Allele Dmel\ap⁴

General Information
Symbol	Dmel\ap⁴	Species	D. melanogaster
Name		FlyBase ID	FBal0000630
Feature type	allele	Associated gene	Dmel\ap

Allele class	hypomorphic allele - genetic evidence
Mutagen	spontaneous
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	hypomorphic allele - genetic evidence (Ringo et al., 1991)
Mutagen	spontaneous (Ringo et al., 1991)
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
Cytology
Phenotypic Data
Phenotypic Class
	courtship behavior defective \| male \| recessive (Ringo et al., 1992) female sterile (Ringo et al., 1991, Altaratz et al., 1991) female sterile \| recessive (Dubrovsky et al., 2002, ) lethal (with Df(2R)M41A4B) (Ringo et al., 1991) lethal \| adult stage (Wilson, 1982) lethal \| adult stage \| recessive (Richard et al., 1998, Richard et al., 1993) male sterile \| recessive (Ringo et al., 1992, ) mating defective \| female (Ringo et al., 1991, Ringo et al., 1992) uncoordinated (Tompkins, 1990) viable \| reduced (Dubrovsky et al., 2002) visible (with ap^{loxP.MCP-PRE.MM}) (Gohl et al., 2008) visible (with Df(2R)DG) (Gohl et al., 2008) visible (with Df(2R)DG-Mcp) (Gohl et al., 2008) visible \| dominant (Shtorch et al., 1995) visible \| recessive
Phenotype Manifest In
	adult corpus allatum (Altaratz et al., 1991, ) basalar muscle 51 (Ghazi et al., 2000) basalar muscle 52 (Ghazi et al., 2000) corpus allatum (Dai and Gilbert, 1993) coxal tergal remotor muscle (Ghazi et al., 2000) direct flight muscle (Ghazi et al., 2000) direct flight muscle, with Scer\GAL4¹¹⁵¹, ap^Scer\UAS.cOa (Ghazi et al., 2000) direct flight muscle, with Scer\GAL4^pnr-MD237, ap^Scer\UAS.cOa (Ghazi et al., 2000) dorsal medial muscle (Ghazi et al., 2000) dorsal medial muscle, with Scer\GAL4^pnr-MD237, ap^Scer\UAS.cOa (Ghazi et al., 2000) dorsocentral bristle egg chamber (Dubrovsky et al., 2002) embryonic/larval fat body embryonic/larval fat body \| adult stage (Richard et al., 1993) haltere hemolymph (Wilson, 1982) indirect flight muscle (Ghazi et al., 2000) indirect flight muscle, with Scer\GAL4¹¹⁵¹, ap^Scer\UAS.cOa (Ghazi et al., 2000) lateral oblique dorsal muscle (Ghazi et al., 2000) muscle of first axillary 53 (Ghazi et al., 2000) muscle of third axillary 54 (Ghazi et al., 2000) nurse cell ovary (Dubrovsky et al., 2002, Richard et al., 2001) peritrophic membrane scutellar bristle tergosternal muscle (Ghazi et al., 2000) wing (Ringo et al., 1991, Ringo et al., 1992, Shtorch et al., 1995, ) wing (with ap^{loxP.MCP-PRE.MM}) (Gohl et al., 2008) wing (with ap^UG-24A) (Shtorch et al., 1995) wing (with Df(2R)DG) (Gohl et al., 2008) wing (with Df(2R)DG-Mcp) (Gohl et al., 2008)
Detailed Description
	Statement Reference Wing defects are seen in ap[loxP.MCP-PRE.MM]/ap[4] mutant flies, but they are significantly less severe than those seen in ap[loxP.MCP-PRE.MM] homozygotes. Wing defects are seen in Df(2R)DG-Mcp/ap[4] mutant flies, but they are less severe than those seen in Df(2R)DG-Mcp homozygotes. Wing defects are seen in Df(2R)DG/ap[4] mutant flies, but they are less severe than those seen in Df(2R)DG homozygotes. (Gohl et al., 2008) Homozygotes have reduced pre-eclosion viability and live for only about 3 days after eclosion. Homozygous females are sterile; the ovaries are poorly developed, fragile and contain no vitellogenic egg chambers. (Dubrovsky et al., 2002) The application of juvenile hormone III has a significant stimulatory effect on the ovary maturity index in mutant females, primarily due to a change in ovary volume. (Richard et al., 2001) Adults show defects in the direct flight muscles. Of the group of four muscles 51-54, only a sliver of muscle fibre develops. IFMs are reduced in width at the posterior attachment site and do not undergo complete splitting. Sometimes only a single fibre is evident. Muscle size and volume is reduced. (Ghazi et al., 2000) Homozygotes generally die 24 to 48 hours after eclosion. (Richard et al., 1998) Homozygotes and hemizygotes exhibit nubbin wings. Transheterozygotes with ap^UG-24A exhibit a much larger wing blade with deep notches. (Shtorch et al., 1995) Ultrastructure of the corpus allatum studied. (Dai and Gilbert, 1993) Histolysis of the larval fat body is not completed in homozygous flies and they die within 48 hours of eclosion. (Richard et al., 1993) Homozygous ap⁴ males are behaviorally sterile but have fertile gametes. For mating success ap⁴ is an amorph. Homozygous ap⁴ males showed significantly less nonwing courtship than wild type, this was a result of weakness and sluggishness, attributable to the ap locus. Homozygous ap⁴ males had higher immature sex appeal and rear displays component of courtship at 2--3 days old than mature wild type flies. (Ringo et al., 1992) Blocks vitellogenesis causing an egg maturation defect. Corpora allata are defective in hormone production. (Altaratz et al., 1991) Homozygous ap⁴ flies have virtually no wing blade and have only 13% female receptivity to mature male ap⁺ flies. Female flies have a high courtship intensity, but low percentage mating: this is due to low receptivity not low 'sex appeal'. Survival of ap⁴/Df(2R)M41A4B is only 1% at 3 days of age. (Ringo et al., 1991) Pleiotropic effects on the morphology and lifespan of adult flies. Mutant males spend less time courting and are less likely to perform some of the courtship behaviours than age-matched controls. Abnormalities are an indirect effect of the mutation which causes partial paralysis, lack of coordination and sluggishness. (Tompkins, 1990) Abnormal haemolymph. (Wilson, 1982) Wings less than 10% normal length, lacking all wing blade structures. Halteres reduced to structureless remnants less than 25% normal size. Scutellar and dorsocentral bristles sometimes missing (FBrf0017003). Wing phenotype disc-autonomous in ap⁴/ap⁺ mosaic flies, although small patches of ap⁴ wing structures are found in ap⁴/ap⁺ mosaic wings. Haltere phenotype disc-autonomous (FBrf0036211). Adults become paralyzed about 30 hr following eclosion and die soon thereafter. Around 1% of adults are long-lived 'escapers' of this phenotype (FBrf0034522). Precocious adult-death phenotype fate-maps to proximity of Malpighian tubules and tubule malfunctioning postulated to result in this phenotype (FBrf0036211). Foregut of females swollen owing to accumulation of peritrophic membrane (FBrf0011908). Female sterile with underdeveloped ovaries; nurse cell nuclei become pycnotic after stage 7 and stage-8 oocytes are the most advanced (FBrf0011400; FBrf0034522). Ovaries develop nonautonomously when transplanted to a wild-type host (FBrf0017349). Application of juvenile hormone analog, ZR-515, to newly eclosed females results in vitellogenic oocytes (FBrf0025594). Membranes of vitellogenic oocytes lack microvilli and pinocytoxic vesicles normally present; development of these structures stimulated by administration of ZR-515 (FBrf0036874). Corpora allata from adult ap⁴ are juvenile-hormone deficient when bioassayed (FBrf0028948). Nonvitellogenic oocyte phenotype fate-maps to same or similar location as precocious adult death phenotype (FBrf0036211). Escaper females develop stage-14 oocytes (FBrf0011908) and are fertile (FBrf0034522). Males show immature sexual behavior and are sterile, but testes appear normal with motile sperm (FBrf0011908). Larval fat body histolysis delayed; this phenotype is nonautonomous as determined by transplantation experiments (FBrf0023756). Application of ZR-515 accelerates larval fat body histolysis in adults (FBrf0032050). Ovarian acid phosphatase level low in females and is restored after application of ZR-515 (FBrf0028948). Ovaries cultured in vitro are capable of yolk protein synthesis (FBrf0064836). ap⁴/Df(2R)M41A4 hemizygote has nearly normal complement of bristles but otherwise resembles ap⁴ homozygote (FBrf0017003). RK2.
External Data
Linkouts
Interactions
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Phenotypic Class
Other
Statement Reference ap[+]/ap⁴, apb¹ has fertile phenotype (Meyer et al., 1950) ap[+]/ap⁴, apb¹ has visible \| dominant phenotype (Meyer et al., 1950) ap⁴, apb[+]/apb¹ has fertile phenotype (Meyer et al., 1950) ap⁴, apb[+]/apb¹ has visible \| dominant phenotype (Meyer et al., 1950)
Phenotype Manifest In
Enhanced by
Statement Reference ap⁴ has indirect flight muscle phenotype, enhanceable by sr[+]/sr^G11 (Ghazi et al., 2000) ap⁴ has indirect flight muscle phenotype, enhanceable by sr⁰³⁹⁹⁹/sr⁰³⁹⁹⁹ (Ghazi et al., 2000) ap⁴ has phenotype, enhanceable by apb¹
Enhancer of
Statement Reference ap⁴/ap⁴ is an enhancer of adult epidermis phenotype of sr⁰³⁹⁹⁹ (Ghazi et al., 2000, Ghazi et al., 2000) ap⁴/ap⁴ is an enhancer of mesothoracic tergum phenotype of sr⁰³⁹⁹⁹ (Ghazi et al., 2000, Ghazi et al., 2000)
Other
Statement Reference ap[+]/ap⁴, apb¹ has wing phenotype (Meyer et al., 1950) ap⁴, apb[+]/apb¹ has wing phenotype (Meyer et al., 1950)
Additional Comments
Genetic Interactions
Statement Reference No wing scalloping is seen when combined with Ssdp^KG03600, Ssdp^BG01663, Ssdp^neo48, Ssdp³¹ or Ssdp¹¹. (Chen et al., 2002) A significant proportion of ap⁴/+, sr^G11/+ flies show defective IFMs. A significant proportion of ap⁴/ap⁴, sr⁰³⁹⁹⁹/sr⁰³⁹⁹⁹ flies show an enhanced IFM mutant phenotype and a dark mid-notal stripe characteristic of strong sr mutants. (Ghazi et al., 2000) apb¹/ap⁴ double heterozygotes show slight notching of the wings and many die within a day, but those that survive are fertile. (Meyer et al., 1950)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	ap^2a1 (Coelho et al., 2005) ap^2a2 (Coelho et al., 2005) ap⁵ (Meyer et al., 1950) ap^41F (Bourgouin et al., 1992) ap^56f (Gohl et al., 2008)
Partially rescued by	ap⁴ is partially rescued by ap^Scer\UAS.cOa/Scer\GAL4¹¹⁵¹ (Ghazi et al., 2000) ap⁴ is partially rescued by ap^Scer\UAS.cOa/Scer\GAL4^pnr-MD237 (Ghazi et al., 2000)
Comments	ap^Scer\UAS.cOa, when expressed via Scer\GAL4¹¹⁵¹, partially rescues the DFM loss phenotype of ap⁴ mutants (though a large single muscle mass develops as opposed to four discrete fibres) but does not rescue the IFM phenotype. When expression of ap^Scer\UAS.cOa is driven by Scer\GAL4^pnr-MD237, the DLM defects of ap⁴ mutants are significantly rescued (though a large single muscle mass develops as opposed to four discrete fibres) while DFM fibres remain disorganised. Muscle volume is significantly restored. (Ghazi et al., 2000)
Stocks ( 3 )
Bloomington	223 ap⁴/SM5 36522 ap⁴/CyO, P{ActGFP}JMR1; P{GawB}EM461
Kyoto	105797 ap⁴/SM5
Notes on Origin
Discoverer	Medvedev, 15th Jan. 1932.

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	ap⁴ (Flatt et al., 2005, Gohl et al., 2008) ap-d
Name Synonym
Secondary FlyBase IDs

References ( 31 )

Generate a list of	All references relating to this allele ( 31 )

List References by type	Research paper ( 25 ) Review ( 3 ) Abstract ( 2 ) Book ( 1 )
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Allele Dmel\ap4

Allele Dmel\ap⁴