Allele Dmel\phl¹¹

General Information
Symbol	Dmel\phl¹¹	Species	D. melanogaster
Name		FlyBase ID	FBal0013779
Feature type	allele	Associated gene	Dmel\phl
Also Known As	D-Raf^11-29, raf^11-29, Draf^11-29, raf¹¹²⁹, l(1)phl^11-29, l(1)ph^11-29

Map ( GBrowse )
Allele class	loss of function allele, amorphic 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	amorphic allele - genetic evidence (Ambrosio et al., 1989, Melnick et al., 1993, Sprenger et al., 1993) loss of function allele (Li et al., 1997)
Mutagen	spontaneous (Kramers et al., 1983, Schalet, 1986, Melnick et al., 1993)
Mutations Mapped to the Genome

Type Location Additional Notes References insertion site X:2,235,672..2,236,448 comment=Intragenic insertion, between amino acids 252 and 466, of 3.6kb, causing truncation of phl protein. evidence=experimental (Melnick et al., 1993)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Intragenic insertion, between amino acids 252 and 466, of 3.6kb, causing truncation of phl protein. (Melnick et al., 1993) Insertion in the 5' coding sequences of phl. (Sprenger et al., 1993) 4kb DNA insertion. (Ambrosio et al., 1989)
Cytology
Phenotypic Data
Phenotypic Class
	lethal \| embryonic stage \| recessive \| rescuable maternal effect (Melnick et al., 1993, Hou et al., 1995) lethal \| larval stage \| recessive (Karim and Rubin, 1998) lethal \| maternal effect \| recessive \| terminal expression pattern (Ambrosio et al., 1989) lethal \| recessive (Schalet, 1986, Kramers et al., 1983) mitotic cell cycle defective \| somatic clone (Yang and Baker, 2003) visible, with phl^wt.hs (Li et al., 2000)
Phenotype Manifest In
	A1-7 ventral acute muscle 2 \| precursor (Johnson Hamlet and Perkins, 2001) abdominal 11 anal tuft (Baek et al., 1996) anal pad (Baek et al., 1996) central nervous system \| maternal effect (Ambrosio et al., 1989) cephalopharyngeal skeleton (Li et al., 1998) chordotonal organ \| lateral \| maternal effect (Ambrosio et al., 1989) dorsal bridge \| maternal effect (Ambrosio et al., 1989) embryo \| posterior (Radke et al., 1997) embryonic/first instar larval cuticle (Radke et al., 2001) embryonic/first instar larval cuticle \| germline clone \| rescuable maternal effect (Hou et al., 1995) embryonic/larval brain \| maternal effect (Ambrosio et al., 1989) embryonic/larval hemocyte (Asha et al., 2003) embryonic/larval labral segment sense organ \| maternal effect (Ambrosio et al., 1989) embryonic/larval optic lobe \| maternal effect (Ambrosio et al., 1989) embryonic/larval pharynx \| embryonic stage \| maternal effect (Ambrosio et al., 1989) embryonic abdominal segment 8 (Li et al., 1998) embryonic abdominal segment 8 \| germline clone \| maternal effect (Ding et al., 2010) embryonic epidermis \| germline clone \| rescuable maternal effect (Hou et al., 1995) filzkorper (Baek et al., 1996, Li et al., 1998) filzkorper \| germline clone \| maternal effect (Ding et al., 2010) gastrula embryo (Li et al., 2003) hemocyte (Luo et al., 2002) hypostomal sclerite \| maternal effect (Ambrosio et al., 1989) labral segment \| maternal effect (Ambrosio et al., 1989) lamellocyte (Luo et al., 2002) leg \| anterior compartment \| ectopic, with phl^wt.hs (Li et al., 2000) leg \| posterior compartment, with phl^wt.hs (Li et al., 2000) peripheral nervous system \| maternal effect (Ambrosio et al., 1989) photoreceptor cell R2 \| somatic clone (Yang and Baker, 2003) photoreceptor cell R3 \| somatic clone (Yang and Baker, 2003) photoreceptor cell R4 \| somatic clone (Yang and Baker, 2003) photoreceptor cell R5 \| somatic clone (Yang and Baker, 2003) photoreceptor cell R8 \| somatic clone \| supernumerary (Yang and Baker, 2001) pole cell \| ectopic \| maternal effect (Li et al., 2003) salivary gland common duct primordium \| maternal effect (Li et al., 2003) spermatocyte (Tran et al., 2000) spermatocyte fusome (Tran et al., 2000) testis (Tran et al., 2000) ventral furrow \| maternal effect (Ambrosio et al., 1989) ventral midline \| dorsal \| maternal effect (Ambrosio et al., 1989) wing \| anterior compartment \| ectopic, with phl^wt.hs (Li et al., 2000) wing \| posterior compartment, with phl^wt.hs (Li et al., 2000) wing disc \| anterior compartment \| ectopic, with phl^wt.hs (Li et al., 2000) wing disc \| posterior compartment, with phl^wt.hs (Li et al., 2000)
Detailed Description
	Statement Reference Embryos derived from females containing homozygous germline clones have an abnormal cuticle pattern with deletion of posterior structures (abdominal segment 8 and Filzkorper). (Ding et al., 2010) Hemizygous larvae have fewer hemocytes than wild-type larvae. (Asha et al., 2003) Embryos derived from phl¹¹ female germ line clones exhibit defective gastrulation and lack the posterior midgut primordium. The majority of pole cells in these mutants are unable to enter the embryo at gastrulation and those that do enter the embryo appear abnormal in both directed migration and aggregation, leading to their random scattering in the embryonic cavity. (Li et al., 2003) In phl¹¹ mutant somatic clones in the 3rd instar eye disc, photoreceptors R2-R5 fail to undergo or differentiation. Within phl¹¹ mutant somatic clones there is a failure of G1 arrest in the furrow: all cells except R8s re-enter the cell cycle. However, none of these cells progress past G2. (Yang and Baker, 2003) The percentage of lamellocytes in the circulating blood cells of third instar larvae is strongly reduced in phl¹¹ mutants compared to wild type. In addition, the total concentration of circulating blood cells is greatly reduced in the mutant larvae compared to wild type. (Luo et al., 2002) Border cells show normal migration during stage 9 when all dorsal follicle cells are mutant for phl¹¹ in females containing homozygous clones. Mutant clusters are rarely recovered at stage 10, but dorsal migration can occur, although in most cases the cells probably die or cause the egg chamber to degenerate at stage 10. (Duchek and Rorth, 2001) phl¹¹/Y embryos show loss of VA2 muscle precursor cells (only 1.2% of hemisegments contain VA2 precursor cells). The embryos have a twisted morphology. (Johnson Hamlet and Perkins, 2001) Paternally rescued embryos derived from females with homozygous germline clones (lacking maternal but not zygotic Egfr function) lack head and tail structures at 24 hours. The embryos appear twisted at 6 hours of development. Hemizygous embryos derived from females with homozygous germline clones (lacking both maternal and zygotic Egfr function) produce what appears to be a small patch of dorsal cuticle at the end of embryonic development. The embryos appear twisted at 10 hours of development. (Radke et al., 2001) Multiple R8 cells are specified in an abnormal pattern in homozygous clones. (Yang and Baker, 2001) phl¹¹ males which have been partially rescued by expression of phl^wt.hs (phl^wt.hs has not been provided during the second and third larval instars) show a mirror-image duplication of the anterior compartment in the posterior of the wing disc. These animals show duplication of anterior pattern elements in the posterior compartment of the wing when allowed to reach adulthood. Comparable pattern duplications are also seen in the legs. (Li et al., 2000) phl¹¹ mutants are lethal in larvae, but such flies carrying phl^hs.PD can survive to fertile adults by using daily heatshocks. If heatshocks are withdrawn on eclosion, testes from 5 day old males are indistinguishable from controls. By day 7 however, almost all mutant testes show great proliferation centre expansion. This is due to excess, early stage germ cells and continues such that on day 15 testes are filled with these cells at the expense of post-mitotic cells. In these mutants fusome structure is not normal. Unbranched fusomes are found in may cells even those located some distance from the hub, where in wild-type testes, only branched fusomes interconnecting secondary spermatogonia are found. Although branches fusomes are found, these fusomes usually appear only after an intervening region containing many excess cells that have only spheroid fusomes. These excess germ-cells do not result from an increased frequency of germline stem cell divisions, as the M-phase index for the tier of cells adjacent to the hub in mutant testes is almost identical to that in wild-type. However the germ-line stem cells as a population appear to remain active longer than seen in wild-type. (Tran et al., 2000) Hemizygotes and homozygotes die as third instar larvae lacking imaginal discs. (Karim and Rubin, 1998) Homozygous embryos exhibit defects in posterior pattern. (Radke et al., 1997) Homozygous embryos lack head and tail structures. (Baek et al., 1996) No effect on the faf eye phenotype. (Huang and Fischer-Vize, 1996) Null embryos from homozygous female germlines differentiate remnants of cuticle with no obvious pattern. Embryos from homozygous germlines that receive a paternal wild type phl have cuticles that resemble those of embryos from tor mutant females. (Hou et al., 1995) Mutation has no effect on rho^hs.sev rough eye phenotype. (Freeman, 1994) Class 1 allele: Unrescued embryos (lacking maternal and zygotic phl activity) fail to differentiate into structured embryos and degenerate around 7 hrs of development. Rescued embryos (carrying wild type phl from their father) miss most structures posterior to abdominal segment 6 or 7 and anteriorly a portion of the cephalopharyngeal skeleton, labral sense organ and medial tooth are deleted. fkh expression (in wild type evident in hindgut, Malpighian tubule(s) and anal pads) entirely absent from class 1 embryos. Expression of tll, hkb, hb, fkh absent from both rescued and unrescued embryos, 7th ftz stripe deleted and 6th ftz stripe variably deleted/expanded. Conclude from 0% to 20--25% EL of blastoderm embryos deleted in class 1 alleles. (Melnick et al., 1993) A hole is seen in the blastoderm layer below the pole cells in embryos. Ventral furrow formation extends posteriorly to approximately 5% egg length. The posterior midgut invagination does not occur. 10% of embryos twist along the dorsal-ventral axis at germband extension to produce corkscrew shaped embryos. Comparison of embryos derived from homozygous germline clones reveal a zygotic requirement for phl: "pole hole" embryos (which have zygotic phl function) undergo germ band shortening and show the "terminal class" phenotype upon completing development. Posterior phenotype: A deletion of terminal identities is observed and cells forming in this region assume more anterior fates. The remaining most posterior segments (typically A5, A6 and A7) are expanded. This altered segmental pattern is first seen at the blastoderm stage. Deformation of the CNS ladder is observed for many embryos after nerve cord condensation. Usually the anterior and posterior commissures are normal in the last complete terminal segment, but breaks in the longitudinal axis of the CNS are seen. In some cases the sensory cells of the PNS are displaced laterally in the expanded abdominal segments. In addition, fusion of the dorsal cell cluster along the dorsal midline is observed, and the positions of the lateral chordotonal organs are shifted dorsally in the most posterior complete segment of the embryo. Anterior phenotype: Several head defects are also apparent in mature "pole hole" embryos, these arise in structures derived from the labrum and acron. Much of the supraoesophageal ganglia (brain) is missing at 16 hours. The optic lobes are also altered. The labral sense organ (epiphysis) is missing. Cell death is seen in the head and tail regions. In contrast, although "null" embryos (which have no zygotic phl function) appear phenotypically equivalent to "pole hole" embryos until the completion of germband extension, development ceases at this elongated germband stage, and they do not undergo germband shortening. After 20 hours of development the embryos consist of a ball of embryonic tissue at the anterior egg pole, and an extruded mass of egg yolk at the posterior pole. The overall organisation and differentiation of organ systems within the embryo is abnormal, due to incomplete development and massive cell death. The development of the nervous system is severely affected. (Ambrosio et al., 1989)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
NOT Enhancer of
Statement Reference phl¹¹ is a non-enhancer of visible phenotype of CanB^GMR.PS, Pp2B-14D^act.GMR (Sullivan and Rubin, 2002) phl¹¹ is a non-enhancer of visible phenotype of os^GMR.PB (Bach et al., 2003) phl¹¹ is a non-enhancer of visible phenotype of Ret^MEN2B.GMR (Read et al., 2005)
Suppressor of
Statement Reference phl¹¹/phl[+] is a suppressor of visible phenotype of PDZ-GEF^EP388, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Lee et al., 2002)
NOT Suppressor of
Statement Reference phl¹¹ is a non-suppressor of melanotic mass phenotype phenotype of hop^Tum (Luo et al., 2002) phl¹¹ is a non-suppressor of visible phenotype of CanB^GMR.PS, Pp2B-14D^act.GMR (Sullivan and Rubin, 2002) phl¹¹ is a non-suppressor of visible phenotype of os^GMR.PB (Bach et al., 2003) phl¹¹ is a non-suppressor of visible phenotype of Ret^MEN2B.GMR (Read et al., 2005)
Phenotype Manifest In
Suppressed by
Statement Reference phl¹¹ has A1-7 ventral acute muscle 2 \| precursor phenotype, suppressible \| partially by Scer\GAL4^twi.PG/csw::Src64B^{src90.Scer\UAS} (Johnson Hamlet and Perkins, 2001) phl¹¹ has abdominal 11 anal tuft phenotype, suppressible \| partially by Hsap\RAF1^Δ305 (Baek et al., 1996) phl¹¹ has abdominal 11 anal tuft phenotype, suppressible by Hsap\RAF1^cFa (Baek et al., 1996) phl¹¹ has anal pad phenotype, suppressible by Hsap\RAF1^cFa (Baek et al., 1996) phl¹¹ has embryonic abdominal segment 8 phenotype, suppressible by phl::tor^13D.hs.sev (Li et al., 1998) phl¹¹ has filzkorper phenotype, suppressible by phl::tor^13D.hs.sev (Li et al., 1998) phl¹¹ has phenotype, suppressible by Hsap\RAF1^cFa (Baek et al., 1996) phl¹¹ has phenotype, suppressible by Hsap\RAF1^Δ305 (Baek et al., 1996)
NOT suppressed by
Statement Reference phl¹¹ has cephalopharyngeal skeleton phenotype, non-suppressible by phl::tor^13D.hs.sev (Li et al., 1998)
Enhancer of
Statement Reference phl¹¹ is an enhancer of eye phenotype of W^GMR.PG/W^GMR.PG (Bergmann et al., 1998) phl¹¹ is an enhancer of ommatidium phenotype of W^GMR.PG/W^GMR.PG (Bergmann et al., 1998)
NOT Enhancer of
Statement Reference phl¹¹ is a non-enhancer of eye phenotype of CanB^GMR.PS, Pp2B-14D^act.GMR (Sullivan and Rubin, 2002) phl¹¹ is a non-enhancer of eye phenotype of os^GMR.PB (Bach et al., 2003) phl¹¹ is a non-enhancer of eye phenotype of Ret^MEN2B.GMR (Read et al., 2005)
Suppressor of
Statement Reference phl¹¹/phl[+] is a suppressor of eye phenotype of PDZ-GEF^EP388, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Lee et al., 2002) phl¹¹/phl[+] is a suppressor of eye photoreceptor cell \| ectopic phenotype of PDZ-GEF^EP388, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Lee et al., 2002) phl¹¹/phl[+] is a suppressor of ommatidium phenotype of PDZ-GEF^EP388, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF (Lee et al., 2002) phl¹¹ is a suppressor \| maternal effect of embryonic/first instar larval cuticle \| maternal effect phenotype of tor^12D (Li and Li, 2003) phl¹¹ is a suppressor of hemocyte \| supernumerary phenotype of hop^Tum (Luo et al., 2002) phl¹¹ is a suppressor of lamellocyte \| supernumerary phenotype of hop^Tum (Luo et al., 2002)
NOT Suppressor of
Statement Reference phl¹¹ is a non-suppressor of embryonic/larval lymph gland phenotype of hop^Tum (Luo et al., 2002) phl¹¹ is a non-suppressor of eye phenotype of CanB^GMR.PS, Pp2B-14D^act.GMR (Sullivan and Rubin, 2002) phl¹¹ is a non-suppressor of eye phenotype of os^GMR.PB (Bach et al., 2003) phl¹¹ is a non-suppressor of eye phenotype of Ret^MEN2B.GMR (Read et al., 2005) phl¹¹ is a non-suppressor of melanotic mass phenotype of hop^Tum (Luo et al., 2002)
Other
Statement Reference Hsap\RAF1^Δ305, phl¹¹ has anal pad phenotype (Baek et al., 1996) phl¹¹, sl¹ has border follicle cell \| somatic clone phenotype (Bianco et al., 2007)
Additional Comments
Genetic Interactions
Statement Reference phl¹¹ sl¹ double mutant border cell clusters (induced as clones) initiate migration, but are delayed later in posterior migration and do not show dorsal migration. phl¹¹ sl¹ double mutant cells are generally located at the rear of the cluster during border cell late posterior migration and dorsal migration in mosaic border cell clusters consisting of both wild-type and mutant cells. (Bianco et al., 2007) Caspase activation is seen posterior to column 5, Within phl¹¹ mutant clones in the eye disc in a BacA\p35^GMR.PH background, caspase activation is seen posterior to column 5 relative to the morphogenetic furrow. There is no increase in caspase activation outside of these clones. (Yang and Baker, 2003) The rough eye phenotype, irregular ommatidial array and increased photoreceptor cell number caused by expression of Gef26^EP388 under the control of Scer\GAL4^GMR.PF are strongly suppressed by phl¹¹/+. (Lee et al., 2002) At 28^oC, the increased differentiation of blood cells into lamellocytes and the increased accumulation of blood cells seen in hop^Tum larvae is completely suppressed by phl¹¹. All hemizygous phl¹¹ hop^Tum larvae have overgrown and melanised lymph glands, similar to that seen in hop^Tum mutants cultured at 28^oC. (Luo et al., 2002) Expression of csw::Src64B^{src90.Scer\UAS} under the control of Scer\GAL4^twi.PG partially suppresses the loss of VA2 muscle precursor cells seen in phl¹¹/Y embryos (10.5% of hemisegments have VA2 cells). (Johnson Hamlet and Perkins, 2001) At 25^oC phl¹¹ dominantly suppresses most of the hyperplastic growth seen in Ras85D^V12.Scer\UAS Scer\GAL4^dpp.blk1 wing discs. (Karim and Rubin, 1998) Filzkorper and eighth abdominal segment are largely rescued by phl::tor^13D.hs.sev whereas the head skeleton is not. (Li et al., 1998)
Xenogenetic Interactions
Statement Reference Injection of Hsap\RAF1^cFa can rescue the terminal structure defects in some phl¹¹ embryos, injection of Hsap\RAF1^Δ305 allows a low percentage rescue of the posterior structures. (Baek et al., 1996)
Complementation & Rescue Data
Rescued by	phl¹¹ is rescued by phl^cBa (Radke et al., 1997) phl¹¹ is rescued by phl^Hsp83.PD (Ding et al., 2010) phl¹¹ is rescued by phl^S388A (Baek et al., 1996) phl¹¹ is rescued by phl^wt.hs (Li et al., 2000) phl¹¹ is rescued by phl^ΔN114.Hsp83 (Ding et al., 2010)
Partially rescued by	phl¹¹ is partially rescued by phl^hs.PR (Radke et al., 2001) phl¹¹ is partially rescued by phl^S388A.hs (Radke et al., 2001) phl¹¹ is partially rescued by phl^S743A.hs (Radke et al., 2001) phl¹¹ is partially rescued by phl^Δ315.hs (Radke et al., 2001) phl¹¹ is partially rescued by phl^Δ315 (Baek et al., 1996) phl¹¹ is partially rescued by phl^Δ370 (Baek et al., 1996) phl¹¹ is partially rescued by phl^Δ445.hs (Radke et al., 2001) phl¹¹ is partially rescued by phl^Δ445 (Baek et al., 1996)
Not rescued by	phl¹¹ is not rescued by phl^hs.PD (Tran et al., 2000) phl¹¹ is not rescued by phl^K497M.hs (Radke et al., 2001) phl¹¹ is not rescued by phl^K497M (Baek et al., 1996) phl¹¹ is not rescued by phl^S743A (Baek et al., 1996)
Comments	phl[ΔN114.Hsp83] can rescue the posterior defects seen in embryos derived from derived from females containing homozygous phl[11] germline clones (rescue is seen if the level of expression from the transgene is equal or greater than the endogenous wild-type level of phl protein), but phl[ΔN114.Hsp83] is less active than phl[Hsp83.PD] in this assay (the latter can rescue the posterior defects at much lower levels of expression). (Ding et al., 2010) Expression of phl^wt.hs using daily heat shocks from embryonic through late pupal stages rescues phl¹¹ males to full viability and fertility. (Li et al., 2000) Injection of wild type RNA rescues the filzkorper, tuft and anal pads. (Radke et al., 1997) Injection of wild type phl transcripts into the posterior region of phl¹¹ embryos rescues the posterior and injection into the head region rescues the anterior terminal structures. Injection of phl^K497M into the anterior or posterior pole of phl¹¹ embryos rescues neither the head nor tail defects. (Baek et al., 1996)
Stocks ( 2 )
Bloomington	5733 phl¹¹/FM6, l(1)FMa¹/Dp(1;Y)B^Sw⁺y⁺
Kyoto	108393 phl¹¹/FM6, l(1)FMa¹/Dp(1;Y)B^Sw⁺y⁺
Notes on Origin
Discoverer	Schalet. Schalet. A. Schalet.

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 15 )
Reported As
Symbol Synonym	11-29 (Kramers et al., 1983) D-Raf^11-29 (Mishra et al., 2005, Li et al., 1998, Li et al., 1997, Hou et al., 1995) Draf^11-29 (Li et al., 2003, Li and Li, 2003, Li et al., 2000, Ding et al., 2010) D-raf^11-29 (Johnson Hamlet and Perkins, 2001, Radke et al., 2001, Baonza and Garcia-Bellido, 1999, Roch et al., 1998, Radke et al., 1997, Hermon and Ambrosio, 1997, Baek et al., 1996, Baek and Ambrosio, 1994, Baek et al., 1995, Sprenger et al., 1993, Melnick et al., 1993) DRaf¹¹²⁹ (Asha et al., 2003) l(1)11-29 (Schalet, 1986, ) l(1)ph^11-29 (Ambrosio et al., 1989, Ambrosio et al., 1989) l(1)phl^11-29 (Scholz et al., 1997, Scholz et al., 1997) phl^1-29 (Sullivan and Rubin, 2002) phl^11.29 (Read et al., 2005) phl¹¹ (Lee et al., 2002, Bianco et al., 2007, Christensen et al., 2008.12.28) raf^11-29 (Bach et al., 2003, Luo et al., 2002, Yang and Baker, 2003, Yang and Baker, 2001, Tran et al., 2000, Bergmann et al., 1998) Raf^11-29 (Karim and Rubin, 1998) raf¹¹²⁹ (Huang and Fischer-Vize, 1996, Schweitzer et al., 1995, Freeman, 1994) Raf^l(1)11-29 (Wilson et al., 2005)
Name Synonym
Secondary FlyBase IDs

References ( 43 )

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

List References by type	Research paper ( 39 ) Erratum ( 1 ) Personal communication to FlyBase ( 1 ) Abstract ( 2 )
Recent research papers (0)
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Allele Dmel\phl11

Allele Dmel\phl¹¹