Allele Dmel\grh^IM

General Information
Symbol	Dmel\grh^IM	Species	D. melanogaster
Name		FlyBase ID	FBal0005208
Feature type	allele	Associated gene	Dmel\grh

Allele class	amorphic allele - genetic evidence
Mutagen	ethyl methanesulfonate
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Ostrowski et al., 2002)
Mutagen	ethyl methanesulfonate (Nusslein-Volhard et al., 1984, Tearle and Nusslein-Volhard, 1987)
Mutations Mapped to the Genome

Type Location Additional Notes References
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Nonsense mutation: TAT to TAA stop-codon introduction in exon seven, within the DNA-binding domain. (Paré et al., 2012)
Cytology
Phenotypic Data
Phenotypic Class
	lethal \| embryonic stage \| recessive (Nusslein-Volhard et al., 1984, Tearle and Nusslein-Volhard, 1987) neuroanatomy defective (with Df(2R)Pcl7B) (Baumgardt et al., 2009) planar polarity defective \| somatic clone (Lee and Adler, 2004) wound healing defective (Mace et al., 2005) wound healing defective \| embryonic stage 17 \| recessive (Kim and McGinnis, 2011)
Phenotype Manifest In
	cephalopharyngeal skeleton (Tearle and Nusslein-Volhard, 1987) cephalopharyngeal skeleton \| embryonic stage (Kim and McGinnis, 2011) chaeta \| somatic clone (Lee and Adler, 2004) chaeta \| somatic clone \| supernumerary (Lee and Adler, 2004) denticle belt (Tearle and Nusslein-Volhard, 1987) dorsal neurohemal organ (with Df(2R)Pcl7B) (Baumgardt et al., 2009) embryonic/first instar larval cuticle (Ostrowski et al., 2002) embryonic/larval cuticle (Kim and McGinnis, 2011) embryonic/larval epidermis \| embryonic stage 17 (Kim and McGinnis, 2011) epicuticle (Gangishetti et al., 2012) eye \| somatic clone (Lee and Adler, 2004) eye photoreceptor cell \| somatic clone (Lee and Adler, 2004) neuroblast NB5-6 of thorax (with Df(2R)Pcl7B) (Baumgardt et al., 2009) ommatidium \| somatic clone (Lee and Adler, 2004) procuticle (Gangishetti et al., 2012) wing \| somatic clone (Lee and Adler, 2004) wing hair \| cell non-autonomous \| somatic clone (Lee and Adler, 2004) wing hair \| cell non-autonomous \| somatic clone \| supernumerary (Lee and Adler, 2004) wing hair \| somatic clone (Lee and Adler, 2004) wing hair \| somatic clone \| supernumerary (Lee and Adler, 2004) wing margin bristle \| somatic clone (Lee and Adler, 2004) wing vein \| ectopic \| somatic clone (Lee and Adler, 2004)
Detailed Description
	Statement Reference The inner two layers of the cuticle (epicuticle and procuticle) appear to have mixed in mutant larvae, resulting in bloated animals that die before hatching. (Gangishetti et al., 2012) The larval cuticle in grh[IM] homozygotes is weak and easily ruptured and the head skeleton is grainy and discontinuous. Stage 17 homozygous grh[IM] embryos display defects in epidermal barrier integrity (an inability to exclude dye from the body cavity). This defect is not observed in grh[IM] heterozygous embryos. grh[IM] homozygous stage 17 embryos do not regain epidermal barrier integrity following wounding. Efficient regeneration of the barrier occurs in grh[IM] heterozygotes. (Kim and McGinnis, 2011) The number of SELK neurons is normal in grh[IM] mutant animals. (Losada-Pérez et al., 2010) grh[IM]/Df(2R)Pcl7B mutants show an increase in the number of ap NB5-6T neuroblasts from 4 to 6. Frequent loss of dorsal neurohemal organ innervation is observed in 75% of animals. (Baumgardt et al., 2009) Mutant embryos show an abnormal wound healing response after wounding with a sterile micropipette. (Mace et al., 2005) When homozygous mutant somatic clones are made in the wing, polarity defects are seen. Multiple wing hairs are sometimes seen, and the hairs are more erect than wild-type. The majority of clones show signs of weak domineering non-autonomy consisting of a small number of multiple wing hairs and/or hairs of abnormal polarity. When pupal wing clones are examined, it is seen that the multiple hair cell phenotype tends to be stronger in larger clones. In some clones not all cells produce multiple hairs. In addition an apparent delay in hair development is seen in many clones. Other phenotypes are also seen in mutant somatic clones. Most wings show ectopic wing veins. There is often disruption to the pattern of marginal bristles and occasionally polyploid cells. Large clones result in bulges of the wing and the region of the clone. Clones in the abdomen have reduced pigmentation and/or multiple hair phenotypes. The reduced pigmentation is most obvious in regions that are darkly pigmented but it could be seen elsewhere. In addition to the reduced pigmentation the clone cells often show a strong multiple hair cell phenotype. In abdominal areas that normally produce few hairs, a strong multiple a strong multiple hair cell phenotype is not seen and in some cases there appears to be a loss of hairs. Clones in the notum have reduced pigmentation. The hair phenotypes of these clones varies from an increased number of hairs to a loss of hairs. Lateral clones usually display a multiple hair cell phenotype, while medial clones often, but not invariably have a reduced number of hairs. Flies carrying mutant clones have rough spots on the eye. This phenotype is associated with ommatidia that show incorrect chirality, incorrect rotation and an abnormal number of photoreceptor cells. (Lee and Adler, 2004) When homozygous embryos are mechanically devitellinised the resulting cuticle preparations stretch to a greater extent than wild-type cuticles, resulting in inflated cuticles ("blimp" phenotype). (Ostrowski et al., 2002) grh^IM mutant embryos have a defective head skeleton and weak denticles. (Tearle and Nusslein-Volhard, 1987) Head skeleton glassy. (Nusslein-Volhard et al., 1984)
External Data
Linkouts
Interactions
Phenotypic Class
NOT Enhanced by
Statement Reference grh^IM has planar polarity defective \| somatic clone phenotype, non-enhanceable by in¹ (Lee and Adler, 2004) grh^IM has planar polarity defective \| somatic clone phenotype, non-enhanceable by mwh¹ (Lee and Adler, 2004)
NOT suppressed by
Statement Reference grh^IM has planar polarity defective \| somatic clone phenotype, non-suppressible by in¹ (Lee and Adler, 2004) grh^IM has planar polarity defective \| somatic clone phenotype, non-suppressible by mwh¹ (Lee and Adler, 2004)
NOT Enhancer of
Statement Reference grh^IM is a non-enhancer of planar polarity defective \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-enhancer of planar polarity defective \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004)
NOT Suppressor of
Statement Reference grh^IM is a non-suppressor of planar polarity defective \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-suppressor of planar polarity defective \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004)
Phenotype Manifest In
NOT Enhanced by
Statement Reference grh^IM has wing hair \| somatic clone phenotype, non-enhanceable by in¹ (Lee and Adler, 2004) grh^IM has wing hair \| somatic clone phenotype, non-enhanceable by mwh¹ (Lee and Adler, 2004) grh^IM has wing hair \| supernumerary \| somatic clone phenotype, non-enhanceable by in¹ (Lee and Adler, 2004) grh^IM has wing hair \| supernumerary \| somatic clone phenotype, non-enhanceable by mwh¹ (Lee and Adler, 2004)
NOT suppressed by
Statement Reference grh^IM has wing hair \| somatic clone phenotype, non-suppressible by in¹ (Lee and Adler, 2004) grh^IM has wing hair \| somatic clone phenotype, non-suppressible by mwh¹ (Lee and Adler, 2004) grh^IM has wing hair \| supernumerary \| somatic clone phenotype, non-suppressible by in¹ (Lee and Adler, 2004) grh^IM has wing hair \| supernumerary \| somatic clone phenotype, non-suppressible by mwh¹ (Lee and Adler, 2004)
NOT Enhancer of
Statement Reference grh^IM is a non-enhancer of wing hair \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-enhancer of wing hair \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004) grh^IM is a non-enhancer of wing hair \| supernumerary \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-enhancer of wing hair \| supernumerary \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004)
NOT Suppressor of
Statement Reference grh^IM is a non-suppressor of wing hair \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-suppressor of wing hair \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004) grh^IM is a non-suppressor of wing hair \| supernumerary \| somatic clone phenotype of in¹ (Lee and Adler, 2004) grh^IM is a non-suppressor of wing hair \| supernumerary \| somatic clone phenotype of mwh¹ (Lee and Adler, 2004)
Additional Comments
Genetic Interactions
Statement Reference When grh^IM mutant clones are made in an in¹ mutant wing the grh^IM phenotype is still seen. When grh^IM mutant clones are made in an mwh¹ mutant wing the grh^IM phenotype is still seen. A slight additive effect is seen on the phenotype. (Lee and Adler, 2004)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Partially rescued by	grh^IM is partially rescued by grh^2A.Scer\UAS/Scer\GAL4^e22c (Kim and McGinnis, 2011) grh^IM is partially rescued by grh^{PanA.Scer\UAS}/Scer\GAL4^e22c (Kim and McGinnis, 2011) grh^IM is partially rescued by Scer\GAL4^e22c/grh^2E.Scer\UAS (Kim and McGinnis, 2011) grh^IM is partially rescued by Scer\GAL4^e22c/grh^Scer\UAS.cKa (Kim and McGinnis, 2011)
Comments	Expression of grh[Scer\UAS.cKa] in the epidermis under the control of Scer\GAL4[e22c] partially rescues the cuticle and head skeleton defects seen in grh[IM] embryos. Expression of grh[2A.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] partially rescues the cuticle and head skeleton defects seen in grh[IM] embryos. Expression of grh[PanA.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] partially rescues the cuticle and head skeleton defects seen in grh[IM] embryos. Expression of grh[2E.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] partially rescues the cuticle and head skeleton defects seen in grh[IM] embryos. Expression of grh[Scer\UAS.cKa] in the epidermis under the control of Scer\GAL4[e22c] rescues the defects in epidermal barrier integrity seen in stage 17 grh[IM] homozygous embryos. Epidermal barrier regeneration following wounding is also rescued. Expression of grh[2A.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] rescues the defects in epidermal barrier integrity seen in stage 17 grh[IM] homozygous embryos. Epidermal barrier regeneration following wounding is only partially rescued. Expression of grh[PanA.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] rescues the defects in epidermal barrier integrity seen in stage 17 grh[IM] homozygous embryos. However epidermal barrier regeneration following wounding is only partially rescued. Expression of grh[2E.Scer\UAS] in the epidermis under the control of Scer\GAL4[e22c] rescues the defects in epidermal barrier integrity seen in stage 17 grh[IM] homozygous embryos. Epidermal barrier regeneration following wounding is also rescued. (Kim and McGinnis, 2011)
Stocks ( 2 )
Bloomington	3270 cn¹ grh^IM bw¹ sp¹/CyO
Kyoto	106949 cn¹ grh^IM bw¹ sp¹/CyO
Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 3 )
Reported As
Symbol Synonym	grh^IM (Luschnig et al., 2006, Mace et al., 2005, Benito-Sipos et al., 2011, Baumgardt et al., 2009, Losada-Pérez et al., 2010, Harrison et al., 2010, Garcia and Stathopoulos, 2011, Gangishetti et al., 2012, Pearson et al., 2012, Kim and McGinnis, 2011, Pearson et al., 2009, Paré et al., 2012) hsk^IM45 l(2)IM45 (Nusslein-Volhard et al., 1984)
Name Synonym
Secondary FlyBase IDs
	FBal0039410
References ( 20 )
Research paper	Gangishetti et al., 2012, Insect Mol. Biol. 21(3): 283--295 The transcription factor Grainy head and the steroid hormone ecdysone cooperate during differentiation of the skin of Drosophila melanogaster. [FBrf0218253] Paré et al., 2012, PLoS ONE 7(5): e36254 The functions of grainy head-like proteins in animals and fungi and the evolution of apical extracellular barriers. [FBrf0218280] Pearson et al., 2012, Dev. Biol. 366(2): 420--432 Drosophila melanogaster Zelda and Single-minded collaborate to regulate an evolutionarily dynamic CNS midline cell enhancer. [FBrf0218349] Benito-Sipos et al., 2011, Development 138(24): 5311--5320 Seven up acts as a temporal factor during two different stages of neuroblast 5-6 development. [FBrf0216799] 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] Kim and McGinnis, 2011, Proc. Natl. Acad. Sci. U.S.A. 108(2): 650--655 Phosphorylation of Grainy head by ERK is essential for wound-dependent regeneration but not for development of an epidermal barrier. [FBrf0214191] Harrison et al., 2010, Dev. Biol. 345(2): 248--255 Grainyhead and Zelda compete for binding to the promoters of the earliest-expressed Drosophila genes. [FBrf0211626] Losada-Pérez et al., 2010, Mech. Dev. 127(9-12): 458--471 Lineage-unrelated neurons generated in different temporal windows and expressing different combinatorial codes can converge in the activation of the same terminal differentiation gene. [FBrf0212045] Baumgardt et al., 2009, Cell 139(5): 969--982 Neuronal subtype specification within a lineage by opposing temporal feed-forward loops. [FBrf0209429] Pearson et al., 2009, Proc. Natl. Acad. Sci. U.S.A. 106(7): 2224--2229 Multiple transcription factor codes activate epidermal wound-response genes in Drosophila. [FBrf0206667] Luschnig et al., 2006, Curr. Biol. 16(2): 186--194 serpentine and vermiform encode matrix proteins with chitin binding and deacetylation domains that limit tracheal tube length in Drosophila. [FBrf0190029] Moussian et al., 2006, Development 133(1): 163--171 Drosophila Knickkopf and Retroactive are needed for epithelial tube growth and cuticle differentiation through their specific requirement for chitin filament organization. [FBrf0190312] Mace et al., 2005, Science 308(5720): 381--385 An epidermal barrier wound repair pathway in Drosophila is mediated by grainy head. [FBrf0188326] Lee and Adler, 2004, Mech. Dev. 121(1): 37--49 The grainy head transcription factor is essential for the function of the frizzled pathway in the Drosophila wing. [FBrf0167844] Mohr and Gelbart, 2002, Genetics 162(1): 165--176 Using the P{wHy} hybrid transposable element to disrupt genes in region 54D-55B in Drosophila melanogaster. [FBrf0152030] Ostrowski et al., 2002, Genetics 161(1): 171--182 Genetic control of cuticle formation during embryonic development of Drosophila melanogaster. [FBrf0149011] Nusslein-Volhard et al., 1984, Rouxs Arch. Dev. Biol. 193: 267--282 Mutations affecting the pattern of the larval cuticle in Drosophila melanogaster. [FBrf0041708]
Supplementary material	Mace et al., 2005, Science 308(5720): Supporting online material. [FBrf0202560]
Personal communication to FlyBase	Bray, 1996.5.13, grh300 and thr. grh300 and thr. [FBrf0086811]
Stock list	Tearle and Nusslein-Volhard, 1987, D. I. S. 66: 209--269 Tubingen mutants and stock list. [FBrf0045941]

Allele Dmel\grhIM

Allele Dmel\grh^IM