Allele Dmel\lwr^4-3

General Information
Symbol	Dmel\lwr^4-3	Species	D. melanogaster
Name		FlyBase ID	FBal0122957
Feature type	allele	Associated gene	Dmel\lwr

Allele class	hypomorphic allele - genetic evidence
Mutagen	Delta2-3
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	hypomorphic allele - genetic evidence (Apionishev et al., 2001, Sun et al., 2003, Huang et al., 2005)
Mutagen	Delta2-3 (Apionishev et al., 2001)
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	lwr⁰⁵⁴⁸⁶ (Apionishev et al., 2001)
Nature of the lesion	Statement Reference Imprecise excision of the P{PZ} element. 34bp of P-element sequences remain, together with 42bp of inserted material, repeats of TAACA. (Apionishev et al., 2001)
Caused by insertion	P{?PZ}lwr^4-3 (Apionishev et al., 2001, Huang et al., 2005, Chiu et al., 2005, Sun et al., 2003)
Cytology
Phenotypic Data
Phenotypic Class
	developmental rate defective (with lwr⁵) (Chiu et al., 2005) enhancer of variegation, with In(1)w^m4h, Scer\GAL4[-]: (Reo et al., 2010) enhancer of variegation, with In(2)Heidi (Reo et al., 2010) immune response defective (with lwr⁵) (Paddibhatla et al., 2010) lethal \| recessive (Apionishev et al., 2001) lethal \| third instar larval stage (Huang et al., 2005) lethal \| third instar larval stage (with lwr¹³) (Sun et al., 2003) melanotic mass phenotype \| larval stage (with lwr⁵) (Chiu et al., 2005) melanotic mass phenotype \| larval stage \| partially (with lwr¹⁴) (Chiu et al., 2005) tumorigenic (with lwr⁵) (Paddibhatla et al., 2010)
Phenotype Manifest In
	crystal cell (with lwr⁵) (Chiu et al., 2005) crystal cell (with lwr¹⁴) (Chiu et al., 2005) embryonic/larval fat body (with lwr⁵) (Paddibhatla et al., 2010) embryonic/larval hemocoel (with lwr⁵) (Paddibhatla et al., 2010) embryonic/larval hemocyte \| supernumerary (with lwr⁵) (Huang et al., 2005) hemocyte & nucleus \| supernumerary (with lwr¹⁴) (Chiu et al., 2005) hemocyte \| supernumerary (with lwr⁵) (Chiu et al., 2005) lamellocyte \| supernumerary (with lwr⁵) (Huang et al., 2005, Chiu et al., 2005) lamellocyte \| supernumerary (with lwr¹⁴) (Chiu et al., 2005) melanotic mass \| larval stage (Huang et al., 2005) plasmatocyte \| supernumerary (with lwr⁵) (Huang et al., 2005)
Detailed Description
	Statement Reference Loss of sumoylation in lwr[4-3]/lwr[5] mutants results in hyperactive immunity with blood cells infiltrating the fat body upon L.victoriae parasitic wasp infection. (Paddibhatla et al., 2010) Over 90% of lwr^4-3/lwr⁵ larvae have melanotic tumours, which are surrounded by lamellocytes, while only ~20% of lwr^4-3/lwr¹⁴ larvae show this phenotype. Four days after egg lay, percentages of circulating hemocytes and lamellocytes are significantly higher in lwr^4-3/lwr⁵ mutants than in controls, while lwr^4-3/lwr¹⁴ mutants show high lamellocyte percentages, but have circulating hemocyte levels that are within control range. Levels of circulating hemocytes continue to increase six days after egg lay in lwr^4-3/lwr⁵ mutants, but do not further increase after eight days. More than 6% of lwr^4-3/lwr¹⁴ hemocytes, in both lymph glands and in circulation, are multinucleate and some hemocytes are abnormally large. lwr^4-3/lwr⁵ mutants also exhibit large hemocytes but show a lower penetrance of multinucleate hemocytes. There is a 5- to 10-fold reduction in crystal cells in both lwr^4-3/lwr⁵ and lwr^4-3/lwr¹⁴ mutants. lwr^4-3/lwr⁵ mutants show a developmental delay and only 25% undergo pupariation on day 8. (Chiu et al., 2005) lwr^4-3 mutants develop melanotic tumours during the third larval stage, which become particularly large in the posterior half of the body. In some cases, hemocytes can invade self-tissues such as fat body cells. The number of hemocytes in the hemolymph of lwr^4-3/lwr⁵ mutant larvae is significantly higher than in wild type and the number of lamellocytes is increased within the hemocyte pool. Most aggregated hemocytes consist of lamellocytes that are partially to fully melanized. Mitotic analyses of hemocytes show that lamellocytes do not divide in circulation so the excess number must be due to proliferation in the lymph gland. However, plasmatocytes and prohemocytes do divide in circulation in lwr^4-3/lwr⁵ mutants, while no such division occurs in wild type. (Huang et al., 2005)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressed by
Statement Reference lwr⁵/lwr^4-3 has immune response defective phenotype, suppressible by cact^E10 (Paddibhatla et al., 2010) lwr⁵/lwr^4-3 has melanotic mass phenotype \| larval stage phenotype, suppressible \| partially by cact^E8/cact^E10 (Chiu et al., 2005) lwr⁵/lwr^4-3 has melanotic mass phenotype \| larval stage phenotype, suppressible \| partially by Df(2L)J4/dl¹ (Chiu et al., 2005) lwr⁵/lwr^4-3 has melanotic mass phenotype \| larval stage phenotype, suppressible \| partially by Df(2L)TW119/Dif¹ (Chiu et al., 2005) lwr⁵/lwr^4-3 has melanotic mass phenotype \| larval stage phenotype, suppressible by Df(2L)TW119/Df(2L)J4 (Chiu et al., 2005) lwr⁵/lwr^4-3 has tumorigenic phenotype, suppressible by cact^E10 (Paddibhatla et al., 2010)
Other
Statement Reference cact^E10, lwr⁵/lwr^4-3 has lethal \| wandering third instar larval stage phenotype (Paddibhatla et al., 2010) lwr⁵/lwr^4-3, spz⁴ has lethal \| wandering third instar larval stage phenotype (Paddibhatla et al., 2010)
Phenotype Manifest In
Suppressed by
Statement Reference lwr⁵/lwr^4-3 has embryonic/larval fat body phenotype, suppressible by cact^E10 (Paddibhatla et al., 2010) lwr⁵/lwr^4-3 has embryonic/larval hemocoel phenotype, suppressible by cact^E10 (Paddibhatla et al., 2010) lwr⁵/lwr^4-3 has embryonic/larval hemocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)J4/Dif² (Huang et al., 2005) lwr⁵/lwr^4-3 has embryonic/larval hemocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)J4/dl¹ (Huang et al., 2005) lwr⁵/lwr^4-3 has embryonic/larval hemocyte \| supernumerary phenotype, suppressible by Df(2L)TW119/Df(2L)J4 (Huang et al., 2005) lwr⁵/lwr^4-3 has hemocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)J4/dl¹ (Chiu et al., 2005) lwr⁵/lwr^4-3 has hemocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)TW119/Dif¹ (Chiu et al., 2005) lwr⁵/lwr^4-3 has hemocyte \| supernumerary phenotype, suppressible by cact^E8/cact^E10 (Chiu et al., 2005) lwr⁵/lwr^4-3 has hemocyte \| supernumerary phenotype, suppressible by Df(2L)TW119/Df(2L)J4 (Chiu et al., 2005) lwr⁵/lwr^4-3 has lamellocyte \| supernumerary phenotype, suppressible by Df(2L)J4/Dif² (Huang et al., 2005) lwr⁵/lwr^4-3 has lamellocyte \| supernumerary phenotype, suppressible by Df(2L)TW119/Df(2L)J4 (Huang et al., 2005, Chiu et al., 2005) lwr⁵/lwr^4-3 has plasmatocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)J4/Dif² (Huang et al., 2005) lwr⁵/lwr^4-3 has plasmatocyte \| supernumerary phenotype, suppressible \| partially by Df(2L)J4/dl¹ (Huang et al., 2005)
NOT suppressed by
Statement Reference lwr⁵/lwr^4-3 has lamellocyte \| supernumerary phenotype, non-suppressible by cact^E8/cact^E10 (Chiu et al., 2005) lwr⁵/lwr^4-3 has lamellocyte \| supernumerary phenotype, non-suppressible by Df(2L)J4/dl¹ (Huang et al., 2005, Chiu et al., 2005) lwr⁵/lwr^4-3 has lamellocyte \| supernumerary phenotype, non-suppressible by Df(2L)TW119/Dif¹ (Chiu et al., 2005)
Additional Comments
Genetic Interactions
Statement Reference A cact[E10] background suppresses blood cell infiltration and tumorigenesis in lwr[4-3]/lwr[5] mutants upon L.victoriae parasitic wasp infection. (Paddibhatla et al., 2010) The penetrance of melanotic tumours is reduced from over 90% in lwr^4-3/lwr⁵ mutants to less than 2% in lwr^4-3 Df(2L)TW119/lwr⁵ Df(2L)J4 triple mutants. The amount of circulating hemocytes and lamellocytes are reduced by day 8 in lwr^4-3 Df(2L)TW119/lwr⁵ Df(2L)J4 triple mutants compared to lwr^4-3/lwr⁵ single mutants. The melanotic tumour penetrance and circulating hemocyte levels are reduced in lwr^4-3 Df(2L)J4/lwr⁵ dl¹ and lwr^4-3 Df(2L)TW119/lwr⁵ Dif¹ double mutants, but to a lower extent than in lwr^4-3 Df(2L)TW119/lwr⁵ Df(2L)J4 triple mutants. However, the amount of lamellocytes is not reduced in the double mutants. The melanotic tumour penetrance is reduced in lwr^4-3 cact^E8/lwr⁵ cact^E10 double mutants compared to lwr^4-3/lwr⁵ single mutants and by day 8, the amount of circulating hemocytes is reduced in the double mutants. In contrast, the amount of lamellocytes remains as high as in the single mutants. (Chiu et al., 2005) In lwr^4-3/lwr⁵; dl¹/Df(2L)J4, total numbers of hemocytes are reduced compared to lwr^4-3/lwr⁵ single mutants. Plasmatocyte levels are reduced by 50% compared to single mutants, while lamellocyte levels are not significantly reduced. In lwr^4-3/lwr⁵; Dif²/Df(2L)J4 double mutants the number of lamellocytes are the same as in wild-type larvae, while the number of plasmatocytes is reduced compared to lwr^4-3/lwr⁵ single mutants, but remain higher than in wild type. In lwr^4-3/lwr⁵; Df(2L)TW119/Df(2L)J4 double mutants, the total number of hemocytes, including lamellocytes, is the same as wild type. (Huang et al., 2005)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Complements	lwr⁰²⁸⁵⁸ (Apionishev et al., 2001) lwr^semi710 (Apionishev et al., 2001)
Fails to complement	lwr⁵ (Apionishev et al., 2001)
Rescued by	lwr⁵/lwr^4-3 is rescued by Scer\GAL4^76B/lwr^Scer\UAS.cAa (Paddibhatla et al., 2010)
Partially rescued by	lwr⁵/lwr^4-3 is partially rescued by lwr^Scer\UAS.cAa/Scer\GAL4^Cg.PA (Chiu et al., 2005) lwr⁵/lwr^4-3 is partially rescued by lwr^Scer\UAS.cAa/Scer\GAL4^e22c (Apionishev et al., 2001) lwr⁵/lwr^4-3 is partially rescued by Scer\GAL4^nos.PG/lwr^Scer\UAS.cAa (Apionishev et al., 2001)
Comments	Expression of lwr[Scer\UAS.cAa] in a subset of the lymph gland cells under the control of Scer\GAL4[76B] significantly reduced the infiltration index in lwr[4-3]/lwr[5] mutants, indicating an anti-inflammatory role for lwr. (Paddibhatla et al., 2010) Expression of lwr^Scer\UAS.cAa under the control of Scer\GAL4^Cg.PA partially rescues the developmental delay of lwr^4-3/lwr⁵ mutants, increasing the amount of animals undergoing pupariation from 25 to 61% at day 8. However, there is no clear rescue of the melanotic tumour phenotype. (Chiu et al., 2005) lwr^Scer\UAS.cAa rescues lwr^4-3/lwr⁵ females to adulthood when expressed using Scer\GAL4^e22c or Scer\GAL4^nos.PG, however the rescued flies are sterile. The ovaries develop to stage 8 or 9 (Scer\GAL4^nos.PG) or stage 2 (Scer\GAL4^e22c). (Apionishev et al., 2001)
Stocks ( 1 )
Bloomington	9321 y¹ w^*; lwr^4-3 P{neoFRT}40A/CyO, y⁺
Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	lwr^4-3 (Miles et al., 2008, Sánchez et al., 2010, Talamillo et al., 2008, Reo et al., 2010) Ubc9^4-3 (Paddibhatla et al., 2010)
Name Synonym
Secondary FlyBase IDs

References ( 9 )
Research paper	Paddibhatla et al., 2010, PLoS Pathog. 6(12): e1001234 Role for sumoylation in systemic inflammation and immune homeostasis in Drosophila larvae. [FBrf0212643] Reo et al., 2010, Nucleic Acids Res. 38(13): 4254--4262 Sumoylation of Drosophila SU(VAR)3-7 is required for its heterochromatic function. [FBrf0211327] Sánchez et al., 2010, J. Biol. Chem. 285(33): 25841--25849 Sumoylation Modulates the Activity of Spalt-like Proteins during Wing Development in Drosophila. [FBrf0211449] Miles et al., 2008, Genes Dev. 22(18): 2578--2590 Medea SUMOylation restricts the signaling range of the Dpp morphogen in the Drosophila embryo. [FBrf0205976] Talamillo et al., 2008, Development 135(9): 1659--1668 Smt3 is required for Drosophila melanogaster metamorphosis. [FBrf0205327] Chiu et al., 2005, Dev. Biol. 288(1): 60--72 dUbc9 negatively regulates the Toll-NF-kappa B pathways in larval hematopoiesis and drosomycin activation in Drosophila. [FBrf0190102] Huang et al., 2005, Dev. Biol. 280(2): 407--420 The lesswright mutation activates Rel-related proteins, leading to overproduction of larval hemocytes in Drosophila melanogaster. [FBrf0187386] Sun et al., 2003, D. I. S. 86: 79--83 Isolation of amorphic alleles of the lesswright gene by P element-mediated male recombination in Drosophila melanogaster. [FBrf0179595] Apionishev et al., 2001, Genes Cells 6(3): 215--224 The Drosophila UBC9 homologue lesswright mediates the disjunction of homologues in meiosis I. [FBrf0134637]

Allele Dmel\lwr4-3

Allele Dmel\lwr^4-3