Allele Dmel\E(spl)m8-HLH¹

General Information
Symbol	Dmel\E(spl)m8-HLH¹	Species	D. melanogaster
Name		FlyBase ID	FBal0003323
Feature type	allele	Associated gene	Dmel\E(spl)m8-HLH
Also Known As	E(spl)^D, E(spl)¹

Map ( GBrowse )
Allele class	gain of function allele
Mutagen	spontaneous
Recent Updates
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FB2013_03
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Nature of the Allele
Allele class	gain of function allele (Knust et al., 1987, Shepard et al., 1989)
Mutagen	spontaneous (Knust et al., 1987, Preiss et al., 1988, Ziemer et al., 1988)
Mutations Mapped to the Genome

Type Location Additional Notes References sequence variant 3R:21,866,415..21,866,898 comment=484 base deletion which removes the region encoding the final 56aa of the E(spl) gene, replacing them with 9 aa. evidence=experimental (Klambt et al., 1989)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Deletion of the C-terminal domain and a putative regulatory element. (Oellers et al., 1993) 0.7-kb deletion in 12 to 14 kb; 5-kb insertion at 16.5 kb
Caused by insertion	P{lacZ}E(spl)m8-HLH¹ (Preiss et al., 1988, Karandikar et al., 2004)
Cytology	Polytene chromosomes normal. (Preiss et al., 1988) Polytene chromosomes normal. (Ziemer et al., 1988)
Phenotypic Data
Phenotypic Class
	visible (Grushko et al., 1998) visible \| dominant (Knust et al., 1987, Alton et al., 1989, Moores et al., 2008)
Phenotype Manifest In
	eye (Oellers et al., 1993) microchaeta & mesothoracic tergum (Moores et al., 2008) wing vein (Sturtevant and Bier, 1995, Grushko et al., 1998)
Detailed Description
	Statement Reference Heterozygous flies have a significantly increased density of microchaetae on the notum compared to controls. (Moores et al., 2008) The number and length of wing vein breaks increases with increasing temperature in homozygous flies. (Grushko et al., 1998) Thickened vein mutant. (Sturtevant and Bier, 1995) Enhances the eye phenotype of N^spl-1. (Oellers et al., 1993) Individuals heterozygous for Bpt¹ and heterozygous for E(spl)¹ exhibit no Bpt¹ phenotype. (Duus et al., 1992) Dominant phenotype increasing the number of microchaetae on the notum. (de Celis et al., 1991) E(spl)¹/E(spl)^rv1 heterozygotes show a homogeneous increase in chaetae density on the tergites. (Mari-Beffa et al., 1991) Homozygous flies have virtually normal eyes, with a very mild roughening at the rear of the eye. (Campos-Ortega and Knust, 1990) Severe reduction in size of compound eye. (Alton et al., 1989) Strongly enhanced expression of "N^Ax" alleles. (Siren and Portin, 1989) P element construct carrying a wild type E(spl) gene copy can rescue the lethality of E(spl)¹ with other E(spl) mutations. (Preiss et al., 1988) Homozygotes displayed neural hypoplasia and enhancement of the N^spl-1 phenotype. (Ziemer et al., 1988) Most homozygotes display slight roughening of the eye. A few exhibit irregular epidermal defects and others show fusion of neighbouring denticle belts. E(spl)¹ has complete penetrance in enhancing the N^spl-1 phenotype. During embryogenesis E(spl)¹ behaves as a dominant allele with incomplete penetrance for cell death in neural primordia. (Knust et al., 1987) unique dominant allele
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference E(spl)m8-HLH¹, N^spl-1 has visible \| dominant phenotype, enhanceable by ato³ (Nagel and Preiss, 1999) E(spl)m8-HLH¹, N^spl-1 has visible \| dominant phenotype, enhanceable by ato⁵ (Nagel and Preiss, 1999) E(spl)m8-HLH¹, N^spl-1 has visible \| dominant phenotype, enhanceable by da⁵ (Nagel and Preiss, 1999) E(spl)m8-HLH¹, N^spl-1 has visible \| suppressible phenotype, enhanceable by sca^unspecified (Nagel and Preiss, 1999)
NOT Enhanced by
Statement Reference E(spl)m8-HLH¹ has visible \| dominant phenotype, non-enhanceable by Hip1^Scer\UAS.cMa/Scer\GAL4^pnr-MD237 (Moores et al., 2008) E(spl)m8-HLH¹ has visible \| dominant phenotype, non-enhanceable by Scer\GAL4^pnr-MD237/Hip1^{ΔANTH.Scer\UAS} (Moores et al., 2008)
Suppressed by
Statement Reference E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl[+]/Dl^BE23 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl[+]/Dl^BX41 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl[+]/Dl^BX43 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl[+]/Dl^CE21 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl[+]/Dl^CE43 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BE21/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BE24/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BE26/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BE32/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BE35/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BX40/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BX45/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^BX46/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^CE23/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^CE33/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^CE34/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, suppressible by Dl^CE37/Dl[+] (Shepard et al., 1989)
NOT suppressed by
Statement Reference E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has visible phenotype, non-suppressible by Dl^HD82/Dl[+] (Shepard et al., 1989) E(spl)m8-HLH¹ has visible \| dominant phenotype, non-suppressible by Hip1^Scer\UAS.cMa/Scer\GAL4^pnr-MD237 (Moores et al., 2008) E(spl)m8-HLH¹ has visible \| dominant phenotype, non-suppressible by Scer\GAL4^pnr-MD237/Hip1^{ΔANTH.Scer\UAS} (Moores et al., 2008)
Enhancer of
Statement Reference E(spl)[+]/E(spl)m8-HLH¹ is an enhancer of visible phenotype of N^spl-1 (Shepard et al., 1989) E(spl)m8-HLH¹ is an enhancer of visible \| dominant phenotype of N^spl-1 (Nagel and Preiss, 1999) E(spl)m8-HLH¹ is an enhancer of visible phenotype of N^spl-1 (Campos-Ortega and Knust, 1990)
Suppressor of
Statement Reference E(spl)m8-HLH¹/E(spl)m8-HLH[+] is a suppressor of visible phenotype of Scer\GAL4^tub.PU, csw^{N308D.Scer\UAS.P\T} (Oishi et al., 2006) E(spl)m8-HLH¹ is a suppressor \| partially of lethal \| recessive \| heat sensitive phenotype of N^Ax-tsl (Grushko et al., 1998)
Other
Statement Reference E(spl)m8-HLH¹, gro^B7 has lethal phenotype (Ziemer et al., 1988, Ziemer et al., 1988) E(spl)m8-HLH¹, gro^B37 has lethal phenotype (Ziemer et al., 1988, Ziemer et al., 1988) E(spl)m8-HLH¹, gro^B48 has lethal phenotype (Ziemer et al., 1988, Ziemer et al., 1988) E(spl)m8-HLH¹, gro^B88 has lethal phenotype (Ziemer et al., 1988, Ziemer et al., 1988) E(spl)m8-HLH¹, gro^B93 has lethal phenotype (Ziemer et al., 1988, Ziemer et al., 1988) E(spl)m8-HLH¹, N^Ax-tsl has visible phenotype (Grushko et al., 1998)
Phenotype Manifest In
NOT Enhanced by
Statement Reference E(spl)m8-HLH¹ has microchaeta & mesothoracic tergum phenotype, non-enhanceable by Hip1^Scer\UAS.cMa/Scer\GAL4^pnr-MD237 (Moores et al., 2008) E(spl)m8-HLH¹ has microchaeta & mesothoracic tergum phenotype, non-enhanceable by Scer\GAL4^pnr-MD237/Hip1^{ΔANTH.Scer\UAS} (Moores et al., 2008)
Suppressed by
Statement Reference E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl[+]/Dl^BE23 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl[+]/Dl^BX41 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl[+]/Dl^BX43 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl[+]/Dl^CE21 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl[+]/Dl^CE43 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BE21/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BE24/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BE26/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BE32/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BE35/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BX40/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BX45/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^BX46/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^CE23/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^CE33/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^CE34/Dl[+] (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, suppressible by Dl^CE37/Dl[+] (Shepard et al., 1989)
NOT suppressed by
Statement Reference E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype, non-suppressible by Dl^HD82/Dl[+] (Shepard et al., 1989) E(spl)m8-HLH¹ has microchaeta & mesothoracic tergum phenotype, non-suppressible by Hip1^Scer\UAS.cMa/Scer\GAL4^pnr-MD237 (Moores et al., 2008) E(spl)m8-HLH¹ has microchaeta & mesothoracic tergum phenotype, non-suppressible by Scer\GAL4^pnr-MD237/Hip1^{ΔANTH.Scer\UAS} (Moores et al., 2008)
Enhancer of
Statement Reference E(spl)[+]/E(spl)m8-HLH¹ is an enhancer of eye phenotype of N^nd-2 (Xu et al., 1990) E(spl)[+]/E(spl)m8-HLH¹ is an enhancer of eye phenotype of N^spl-1 (Shepard et al., 1989) E(spl)[+]/E(spl)m8-HLH¹ is an enhancer of macrochaeta phenotype of CkIIα^{dsRNA.Scer\UAS}, Scer\GAL4^sca-537.4 (Bose et al., 2006) E(spl)m8-HLH¹ is an enhancer of eye phenotype of N^spl-1 (Nagel and Preiss, 1999, Campos-Ortega and Knust, 1990) E(spl)m8-HLH¹ is an enhancer of ommatidium phenotype of N^spl-1 (Nagel and Preiss, 1999, Campos-Ortega and Knust, 1990) E(spl)m8-HLH¹ is an enhancer of wing vein phenotype of N^Ax-E2 (Grushko et al., 1998)
NOT Enhancer of
Statement Reference E(spl)[+]/E(spl)m8-HLH¹ is a non-enhancer of wing vein \| ectopic phenotype of Snr1^E1 (Marenda et al., 2004)
Suppressor of
Statement Reference E(spl)m8-HLH¹/E(spl)m8-HLH[+] is a suppressor of wing vein \| ectopic phenotype of Scer\GAL4^tub.PU, csw^{N308D.Scer\UAS.P\T} (Oishi et al., 2006) E(spl)m8-HLH¹ is a suppressor of macrochaeta phenotype of N^Ax-E2 (Grushko et al., 1998) E(spl)m8-HLH¹ is a suppressor of wing vein phenotype of N^Ax-tsl (Grushko et al., 1998)
NOT Suppressor of
Statement Reference E(spl)[+]/E(spl)m8-HLH¹ is a non-suppressor of wing vein \| ectopic phenotype of Snr1^E1 (Marenda et al., 2004)
Other
Statement Reference E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has eye phenotype (Karandikar et al., 2004) E(spl)[+]/E(spl)m8-HLH¹, N^spl-1 has ommatidium phenotype (Karandikar et al., 2004) E(spl)m8-HLH¹, N^nd-2 has eye phenotype (Xu et al., 1990) E(spl)m8-HLH¹, N^spl-1 has eye phenotype (Karandikar et al., 2004, Karandikar et al., 2004, Kahali et al., 2010) E(spl)m8-HLH¹, N^spl-1 has interommatidial bristle phenotype (Kahali et al., 2010) E(spl)m8-HLH¹, N^spl-1 has ommatidium phenotype (Karandikar et al., 2004, Karandikar et al., 2004, Kahali et al., 2010)
Additional Comments
Genetic Interactions
Statement Reference N[spl-1]/Y; E(spl)[1]/+ mutants exhibit a severely reduced eye. The residual eye field is highly disorganised; the clustering of the interommatidial bristles might well reflect facet loss. The severe reduced eye found in N[spl-1]/Y; E(spl)[1]/+ mutants is partially suppressed upon overexpression of E(spl)[123-179.CtD.Scer\UAS] immediately anterior to the morphogenetic furrow under the control of Scer\GAL4[h-H10]. Expression of E(spl)[123-179.CtD.Scer\UAS] not only increases eye size (facet numbers), but also restores the patterning of the facets. This patterning is, however, lost toward the anterior margin of the residual eye. The severe reduced eye found in N[spl-1]/Y; E(spl)[1]/+ mutants is unaffected by overexpression of E(spl)[S159D.123-179.CtD.Scer\UAS] under the control of Scer\GAL4[h-H10]. Expression of E(spl)[123-179.CtD.Scer\UAS] in trans, in all cells posterior to the morphogenetic furrow, under the control of Scer\GAL4[GMR.PF] does not rescue the severe reduced eye phenotype found in N[spl-1]/Y; E(spl)[1]/+ mutants. The absence of rescue is also supported by facet numbers, which upon expression of three independent E(spl)[123-179.CtD.Scer\UAS] lines closely mimics control levels in N[spl-1]/Y; E(spl)[1]/+ mutants. Rescue of the N[spl-1]/Y; E(spl)[1]/+ retinal defects by E(spl)[123-179.CtD.Scer\UAS] therefore requires expression in a region of the developing eye where funding R8 photoreceptors are patterned and specified. (Kahali et al., 2010) Expression of either Hip1[Scer\UAS.cMa] or Hip1[ΔANTH.Scer\UAS] under the control of Scer\GAL4[pnr-MD237] has no effect on the increased density of microchaetae on the notum seen in E(spl)[1]/+ flies. (Moores et al., 2008) CkIIα^Tik, E(spl)¹ double mutant flies do not show ommatidial or interommatidial bristle defects. Expression of one copy of CkIIα^{dsRNA.Scer\UAS} under the control of one copy of Scer\GAL4^sca-537.4 in an E(spl)¹/+ background results in a split bristle phenotype, while this level of CkIIα^{dsRNA.Scer\UAS} dosage is not sufficient to induce split bristles in a wild-type background. (Bose et al., 2006) E(spl)¹/+, N^spl-1/Y flies have severely reduced eyes. They typically exhibit 4-8 residual ommatidia. (Karandikar et al., 2004) Has no effect on bristle phenotype of N^spl-1 but enhances the eye phenotype. (Nagel and Preiss, 1999) The variation of the number of missing bristles and number and length of wing vein breaks with temperature in N^Ax-1; E(spl)¹ double mutants is the same as in E(spl)¹ single mutants. The number of bristles is rescued almost to wild type in N^Ax-E2 ; E(spl)¹ double mutants, while the number and length of wing vein breaks are higher in N^Ax-E2; E(spl)¹ double mutants than in N^Ax-E2 single mutants. N^Ax-tsl homozygotes and hemizygotes do not survive at 29^oC. N^Ax-tsl; E(spl)¹ double mutants show semi-lethality at 29^oC. The number and length of wing vein breaks are rescued to wild type in N^Ax-tsl; E(spl)¹ double mutants. (Grushko et al., 1998) Strong enhancer of the N^spl-1 phenotype; N^spl-1/Y E(spl)¹/+ flies have a severely reduced compound eye with only 15-30 widely spaced facets. The ommatidia contain fewer photoreceptors than normal, and the rhabdomeres are often distorted. A large number of secondary pigment cells fill the space between the scattered photoreceptor cell clusters. Photoreceptor cell axons follow an aberrant course, and usually fail to enter the eye stalk. The pattern of cell proliferation is qualitatively normal in N^spl-1/Y E(spl)¹/+ flies as assayed by BrdU incorporation. There is abundant cell death in the eye imaginal discs of these flies during the third instar larval stage, dead cells are more abundant behind the morphogenetic furrow. (Campos-Ortega and Knust, 1990) No phenotypic interaction with N^Ax-E2 or Dl^9P. Causes a rough eye phenotype when heterozygous with N^nd-2, more severe at room temperature. (Xu et al., 1990)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	E(spl)m8-HLH^rv29 (Ziemer et al., 1988) E(spl)m8-HLH^rv30 (Ziemer et al., 1988) E(spl)m8-HLH^rv31 (Ziemer et al., 1988)
Comments	The severe reduced eye found in N[spl-1]/Y; E(spl)[1]/+ mutants is partially suppressed upon overexpression of E(spl)[123-179.CtD.Scer\UAS] immediately anterior to the morphogenetic furrow under the control of Scer\GAL4[h-H10]. Expression of E(spl)[123-179.CtD.Scer\UAS] not only increases eye size (facet numbers), but also restores the patterning of the facets. This patterning is, however, lost toward the anterior margin of the residual eye. The severe reduced eye found in N[spl-1]/Y; E(spl)[1]/+ mutants is unaffected by overexpression of E(spl)[S159D.123-179.CtD.Scer\UAS] under the control of Scer\GAL4[h-H10]. Expression of E(spl)[123-179.CtD.Scer\UAS] in trans, in all cells posterior to the morphogenetic furrow, under the control of Scer\GAL4[GMR.PF] does not rescue the severe reduced eye phenotype found in N[spl-1]/Y; E(spl)[1]/+ mutants. The absence of rescue is also supported by facet numbers, which upon expression of three independent E(spl)[123-179.CtD.Scer\UAS] lines closely mimics control levels in N[spl-1]/Y; E(spl)[1]/+ mutants. Rescue of the N[spl-1]/Y; E(spl)[1]/+ retinal defects by E(spl)[123-179.CtD.Scer\UAS] therefore requires expression in a region of the developing eye where funding R8 photoreceptors are patterned and specified. (Kahali et al., 2010)
Stocks ( 2 )
Bloomington	2447 e⁴ E(spl)m8-HLH¹
Kyoto	107334 e⁴ E(spl)m8-HLH¹
Notes on Origin
Discoverer	Green.

Comments
	Chromosome carries two molecular lesions: 6.4kb insertion in the third intron of gro (gro^E(spl)-1) and a small deletion truncating the adjacent E(spl) transcript at the 3' end (E(spl)¹). (Nagel et al., 1997)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 5 )
Reported As
Symbol Synonym	E(spl)¹ (Moores et al., 2008, Oishi et al., 2006) E(spl)^D (Nagel et al., 1999, Gurganus et al., 1999, Nagel and Preiss, 1999, Grushko et al., 1998, Lai et al., 1998, Fisher and Caudy, 1998, Mackay and Fry, 1996, Oellers et al., 1993, Sturtevant and Bier, 1995, de Celis et al., 1991, Coyle-Thompson and Banerjee, 1993, Campos-Ortega, 1993, Schrons et al., 1992, Duus et al., 1992, Woods and Bryant, 1992, Mari-Beffa et al., 1991, Basler and Hafen, 1991, Baker et al., 1990, Campos-Ortega and Knust, 1990, Xu et al., 1990, Campos-Ortega and Knust, 1990, Shepard et al., 1989, Siren and Portin, 1989, Klambt et al., 1989, Alton et al., 1989, Ziemer et al., 1988, Preiss et al., 1988, Knust et al., 1987, Bose et al., 2006) E(spl)D (Kahali et al., 2010) E(spl)m8-HLH¹ En-spl
Name Synonym
Secondary FlyBase IDs

References ( 41 )

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List References by type	Research paper ( 31 ) Review ( 5 ) Abstract ( 4 ) Book ( 1 )
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Allele Dmel\E(spl)m8-HLH1

Allele Dmel\E(spl)m8-HLH¹