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General Information
Symbol
Dmel\dpphr4
Species
D. melanogaster
Name
FlyBase ID
FBal0003066
Feature type
allele
Associated gene
Associated Insertion(s)
Carried in Construct
Also Known As
dpp4, dpphin-r4
Nature of the Allele
Mutations Mapped to the Genome
 
Type
Location
Additional Notes
References
point mutation
Nucleotide change:

G2457998A

Reported nucleotide change:

G2392A

Amino acid change:

G402E | dpp-PA; G402E | dpp-PB; G402E | dpp-PC; G402E | dpp-PE

Reported amino acid change:

G402E

Associated Sequence Data
DNA sequence
Protein sequence
 
 
Progenitor genotype
Cytology
Nature of the lesion
Statement
Reference

Amino acid replacement: G402E.

Mutation falls in the N-terminal pro region of the dpp protein.

Nucleotide substitution: G2392A.

Point mutation.

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

cytoneme & dorsal mesothoracic disc | somatic clone (with dpphr56)

denticle belt & thorax

Detailed Description
Statement
Reference

dpphr4 homozygous, but not heterozygous, embryos present a ventralized phenotype, as compared to controls.

Homozygous embryos have a ventralised cuticle with a short curved body, dorsally extended denticles, herniated head and defective Filzkorper.

Crossing dpphr4/+ males to wild-type females results in viable heterozygous dpphr4/+ progeny.

7-day old dpphr4/+ germaria have normal numbers of germline stem cells (GSCs), but at 63-days old, these germaria have significantly lower GSC numbers compared to controls - most have only one or no GSCs. 35-day old and 63-day old heterozygous germaria also contain significantly fewer cysts than controls, although cap cells are maintained at normal numbers.

dppd5/dpphr4 flies show reductions in wing vein spacing and loss of wing veins.

dppd5/dpphr4 wings show a truncation of both the L2 and L4 veins, while dppd6/dpphr4 wings have only two veins, L3 and L5 and are missing L2 and L4 completely.

Wing size is reduced in dpphr4/dppd6 flies.

When single cell dpphr4/dpphr56 clones are made in the wing imaginal disc, cytonemes in the wing primordium are affected. At the non-permissive temperature (29oC) cytonemes emanating from cells at the lateral flanks of discs do not orient themselves towards the anterior posterior organiser (as those see in wild-type do). Cytonemes in these discs are also more numerous than in wild-type, and unlike in the wild-type, are sometimes seen to be bent, curved or crossing over each other. These effects are not observed if mutant larvae are heatshocked and returned to permissive temperatures.

dpps6/dpphr4 transheterozygotes are adult viable and exhibit occasional wing vein defects. Truncation of longitudinal vein 5 (L5) is observed in approximately 11% of adults, while 2% display truncations of L4 and L5.

dpps6/dpphr4 Df(2L)C28 flies exhibit an increased frequency of L5 wing vein defects (95% compared to 11% in dpps6/dpphr4 flies). Approximately 20% show defects in L4 and L5 wing veins, and 5% exhibit defects in L5 and L2, or L5 and the posterior crossvein, indicating an enhancement of the dpps6/dpphr4 wing vein phenotype.

No lethality is associated with either the dpps6/dpphr4 or dpps6/dpphr4 Df(2L)C28 genotypes.

dpps6/dpphr4 Df(3R)E40 flies exhibit an increased frequency of L5 wing vein defects (28% compared to 11% in dpps6/dpphr4 flies).

dpphr4/dpphr56 males raised at 18oC to eclosion and then shifted to 29oC, show no significant loss of germline stem cells after 1 week.

Approximately 50% of eggs laid by heterozygous females have shortened and abnormally shaped respiratory appendages.

dpphr4/dpps6 causes relatively few wings to have abnormal venation.

Females carrying temperature sensitive combinations dppe90/dpphr56 or dpphr4/dpphr56 after shifting to the restrictive temperature for one week show 40-50% of germaria that are significantly smaller than wild type. The effect is more pronounced in older females. Stem cell number is reduced, stem cells have differentiated into cysts. Some effect on egg chamber budding is seen.

Eye discs from dpphr4/dpphr56 larvae grown at 29oC for 48 hrs show several phenotypes including failure of retinal differentiation or partial ommatidial differentiation, ommatidia are retricted to the posterior part of the disc and absent from the lateral margins. Occasionally eye discs show normal size and morphology. These eye discs show anterior-most ommatidia that are abnormal mature in comparison to wild type ommatidia at similar positions. Also, cell death is observed ahead of the differentiated region. Surviving adults show slightly roughened eyes that are reduced along the anterior-posterior axis, consistent with a failure of normal morphgenetic progression.

Embryos are ventralised. dpphr4/dppd6 transheterozygotes exhibit reduced wings to approximately half wild type size and no defects in eyes or legs.

Homozygotes produce approximately 30 amnioserosa cells. This phenotype is not altered if the embryos are also carrying 2 copies of tsgtwi.PM.

Embryos show cuticular holes, disorganization of the head skeleton, internalization of the seventh and eighth abdominal segments and the filzkorper and a thoracic constriction. The first and second ventral denticle belts are expanded. A significant percentage of surviving transheterozygotes between dpphr56 and dpphr4 or dpphr27 have a split notum.

Homozygous embryos have 6 +/- 10 amnioserosa cells (compare to 164 +/- 23 cells in wild type embryos), many embryos lack amnioserosa entirely. Heterozygotes have 124 +/- 9 amnioserosa cells, a fraction of embryos die.

dpphr4/dppd-ho transheterozygotes show a wild type wing phenotype in an otherwise normal genetic background.

Heterozygotes are viable, heterozygotes derived from Mad-/Mad+ mothers frequently die. Transheterozygotes with dppd6 have wings approximately half normal size and normal legs and eyes (less severe than dppd6 homozygous phenotype). When in combination with Df(2L)C28/Mad+ the transheterozygotes exhibit more severe phenotype, further reduction in wing blade, slight reduction in the eye and loss of tarsal claws. Trisomy for Mad (Dp(2;3)JS20) partially ameliorates transheterozygote phenotype, wings are of normal size and show residual vein fusions.

dppd6/dpphr4 individuals have a nearly full size wing, except for the absence of the anterior crossvein, a reduction in the distance between L4 and L5 and a small amount of ectopic vein between L2 and L3. If the flies are also heterozygous for (paternally contributed) sax1 or sax2, L2 is lost and L4 and L5 fused.

Survival of heterozygotes is reduced if also heterozygous for certain alleles of tld.

Embryos display a moderately ventralised phenotype, maternally contributed tkv alleles cause lethality.

Clonal analysis in the developing eye (using the FLP/FRT system) revealed nonautonomous phenotypes with large clones showing posterior-lateral eye parts missing.

Dominant lethality less than 50%. Homozygous and transheterozygous embryos were examined with respect to 25 cuticular markers, results demonstrate a graded requirement for dpp along the dorso-ventral axis.

Weak ventralised phenotype. Rings of ventral denticle belts differentiate around the entire dorsoventral axis, almost no dorsal hairs are seen and the antennal and maxillary sense organs are missing. Defective movements of the germ band: due to loss of the amnioserosa and because the dorsalmost cells have acquired the lateral fate of the dorsal ectoderm. Dorsal cell fates are deleted and ventrolateral mitotic domains are expanded.

Fully rescued to adulthood by two wild type copies of dpp.

z1 product subliminally inhibits the complementation of dpphr4 and dppd-ho. The effect is only seen when compounded by the particular effects of rearrangements on synapsis.

Abnormal development begins after germ band extension.

Heterozygotes exhibit wings that are stumps. The wings are held out laterally instead of in line with the longitudinal axis of the body. Transheterozygotes with dppd-ho exhibit normally oriented wings.

Heterozygotes with class I dpp alleles have a wild type wing phenotype. Heterozygotes with class II allele dppd5 exhibit wings with a deletion of the anterior crossvein. Heterozygotes with class II allele dppd28 exhibit held out wings that are missing longitudinal and cross veins, deletion of tarsal claws, male terminalia and female analia. Heterozygotes with class III dpp alleles display reduced held out wings that are missing longitudinal and cross veins, capitella-less halteres, reduced eyes, legs missing claws and missing or rotated male terminalia or female analia. Heterozygotes with class IV dpp alleles display the same phenotype as class III heterozygotes with duplication of scutellar bristles, medial cleft of the dorsal notum, duplication of sex combs and partial duplication of the third antennal segment. Heterozygotes with class V dpp alleles rarely eclose, the surviors exhibit the wing, haltere, eye and terminalia defects. At 18oC the heterozygotes have less severe phenotypes.

External Data
Interactions
Show genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement
Reference
Suppressed by
Statement
Reference

dpphr4/dpp[+], sax1 has lethal | dominant | maternal effect phenotype, suppressible by Dp(2;2)DTD48/+

dpphr4/dpp[+], sax1 has lethal | dominant | maternal effect phenotype, suppressible by Dp(2;2)B16/+

dpphr4/dpp[+], sax2 has lethal | dominant | maternal effect phenotype, suppressible by Dp(2;2)DTD48/+

dpphr4/dpp[+], sax2 has lethal | dominant | maternal effect phenotype, suppressible by Dp(2;2)B16/+

NOT Enhancer of
Statement
Reference
NOT Suppressor of
Statement
Reference
Other
Statement
Reference
Phenotype Manifest In
Enhanced by
Statement
Reference

dppd5/dpphr4 has wing vein L2 phenotype, enhanceable by gbb1/gbb[+]

dpps6/dpphr4 has wing vein phenotype, enhanceable by Mad1

dpps6/dpphr4 has wing vein phenotype, enhanceable by Med7

dpps6/dpphr4 has wing vein L5 phenotype, enhanceable by Med7

dpps6/dpphr4 has wing vein L4 phenotype, enhanceable by Med7

dpphr4 has phenotype, enhanceable by Med11/Med[+]

dpphr4 has phenotype, enhanceable by Med[+]/Med12

dpphr4 has phenotype, enhanceable by Med13/Med[+]

dpphr4 has phenotype, enhanceable by Med14/Med[+]

dpphr4 has phenotype, enhanceable by Med[+]/Med15

dpphr4 has phenotype, enhanceable by Med16/Med[+]

dpphr4 has phenotype, enhanceable by Med17/Med[+]

dpphr4 has phenotype, enhanceable by Med[+]/Med18

dpphr4 has phenotype, enhanceable by Df(3L)66C-G28/+

dpphr4 has wing vein phenotype, enhanceable by Df(2R)Pcl11B/+

dpphr4 has wing vein phenotype, enhanceable by Df(2R)Pcl7B/+

dpphr4/dppd6 has wing phenotype, enhanceable by vri[+]/vri1

dpphr4/dppd6 has wing phenotype, enhanceable by vri2/vri[+]

dpphr4 has phenotype, enhanceable by vri5R4.11

dpphr4 has phenotype, enhanceable by vri5R5.24

dpphr4 has phenotype, enhanceable by vri5R5.7

dpphr4 has phenotype, enhanceable by vri5R7.2

dpphr4 has phenotype, enhanceable by vri5R8.4

dpphr4 has phenotype, enhanceable by vrik03801

dpphr4 has phenotype, enhanceable by vrik05901

dpphr4 has phenotype, enhanceable by vrik09602

dpphr4 has phenotype, enhanceable by vrik09713

dpphr4 has phenotype, enhanceable by vrik11805

dpphr4 has phenotype, enhanceable by Mad2

dpphr4 has phenotype, enhanceable by Mad3

dpphr4 has phenotype, enhanceable by Mad4

dpphr4 has phenotype, enhanceable by Med1

dpphr4 has phenotype, enhanceable by Med2

dpphr4 has phenotype, enhanceable by Med3

dpphr4 has phenotype, enhanceable by Med4

dpphr4 has phenotype, enhanceable by scwE1

dpphr4 has phenotype, enhanceable by scwE2

dpphr4 has phenotype, enhanceable by tld14

dpphr4 has phenotype, enhanceable by tldE1

NOT Enhanced by
Statement
Reference

dppSal20, dpphr4 has phenotype, non-enhanceable by Df(3L)66C-G28/+

Suppressed by
Statement
Reference

dpphr4, lolal[+]/lolal1122 has embryo | maternal effect phenotype, suppressible | partially by lolalUASp.cQa

dpphr4, lolal[+]/lolal1122 has embryo | maternal effect phenotype, suppressible | partially by tkvQ253D.UASp

dpphr4, lolal[+]/lolal1122 has embryo | maternal effect phenotype, suppressible | partially by MedUASp.cSa

dpphr4, lolal[+]/lolal1122 has embryo phenotype, suppressible | partially by MedK738R.UASp

NOT suppressed by
Statement
Reference
Enhancer of
Statement
Reference
NOT Enhancer of
Statement
Reference
Suppressor of
Statement
Reference

dpphr4/dpp[+] is a suppressor | partially of denticle phenotype of sogY506

dppd-ho/dpphr4 is a suppressor of eye phenotype of CycEJP

NOT Suppressor of
Statement
Reference
Other
Statement
Reference
Additional Comments
Genetic Interactions
Statement
Reference

The progeny from dpphr4/+ fathers exhibit near lethality when mothers are either Df(2R)Pcl11B/+, lolalk02512/+ or lolal1122/+, and exhibit increased lethality when mothers are either lolal1722/+ or lolalG9603/+, as compared to controls mothers; there is no obvious increased lethality as compared to controls if these parental genders are reversed. The progeny from dpphr4/+ fathers exhibit near lethality when mothers are Med15/+, and semi-lethality when mothers are Mad12/+, but not when mothers are Df(2R)12/+, as compared to controls mothers.

The near lethality of the progeny from Df(2R)Pcl11B/+ mothers and dpphr4/+ fathers is weakly rescued by the maternal expression of either lolalScer\UAS.P\T.cQa or lolalScer\UAS.P\T.T:Ivir\HA1, either under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 or in the absence of the Gal4 driver.

The near lethality of the progeny from lolal1122/+ mothers and dpphr4/+ fathers is fully rescued by the maternal expression of either tkvQ253D.Scer\UAS.P\T or MedK738R.Scer\UAS.P\T, either under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 or in the absence of the Gal4 driver, is fully rescued by the maternal expression of either MedScer\UAS.P\T.cSa or TrlEY04024, under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 but only partially rescued in the absence of the Gal4 driver, and is partially rescued by the maternal expression of either lolalScer\UAS.P\T.cQa, lolalScer\UAS.P\T.T:Ivir\HA1 or saxQ263D.Scer\UAS.cDa, either under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 or in the absence of the Gal4 driver.

The progeny from dpphr4/+ fathers present embryo ventralization in most cases if mothers are Med15/+, and present embryo ventralization in less than half of cases if mothers are either Mad12/+, Df(2R)Pcl11B/+, lolalk02512/+, lolal1122/+, lolal1722/+, lolalG9603/+, msk5/+ or mskB185/+, as compared to controls; the embryo ventralization of the progeny from dpphr4/+ fathers and lolal1122/+ mothers is fully rescued by the maternal expression of tkvQ253D.Scer\UAS.P\T under the control of Scer\GAL4nos.UTR.T:Hsim\VP16, but only partially in the absence of the Gal4 driver, and is partially rescued by the maternal expression of either lolalScer\UAS.P\T.cQa, lolalScer\UAS.P\T.T:Ivir\HA1, saxQ263D.Scer\UAS.cDa, MedScer\UAS.P\T.cSa or MedK738R.Scer\UAS.P\T, either under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 or in the absence of the Gal4 driver.

Embryos derived from fafFO8/+ females mated to dpphr4/+ males show strong lethality (only 10% of the expected number survive). The embryos have a ventralised cuticle phenotype.

Animals derived from homozygous fafEP381 females mated to dpphr4/+ males show strong lethality (only 26% of the expected number survive). The embryos have a ventralised cuticle phenotype.

Animals derived from Med8/+ females mated to dpphr4/+ males show strong lethality (5% of the expected number survive). The embryos have a ventralised cuticle phenotype.

Animals derived from homozygous Med15 or Med17 females mated to dpphr4/+ males show strong lethality (less than 5% of the expected number survive).

dpphr4/+ partially suppresses the cuticle phenotype of sogY506 hemizygous embryos, partially restoring denticles and producing a curved body.

The survival of the progeny of fafFO8/+ females mated to dpphr4/+ males is increased 1.6-fold by basal expression of MedScer\UAS.P\T.cSa (in the absence of a Scer\GAL4 driver) in the females, while basal expression of MedK738R.Scer\UAS.P\T in the females increases survival of the progeny 4.2-fold. Expression of either MedScer\UAS.P\T.cSa or MedK738R.Scer\UAS.P\T under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 in the females increases survival of the progeny to approximately 80% in both cases.

The ventralised cuticle phenotype of embryos derived from fafFO8/+ females mated to dpphr4/+ males is not rescued by basal expression of MedScer\UAS.P\T.cSa (in the absence of a Scer\GAL4 driver) in the females, while basal expression of MedK738R.Scer\UAS.P\T in the females results in the rescue of the ventralised cuticle phenotype in some embryos. Expression of either MedScer\UAS.P\T.cSa or MedK738R.Scer\UAS.P\T under the control of Scer\GAL4nos.UTR.T:Hsim\VP16 in the females effectively rescues the ventralised cuticle phenotype of the embryos.

icoGA1/+, dpphr4/+ animals die as embryos.

icoGA25/+, dpphr4/+ animals die as embryos.

When dpphr4/+ males are crossed to sax1/+ or sax2/+ females, all the resulting dpphr4/+ progeny die. Similarly, any one of the paternal dppe87/+, dpphr56/+, dpphr90/+ or dpphr27/+ genotypes in combination with either one of the maternal sax1/+ or sax2/+ genotypes results in increased dpp zygotic haplolethality in the progeny.

The maternal enhancement of the dpphr4/+ zygotic haplolethality by sax1/+ is suppressed by either Dp(2;2)DTD48/+ or Dp(2;2)B16/.

The maternal enhancement of the dpphr4/+ zygotic haplolethality by sax2/+ is suppressed by either Dp(2;2)DTD48/+ or Dp(2;2)B16/.

When dpphr4/+ males are crossed to females of any one of the Df(2R)ST1/+, Df(2R)P32a/+ or Df(2R)H23/+ genotypes, similarly to wild-type flies, most of the resulting dpphr4/+ progeny survive.

When saxP/+ females are crossed to dpphr4/+ males, most of the resulting dpphr4/+ progeny die.

When sax1rv1/+ females are crossed to dpphr4/+ males, the resulting dpphr4/+ progeny show a moderate drop in viability compared to the dpphr4/+ progeny of wild-type mothers in control crosses.

When sax6/+ females are crossed to dpphr4/+ males, the resulting dpphr4/+ progeny show a moderate drop in viability compared to the dpphr4/+ progeny of wild-type mothers in control crosses.

When dpp[hr4]/+ males are crossed to females of any one of the genotypes sax1rv5/+, sax3/+, sax4/+ and sax5/+, the resulting dpphr4/+ progeny do not show any significant drop in viability compared to the dpphr4/+ progeny of wild-type mothers in control crosses.

Flies that express tkvScer\UAS.cNa, under the control of Scer\GAL4tub, in a dpphr4/dppd6 background, have smaller wings than flies expressing the transgene in a wild-type background and than dpphr4/dppd6 flies that do not express tkvScer\UAS.cNa.

bamhs.PO animals cultured at 18oC and heat shocked as late stage pupae (37oC, 4 times for 2 hours each with an 8-12 hour interval) produce adult females with agametic germaria. If these animals also carry the temperature sensitive allelic combination dpphr56/dppe90, and are shifted to 29oC (the non-permissive temperature) for 10 days, the resulting agametic germaria contain only slightly less follicle cells than controls (bamhs.PO only).

dpps6/dpphr4 Mad1 flies exhibit L4 and L5 wing vein defects in almost 100% of cases. Approximately 10% of wings with L4 and L5 defects exhibit missing crossveins (anterior, posterior, or both) and small wing margin notches. This indicates an enhancement of the dpps6/dpphr4 phenotype by Mad1.

Only 71% of the expected dpps6/dpphr4 Mad1 progeny are recovered, indicating a statistically significant amount of lethality.

dpps6/dpphr4 Med7 flies exhibit L4 and L5 wing vein defects in approximately 42% of cases, with 15% of these wings also exhibiting defects in L2, or the posterior crossvein or margin notches.

dpphr4/dpphr56; gbbD4/+ males raised at 18oC to eclosion and then shifted to 29oC for one week have an average of 3.1 germ-line stem cells per testis, compared to 8.6 in gbbD4/+.

In(2LR)DTD52, dppd-ho/In(2LR)DTD11, dpphr4 flies have wild-type wing posture. Df(2L)DTD52-D51, dppd-ho/In(2LR)DTD11, dpphr4 flies have a heldout wing posture.

71% of the expected cype03771/dpphr4 adult progeny are recovered from a cross of dpphr4/+ females to cype03771/+ males. Only 38% of the expected cype03771/dpphr4 adult progeny are recovered from a cross of cype03771/+ females to dpphr4/+ males. The lethality is embryonic and the embryos are weakly to moderately ventralised.

Enhanced lethality in heterozygous individuals from crosses involving Df(2R)Pcl11B or Df(3L)66C-G28 acts during embryonic and larval stages, though cuticle preparations from affected embryos reveal no consistent abnormalities. The incidence of head skeleton herniation is increased compared to that seen for singly dpphr4 mutant embryos. The incidence of abnormal venation in dpphr4/dpps6 is increased by Df(2R)Pcl11B or Df(2R)Pcl7B.

When in combination with vri mutations, vrik05901 lethal revertant lines or Df(2L)tkv2 lethality is greatly increased and embryos are more strongly ventralised. vri1/+ or vri2/+ causes a further reduction in wing size and wing veins of dpphr4/dppd6 transheterozygotes. Eyes are smaller and rough and legs are truncated.

Homozygous lethal phenotype can be enhanced by Med, Mad, tld and scw enhancer mutations.

The dpphr4/dppd-ho phenotype is dominantly enhanced by en96 (In(2R)DTD96), to cause heldout wings.

Increased dose of tld+ does not suppress the mutant phenotype.

Enhancement of tor13D embryos: decrease in proportion of embryos that formed cuticle, or cuticle with denticles and increase in proportion of embryos that formed empty sacs.

Xenogenetic Interactions
Statement
Reference
Complementation and Rescue Data
Partially complements
Comments

Lethal interaction between dpphr4 and Df(2R)Pcl11B, Df(3L)66C-G28 and Mad12 can be rescued by dppSal20.

The transheterozygous combination dpphr4/dpphr56 allows development to adult-hood in a temperature-sensitive manner. The permissive temperature is 17oC and the restrictive temperature is 29oC.

Allele class: hin-r

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Mutant
Wild-type
Stocks (1)
Notes on Origin
Discoverer
Comments
Comments

Based on considerations of degree of dominant lethality, dpp alleles can be placed in an allelic series. Progressing from weakest to most severe the series is: dppe87 < dppe90 < dpphr56 < dpphr4 < dpphr92 < dpphr27 < dpphr93 < dppH94 < dppH95 < dppH96 = dppH91 = dppH46.

dpphr4 shows interallelic complementation at 18oC with dpphr56 and dppe90.

External Crossreferences and Linkouts ( 0 )
Synonyms and Secondary IDs (8)
References (64)