Allele Dmel\ed^1X5

General Information
Symbol	Dmel\ed^1X5	Species	D. melanogaster
Name		FlyBase ID	FBal0086650
Feature type	allele	Associated gene	Dmel\ed

Map ( GBrowse )
Allele class	loss of function allele
Mutagen
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	loss of function allele (Hsu et al., 1998, Bai et al., 2001)
Mutagen
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation 2L:4,098,668..4,098,668 comment=Site of nucleotide substitution in mutant inferred by FlyBase based on reported amino acid change. evidence=experimental na_change=C4098668T pr_change=Q524@\|ed-PA reported_pr_change=?524@ (Escudero et al., 2003)
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: ?524@. (Escudero et al., 2003)
Cytology
Phenotypic Data
Phenotypic Class
	cell shape defective \| cell autonomous \| somatic clone (Chang et al., 2011) cell shape defective \| cell non-autonomous \| somatic clone (Chang et al., 2011) cell shape defective \| somatic clone (Wei et al., 2005) lethal \| embryonic stage (Swan et al., 2006) lethal \| pupal stage \| recessive (Bai et al., 2001) lethal \| recessive (Escudero et al., 2003) size defective \| adult stage \| somatic clone (Yue et al., 2012) viable (with ed^slH8) (Escudero et al., 2003) visible (with ed^slH8) (Escudero et al., 2003, Bai et al., 2001) visible \| adult stage \| somatic clone (Yue et al., 2012) visible \| recessive \| somatic clone (Escudero et al., 2003)
Phenotype Manifest In
	apical plasma membrane & cell & dorsal mesothoracic disc & third instar larva \| somatic clone (Wei et al., 2005) cone cell (Hsu et al., 1998) cone cell (with ed^slH8) (Bai et al., 2001) eye \| somatic clone (Yue et al., 2012) follicle cell \| somatic clone (Yue et al., 2012) lateral transverse muscle (Swan et al., 2006) lateral transverse muscle (with ed^slH8) (Swan et al., 2006) macrochaeta \| ectopic (with ed^slH8) (Escudero et al., 2003) macrochaeta \| ectopic \| somatic clone (Escudero et al., 2003) microchaeta \| ectopic (with ed^slH8) (Escudero et al., 2003) microchaeta \| ectopic \| somatic clone (Escudero et al., 2003) mystery cell (with ed^slH8) (Bai et al., 2001) ommatidium (with ed^slH8) (Bai et al., 2001) ommatidium \| somatic clone (Fetting et al., 2009) oocyte \| somatic clone (Yue et al., 2012) photoreceptor cell (Hsu et al., 1998) photoreceptor cell R1 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R1 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R2 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R2 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R3 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R3 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R4 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R4 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R5 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R5 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R6 (with ed^slH8) (Bai et al., 2001) photoreceptor cell R6 \| supernumerary (with ed^slH8) (Bai et al., 2001) photoreceptor cell R7 \| supernumerary (with ed^slH8) (Bai et al., 2001) sensillum campaniformium & wing \| supernumerary \| somatic clone (Wei et al., 2005) sensory mother cell \| ectopic \| somatic clone (Escudero et al., 2003) ventral longitudinal muscle (Swan et al., 2006) ventral longitudinal muscle (with ed^slH8) (Swan et al., 2006) wing (with ed^slH8) (Bai et al., 2001) wing cell \| somatic clone (Wei et al., 2005) wing disc \| cell autonomous \| somatic clone (Chang et al., 2011) wing disc \| cell non-autonomous \| somatic clone (Chang et al., 2011) wing disc \| somatic clone (Escudero et al., 2003) wing disc \| somatic clone \| third instar larval stage (Wei et al., 2005) wing hair \| somatic clone \| supernumerary (Wei et al., 2005) wing vein \| somatic clone (Wei et al., 2005) wing vein \| supernumerary (with ed^slH8) (Bai et al., 2001)
Detailed Description
	Statement Reference ed[1X5] mutant eye clones (generated using the ey-FLP method) are overgrown compared with control clones. In ed[1X5] mutant posterior follicle cell clones multiple layers are formed around the wild type oocyte, rather than the single layer of cells seen in wild type. Mislocalisation of the oocyte nucleus is also seen. (Yue et al., 2012) ed[1X5] mutant wing disc clones sort out from the surrounding wild type cells, exhibiting both rounded and smooth contours rather than the jagged border seen in controls. The wild type cells surrounding the clone assemble a ring of actomyosin cable at the interface and the clone cells themselves develop an apical constriction that is inversely proportional to the size of the clone. Similar apical constriction is seen in wild type cells when they are surrounded by a large number of ed[1X5] mutant cells. (Chang et al., 2011) Mosaic ommatidia containing wild-type photoreceptors and between one and four mutant cone cells misrotate. (Fetting et al., 2009) ed^1X5 animals predominantly die as embryos although some develop into second instar larvae. In these larvae, defects in ventrolateral muscles and lateral transverse muscles are evident with both muscle groups forming aberrant projections and ectopic adhesion points. Consistent and prominent defects in muscle morphology can also be observed at the embryonic stage. The same defects are present in ed^1X5/ed^slH8 transheterozygotes. Epidermal tendon cells are positioned correctly in ed^1X5 mutants. (Swan et al., 2006) ed^1X5 mutant clones in the third instar wing disc exhibit round and smooth contours in contrast to clones of wild type cells that show wiggly shapes. Both large and small ed^1X5 clones are simultaneously observed, and often straddle either the dorsal-ventral or anterior-posterior compartment boundary. When ed^1X5 clones are small, the mutant cells have a reduced apical surface: this apical constriction continues through the levels of the adherens and septate junctions, ending at the plane just below the gap junctions. Hence, these ed^1X5 cells adopt a bottle shape. In contrast, ed^1X5 cells within larger mutant clones (over hundreds of cells) have an apical surface more similar to those of wild type cells. Apposed ed^1X5/ed^1X5 and ed^1X5/+ cells at the border of mutant clones are enlarged and adopt a rectangular shape, elongating towards the mutant clone itself. Large ed^1X5 clones that are close to or touch the dorsal-ventral boundary of the wing disc displace the boundary towards the mutant clone. In contrast, ed^1X5 clones that straddle this boundary do not overtly distort it, although the boundary can be less smooth within the clone. Adult wings bearing ed^1X5 clones display rounded areas with altered patterns of trichomes. When these areas are small, the density of trichomes is increased and/or the pointing of trichomes is altered, while larger areas show these effects to a lesser or no extent. Appropriately located putative clones can develop extra sensilla campaniformia, extra wing vein material or a partial loss of wing vein. A crease of cuticle often surrounds clones, especially small ones. (Wei et al., 2005) The number of macrochaetae/heminotum in ed^1X5/ed^slH8 mutants is - anterior+posterior notopleural: 2.10, presutural: 1.28, anterior supraalar: 1.13, posterior supraalar: 1.05, anterior postalar: 1.80, posterior postalar: 1.85, anterior+posterior dorsocentral: 2.80 and anterior+posterior scutellar: 2.63. The density of microchaetae on the notum is also increased in these flies. Homozygous clones in imaginal wing discs are much smaller than wild-type twin clones, and many twins have no associated mutant clone. Within mutant clones, extra sensory organ precursor (SOP) cells are detected near extant SOPs. These clones give rise to an extra macrochaeta in the adult. Homozygous clones in the wing disc induced a Minute background are viable and include extra SOPs when they include regions from which extant SOPs arise. These clones give rise to groups of macrochaetae or areas of increased density of microchaetae in the adult. (Escudero et al., 2003) 33% of ommatidia of ed^1X5/ed^slH8 flies have extra R7-like cells. 26% of ommatidia have extra outer photoreceptors and 6% have fewer than normal outer photoreceptors. Mystery cells are transformed into neuronal photoreceptor cells. 69% of ommatidia have 5-6 instead of 4 cone cells, and 10% have three primary pigment cells. ed^1X5/ed^slH8 flies show an enlarged wing with extra wing vein material. (Bai et al., 2001)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference ed^slH8/ed^1X5 has visible phenotype, enhanceable by Dl^RevF10/Dl[+] (Escudero et al., 2003)
Suppressed by
Statement Reference ed^slH8/ed^1X5 has visible phenotype, suppressible \| partially by H²/H[+] (Escudero et al., 2003) ed^slH8/ed^1X5 has visible phenotype, suppressible \| partially by Scer\GAL4^ap-md544/Egfr^{DN.Scer\UAS.cBa} (Escudero et al., 2003)
NOT suppressed by
Statement Reference ed^1X5 has cell shape defective \| somatic clone \| cell autonomous phenotype, non-suppressible by shg^αTub84B.PP (Chang et al., 2011) ed^1X5 has cell shape defective \| somatic clone \| cell non-autonomous phenotype, non-suppressible by shg^αTub84B.PP (Chang et al., 2011)
Enhancer of
Statement Reference ed[+]/ed^1X5 is an enhancer of visible \| dominant phenotype of Egfr^E3 (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of visible phenotype of Egfr::tor^{t4021E.hs.sev} (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of visible phenotype of phl::tor^13D.hs.sev (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of visible phenotype of Ras85D^V12.sev (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of visible phenotype of rho^hs.PSt (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of visible phenotype of rl^Sem (Bai et al., 2001)
Suppressor of
Statement Reference ed[+]/ed^1X5 is a suppressor \| partially of visible phenotype of Ras85D^N17.sev (Bai et al., 2001) ed[+]/ed^1X5 is a suppressor of visible phenotype of aos^hs.PF (Bai et al., 2001)
NOT Suppressor of
Statement Reference ed^slH8/ed^1X5 is a non-suppressor of visible \| dominant phenotype of N^Ax-M1 (Escudero et al., 2003) ed^slH8/ed^1X5 is a non-suppressor of visible \| dominant phenotype of N^Mcd1 (Escudero et al., 2003)
Other
Statement Reference ed[+]/ed^1X5, sty^S88 has visible phenotype (Bai et al., 2001) ed^1X5, sty^S88 has visible phenotype (Bai et al., 2001) N[+]/N^55e11, ed^slH8/ed^1X5 has visible phenotype (Escudero et al., 2003) N^55e11, ed^slH8/ed^1X5 has visible phenotype (Escudero et al., 2003) RasGAP1^B2, ed[+]/ed^1X5 has visible phenotype (Bai et al., 2001) RasGAP1^B2, ed^1X5 has visible phenotype (Bai et al., 2001)
Phenotype Manifest In
Enhanced by
Statement Reference ed^slH8/ed^1X5 has macrochaeta \| ectopic phenotype, enhanceable by Dl^RevF10/Dl[+] (Escudero et al., 2003)
Suppressed by
Statement Reference ed^1X5 has wing disc \| somatic clone \| cell autonomous phenotype, suppressible \| partially by Scer\GAL4^C-765/ed::shg^Scer\UAS.cCa (Chang et al., 2011) ed^slH8/ed^1X5 has macrochaeta \| ectopic phenotype, suppressible \| partially by Scer\GAL4^ap-md544/Egfr^{DN.Scer\UAS.cBa} (Escudero et al., 2003) ed^slH8/ed^1X5 has macrochaeta \| ectopic phenotype, suppressible by H²/H[+] (Escudero et al., 2003)
NOT suppressed by
Statement Reference ed^1X5 has wing disc \| somatic clone \| cell autonomous phenotype, non-suppressible by shg^αTub84B.PP (Chang et al., 2011) ed^1X5 has wing disc \| somatic clone \| cell non-autonomous phenotype, non-suppressible by shg^αTub84B.PP (Chang et al., 2011) ed^1X5 has wing disc \| somatic clone phenotype, non-suppressible by shg^αTub84B.PP (Chang et al., 2011) ed^slH8/ed^1X5 has microchaeta phenotype, non-suppressible by H²/H[+] (Escudero et al., 2003)
Enhancer of
Statement Reference ed[+]/ed^1X5 is an enhancer of eye phenotype of Egfr::tor^{t4021E.hs.sev} (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of eye phenotype of Egfr^E3 (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of eye phenotype of phl::tor^13D.hs.sev (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of eye phenotype of Ras85D^V12.sev (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of eye phenotype of rho^hs.PSt (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of cno^mis1/cno² (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of dgo³⁸⁰ (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of dsh¹ (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of fz²⁰/fz¹⁹ (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of pnt¹²⁷⁷/pnt^Δ88 (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ommatidium phenotype of stan^frz3 (Fetting et al., 2009) ed[+]/ed^1X5 is an enhancer of ventral longitudinal muscle phenotype of Grip^ex36 (Swan et al., 2006) ed[+]/ed^1X5 is an enhancer of wing vein \| supernumerary phenotype of Egfr^E3 (Bai et al., 2001) ed[+]/ed^1X5 is an enhancer of wing vein \| supernumerary phenotype of rl^Sem (Bai et al., 2001) ed^1X5 is an enhancer of eye phenotype of Egfr^E1 (Hsu et al., 1998) ed^1X5 is an enhancer of eye phenotype of Ras85D^V12 (Hsu et al., 1998) ed^1X5 is an enhancer of wing vein phenotype of Egfr^E3 (Hsu et al., 1998) ed^1X5 is an enhancer of wing vein phenotype of rl^Sem (Hsu et al., 1998)
Suppressor of
Statement Reference ed[+]/ed^1X5 is a suppressor \| partially of eye phenotype of Ras85D^N17.sev (Bai et al., 2001) ed[+]/ed^1X5 is a suppressor of eye phenotype of aos^hs.PF (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of ommatidium phenotype of aop^ACT.sev (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of ommatidium phenotype of pnt¹²⁷⁷/pnt^Δ88 (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of ommatidium phenotype of sina³/sina² (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of photoreceptor cell R7 phenotype of aop^ACT.sev (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of photoreceptor cell R7 phenotype of pnt¹²⁷⁷/pnt^Δ88 (Bai et al., 2001) ed^slH8/ed^1X5 is a suppressor of photoreceptor cell R7 phenotype of sina³/sina² (Bai et al., 2001)
NOT Suppressor of
Statement Reference ed^slH8/ed^1X5 is a non-suppressor of macrochaeta phenotype of N^Ax-M1 (Escudero et al., 2003) ed^slH8/ed^1X5 is a non-suppressor of microchaeta phenotype of N^Mcd1 (Escudero et al., 2003) ed^slH8/ed^1X5 is a non-suppressor of photoreceptor cell phenotype of ttk^{88.GMR.T:Hsap\MYC} (Bai et al., 2001)
Other
Statement Reference ed[+]/ed^1X5, sty^S88 has wing vein \| supernumerary phenotype (Bai et al., 2001) ed^1X5, smo³ has wing disc \| somatic clone \| third instar larval stage phenotype (Wei et al., 2005) ed^1X5, sty^S88 has wing vein \| supernumerary phenotype (Bai et al., 2001) ed^slH8/ed^1X5, sev¹⁴ has photoreceptor cell R7 \| supernumerary phenotype (Bai et al., 2001, Bai et al., 2001) Grip^ex36, ed[+]/ed^1X5 has lateral transverse muscle phenotype (Swan et al., 2006) N[+]/N^55e11, ed^slH8/ed^1X5 has microchaeta phenotype (Escudero et al., 2003) N^55e11, ed^slH8/ed^1X5 has microchaeta phenotype (Escudero et al., 2003) RasGAP1^B2, ed[+]/ed^1X5 has wing vein \| supernumerary phenotype (Bai et al., 2001) RasGAP1^B2, ed^1X5 has wing vein \| supernumerary phenotype (Bai et al., 2001)
Additional Comments
Genetic Interactions
Statement Reference Ubiquitous expression of ed::shg[Scer\UAS.cCa] under the control of Scer\GAL4[C-765] is unable to suppress the cell sorting phenotypes seen in ed[1X5] mutant wing disc clones. An actomyosin cable still forms around the clone and the boundary between the mutant clone and neighbouring wild type cells remains smooth. Apical constriction of the clone cells is still seen. Expression of shg[αTub84B.PP] is unable to rescue the cell sorting phenotypes seen in ed[1X5] mutant wing disc clones. (Chang et al., 2011) ed[1X5] dominantly enhances the phenotype of mis-rotation of ommatidia that is seen in dgo[380], dsh[1], stan[frz3], cno[2]/cno[mis1] and pnt[Δ88]/pnt[1277] animals. ed[1X5] dominantly enhances the phenotype of mis-rotation of ommatidia that is seen in fz[19]/fz[20] animals, without affecting the chirality phenotype. (Fetting et al., 2009) Grip^ex36, ed^1X5/+ embryos exhibit more severe VLM defects than Grip^ex36 embryos. In severe cases, the double mutants exhibit completely deranged somatic musculature, where muscle identification is no longer possible. Lateral transverse muscles are defective in Grip^ex36/Y; ed^1X5/+ embryos, while these muscles appear normal in both single heterozygote. (Swan et al., 2006) Independent ed^1X5, smo³ double mutant clones that originate in different compartments (as evidenced by ci or en expression) can fuse together to form composite clones which have roundish, smooth shapes. Within such composite clones, the characteristic segregation of 'anterior' and 'posterior' cells is maintained - they do not intermix. (Wei et al., 2005) Expression of Egfr^{DN.Scer\UAS.cBa} under the control of Scer\GAL4^ap-md544 partially suppresses the extra bristle phenotype seen in ed^1X5/ed^slH8 flies. The number of macrochaetae/heminotum in animals expressing Egfr^{DN.Scer\UAS.cBa} under the control of Scer\GAL4^ap-md544 in a ed^1X5/ed^slH8 background is - anterior+posterior notopleural: 2.15, presutural: 1.05, anterior supraalar: 0.80, posterior supraalar: 0, anterior postalar: 0.60, posterior postalar: 0, anterior+posterior dorsocentral: 1.30 and anterior+posterior scutellar: 2.25. N^55e11/+ ; ed^1X5/ed^slH8 adults show an almost complete absence of microchaetae on the notum. Dl^RevF10/+ increases the number of extra macrochaetae on the notum seen in ed^1X5/ed^slH8 adults. H²/+ almost completely eliminates the extra macrochaetae seen on the notum of ed^1X5/ed^slH8 animals, although the increased density of microchaetae seen in ed^1X5/ed^slH8 flies is not reduced by H²/+. The low density of microchaetae on the notum seen in N^Mcd1/+ flies is not rescued by ed^1X5/ed^slH8. (Escudero et al., 2003) ed^1X5 suppresses the rough eye phenotype caused by overexpression of argos. (Hsu et al., 1998)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	ed^k01102 (Bai et al., 2001) ed^lF20 (Bai et al., 2001)
Rescued by	ed^1X5 is rescued by ed^Scer\UAS.cBa/Scer\GAL4^C-765 (Escudero et al., 2003, Chang et al., 2011) ed^1X5 is rescued by ed^{TMC.Scer\UAS.T:Hsap\MYC}/Scer\GAL4^tub.PU (Yue et al., 2012)
Not rescued by	ed^1X5 is not rescued by Scer\GAL4^tub.PU/ed^{ΔC.Scer\UAS.T:Hsap\MYC} (Yue et al., 2012)
Comments	Expression of ed[ΔC.Scer\UAS.T:Hsap\MYC] under the control of Scer\GAL4[tub.PU] is unable to rescue the multiple cell layer phenotype seen in ed[1X5] mutant follicle cell clones. Expression of ed[TMC.Scer\UAS.T:Hsap\MYC] under the control of Scer\GAL4[tub.PU] rescues the multiple cell layer phenotype seen in ed[1X5] mutant follicle cell clones. (Yue et al., 2012) Ubiquitous expression of ed[Scer\UAS.cBa] under the control of Scer\GAL4[C-765] rescues the cell sorting phenotypes seen in ed[1X5] mutant wing disc clones. The boundary between the clone and the surrounding wild type cells is no longer smooth edged, instead appearing jagged as in wild type. Normal levels of shg are seen. (Chang et al., 2011)
Stocks ( 0 )

Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	ed^1x5 (Escudero et al., 2003, Swan et al., 2006, Swan et al., 2006, Wei et al., 2005, Fetting et al., 2009) ed^1X5 (Yue et al., 2012, Chang et al., 2011)
Name Synonym
Secondary FlyBase IDs

References ( 9 )
Research paper	Yue et al., 2012, Dev. Cell 22(2): 255--267 The cell adhesion molecule echinoid functions as a tumor suppressor and upstream regulator of the hippo signaling pathway. [FBrf0217474] Chang et al., 2011, Development 138(17): 3803--3812 Differential adhesion and actomyosin cable collaborate to drive Echinoid-mediated cell sorting. [FBrf0214612] Fetting et al., 2009, Development 136(19): 3323--3333 The cell adhesion molecules Echinoid and Friend of Echinoid coordinate cell adhesion and cell signaling to regulate the fidelity of ommatidial rotation in the Drosophila eye. [FBrf0208836] Swan et al., 2006, EMBO J. 25(15): 3640--3651 Complex interaction of Drosophila GRIP PDZ domains and Echinoid during muscle morphogenesis. [FBrf0192056] Wei et al., 2005, Dev. Cell 8(4): 493--504 Echinoid is a component of adherens junctions that cooperates with DE-Cadherin to mediate cell adhesion. [FBrf0187414] Escudero et al., 2003, Development 130(25): 6305--6316 Echinoid synergizes with the Notch signaling pathway in Drosophila mesothorax bristle patterning. [FBrf0167516] Bai et al., 2001, Development 128(4): 591--601 The cell adhesion molecule Echinoid defines a new pathway that antagonizes the Drosophila EGF receptor signaling pathway. [FBrf0134572]
Supplementary material	Swan et al., 2006, EMBO J. 25(15): Supplementary Methods. [FBrf0198715]
Abstract	Hsu et al., 1998, A. Dros. Res. Conf. 39: 193A echinoid,a negative regulator of the Drosophila EGF receptor signaling pathway. [FBrf0101230]

Allele Dmel\ed1X5

Allele Dmel\ed^1X5