The gene gurken is referred to in FlyBase by the symbol Dmel\grk (CG17610, FBgn0001137). It is a protein_coding_gene from Drosophila melanogaster. It is reported to have molecular function: gurken receptor binding; epidermal growth factor receptor binding. There is experimental evidence that it is involved in the biological process: maternal determination of dorsal/ventral axis, ovarian follicular epithelium, germ-line encoded; establishment of oocyte nucleus localization involved in oocyte dorsal/ventral axis specification; oocyte microtubule cytoskeleton organization; border follicle cell migration; chorion-containing eggshell pattern formation. 85 alleles are reported. The phenotypes of these alleles are annotated with: organ system; multicellular structure; anatomical structure; acellular anatomical structure; adult segment; organ system subdivision; portion of tissue; thoracic segment; epithelial cell; epithelial furrow; cell part. It has one annotated transcript and one annotated polypeptide. Protein features are: EGF-like, conserved site; Epidermal growth factor-like domain. Summary of modENCODE Temporal Expression Profile: Temporal profile ranges from a peak of moderately high expression to a trough of low expression. Peak expression observed during late pupal stages. Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult crop, adult hindgut, adult heart, adult ovary, larval/adult carcass. Comments on Affy2 ProbeSet: ProbeSet 1627656_at completely aligns to an exonic region of the only FlyBase-annotated transcript isoform of grk. Gene sequence location is 2L:8431086..8433598.
User Contributed Data
External Summaries
Phenotypic Description from the Red Book (Lindsley
& Zimm 1992)
Gene/Allele symbols may differ
from current usage
grk: gurken
Maternal-effect lethal. Wild-type allele required
for normal dorsoventral pattern in the egg. In mutants, the
ventral regions of the chorion and the embryo are expanded at
the expense of the dorsal regions. The pattern of the chorion
is altered, a second micropyle and a small patch of
operculum-like material often forming at the posterior pole in
extremely mutant eggs; fewer cells contribute to the dorsal
appendage, which is usually shifted posteriorly, but more follicle cells contribute to the main body of the chorion. In
the embryo the major increase in cell mass occurs in the mesoderm as an invagination on the ventral side during early gastrulation. Analysis of mosaic females in which germ cells and
sister nurse cells are of different genotype indicate that grk
mutations act only in the germ line.
Recent Updates
Description
What does this section display?
This section contains items that were added to this record for each release.
It currently only tracks new links between this FlyBase report and other
FlyBase data classes (e.g. genes, references, stocks) or controlled
vocabulary terms (e.g. GO, anatomy terms).
What does this section not display?
This section does not currently display links that were removed or gene model changes.
Update Feed
Click the icon below to subscribe to this FlyBase record and receive updates automatically through your
feed reader.
FB2013_03
FB2013_02
All updates
Click here to see a list of all updates to this record from FB2010_08 and on.
grk transcripts are first detected in region 2B of the germarium. In oogenesis stages S1-S7, transcripts are predominantly found in a crescent lining the posterior end of the oocyte. During oogenesis stages S8 and S9, transcripts become localized to the dorsal-anterior corner of the oocyte nucleus. In fs(1)K10, sqd, spir, and capu mutants, grk transcripts are no longer restricted to the dorsal side of the oocyte at later stages of oogenesis but are also abundant along the anterior margin of the oocyte. In Egfr and cni mutants, the grk transcript distribution is unaffected.
Transcript as detected by in situ hybridization and injection of labeled grk RNA into cultured oocytes show that at stage 8 grk transcript is localized in an anterior ring and by stage 9 the transcript has been concentrated in the dorsoanterior region of the oocyte. A less enriched concentration of grk transcript can be detected in the ventral anterior region of the oocyte. Videomicroscopy showed that this localization occurs in a 2 step process in which the transcript first moves anteriorly and then is concentrated dorsally.
grk transcripts are detected at very low levels on northern blots in grk2, grk3, and grk4 mutants. No transcripts are detected in egg chambers of these mutants.
Transcript is localized in the posterior of stage 6 oocytes and relocalizes to the anterior of stage 8 oocytes. Further refinement of transcript localization to the anterodorsal corner of the oocyte is observed in stage 9-10 oocytes.
grk protein is detected starting in early oogenesis, in germarium region 2b. Like grk transcript, grk protein localizes to the oocyte in early egg chambers, and starting at stage S8, becomes localized to the future dorsal cortex of the oocyte. At stage S10-S12, the grk protein localization differs from grk transcript localization. grk protein forms an elongated anterior-posterior stripe over about half the length of the oocyte\'s dorsal midline.
Summary of FlyAtlas Anatomical Expression Data: Expression at moderate levels in the following post-embryonic organs or tissues: adult crop, adult hindgut, adult heart, adult ovary, larval/adult carcass.
[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
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
Linear, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
High
Linear, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
High
Very high
Linear, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
Very high
log, scaled to maximum expression level
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
High
log, scaled to Moderate expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
High
log, scaled to High level expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
Expression Level Scale
None
Low
Moderate
High
Very high
log, scaled to Very high expression
Tissue
Expression Level
Larval Central Nervous System
16.55
Larval Midgut
20.7
Larval Hindgut
96.5
Larval Malpighian Tubules
22
Larval Fat Body
15.5
Larval Salivary Gland
14.3
Larval Trachea
9.975
Larval Carcass
273.625
Adult Head
86.2
Adult Eye
10.975
Adult Brain
6.3
Adult Thoracic-Abdominal Ganglion
9.4
Adult Crop
328.7
Adult Midgut
19.3
Adult Hindgut
169.2
Adult Malpighian Tubules
5.8
Adult Fat Body
24.6
Adult Salivary Gland
17.2
Adult Heart
237.825
Adult VirginFemale Spermatheca
28.4
Adult InseminatedFemale Spermatheca
44.4
Adult Ovary
104.7
Adult Testis
16.5
Adult Male Accessory Gland
62.5
Adult Carcass
179.9
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 low expression. Peak expression observed during late pupal stages.
[download data (TSV)]
Please Note FlyBase no
longer curates genomic clone accessions so this list
may not be complete
cDNA Clones ( 8 )
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.
grk RNA is transported as particles from nurse cells to the oocyte by a Dhc64C/BicD dependent mechanism distinct from the general flow of cytoplasm from nurse cell to oocyte.
Different dorsal follicle cell fates are not determined by a grk morphogen gradient. They are specified by secondary signal amplification and refinement processes that integrate the grk signal with positive and negative feedback mechanisms generated by target genes of the Egfr pathway.
grk plays a role in the development of follicular epithelium by cooperating with brn for the Egfr-dependent migration of the prefollicular cels around each nurse cell-oocyte complex. brn may help provide specificity to and/or facilitate the multiplicity of grk-Egfr functions during oogenesis.
Mutations in grk cause a ventralized phenotype in egg and embryo. Changing grk dosage in otherwise wild type ovaries is sufficicent to alter the number of somatic follicle cells directed to the dorsal fate. The grk-Egfr signalling process plays an instructive role in oogenesis, inducing dorsal cell fates in the follicle cell epithelium and controlling the production of maternal components that will direct the embryonic dorsoventral pattern.
grk, unlike other localized cytoplasmic determinants, is not directly responsible for the establishment of cell fates along a body axis, but restricts and orients an active axis-forming process which occurs later in the follicular epithelium or in the early embryo.
Mutations at the grk locus cause defects in midoogenesis. Double mutant analysis indicates that rho acts upstream of Tl in dorsal-ventral axis formation, and the action of rho requires the grk-Egfr signaling pathway.
maternal-effect lethal. Wild-type allele required for normal dorsoventral pattern in the egg. In mutants, the ventral regions of the chorion and the embryo are expanded at the expense of the dorsal regions. The pattern of the chorion is altered, a second micropyle and a small patch of operculum-like material often forming at the posterior pole in extremely mutant eggs; fewer cells contribute to the dorsal appendage, which is usually shifted posteriorly, but more follicle cells contribute to the main body of the chorion. In the embryo the major increase in cell mass occurs in the mesoderm as an invagination on the ventral side during early gastrulation. Analysis of mosaic females in which germ cells and sister nurse cells are of different genotype indicate that grk mutations act only in the germ line.
Leibfried et al., 2013, Development 140(2): 362--371
A Cdc42-regulated actin cytoskeleton mediates Drosophila oocyte polarization. [FBrf0220348]
Anand and Kai, 2012, EMBO J. 31(4): 870--882
The tudor domain protein Kumo is required to assemble the nuage and to generate germline piRNAs in Drosophila. [FBrf0217541]
Baffet et al., 2012, Mol. Biol. Cell 23(18): 3591--3601
Drosophila tubulin-binding cofactor B is required for microtubule network formation and for cell polarity. [FBrf0219447]
Ferguson et al., 2012, J. Cell Sci. 125(6): 1407--1419
Modulation of gurken translation by insulin and TOR signaling in Drosophila. [FBrf0218097]
Ganguly et al., 2012, Proc. Natl. Acad. Sci. U.S.A. 109(38): 15109--15114
Cytoplasmic streaming in Drosophila oocytes varies with kinesin activity and correlates with the microtubule cytoskeleton architecture. [FBrf0219481]
Hartswood et al., 2012, RNA 18(4): 729--737
RNA:RNA interaction can enhance RNA localization in Drosophila oocytes. [FBrf0217830]
Ji and Tulin, 2012, Nat. Commun. 3: 760
Poly(ADP-ribose) controls DE-cadherin-dependent stem cell maintenance and oocyte localization. [FBrf0217856]
McDermott et al., 2012, Biol. Open 1(5): 488--497
Drosophila Syncrip binds the gurken mRNA localisation signal and regulates localised transcripts during axis specification. [FBrf0218722]
Naylor and Diantonio, 2012, Dev. Neurobiol. 72(9): 1229--1242
EGFR signaling modulates synaptic connectivity via Gurken. [FBrf0219184]
Simakov et al., 2012, Development 139(15): 2814--2820
EGFR-dependent network interactions that pattern Drosophila eggshell appendages. [FBrf0218814]
Technau et al., 2012, Dev. Genes Evol. 222(1): 1--17
Molecular mechanisms of EGF signaling-dependent regulation of pipe, a gene crucial for dorsoventral axis formation in Drosophila. [FBrf0217623]
Weber et al., 2012, PLoS ONE 7(4): e34745
Genome-Wide Association Analysis of Oxidative Stress Resistance in Drosophila melanogaster. [FBrf0218073]
Weil et al., 2012, Nat. Cell Biol. 14(12): 1305--1315
Drosophila patterning is established by differential association of mRNAs with P bodies. [FBrf0220085]
Yue et al., 2012, Dev. Cell 22(2): 255--267
The cell adhesion molecule echinoid functions as a tumor suppressor and upstream regulator of the hippo signaling pathway. [FBrf0217474]
Avery et al., 2011, RNA 17(4): 624--638
Drosophila Upf1 and Upf2 loss of function inhibits cell growth and causes animal death in a Upf3-independent manner. [FBrf0213282]
Dubin-Bar et al., 2011, Development 138(21): 4661--4671
Drosophila javelin-like encodes a novel microtubule-associated protein and is required for mRNA localization during oogenesis. [FBrf0216406]
Fan et al., 2011, PLoS ONE 6(5): e20612
Drosophila Ge-1 Promotes P Body Formation and oskar mRNA Localization. [FBrf0213914]
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]
Horn et al., 2011, Nat. Methods 8(4): 341--346
Mapping of signaling networks through synthetic genetic interaction analysis by RNAi. [FBrf0213352]
Joyce et al., 2011, J. Cell Biol. 195(3): 359--367
Drosophila ATM and ATR have distinct activities in the regulation of meiotic DNA damage and repair. [FBrf0216489]
Poukkula et al., 2011, J. Cell Biol. 192(3): 513--524
Cell behaviors regulated by guidance cues in collective migration of border cells. [FBrf0212926]
Sato et al., 2011, Genes Dev. 25(22): 2361--2373
Maelstrom coordinates microtubule organization during Drosophila oogenesis through interaction with components of the MTOC. [FBrf0216748]
Serbus et al., 2011, J. Cell Sci. 124(24): 4299--4308
A feedback loop between Wolbachia and the Drosophila gurken mRNP complex influences Wolbachia titer. [FBrf0217227]
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]
Sun et al., 2011, Development 138(10): 1991--2001
Regulation of somatic myosin activity by Protein Phosphatase 1{beta} controls Drosophila oocyte polarization. [FBrf0213586]
Vazquez-Pianzola et al., 2011, Dev. Biol. 357(2): 404--418
Pabp binds to the osk 3'UTR and specifically contributes to osk mRNA stability and oocyte accumulation. [FBrf0214798]
Wang et al., 2011, Anal. Bioanal. Chem. 400(2): 335--341
Lie group study of Raman spectra of the Gurken gradient in Drosophila oogenesis. [FBrf0214310]
Wang and Pai, 2011, PLoS ONE 6(2): e17097
D-Cbl Binding to Drk Leads to Dose-Dependent Down-Regulation of EGFR Signaling and Increases Receptor-Ligand Endocytosis. [FBrf0213073]
Wang and Elgin, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(52): 21164--21169
Drosophila Piwi functions downstream of piRNA production mediating a chromatin-based transposon silencing mechanism in female germ line. [FBrf0217081]
Wong et al., 2011, PLoS ONE 6(9): e24355
The Functioning of the Drosophila CPEB Protein Orb Is Regulated by Phosphorylation and Requires Casein Kinase 2 Activity. [FBrf0216252]
Yan et al., 2011, Development 138(9): 1697--1703
Drosophila PI4KIIIalpha is required in follicle cells for oocyte polarization and Hippo signaling. [FBrf0213488]
Zettl et al., 2011, Cell 145(1): 79--91
Rhomboid Family Pseudoproteases Use the ER Quality Control Machinery to Regulate Intercellular Signaling. [FBrf0213387]