisher et al., 2012)

embryonic/larval nervous system

(Fisher et al., 2012)

ventral nerve cord

(Fisher et al., 2012)

embryonic stage 5 -- 6

posterior midgut primordium

(Hatton-Ellis et al., 2007)

Additional Descriptive Data

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

(Urbach and Technau, 2003)

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.

(Myat and Andrew, 2002)

hkb transcript is expressed in two domains in the cellular balstoderm, forming an anterior and a posterior cap.

(Gao et al., 1996)

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.

(Bronner et al., 1994)

Marker for

stage 3 neuroblast

(Chang et al., 2000)

Subcellular Localization

CV Term

Polypeptide Expression

immunolocalization

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 11

neuroblast NB1-1

(McDonald and Doe, 1997)

neuroblast NB7-3

Comment: late stage 11

(McDonald and Doe, 1997)

neuroblast NB5-4

(McDonald and Doe, 1997)

neuroblast NB4-2

(McDonald and Doe, 1997)

blastoderm stage

organism | 90-100% egg length

Comment: ~95-100% egg length

(McDonald and Doe, 1997)

organism | 0-10% egg length

Comment: ~0-5% egg length

(McDonald and Doe, 1997)

embryonic stage 9

neuroblast NB2-2

(McDonald and Doe, 1997)

embryonic stage 10

intermediate ventral neurectoderm | restricted

Comment: neuroblast row 1/2

(McDonald and Doe, 1997)

Additional Descriptive Data

At the blastoderm stage, hkb protein is expressed in the anterior and posterior tips of the embryo, and at stage 7, it is expressed in the anterior and posterior midgut primordia. Starting at stage 8, until late stage 11, hkb protein is detected in the neurectoderm. hkb protein appears first in a neurectodermal cluster, then in the neuroblasts which laminate from that position, and finally in the early progenitors of these neuroblasts. The exception is neuroblast NB1-1, which begins expressing hkb protein partway through its development. hkb protein is detected in the central nervous system until the end of stage 16. hkb protein expression pattern closely matches the hkb transcript expression pattern, but the protein persists slightly longer than the RNA.

(McDonald and Doe, 1997)

Marker for

Subcellular Localization

CV Term

Evidence

References

inferred from direct assay

(McDonald and Doe, 1997)

Expression Deduced from Reporters

Reporter: P{lacW}hkb^s5953

Stage

Tissue/Position (including subcellular localization)

Reference

embryonic stage 11

neuroblast NB2-4

(Doe, 1992)

neuroblast NB5-5

Comment: late stage 11

(Doe, 1992)

neuroblast NB5-4

(Doe, 1992)

embryonic stage 10 -- 11

neuroblast NB2-2

(Doe, 1992)

embryonic stage 9 -- 10

ventral neurectoderm

Comment: row 5

(Chu-LaGraff and Doe, 1993)

embryonic stage 9 -- 10

ventral neurectoderm

Comment: row 2/3

(Chu-LaGraff and Doe, 1993)

neuroblast NB2-3

(Chu-LaGraff and Doe, 1993)

embryonic stage

neuroblast of ventral nervous system | subset

(Higashijima et al., 1996)

embryonic stage 11

neuroblast NB4-4

(Doe, 1992)

neuroblast NB7-3

Comment: late stage 11

(Doe, 1992)

neuroblast NB4-3

Comment: late stage 11

(Doe, 1992)

neuroblast NB2-1

(Doe, 1992)

embryonic stage 10 -- 11

neuroblast NB1-1

Comment: reference states 5-5.5 hr AEL

(Bhat et al., 2000)

neuroblast NB2-2

Comment: reference states 5-5.5 hr AEL

(Bhat et al., 2000)

neuroblast NB2-5

Comment: reference states 5-5.5 hr AEL

(Bhat et al., 2000)

neuroblast NB4-2

Comment: reference states 5-5.5 hr AEL

(Bhat et al., 2000)

embryonic stage 17

A-subperineurial glial cell of abdomen | faint

(Bossing et al., 1996)

A-subperineurial glial cell of thorax | faint

(Bossing et al., 1996)

B-subperineurial glial cell of abdomen

(Bossing et al., 1996)

RP2 motor neuron

(Bossing et al., 1996)

larval serotonergic VNC neuron

(Lundell et al., 1996)

first instar larval stage

larval abdominal ganglion Leucokinin neuron

(Benito-Sipos et al., 2010)

neuroblast NB5-3 of abdomen

(Estacio-Gómez et al., 2013)

neuroblast NB3-5 of abdomen

(Estacio-Gómez et al., 2013)

adult abdominal ganglion Leucokinin neuron

(Estacio-Gómez et al., 2013)

embryonic stage 11 -- 16

B-subperineurial glial cell of abdomen

(Beckervordersandforth et al., 2008)

A-subperineurial glial cell

(Beckervordersandforth et al., 2008)

lateral ventral subperineurial glial cell

(Beckervordersandforth et al., 2008)

embryonic/larval neuron

(Beckervordersandforth et al., 2008)

ventral interface glial cell

(Beckervordersandforth et al., 2008)

lateral interface glial cell

(Beckervordersandforth et al., 2008)

High-Throughput Expression Data

Associated Tools

GBrowse - Visual display of RNA-Seq signals

View Dmel\hkb in GBrowse 2

RNA-Seq by Region - Search RNA-Seq expression levels by exon or genomic region

View exonic expression by developmental stage for Dmel\hkb

View exonic expression by tissue for Dmel\hkb

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

FBgn0261434

FLIGHT - Cell culture data for RNAi and other high-throughput technologies

FBgn0261434

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG9768

Images

Image Based Resources

Alleles, Insertions, and Transgenic Constructs

Classical and Insertion Alleles ( 17 )

For All Classical and Insertion Alleles Show

Other relevant insertions

Transgenic Constructs ( 12 )

For All Alleles Carried on Transgenic Constructs Show

Transgenic constructs containing/affecting coding region of hkb

Transgenic constructs containing regulatory region of hkb

Deletions and Duplications ( 12 )

Not disrupted in

(Irion and Leptin, 1999)

Phenotypes

For more details about a specific phenotype click on the relevant allele symbol.

Lethality

Allele

lethal, with Scer\GAL4^sca-4512

lethal | pupal stage, with Scer\GAL4^pnr-MD237

hkb^GD4139

lethal - all die before end of embryonic stage

partially lethal | male limited, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

partially lethal | male limited | cold sensitive, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

partially lethal - majority die | pupal stage, with Scer\GAL4^pnr-MD237

hkb^GD4139

Other Phenotypes

Allele

neuroanatomy defective

hkb¹

neuroanatomy defective | embryonic stage

hkb¹

some die during embryonic stage

visible, with Scer\GAL4^pnr-MD237

hkb^GD4139

visible | female limited, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

visible | female limited | cold sensitive, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

hkb^tBa

Phenotype manifest in

Allele

adult labral sense organ

hkb²

anterior midgut primordium

hkb^gurt

A-subperineurial glial cell

hkb²

B-subperineurial glial cell of abdomen

hkb²

central nervous system

hkb¹

cephalopharyngeal skeleton

chaeta | supernumerary, with Scer\GAL4^pnr-MD237

hkb^GD4139

circular visceral muscle fiber

hkb^gurt

hkb²

dorsopharyngeal organ

hkb²

embryonic/larval central nervous system

hkb¹

embryonic/larval esophagus | embryonic stage

hkb²

embryonic/larval labral nerve

hkb²

embryonic/larval midgut | anterior | embryonic stage

embryonic/larval midgut | embryonic stage

hkb^unspecified

embryonic/larval midgut | embryonic stage | posterior

embryonic/larval posterior spiracle

embryonic/larval salivary gland | embryonic stage

embryonic Malpighian tubule

hkb^gurt

embryonic visceral muscle

ganglion mother cell GMC4-2a

hkb^hs.PB

hkb^A

hkb²

lateral ventral subperineurial glial cell

hkb²

longitudinal connective, with Scer\GAL4^sca-4512

longitudinal visceral muscle primordium

hkb^XM9

Malpighian tubule primordium

hkb^unspecified

mesothoracic tergum, with Scer\GAL4^pnr-MD237

hkb^GD4139

hkb²

neuroblast NB4-2

pharyngeal monoscolopidial chordotonal organ

hkb²

presumptive embryonic salivary gland

hkb²

presumptive embryonic salivary gland, with Scer\GAL4^fkh.PH

hkb^UAS.cMa

RP2 motor neuron

hkb^hs.PB

RP2 neuron & axon

hkb¹

salivary gland common duct primordium

hkb^gurt

serotonergic neuron | embryonic stage

hkb⁶²

stomatogastric nervous system

subperineurial glial cell | supernumerary, with Scer\GAL4^sca-4512

hkb^UAS.cMa

ventral nerve cord commissure, with Scer\GAL4^sca-4512

visceral mesoderm

hkb^gurt

visceral mesoderm (with hkb²)

hkb^gurt

visceral mesoderm (with hkb^gurt)

hkb²

wing, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

wing | cold sensitive, with Scer\GAL4^Bx-MS1096

hkb^{UAS.ORF.GW.Tag:HA}

Orthologs

Downloads

Download All DIOPT Orthologs

Download All OrthoDB Orthologs

Human Orthologs (via DIOPT v8.0)

Homo sapiens (Human) (20)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

2 of 15

Yes

No

1 of 15

No

No

1 of 15

No

No

1 of 15

No

No

1 of 15

No

No

1 of 15

No

No

1 of 15

No

No

1 of 15

No

No

1

1 of 15

No

Yes

1 of 15

No

No

0

1 of 15

No

No

1 of 15

No

No

0

1 of 15

No

No

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

1 of 15

No

Yes

1 of 15

No

Yes

Model Organism Orthologs (via DIOPT v8.0)

Mus musculus (laboratory mouse) (11)

Species\Gene Symbol

Score

Best Score

Best Reverse Score

Source

Alignment

Complementation?

Transgene?

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1

1 of 15

Yes

No

1 of 15

Yes

Yes

Rattus norvegicus (Norway rat) (12)

1 of 13

Yes

No

1 of 13

Yes

No

Rnor\LOC102546394

1 of 13

Yes

Yes

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

No

1 of 13

Yes

No

1 of 13

Yes

No

1 of 13

Yes

No

Xenopus tropicalis (Western clawed frog) (4)

1 of 12

Yes

Yes

1 of 12

Yes

No

1 of 12

Yes

No

1 of 12

Yes

No

Danio rerio (Zebrafish) (18)

1 of 15

Yes

Yes

Drer\LOC101884382

1 of 15

Yes

Yes

Drer\LOC110437822

1 of 15

Yes

Yes

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

No

Drer\wu:fe14d05

1 of 15

Yes

Yes

Drer\zgc:113102

1 of 15

Yes

Yes

Caenorhabditis elegans (Nematode, roundworm) (7)

2 of 15

Yes

No

1 of 15

No

Yes

1 of 15

No

Yes

1 of 15

No

No

1 of 15

No

Yes

1 of 15

No

No

1 of 15

No

No

Arabidopsis thaliana (thale-cress) (0)

No records found.

Saccharomyces cerevisiae (Brewer's yeast) (3)

1 of 15

Yes

No

1 of 15

Yes

No

1 of 15

Yes

Yes

Schizosaccharomyces pombe (Fission yeast) (0)

No records found.

Orthologs in Drosophila Species (via OrthoDB v9.1) ( EOG09190D5D )

Organism

Common Name

Gene

AAA Syntenic Ortholog

Multiple Dmel Genes in this Orthologous Group

Drosophila suzukii

Spotted wing Drosophila

Dsuz\DS10_00007611

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) ( EOG09150CLW )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Musca domestica

House fly

Glossina morsitans

Tsetse fly

Gmor\GMOY003244

Lucilia cuprina

Australian sheep blowfly

Lcup\FF38_12818

Mayetiola destructor

Hessian fly

Mdes\Mdes000741

Aedes aegypti

Yellow fever mosquito

Aaeg\AAEL014006

Anopheles darlingi

American malaria mosquito

Adar\ADAC010229

Anopheles gambiae

Malaria mosquito

Agam\AGAP004517

Culex quinquefasciatus

Southern house mosquito

Cqui\CPIJ008311

Orthologs in non-Dipteran Insects (via OrthoDB v9.1) ( EOG090W08U6 )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Bombyx mori

Silkmoth

Bmor\BGIBMGA002238

Danaus plexippus

Monarch butterfly

Dple\DPOGS213224-PA

Heliconius melpomene

Postman butterfly

Hmel\HMEL016959

Apis florea

Little honeybee

Aflo\2749453396

Apis mellifera

Western honey bee

Amel\LOC100578561

Bombus impatiens

Common eastern bumble bee

Bombus terrestris

Buff-tailed bumblebee

Linepithema humile

Argentine ant

Nasonia vitripennis

Parasitic wasp

Nvit\Nasvi2EG005777

Dendroctonus ponderosae

Mountain pine beetle

Tribolium castaneum

Red flour beetle

Pediculus humanus

Human body louse

Phum\PHUM127400

Rhodnius prolixus

Kissing bug

Rpro\RPRC013233

Rhodnius prolixus

Kissing bug

Rpro\RPRC014216

Cimex lectularius

Bed bug

Clec\CLEC005464

Orthologs in non-Insect Arthropods (via OrthoDB v9.1) ( EOG090X08Q3 )

Organism

Common Name

Gene

Multiple Dmel Genes in this Orthologous Group

Strigamia maritima

European centipede

Smar\SMAR000785

Strigamia maritima

European centipede

Smar\SMAR005401

Ixodes scapularis

Black-legged tick

Isca\ISCW000054

Stegodyphus mimosarum

African social velvet spider

Smim\KFM74079.1

Tetranychus urticae

Two-spotted spider mite

Turt\tetur29g01340

Tetranychus urticae

Two-spotted spider mite

Turt\tetur02g06400

Tetranychus urticae

Two-spotted spider mite

Turt\tetur02g06440

Daphnia pulex

Water flea

Dpux\DAPPUDRAFT_331337

Orthologs in non-Arthropod Metazoa (via OrthoDB v9.1) ( None identified )

No non-Arthropod Metazoa orthologies identified

Paralogs

Downloads

Download All DIOPT Paralogs

Paralogs (via DIOPT v8.0)

Drosophila melanogaster (Fruit fly) (21)

Gene Symbol

Score

Source

Alignment

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 ( 0 )

Allele

Disease

Interaction

References

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?

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

esyN Network Diagram

Interactions Browser

View in Interactions Browser

Please see the Physical Interaction reports below for full details

protein-protein

Physical Interaction

Assay

References

pull down, tag visualisation, competition binding

(Kazemian et al., 2013)

two hybrid array

(Shokri et al., 2019)

two hybrid array

(Shokri et al., 2019)

two hybrid array

(Shokri et al., 2019)

two hybrid array

(Shokri et al., 2019)

pull down, tag visualisation

(Kazemian et al., 2013)

two hybrid, pull down, autoradiography

(Bejarano et al., 2007, Goldstein et al., 1999)

two hybrid array

(Shokri et al., 2019)

two hybrid array

(Shokri et al., 2019)

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

enhanceable

(De Iaco et al., 2006)

suppressible

(Myat and Andrew, 2002)

Starting gene(s)

Interaction type

Interacting gene(s)

Reference

suppressible

(Dittrich et al., 1997)

enhanceable

(De Iaco et al., 2006)

suppressible

(Popkova et al., 2012)

enhanceable

(Zhang et al., 2002)

suppressible

(Li and Li, 2003)

External Data

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0261434

InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.

40549

MIST (genetic) - An integrated Molecular Interaction Database

hkb

MIST (protein-protein) - An integrated Molecular Interaction Database

hkb

Pathways

Signaling Pathways (FlyBase)

Metabolic Pathways

External Data

Linkouts

KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.

MAPK signaling pathway - fly

Genomic Location and Detailed Mapping Data

Chromosome (arm)

3R

Recombination map

3-47.1

Cytogenetic map

Sequence location

3R:4,345,015..4,346,646 [-]

FlyBase Computed Cytological Location

Cytogenetic map

Evidence for location

82A4-82A4

Limits computationally determined from genome sequence between P{PZ}l(3)02733⁰²⁷³³ and P{EP}EP974

Experimentally Determined Cytological Location

Cytogenetic map

Notes

References

82A2-82A6

(Bronner, 1994.9.22)

82B3-82B3

(determined by in situ hybridisation)

(Doe, 1992)

Experimentally Determined Recombination Data

Location

3-47.1

(FlyBase, 2016)

3-44.1

(Comeron et al., 2012)

Left of (cM)

Right of (cM)

Notes

Stocks and Reagents

Stocks (12)

31976

y¹ v¹; P{TRiP.JF03410}attP2

2059

hkb^A321R1/TM3, Sb¹ Ser¹

83668

y¹ w^*; PBac{hkb-GFP.FPTB}VK00037

5457

hkb² p^p/TM3, p⁺ Sb¹

83597

y¹ sc^* v¹ sev²¹; P{TOE.GS05258}attP40

34736

y¹ sc^* v¹ sev²¹; P{TRiP.HMS01216}attP2/TM3, Sb¹

F003349

M{UAS-hkb.ORF-CC}ZH-21F

F004747

M{UAS-hkb.ORF-VN}ZH-86Fb

F000048

M{UAS-hkb.ORF.3xHA.GW}ZH-86Fb

106642

hkb^A321R1/TM3, Sb¹

108249

hkb² p^p/TM3, p⁺ Sb¹

v13779

w¹¹¹⁸ P{GD4139}v13779

Genomic Clones (10)

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

cDNA Clones (52)

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.

FBgn0261434

cDNA Clones, End Sequenced (ESTs)

BDGP DGC clones

RNAi and Array Information

Linkouts

DRSC - Results frm RNAi screens

FBgn0261434

GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents

40549

Antibody Information

Laboratory Generated Antibodies

polyclonal

(Ashyraliyev et al., 2009, McDonald and Doe, 1997)

Commercially Available Antibodies

Other Information

Relationship to Other Genes

Source for database identify of

Source for database merge of

Source for merge of: hkb gurt

(Wolfstetter et al., 2009)

Additional comments

Other Comments

DNA-protein interactions: genome-wide binding profile assayed for hkb protein in 0-12 hr embryos; see mE1_TFBS_hkb collection report.

(Negre et al., 2011)

hkb triggers gcm autoregulation via direct physical interaction.

(De Iaco et al., 2006)

hkb controls polarised cell shape change and apical membrane growth during salivary gland cell invagination in the embryo, via crb and klar.

(Myat and Andrew, 2002)

Internalisation of the salivary gland and salivary gland shape are altered in hkb mutant embryos.

(Myat and Andrew, 2000)

gro protein mediates repressor activity of the hkb protein.

(Goldstein et al., 1999)

hkb is required zygotically for the migration of the germ cells through the posterior midgut wall.

(Moore et al., 1998)

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.

(Moore et al., 1998)

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.

(Wu and Lengyel, 1998)

eg and hkb act in concert to determine serotonergic cell fate in the developing central nervous system.

(Dittrich et al., 1997)

The expression pattern of hkb protein during embryogenesis has been studied.

(McDonald and Doe, 1997)

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.

(Gao et al., 1996)

hkb is necessary and sufficient for the expression of eve in the identified neural NB4-2 lineage.

(Chu-LaGraff et al., 1995)

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.

(Chu-LaGraff et al., 1995)

hkb is involved in regulation of serotonin cell development and has a role in neuronal identity subsequent to formation of neuroblast 7-3.

(Lundell and Hirsh, 1995)

hkb is necessary for endoderm development and its activity defines the spatial limits within the blastoderm embryo in which the germ layers are established.

(Bronner et al., 1994)

hkb is a repressor of byn expression.

(Kispert et al., 1994)

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.

(Leptin, 1994)

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.

(Reuter and Leptin, 1994)

srp acts downstream of hkb to promote morphogenesis and differentiation of anterior and posterior midgut.

(Reuter, 1994)

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

(Klingler and Gergen, 1993)

In mutant embryos, which do not form endodermal derivatives, germ band retraction is properly executed.

(Reuter et al., 1993)

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.

(Ronchi et al., 1993)

Expression analysed in CNS study of neuroblasts and ganglion mother cells, using an enhancer trap to reveal the expression pattern.

(Doe, 1992)

In csw^- embryos hb remains as a posterior cap and the seventh ftz stripe expands posteriorly, both due to lack of hkb repressing activity.

(Perkins et al., 1992)

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.

(Bronner and Jäckle, 1991)

At the anterior end of the embryo tor effects on gt are mediated primarily through hkb.

(Eldon and Pirrotta, 1991)

Mutations in zygotic cardinal gene hkb do not interact with RpII140^wimp.

(Parkhurst and Ish-Horowicz, 1991)

The effect of hkb on hb and ftz expression has been studied.

(Casanova, 1990)

hkb represses Kr expression in the central segmentation domain and activates Kr expression in the posterior Malpighian tubule domain.

(Gaul and Weigel, 1990)

Origin and Etymology

Discoverer

Etymology

Identification

External Crossreferences and Linkouts ( 28 )

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.

40549

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/TrEMBL - Automatically annotated and unreviewed records of protein sequence and functional information

Q9VN10

Other crossreferences

BDGP expression data - Patterns of gene expression in Drosophila embryogenesis

FBgn0261434

Drosophila Genomics Resource Center - Drosophila Genomics Resource Center (DGRC) cDNA clones

FBgn0261434

Eukaryotic Promoter Database - A collection of databases of experimentally validated promoters for selected model organisms.

hkb_1

Fly-FISH - A database of Drosophila embryo and larvae mRNA localization patterns

CG9768

FlyMine - An integrated database for Drosophila genomics

FBgn0261434

GenomeRNAi - A database for cell-based and in vivo RNAi phenotypes and reagents

40549

iBeetle-Base - RNAi phenotypes in the red flour beetle (Tribolium castaneum)

TC010992

InterPro - A database of protein families, domains and functional sites

KEGG Genes - Molecular building blocks of life in the genomic space.

Dmel_CG9768

modMine - A data warehouse for the modENCODE project

FBgn0261434

SignaLink - A signaling pathway resource with multi-layered regulatory networks.

Q9VN10

Linkouts

DroID - A comprehensive database of gene and protein interactions.

FBgn0261434

DRSC - Results frm RNAi screens

FBgn0261434

FLIGHT - Cell culture data for RNAi and other high-throughput technologies

FBgn0261434

Interactive Fly - A cyberspace guide to Drosophila development and metazoan evolution

Interactive Fly

InterologFinder - Protein-protein interactions (PPI) from both known and predicted PPI data sets.

40549

KEGG Pathways - Wiring diagrams of molecular interactions, reactions and relations.

MAPK signaling pathway - fly

MIST (genetic) - An integrated Molecular Interaction Database

hkb

MIST (protein-protein) - An integrated Molecular Interaction Database

hkb

Synonyms and Secondary IDs (12)

Reported As

Symbol Synonym

5953

(Chu-LaGraff and Doe, 1993)

CG9768

(Ni et al., 2010.12.1, Perkins et al., 2009)

HKB

(Dresch et al., 2016, Cheng et al., 2013, Dilão and Muraro, 2010, MacArthur et al., 2009)

Hkb

(Chertkova et al., 2017, Altenhein et al., 2016, Villaverde et al., 2015, Ferrero et al., 2014, Slattery et al., 2014, Hartmann et al., 2013, Crombach et al., 2012, Noyes et al., 2008, Bejarano et al., 2007, Karcavich and Doe, 2005, McGovern et al., 2003, Nelson, 2003)

anon-82B3

gurt

(Wolfstetter et al., 2009, Wolfstetter et al., 2007)

hkb

(Johnson and Toettcher, 2019, Kwasnieski et al., 2019, Shokri et al., 2019, Bischof et al., 2018, Goyal et al., 2018, Forés et al., 2017, Goyal et al., 2017, Johnson et al., 2017, Karaiskos et al., 2017, Transgenic RNAi Project members, 2017-, Wolfstetter et al., 2017, Urbach et al., 2016, Cicin-Sain et al., 2015, Forés et al., 2015, Kozlov et al., 2015, model organism Encyclopedia of Regulatory Network (modERN) Project, 2015-, Nadimpalli et al., 2015, Schertel et al., 2015, Boyle et al., 2014, Jiang and Singh, 2014, Navarro et al., 2014, Aleksic et al., 2013, Combs and Eisen, 2013, Kim et al., 2013, Li and Gilmour, 2013, Peters et al., 2013, Saunders et al., 2013, Webber et al., 2013, Andrioli et al., 2012, Aswani et al., 2012, Crombach et al., 2012, Japanese National Institute of Genetics, 2012.5.21, Kvon et al., 2012, Popkova et al., 2012, Turki-Judeh and Courey, 2012, Ajuria et al., 2011, Dunipace et al., 2011, Fowlkes et al., 2011, Garcia and Stathopoulos, 2011, Grillo et al., 2011, Helman et al., 2011, Kaplan et al., 2011, Kim et al., 2011, Kim et al., 2011, Kuzin et al., 2011, Li et al., 2011, Nègre et al., 2011, Seong et al., 2011, Tio et al., 2011, Tsurumi et al., 2011, Uddin et al., 2011, Frise et al., 2010, Ismat et al., 2010, Kazemian et al., 2010, Kechris et al., 2010, The modENCODE Consortium, 2010, The modENCODE Consortium, 2010, Tipping et al., 2010, Ashyraliyev et al., 2009, Huh et al., 2009, Löhr et al., 2009, Moore et al., 2009, Wolfstetter et al., 2009, Anaka et al., 2008, Beckervordersandforth et al., 2008, Blanco and Gehring, 2008, Christensen et al., 2008.4.15, Cinnamon et al., 2008, Dougherty et al., 2008, Florence and Faller, 2008, Frise et al., 2008, Jennings et al., 2008, Miura et al., 2008, Sanders et al., 2008, Segal et al., 2008, Astigarraga et al., 2007, Coppey et al., 2007, Harris and Beckendorf, 2007, Hatton-Ellis et al., 2007, Sandmann et al., 2007, Zeitlinger et al., 2007, Abrams et al., 2006, Gresens and Cook, 2006.8.29, Jaeger and Reinitz, 2006, Jennings et al., 2006, Wang et al., 2006, Abrams and Andrew, 2005, Brown et al., 2005, McDonald et al., 2003, Myat and Andrew, 2002, Hayashi and Murakami, 2001, Myat et al., 2000, Lundell et al., 1996)

Name Synonyms

Huckebein

(Gilmour et al., 2017, Li et al., 2014, Mannervik, 2014, Slattery et al., 2014, Bieler et al., 2011, Kuzin et al., 2011, Dilão and Muraro, 2010, MacArthur et al., 2009, Jennings et al., 2008, Zinzen and Papatsenko, 2007, Karcavich and Doe, 2005, McDonald et al., 2003)

gurtelchen

(Wolfstetter et al., 2009, Wolfstetter et al., 2007)

huckbein

(Moore et al., 2009)

huckebein

(Urbach et al., 2016, Cicin-Sain et al., 2015, Forés et al., 2015, Kozlov et al., 2015, Peters et al., 2013, Shen et al., 2013, Andrioli et al., 2012, Crombach et al., 2012, Ajuria et al., 2011, Grillo et al., 2011, Helman et al., 2011, Singh et al., 2011, Tsurumi et al., 2011, Biehs et al., 2010, Ashyraliyev et al., 2009, Chung et al., 2009, Löhr et al., 2009, Wolfstetter et al., 2009, Anaka et al., 2008, Beckervordersandforth et al., 2008, Blanco and Gehring, 2008, Cinnamon et al., 2008, Florence and Faller, 2008, Kim et al., 2008, Miura et al., 2008, Moore and Lasko, 2008, Sanders et al., 2008, Astigarraga et al., 2007, Coppey et al., 2007, Hatton-Ellis et al., 2007, Munoz-Descalzo et al., 2007, Abrams et al., 2006, Jaeger and Reinitz, 2006, Jones et al., 2006, McGregor, 2006, Yucel and Small, 2006, Hariharan, 2005, Lundell et al., 1996)

hucklebein

(Kim et al., 2011)

Secondary FlyBase IDs

FBgn0086720
FBgn0001204
FBan0009768
FBgn0014111

Datasets (2)

Study focus (2)

Experimental Role

Project

Project Type

Title

bait_protein

genome binding

ChIP characterization of transcription factor genome binding, Berkeley Drosophila Transcription Factor Network Project.

bait_protein

modENCODE_regulation_TFs

genome binding

Genome-wide localization of transcription factors by ChIP-chip and ChIP-Seq.

References (406)