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General Information
D. melanogaster
FlyBase ID
Feature type
Associated gene
Associated Insertion(s)
Carried in Construct
Also Known As
Nature of the Allele
Mutations Mapped to the Genome
Additional Notes
Associated Sequence Data
DNA sequence
Protein sequence
Progenitor genotype
Nature of the lesion

Imprecise excision of the P{PZ} element. 34bp of P-element sequences remain, together with 42bp of inserted material, repeats of TAACA.

Insertion components
Expression Data
Reporter Expression
Additional Information
Marker for
Reflects expression of
Reporter construct used in assay
Human Disease Associations
Disease Ontology (DO) Annotations
Models Based on Experimental Evidence ( 1 )
Modifiers Based on Experimental Evidence ( 0 )
Comments on Models/Modifiers Based on Experimental Evidence ( 0 )
Phenotypic Data
Phenotypic Class
Phenotype Manifest In
Detailed Description

Loss of sumoylation in lwr4-3/lwr5 mutants results in hyperactive immunity with blood cells infiltrating the fat body upon L.victoriae parasitic wasp infection.

Over 90% of lwr4-3/lwr5 larvae have melanotic tumours, which are surrounded by lamellocytes, while only ~20% of lwr4-3/lwr14 larvae show this phenotype. Four days after egg lay, percentages of circulating hemocytes and lamellocytes are significantly higher in lwr4-3/lwr5 mutants than in controls, while lwr4-3/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 lwr4-3/lwr5 mutants, but do not further increase after eight days. More than 6% of lwr4-3/lwr14 hemocytes, in both lymph glands and in circulation, are multinucleate and some hemocytes are abnormally large. lwr4-3/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 lwr4-3/lwr5 and lwr4-3/lwr14 mutants. lwr4-3/lwr5 mutants show a developmental delay and only 25% undergo pupariation on day 8.

lwr4-3 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 lwr4-3/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 lwr4-3/lwr5 mutants, while no such division occurs in wild type.

External Data
Show genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Phenotype Manifest In
Suppressed by
NOT suppressed by
Enhancer of

lwr[+]/lwr4-3 is an enhancer of wing phenotype of tna1

Additional Comments
Genetic Interactions

lwr4-3/+ enhances both the penetrance and expressivity of the held-out wing phenotype of tna1/+ flies.

A cactE10 background suppresses blood cell infiltration and tumorigenesis in lwr4-3/lwr5 mutants upon L.victoriae parasitic wasp infection.

The penetrance of melanotic tumours is reduced from over 90% in lwr4-3/lwr5 mutants to less than 2% in lwr4-3 Df(2L)TW119/lwr5 Df(2L)J4 triple mutants. The amount of circulating hemocytes and lamellocytes are reduced by day 8 in lwr4-3 Df(2L)TW119/lwr5 Df(2L)J4 triple mutants compared to lwr4-3/lwr5 single mutants. The melanotic tumour penetrance and circulating hemocyte levels are reduced in lwr4-3 Df(2L)J4/lwr5 dl1 and lwr4-3 Df(2L)TW119/lwr5 Dif1 double mutants, but to a lower extent than in lwr4-3 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 lwr4-3 cactE8/lwr5 cactE10 double mutants compared to lwr4-3/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.

In lwr4-3/lwr5; dl1/Df(2L)J4, total numbers of hemocytes are reduced compared to lwr4-3/lwr5 single mutants. Plasmatocyte levels are reduced by 50% compared to single mutants, while lamellocyte levels are not significantly reduced. In lwr4-3/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 lwr4-3/lwr5 single mutants, but remain higher than in wild type. In lwr4-3/lwr5; Df(2L)TW119/Df(2L)J4 double mutants, the total number of hemocytes, including lamellocytes, is the same as wild type.

Xenogenetic Interactions
Complementation and Rescue Data
Fails to complement
Partially rescued by

Expression of lwrScer\UAS.cAa in a subset of the lymph gland cells under the control of Scer\GAL476B significantly reduced the infiltration index in lwr4-3/lwr5 mutants, indicating an anti-inflammatory role for lwr.

Expression of lwrScer\UAS.cAa under the control of Scer\GAL4Cg.PA partially rescues the developmental delay of lwr4-3/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.

lwrScer\UAS.cAa rescues lwr4-3/lwr5 females to adulthood when expressed using Scer\GAL4e22c or Scer\GAL4nos.PG, however the rescued flies are sterile. The ovaries develop to stage 8 or 9 (Scer\GAL4nos.PG) or stage 2 (Scer\GAL4e22c).

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Stocks (1)
Notes on Origin
External Crossreferences and Linkouts ( 0 )
Synonyms and Secondary IDs (2)
References (12)