Allele Dmel\N^Ax-1

General Information
Symbol	Dmel\N^Ax-1	Species	D. melanogaster
Name		FlyBase ID	FBal0012850
Feature type	allele	Associated gene	Dmel\N
Also Known As	Ax²⁸, Ax¹, Ax^28a, N^Ax1, N^Ax, N^Ax28, Ax

Map ( GBrowse )
Allele class	antimorphic allele - genetic evidence, gain of function allele
Mutagen	spontaneous
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	antimorphic allele - genetic evidence (Mari-Beffa et al., 1991, de Celis and Garcia-Bellido, 1994, de Celis et al., 1991) gain of function allele (Mari-Beffa et al., 1991, de Celis et al., 1996, de Celis, 1997, de Celis et al., 1997, Royet et al., 1998)
Mutagen	spontaneous (Lindsley and Zimm, 1992)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation X:3,059,618..3,059,618 comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. evidence=experimental na_change=A3059618T pr_change=N986I\|N-PA reported_pr_change=N986I (Lindsley and Zimm, 1992)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Amino acid replacement: N986I. N986 lies in EGF repeat 25. Has Gly at residue 2057 characteristic of Oregon R rather than Ser of Canton S (FBrf0045770). (Lindsley and Zimm, 1992) Amino acid replacements in the EGF-homologous regions of the extracellular domain of the N protein. (de Celis et al., 1991)
Cytology	A single-band duplication, presumed to be for 3C7 by Schultz (FBrf0005574). Lefevre et al., (FBrf0009019), on the other hand, argue against a duplication for 3C7 on the basis of equal X-ray-induced mutability to N in 'Ax' and +. Molecular information also incompatible with the presence of a duplication. (Lindsley and Zimm, 1992)
Phenotypic Data
Phenotypic Class
	lethal \| partially (with N^Ax-16) (Grushko et al., 1998) lethal \| partially (with N^Ax-71d) (Grushko et al., 1998) lethal \| partially (with N^Ax-E2) (Grushko et al., 1998) lethal \| recessive (Kramers et al., 1983) lethal \| recessive \| heat sensitive (Lindsley and Zimm, 1992, Portin and Siren, 1976) planar polarity defective (Fanto and Mlodzik, 1999) viable (de Celis and Bray, 2000, de Celis et al., 1996, Grushko et al., 1998) viable (with N^Ax-9) (Grushko et al., 1998) viable \| reduced (with N^Ax-16) (Grushko et al., 1998) viable \| reduced (with N^Ax-71d) (Grushko et al., 1998) viable \| reduced (with N^Ax-E2) (Grushko et al., 1998) visible (Grushko et al., 1998, Palka et al., 1990, Adachi-Yamada et al., 1999, Thumm and Kadowaki, 2001) visible \| dominant (Royet et al., 1998, Guo et al., 1999) visible \| heat sensitive (Thompson et al., 1988) visible \| recessive \| heat sensitive (Lindsley and Zimm, 1992)
Phenotype Manifest In
	dorsocentral bristle (Royet et al., 1998) macrochaeta (de Celis and Garcia-Bellido, 1994, Grushko et al., 1998, Lindsley and Zimm, 1992) ommatidium (Fanto and Mlodzik, 1999) orbital bristle (Royet et al., 1998) photoreceptor cell R3 (Fanto and Mlodzik, 1999) sensillum campaniformium of anterior crossvein (Palka et al., 1990) twin sensillum of margin 1 (Palka et al., 1990) twin sensillum of margin 2 (Palka et al., 1990) wing (Adachi-Yamada et al., 1999) wing disc (Go et al., 1998) wing vein (Grushko et al., 1998, Royet et al., 1998, Sturtevant and Bier, 1995, Miyamoto et al., 1995, Lindsley and Zimm, 1992, Thumm and Kadowaki, 2001) wing vein L2 (Guo et al., 1999) wing vein L4 (Morey et al., 2001) wing vein L5 (Morey et al., 2001, Guo et al., 1999) wing vein L5 \| heat sensitive (Thompson et al., 1988)
Detailed Description
	Statement Reference When grown at 25^oC, mutants lack the distal most parts of wing vein L5 band L4. (Morey et al., 2001) N^Ax-1 flies have mild eye-polarity defects; ommatidia are sometimes symmetrical, having an R4/R4-like phenotype. (Fanto and Mlodzik, 1999) Wing vein L2 is absent and wing vein L5 is interrupted. (Guo et al., 1999) N^Ax-1/N^Ax-16 larvae have enlarged wing discs compared to wild-type. This phenotype is associated with elevated mitotic activity in the disc, particularly in the peripheral region of the wing pouch. (Go et al., 1998) Viable in combination with N^Ax-9. (Grushko et al., 1998) The number of missing bristles increases with increasing temperature in homozygous flies, while the number and length of wing vein breaks increases from 18^oC to 25^oC and then decreases again from 25^oC to 29^oC. (Grushko et al., 1998) Reduced number of bristles on the head and thorax and shortening of wing veins. (Royet et al., 1998) Homozygotes form wing vein gaps. (Miyamoto et al., 1995) Thickened vein mutant. (Sturtevant and Bier, 1995) Length of macrochaetae is reduced with respect to that of wild type. Suppresses the N haplo-insufficient phenotype of loss of the wing margin. (de Celis and Garcia-Bellido, 1994) Neural precursors never form in the Ax class of N alleles. Cells mutant for Ax class alleles but with some neural potential are inhibited from becoming neural by their neighbours but do not themselves affect their neighbours, which can become neural. (Heitzler and Simpson, 1993) sgg³² is epistatic to N^Ax-1. (Ruel et al., 1993) Male viable at 25^oC but nearly lethal at 29^oC. N^Ax-1/N^Ax-E2 semi-lethal at 25^oC and lethal at 29^oC. N^Ax-1/+ females show short L5 in half of the flies and sparse hair pattern on thorax. Lower temperature (19^oC) markedly decreases expression, and higher temperature enhances it. Some Ax alleles enhance N expression in N^Ax-1/N heterozygotes, but others suppress the dominant N phenotype. For example, N^Ax-1/Df(1)N-8 approaches wild type in all characteristics. No wing-vein interruption in N^Ax-1/+ at 18^oC and 26^oC and enhancement by H¹ occurs. N^Ax-1/N^Ax-1; ci^D/+ and N^Ax-1/Y; ci^D/+ are lethal or nearly so at 26^oC. At 22^oC, males survive and show enhanced wing-vein interruption and more missing bristles. Wing nicking is suppressed in N^Ax-1/N^55e11 at 25^oC and Ax venation is weakly expressed. In heterozygotes of N^Ax-1 with the recessives at N at 18^oC and 25^oC, there is neither expression of the recessive nor Ax-type venation. Temperature-sensitive period for lethality of N^Ax-1 at beginning of pupal stage; of N^Ax-1/N^Ax-E2 at end of third instar and into early pupal stage. (Lindsley and Zimm, 1992) Lack of chaetae. N^Ax-1 and N^Ax-16 correct Hw phenotypes in both ectopic and normal positions. (de Celis et al., 1991) Phenotypes have topographic pattern specificities. (Mari-Beffa et al., 1991) Wing phenotype is enhanced by mutations at da. (Brand and Campos-Ortega, 1990) Some hemizygous males occasionally lack the twin sensilla of the wing margin, the ventral sensillum of wing vein L3, and the anterior cross vein sensillum. (Palka et al., 1990) The L5 wing vein phenotype is more severe at 25^oC than 20^oC, the temperature-sensitive period is between 10 and 15% of pupal development time. (Thompson et al., 1988) All heterozygotes with facet or notchoid alleles except N^Ax-1/N^nd-2 show some Ax venation. (Portin, 1977) Nearly lethal when reared at 29^oC; temperature-sensitive period early pupa. (Portin and Siren, 1976) N^Ax-1/N^Ax-1; Dp(1;2)51b/+ shows weak Ax venation. (Portin, 1975)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference N^Ax-1/N^Ax-16 has lethal \| partially phenotype, enhanceable by Dp(3;3)bxd¹¹⁰ (Grushko et al., 1998) N^Ax-1/N^Ax-16 has lethal \| partially phenotype, enhanceable by Dp(3;3)MKRS-D2 (Grushko et al., 1998) N^Ax-1/N^Ax-E2 has lethal \| partially phenotype, enhanceable by Dp(3;3)bxd¹¹⁰ (Grushko et al., 1998) N^Ax-1/N^Ax-E2 has lethal \| partially phenotype, enhanceable by Dp(3;3)MKRS-D2 (Grushko et al., 1998) N^Ax-71d/N^Ax-1 has lethal \| partially phenotype, enhanceable by Dp(3;3)bxd¹¹⁰ (Grushko et al., 1998) N^Ax-71d/N^Ax-1 has lethal \| partially phenotype, enhanceable by Dp(3;3)MKRS-D2 (Grushko et al., 1998)
Suppressed by
Statement Reference N^Ax-1 has visible phenotype, suppressible by CG6194^P0997/CG6194^P0997 (Thumm and Kadowaki, 2001)
NOT Enhancer of
Statement Reference N^Ax-1 is a non-enhancer of visible phenotype of Scer\GAL4^455.2, E(spl)mα-BFM^Scer\UAS.cAa (Apidianakis et al., 1999)
Suppressor of
Statement Reference N^Ax-1/N[+] is a suppressor \| partially of visible \| dominant phenotype of fng^D4 (Dansereau et al., 2002) N^Ax-1 is a suppressor of visible phenotype of Scer\GAL4^455.2, E(spl)m4-BFM^Scer\UAS.cAa (Apidianakis et al., 1999)
NOT Suppressor of
Statement Reference N^Ax-1 is a non-suppressor of visible phenotype of Scer\GAL4^455.2, E(spl)mα-BFM^Scer\UAS.cAa (Apidianakis et al., 1999)
Other
Statement Reference Dp(3;3)bxd¹¹⁰, N^Ax-1 has lethal \| partially phenotype (Grushko et al., 1998) Dp(3;3)MKRS-D2, N^Ax-1 has lethal \| partially phenotype (Grushko et al., 1998)
Phenotype Manifest In
Enhanced by
Statement Reference N^Ax-1 has dorsocentral bristle phenotype, enhanceable by Nle[+]/Nle^k13714 (Royet et al., 1998) N^Ax-1 has dorsocentral bristle phenotype, enhanceable by Nle[+]/Nle^Δ8 (Royet et al., 1998) N^Ax-1 has dorsocentral bristle phenotype, enhanceable by Scer\GAL4^ap-md544/Nle^Scer\UAS.cRa (Royet et al., 1998) N^Ax-1 has orbital bristle phenotype, enhanceable by Nle[+]/Nle^k13714 (Royet et al., 1998) N^Ax-1 has orbital bristle phenotype, enhanceable by Nle[+]/Nle^Δ8 (Royet et al., 1998) N^Ax-1 has orbital bristle phenotype, enhanceable by Scer\GAL4^ap-md544/Nle^Scer\UAS.cRa (Royet et al., 1998) N^Ax-1 has wing vein L5 phenotype, enhanceable by ab¹ (Thompson et al., 1988) N^Ax-1 has wing vein phenotype, enhanceable by Nle[+]/Nle^k13714 (Royet et al., 1998) N^Ax-1 has wing vein phenotype, enhanceable by Nle[+]/Nle^Δ8 (Royet et al., 1998) N^Ax-1 has wing vein phenotype, enhanceable by Scer\GAL4^ap-md544/Nle^Scer\UAS.cRa (Royet et al., 1998)
NOT Enhanced by
Statement Reference N^Ax-1 has wing vein L4 phenotype, non-enhanceable by SelD^k11320 (Morey et al., 2001) N^Ax-1 has wing vein L5 phenotype, non-enhanceable by SelD^k11320 (Morey et al., 2001)
Suppressed by
Statement Reference N^Ax-1 has wing vein L2 phenotype, suppressible \| partially by Psn^W6 (Guo et al., 1999) N^Ax-1 has wing vein L2 phenotype, suppressible \| partially by Psn^W11 (Guo et al., 1999) N^Ax-1 has wing vein L5 phenotype, suppressible \| partially by Psn^W6 (Guo et al., 1999) N^Ax-1 has wing vein L5 phenotype, suppressible \| partially by Psn^W11 (Guo et al., 1999) N^Ax-1 has wing vein phenotype, suppressible by CG6194^P0997/CG6194^P0997 (Thumm and Kadowaki, 2001) N^Ax-1 has wing vein phenotype, suppressible by cno^mis1 (Miyamoto et al., 1995)
NOT suppressed by
Statement Reference N^Ax-1 has wing phenotype, non-suppressible by p38b^DN.Scer\UAS/Scer\GAL4^unspecified (Adachi-Yamada et al., 1999) N^Ax-1 has wing vein L4 phenotype, non-suppressible by SelD^k11320 (Morey et al., 2001) N^Ax-1 has wing vein L5 phenotype, non-suppressible by SelD^k11320 (Morey et al., 2001)
Enhancer of
Statement Reference N^Ax-1 is an enhancer of phenotype of emc^D (Mari-Beffa et al., 1991) N^Ax-1 is an enhancer of phenotype of H^unspecified (Mari-Beffa et al., 1991)
Suppressor of
Statement Reference N^Ax-1/N[+] is a suppressor \| partially of wing margin phenotype of fng^D4 (Dansereau et al., 2002) N^Ax-1/N[+] is a suppressor of wing margin phenotype of Scer\GAL4^Bx-MS1096, rho^Neo.Scer\UAS (Guichard et al., 2002) N^Ax-1/N[+] is a suppressor of wing phenotype of Scer\GAL4^Bx-MS1096, rho^Neo.Scer\UAS (Guichard et al., 2002) N^Ax-1/N[+] is a suppressor of wing vein phenotype of Scer\GAL4^Bx-MS1096, rho^Neo.Scer\UAS (Guichard et al., 2002) N^Ax-1 is a suppressor of phenotype of emc^unspecified (Mari-Beffa et al., 1991) N^Ax-1 is a suppressor of scutellar bristle phenotype of Scer\GAL4^455.2, E(spl)m4-BFM^Scer\UAS.cAa (Apidianakis et al., 1999) N^Ax-1 is a suppressor of wing vein phenotype of tkv¹/tkv⁸ (de Celis, 1997)
Other
Statement Reference N^Ax-1, dpp^s4/dpp^s8 has wing vein phenotype (de Celis, 1997, de Celis, 1997)
Additional Comments
Genetic Interactions
Statement Reference CG6194^P0997/CG6194^P0997 suppresses the loss of wing vein seen in N^Ax-1/Y flies. (Thumm and Kadowaki, 2001) Expression of p38b^DN.Scer\UAS under the control of Scer\GAL4^unspecified has no significant effect on the wing phenotype caused by N^Ax-1. (Adachi-Yamada et al., 1999) N^Ax-1 suppresses the supernumerary scutellar bristle phenotype seen in flies with m4^Scer\UAS.cAa driven by Scer\GAL4^455.2. The addition of N^Ax-1 reduces the number of supernumerary scutellar bristles seen in flies with mα^Scer\UAS.cAa driven by Scer\GAL4^455.2, though there are still more bristles seen than in wild-type. (Apidianakis et al., 1999) The wing vein phenotypes of heterozygotes are partially suppressed by Psn^W6 and Psn^W11. (Guo et al., 1999) Homozygous viability is decreased if the flies also carry Dp(3;3)MKRS-D2 or Dp(3;3)bxd¹¹⁰. (Grushko et al., 1998) 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. (Grushko et al., 1998) dpp^s4/dpp^s8 N^Ax-1 or tkv¹/tkv⁸ N^Ax-1 double mutant flies show pronounced wing vein loss. The thickening of veins seen in tkv¹/tkv⁸ flies is suppressed in combination with N^Ax-1. (de Celis, 1997) The wing vein gaps of N^Ax-1 homozygotes are dominantly suppressed by cno^mis1. (Miyamoto et al., 1995) The L5 wing vein phenotype is more extreme in N^Ax-1 ab¹ double mutants, and there is a deep middorsal furrowing of the thorax (a trait characteristic of some Abruptex alleles), suggesting that ab¹ acts as an enhancer of the N^Ax-1 phenotype. (Thompson et al., 1988)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	N^264-8 (Slizynska, 1938) N^264-39 (Slizynska, 1938)
Comments
Stocks ( 0 )

Notes on Origin
Discoverer	Nazarenko, Jan. 1928.

Comments
	N^Ax-1 shows negative complementation with N^Ax-M1 and N^Ax-E2. (de Celis and Garcia-Bellido, 1994)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 11 )
Reported As
Symbol Synonym	Ax (Kramers et al., 1983, Slizynska, 1938) Ax¹ (Adachi-Yamada et al., 1999, Matsuo et al., 1995, Brand and Campos-Ortega, 1990, Thompson et al., 1988, ) Ax²⁸ (de Celis and Bray, 2000, Grushko et al., 1998, Royet et al., 1998, Grushko et al., 1998, de Celis et al., 1997, de Celis, 1997, Gomez-Skarmeta and Modolell, 1996, de Celis et al., 1996, de Celis et al., 1995, de Celis et al., 1991, de Celis and Garcia-Bellido, 1994, de Celis et al., 1993, Heitzler and Simpson, 1993, ) Ax^28a (Go et al., 1998, Mari-Beffa et al., 1991, Palka et al., 1990) Dp(1;1)Ax28a Dp(1;1)Ax^28a N^Ax1 (Thumm and Kadowaki, 2001, Guichard et al., 2002) N^Ax-1 N^Ax28 (Apidianakis et al., 1999, Fanto and Mlodzik, 1999) N^Ax-28a N^Ax (Apidianakis et al., 1999, Sturtevant and Bier, 1995)
Name Synonym
Secondary FlyBase IDs
	FBab0003034 FBal0029937
References ( 43 )

Generate a list of	All references relating to this allele ( 43 )

List References by type	Research paper ( 39 ) Abstract ( 2 ) Book ( 1 ) Letter ( 1 )
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Allele Dmel\NAx-1

Allele Dmel\N^Ax-1