Allele Dmel\sax⁴

General Information
Symbol	Dmel\sax⁴	Species	D. melanogaster
Name		FlyBase ID	FBal0050458
Feature type	allele	Associated gene	Dmel\sax

Map ( GBrowse )
Allele class	loss of function allele, amorphic allele - genetic evidence, amorphic allele
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 (Fritsch et al., 2010) amorphic allele - genetic evidence (Singer et al., 1997) loss of function allele (Horsfield et al., 1998)
Mutagen	ethyl methanesulfonate (Twombly et al., 1996, Singer et al., 1997, Twombly et al., 2009)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 2R:3,695,285..3,695,285 na_change=C3695285T reported_na_change=C?T pr_change=Q114 \| sax-PA; Q79 \| sax-PB; Q126@ \| sax-PC reported_pr_change=Q114@ (Twombly et al., 2009)
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: Q114@. (Twombly et al., 2009) A premature stop codon in the extracellular domain. (Twombly et al., 2009)
Cytology
Phenotypic Data
Phenotypic Class
	developmental rate defective \| larval stage (with Df(2R)H23) (Twombly et al., 2009) lethal (with Df(2R)sax^1rv2) (Twombly et al., 2009) lethal (with Df(2R)sax^PE7) (Twombly et al., 2009) lethal (with Df(2R)sax-H9) (Twombly et al., 2009) lethal (with sax^1rv5) (Twombly et al., 2009) lethal (with sax^PE5) (Twombly et al., 2009) lethal (with sax^PE10) (Twombly et al., 2009) lethal \| larval stage (with Df(2R)H23) (Twombly et al., 2009) lethal \| larval stage (with sax^1rv1) (Twombly et al., 2009) lethal \| larval stage (with sax³) (Twombly et al., 2009) lethal \| larval stage (with sax⁵) (Twombly et al., 2009) lethal \| larval stage (with sax⁶) (Twombly et al., 2009) lethal \| P-stage (with Df(2R)H23) (Twombly et al., 2009) lethal \| P-stage (with sax^1rv1) (Twombly et al., 2009) lethal \| P-stage (with sax³) (Twombly et al., 2009) lethal \| P-stage (with sax⁵) (Twombly et al., 2009) lethal \| P-stage (with sax⁶) (Twombly et al., 2009) lethal \| recessive (Singer et al., 1997, Dworkin and Gibson, 2006) neuroanatomy defective (with Df(2R)cn7969) (Sweeney and Davis, 2002) neuroanatomy defective (with Df(2R)sax-H9) (McCabe et al., 2004) neurophysiology defective (with sax⁵) (McCabe et al., 2004) partially lethal - majority die (with sax^P) (Twombly et al., 2009) size defective \| larval stage (with Df(2R)H23) (Twombly et al., 2009) viable (with sax¹) (Twombly et al., 2009) viable (with sax²) (Twombly et al., 2009) visible \| dominant (Dworkin and Gibson, 2006) visible \| recessive \| somatic clone (Ray and Wharton, 2001)
Phenotype Manifest In
	amnioserosa (Dorfman and Shilo, 2001) bouton (with Df(2R)cn7969) (Sweeney and Davis, 2002) brain \| larval stage (with Df(2R)H23) (Twombly et al., 2009) distal costal vein \| ectopic (Singer et al., 1997) dorsal appendage \| germline clone (Twombly et al., 1996) dorsal double row (Singer et al., 1997) egg \| germline clone (Twombly et al., 1996) imaginal disc (with Df(2R)H23) (Twombly et al., 2009) male germline stem cell \| germline clone (Kawase et al., 2004) neuromuscular junction (with Df(2R)cn7969) (Sweeney and Davis, 2002) neuromuscular junction (with Df(2R)sax-H9) (McCabe et al., 2004) ovary \| germline clone (Twombly et al., 1996) trachea \| larval stage (with Df(2R)H23) (Twombly et al., 2009) trichome of the posterior wing margin (Singer et al., 1997) triple row \| ectopic (Singer et al., 1997) ventral double row (Singer et al., 1997) wing (Singer et al., 1997, Dworkin and Gibson, 2006) wing \| somatic clone (Ray and Wharton, 2001) wing vein \| cell non-autonomous \| ectopic \| somatic clone (Bangi and Wharton, 2006) wing vein \| ectopic (Singer et al., 1997) wing vein \| ectopic \| somatic clone (Ray and Wharton, 2001)
Detailed Description
	Statement Reference Large posterior clones of sax[4] show no wing patterning abnormalities. Less than 5% of sax[4]/Df(2R)H23 hemizygous mutant larvae exhibit developmental delay, lethargy, a reduction in the size of imaginal discs, brain \| larval stage, and midgut structures, as well as trachea \| larval stagel truncations. (Twombly et al., 2009) When the anterior border of a sax⁴ clone falls between L2 and L3 in the wing, an ectopic L2 may form at the anterior boundary of the clone. sax⁴ clones near but anterior to L2 result in the formation of an ectopic wing vein adjacent to their anterior border. These clones often exhibit non-autonomous effects such as ectopic veins forming in wild-type tissue outside the boundaries of the clone. (Bangi and Wharton, 2006) Heterozygotes show a direct effect on the shape of the wing. (Dworkin and Gibson, 2006) When neutral marked clones are induced in the ovary, the proportion of germaria carrying marked somatic stem cells 3 weeks after clone induction is around 70% of that seen one week after clone induction. For sax⁴ homozygous clones, the equivalent figure is around 60%. (Kirilly et al., 2005) Clones of male sax⁴ homozygous germline stem cells are still present in 32% of testes one week after clone induction and 6.3% two weeks after clone induction. This is in contrast to wild-type control clones, which are present in 82% of testes one week after clone induction and 64% two weeks after clone induction. (Kawase et al., 2004) sax⁴/Df(2R)sax-H9 larvae show a reduction in size of the neuromuscular junction (NMJ) compared to wild type; the number of synaptic boutons/muscle surface area at muscle 6/7 is 44.1 +/- 1.0% of wild type. The evoked excitatory junctional potential (EJP) (measured at muscle 6 of segment A3) shows a decrease in amplitude in sax⁴/sax⁵ animals compared to wild type. Quantal content is reduced compared to wild type. (McCabe et al., 2004) Mutant stage 13-14 embryos contain the normal number of crystal cells per embryo. (Milchanowski et al., 2004) sax⁴/Df(2R)cn7969 larvae show a significant decrease in bouton number at the neuromuscular junction. (Sweeney and Davis, 2002) Mutant embryos lacking both maternal and zygotic sax function have a reduced number of amnioserosa cells. (Dorfman and Shilo, 2001) Homozygous clones that occupy the entire posterior compartment of the wing have no effect on venation. Homozygous clones that occupy the entire anterior compartment of the wing have no effect on venation, but the wing is reduced in size. Homozygous wing clones where the clone boundary subdivides a compartment result in an ectopic wing vein at the clone boundary. (Ray and Wharton, 2001) Clones induced in the developing eye that span the morphogenetic furrow have condensed chromosomes indicative of early stages in mitosis at the interface between the CycB-expressing and non-expressing cells. Clones at the anterior edge of the furrow show mislocalized nuclei, they fail to reach the apical surface where mitosis normally takes place. Nuclei are also misplaced when the clone is within the furrow and posterior to it. Cell fate specification is not, however, affected. (Penton et al., 1997) A large anterior clone in the wing (induced early in larval development) reduces the size of the wing, blunts the wing tip and causes ectopic venation. At the anterior clone boundary cells shift to a more anterior cell fate. At the anterior edge of clones that intersect the margin several margin cells posterior to wing vein L3 produce double row bristles (and not the posterior row of hairs), at other points of the wing margin double row cells fates are transformed to triple row. At the anterior edge of clones adjacent to the naked stretch of margin, ectopic distal costa are produced in the naked region. A large posterior clone in the wing (induced early in larval development) variably disrupts wing venation. Dorsal or ventral clones located anterior to wing vein L2 or dorsal clones posterior to wing vein L5 do not cause ectopic venation. Clones posterior to L2 or anterior of L5 do allow ectopic venation. Where the clone diverges from the line of the normal vein the ectopic vein is produced as a ridge on the dorsal surface. (Singer et al., 1997) In germline clones, only 10-25% of normal number of eggs is produced, over a much briefer time period than for wild type. Most are normal though 20% show aberrant dorsal appendages and short length. (Twombly et al., 1996)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressed by
Statement Reference sax⁴/Df(2R)BSC265 has lethal phenotype, suppressible by sax::Agam\sax^{AgamEX.DmelIN.sax} (Fritsch et al., 2010)
Enhancer of
Statement Reference sax⁴/sax[+] is an enhancer of visible phenotype of Scer\GAL4^A9, gbb^Scer\UAS.cKa (Bangi and Wharton, 2006)
NOT Enhancer of
Statement Reference sax⁴/sax[+] is a non-enhancer of visible phenotype of I-2^{Scer\UAS.T:Zzzz\His6,T:Hsap\MYC}, Scer\GAL4^vg.PM, put^Scer\UAS.cMa (Bennett et al., 2003)
Suppressor of
Statement Reference sax⁴/Df(2R)cn7969 is a suppressor of neuroanatomy defective phenotype of spict^mut (Wang et al., 2007) sax⁴/Df(2R)cn7969 is a suppressor of neuroanatomy defective phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴/sax[+] is a suppressor \| partially of female sterile phenotype of mir-184^Δ (Iovino et al., 2009) sax⁴/sax[+] is a suppressor \| partially of neuroanatomy defective phenotype of spict^mut (Wang et al., 2007) sax⁴/sax[+] is a suppressor \| partially of visible phenotype of Scer\GAL4^A9, dpp^Scer\UAS.cHa (Bangi and Wharton, 2006) sax⁴/sax[+] is a suppressor of neuroanatomy defective phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴ is a suppressor of visible phenotype of os^GMR.PB (Bach et al., 2003)
NOT Suppressor of
Statement Reference sax⁴/sax[+] is a non-suppressor of visible phenotype of I-2^{Scer\UAS.T:Zzzz\His6,T:Hsap\MYC}, Scer\GAL4^vg.PM, put^Scer\UAS.cMa (Bennett et al., 2003)
Other
Statement Reference gbb⁴, sax⁴ has viable \| poor phenotype (Khalsa et al., 1998) gbb⁴/gbb¹, sax⁴ has viable \| poor phenotype (Khalsa et al., 1998) sax⁴, wit^HA3/wit[+] has neuroanatomy defective \| dominant phenotype (McCabe et al., 2004) sax⁴/sax[+], wit^HA3 has neuroanatomy defective phenotype (McCabe et al., 2004)
Phenotype Manifest In
Suppressed by
Statement Reference sax⁴ has wing vein \| cell non-autonomous \| ectopic \| somatic clone phenotype, suppressible by gbb⁴/gbb⁴ (Bangi and Wharton, 2006)
Enhancer of
Statement Reference sax⁴/sax[+] is an enhancer of wing vein phenotype of Scer\GAL4^A9, gbb^Scer\UAS.cKa (Bangi and Wharton, 2006)
NOT Enhancer of
Statement Reference sax⁴/sax[+] is a non-enhancer of wing phenotype of I-2^{Scer\UAS.T:Zzzz\His6,T:Hsap\MYC}, Scer\GAL4^vg.PM, put^Scer\UAS.cMa (Bennett et al., 2003)
Suppressor of
Statement Reference sax⁴/Df(2R)cn7969 is a suppressor of bouton \| supernumerary phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴/Df(2R)cn7969 is a suppressor of embryonic/larval neuromuscular junction phenotype of spict^mut (Wang et al., 2007) sax⁴/Df(2R)cn7969 is a suppressor of neuromuscular junction phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴/sax[+] is a suppressor \| partially of embryonic/larval neuromuscular junction phenotype of spict^mut (Wang et al., 2007) sax⁴/sax[+] is a suppressor \| partially of wing phenotype of Scer\GAL4^A9, dpp^Scer\UAS.cHa (Bangi and Wharton, 2006) sax⁴/sax[+] is a suppressor \| partially of wing vein phenotype of Dd^G0269 (Liu et al., 2011) sax⁴/sax[+] is a suppressor of bouton \| supernumerary phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴/sax[+] is a suppressor of neuromuscular junction phenotype of spin^k09905/spin¹⁰⁴⁰³ (Sweeney and Davis, 2002) sax⁴ is a suppressor \| partially of wing phenotype of dpp^d5/dpp^hr56 (Twombly et al., 2009) sax⁴ is a suppressor of eye phenotype of os^GMR.PB (Bach et al., 2003)
NOT Suppressor of
Statement Reference sax⁴/sax[+] is a non-suppressor of wing phenotype of I-2^{Scer\UAS.T:Zzzz\His6,T:Hsap\MYC}, Scer\GAL4^vg.PM, put^Scer\UAS.cMa (Bennett et al., 2003)
Other
Statement Reference sax⁴, wit^HA3/wit[+] has bouton phenotype (McCabe et al., 2004) sax⁴, wit^HA3/wit[+] has neuromuscular junction phenotype (McCabe et al., 2004) sax⁴, wit^HA3 has bouton phenotype (McCabe et al., 2004) sax⁴, wit^HA3 has neuromuscular junction phenotype (McCabe et al., 2004)
Additional Comments
Genetic Interactions
Statement Reference The aberrant wing vein phenotype seen in Dd[G0269]/Y animals is suppressed by sax[4]/+. (Liu et al., 2011) One copy of sax[4] substantially rescues the infertility of mir-184[Δ] females; they produce many more eggs and do not become sterile over time. (Iovino et al., 2009) When dpp[hr4]/+ males are crossed to sax[4]/+ females, the resulting dpp[hr4]/+ progeny do not show any significant drop in viability compared to the dpp[hr4]/+ progeny of wild-type mothers in control crosses. sax[4] slightly suppresses the dpp[d5]/dpp[hr56] wing phenotype. (Twombly et al., 2009) spict^mut neuromuscular junction overgrowth phenotypes are fully suppressed in sax⁴/Df(2R)cn7969 mutants. The synaptic undergrowth phenotypes in larvae homozygous for spict^mut in a sax⁴/Df(2R)cn7969 background are indistinguishable from that of sax⁴/Df(2R)cn7969 mutants alone. In addition, a heterozygous sax⁴ background partially suppresses the neuromuscular junction expansion of spict^mut larvae. (Wang et al., 2007) Expression of gbb^Scer\UAS.cKa, under the control of Scer\GAL4^A9, in a sax⁴/+ background partially suppresses the extra wing vein phenotype; over 90% of flies expressing both UAS transgenes show the most severe form of the Scer\GAL4^A9>gbb^Scer\UAS.cKa phenotype, compared to about 20% of Scer\GAL4^A9>gbb^Scer\UAS.cKa flies. A sax⁴/+ background causes a mild suppression of the Scer\GAL4^A9>dpp^Scer\UAS.cHa wing phenotype; with fewer wings showing the more severe forms of the phenotype. sax⁴ clones induced in a gbb⁴ background that are posterior to L5 or anterior to L2 never lead to ectopic wing vein formation. (Bangi and Wharton, 2006) The number of synaptic boutons/muscle surface area at muscle 6/7 in sax⁴/wit^HA3 double heterozygotes is 85.2 +/- 3.6 % of wild type. (McCabe et al., 2004) One copy of sax⁴ suppresses the increased bouton number seen at the neuromuscular junction in spin¹⁰⁴⁰³/spin^k09905 animals. spin¹⁰⁴⁰³/spin^k09905 animals which are also mutant for sax⁴/Df(2R)cn7969 show a further decrease in bouton number. (Sweeney and Davis, 2002) Dominantly reduces the viability of gbb¹/gbb⁴ flies. (Khalsa et al., 1998)
Xenogenetic Interactions
Statement Reference sax::Agam\sax[AgamEX.DmelIN.sax] completely rescues the lethality of sax[4]/Df(2R)BSC265 animals. (Fritsch et al., 2010)
Complementation & Rescue Data
Fails to complement	sax^unspecified (Dworkin and Gibson, 2006)
Rescued by	sax⁴/Df(2R)BSC265 is rescued by sax^+t8.7 (Fritsch et al., 2010) sax⁴/Df(2R)sax-H9 is rescued by sax^hs.PB (Twombly et al., 2009) sax⁴/Df(2R)sax-H9 is rescued by sax^{Scer\UAS.T:Ivir\HA1}/Scer\GAL4^elav.PLu (McCabe et al., 2004)
Comments
Stocks ( 1 )
Bloomington	5404 y¹ w^*; P{FRT(w^hs)}G13 sax⁴/SM6a
Notes on Origin
Discoverer

Comments
	Null allele. (Horsfield et al., 1998)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 3 )
Reported As
Symbol Synonym	sax4 (Dworkin and Gibson, 2006) sax⁴ (Dworkin and Gibson., 2006, Bangi and Wharton, 2006, Kirilly et al., 2005, Wang et al., 2007, Iovino et al., 2009, Fritsch et al., 2010, Twombly et al., 2009, Pilgram et al., 2011, Ball et al., 2010, Quijano et al., 2011) Sax⁴ (Liu et al., 2011)
Name Synonym
Secondary FlyBase IDs

References ( 27 )

Generate a list of	All references relating to this allele ( 27 )

List References by type	Research paper ( 26 ) Supplementary material ( 1 )
Recent research papers ( 3 )
	Liu et al., 2011, Dev. Growth Differ. 53(6): 822--841 Negative modulation of bone morphogenetic protein signaling by Dullard during wing vein formation in Drosophila. [FBrf0214664] Pilgram et al., 2011, J. Neurosci. 31(2): 492--500 The RhoGAP crossveinless-c Interacts with Dystrophin and Is Required for Synaptic Homeostasis at the Drosophila Neuromuscular Junction. [FBrf0212767] Quijano et al., 2011, Genetics 189(3): 809--824 Wg Signaling via Zw3 and Mad Restricts Self-Renewal of Sensory Organ Precursor Cells in Drosophila. [FBrf0216675] Show all research papers ...

Allele Dmel\sax4

Allele Dmel\sax⁴