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General Information
Symbol
Dmel\zip2
Species
D. melanogaster
Name
FlyBase ID
FBal0018863
Feature type
allele
Associated gene
Associated Insertion(s)
Carried in Construct
Also Known As
zipIIF107
Nature of the Allele
Mutations Mapped to the Genome
 
Type
Location
Additional Notes
References
point mutation
Nucleotide change:

C24995643T

Reported nucleotide change:

C15798T

Amino acid change:

Q835term | zip-PA; Q790term | zip-PB; Q750term | zip-PC; Q795term | zip-PD; Q750term | zip-PE; Q758term | zip-PF; Q750term | zip-PG; Q758term | zip-PH; Q803term | zip-PI

Reported amino acid change:

Q751term

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

Amino acid replacement: Q751term.

The mutation results in a protein that is truncated by 1222 amino acid residues compared to wild-type.

Nucleotide substitution: C15798T.

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
Detailed Description
Statement
Reference

In the larval eye-antennal disc, zip2 mutant clones at the A1 and Ar epithelial folds show reduced folding, and larger clones exhibit apical swelling and/or delamination, as compared to control clones. Clones that span the A1 fold show a breakdown of the field boundary at the A1 fold. Adult heads with zip2 clones show mislocalized ommatidia in head cuticle and antennae, and antennal-like tissue at the borders of compound eyes, in contrast to controls.

zip2 mutant adults occasionally show necrotic scar-like cells on the head, in contrast to controls.

Zygotic mutant embryos for zip2 show normal dorsal closure.

zip2 homozygous mutants show laterality defects in the proventriculus and anterior midgut. The laterality of the other parts of the embryonic gut, including the hindgut and the posterior part of the midgut is normal in these mutants. The proventriculus and the anterior midgut do not rotate (as in wild-type) in these mutants at stages 15 to 17.

zip3/zip2 embryos show laterality defects in the proventriculus and anterior midgut. The laterality of the other parts of the embryonic gut, including the hindgut and the posterior part of the midgut is normal in these mutants. The proventriculus and the anterior midgut do not rotate (as in wild-type) in these mutants at stages 15 to 17.

zip4/zip2 embryos show laterality defects in the proventriculus and anterior midgut. The laterality of the other parts of the embryonic gut, including the hindgut and the posterior part of the midgut is normal in these mutants. The proventriculus and the anterior midgut do not rotate (as in wild-type) in these mutants at stages 15 to 17.

zip2/zip1 embryos show laterality defects in the proventriculus and anterior midgut. The laterality of the other parts of the embryonic gut, including the hindgut and the posterior part of the midgut is normal in these mutants. The proventriculus and the anterior midgut do not rotate (as in wild-type) in these mutants at stages 15 to 17.

Approximately 20% of zip4 stage 17 embryos show left-right inversion in the looping of the esophagus.

zip1/zip2 stage 16 embryos have longer dorsal trunks than normal.

zip1/zip2 embryos show a dorsal closure phenotype with low penetrance. In transgenic mosaic embryos in which some cells express zipFL.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4e22c in a zip1/zip2 background, leading-edge cells that do not express zipFL.Scer\UAS.T:Avic\GFP cannot maintain tension and are stretched by neighbouring zipFL.Scer\UAS.T:Avic\GFP expressing cells. The length of contiguous regions of Scer\GAL4e22c>zipFL.Scer\UAS.T:Avic\GFP expressing leading-edge cells along the supracellular purse string decrease in length over time, indicative of contraction. In contrast, non zipFL.Scer\UAS.T:Avic\GFP-expressing contiguous regions fail to contract, increasing in length over time. In transgenic mosaic embryos in which some cells express zipFL.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW in a zip1/zip2 background, Scer\GAL4sqh.PW>zipFL.Scer\UAS.T:Avic\GFP expressing leading-edge cells that contact Scer\GAL4sqh.PW>zipFL.Scer\UAS.T:Avic\GFP expressing amnioserosa cells are severely contracted, and the scalloped morphology of the early leading edge is enhanced. In contrast, when non-expressing leading-edge cells contact non-expressing amnioserosa cells the leading edge cells fail to contract and become stretched. When non-expressing leading-edge cells contact Scer\GAL4sqh.PW>zipFL.Scer\UAS.T:Avic\GFP-expressing amnioserosa cells, leading-edge cells show some contraction. Finally, Scer\GAL4sqh.PW>zipFL.Scer\UAS.T:Avic\GFP-expressing cells show some contraction when they contact non-expressing amnioserosa cells, although this is to a lesser extent than when both cell types express zipFL.Scer\UAS.T:Avic\GFP. Nonexpressing leading-edge cells fail to incorporate into the canthus and cause inward progression of the canthus to stall. These nonexpressors never get fully incorporated into the seam - instead, a small gap is formed when zipping fails and closure bypasses the nonexpressing cells to initiate a new seam. In contrast, Scer\GAL4sqh.PW>zipFL.Scer\UAS.T:Avic\GFP-expressing cells are incorporated into each canthus at a similar and nearly constant rate. In transgenic mosaic embryos in which some amnioserosa cells express zipFL.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4c381 in a zip1/zip2 background, nonexpressing amnioserosa cells fail to contract apically and remain rounded. These cells do show an apical shape change at later stages but this may be due to forces generated by surrounding Scer\GAL4c381>zipFL.Scer\UAS.T:Avic\GFP-expressing amnioserosa cells and the approaching lateral-epidermis sheet. Embryos that express zipFL.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4en-e16E in a zip1/zip2 background display stripe-alignment defects in 65% of cases.

Mutant embryos exhibit defects in germ band elongation. Cell intercalation is severely disrupted, leading to quasi-'frozen' tissue. Junction remodelling does not occur, though the epithelium remains intact.

zip2 mutants fail to complete dorsal closure leading to the extrusion of the gut. Mutants also fail to complete head involution. In mutant embryos the muscle VA3 is missing from each segment and VA1 and VA2 are often both not present in each segment. When individual sarcomere are examined the I bands seen in wild-type muscles are absent, instead actin is organised into continuous filaments that run the length of the muscle.

The salivary glands of mutant embryos are wider and shorter than normal, appearing to have bulges and bubbles in their lumens. Malpighian tubules are shorter and slightly thicker than wild type. The tubules have short branches and the lumens widen in some areas.

Homozygous embryos do not complete head involution. The salivary glands are enlarged, with lumens distended by bulges and bubbles. Malpighian tubules are abnormal, remaining coiled near their juncture with the hindgut rather than extending anteriorly as in the wild-type. Abnormal branching of the tubules, consisting of short dead-ends extending only the length of one or two cells, is seen. The diameter of the Malpighian tubules is similar to wild-type. The hindgut appears normal and the midgut constrictions appear substantially similar to wild-type. Abnormalities in the peripheral nervous system cannot be detected.

Does not produce malformed phenotype when in double heterozygosis with second site zip non-complementing mutants.

Mutant phenotype can be rescued to adulthood by P-element mediated introduction of a genomic fragment spanning the entire transcription unit. Rescue by zip cDNA driven from a minimal Hsp70 promoter allows rescue to larvae and intermittant heat shocks allows some rescue to adulthood. During the late dorsal closure stages myosin becomes improperly localised in or absent from the leading edge causing a prominent, irregularly shaped dorsal opening in the cuticle.

The nervous system develops abnormally in homozygotes, with brain tissue located more anteriorly and being larger than normal in the fully developed embryo. It is partially surrounded by the cuticle which never encloses the brain completely. The lateral fascicles of different abdominal segments do not respect segment boundaries in a variable manner. Sensory neurons of the gnathal sensory organs remain in the position of the progenitor cells, and their axons fasiculate in a single, thick axon bundle. Homozygous embryos also have an abnormal cuticle pattern.

weaker allele than zip1

External Data
Interactions
Show genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhancer of
Statement
Reference

zip2/zip[+] is an enhancer of visible phenotype of Drakdel

zip2/zip[+] is an enhancer of visible phenotype of DrakKO

Suppressor of
Statement
Reference
Other
Phenotype Manifest In
Enhanced by
Statement
Reference
Enhancer of
Statement
Reference

zip2/zip[+] is an enhancer of wing phenotype of Drakdel

zip2/zip[+] is an enhancer of wing phenotype of DrakKO

zip2/zip[+] is an enhancer of eye phenotype of Ppt1UAS.cKa, Scer\GAL4GMR.PF/Scer\GAL4GMR.PF

Suppressor of
Statement
Reference

zip2/zip[+] is a suppressor | partially of head | embryonic stage phenotype of stepKG09493/stepk08110

zip2/zip[+] is a suppressor of wing disc phenotype of CskGD9345, Scer\GAL4ptc-559.1

zip2 is a suppressor of dorsal hair phenotype of raw1

zip2 is a suppressor of embryo | dorsal closure stage phenotype of raw1

zip2 is a suppressor of Malpighian tubule phenotype of rib1

zip2 is a suppressor of denticle phenotype of rib1

zip2 is a suppressor of dorsal hair phenotype of rib1

zip2 is a suppressor of Malpighian tubule phenotype of raw1

zip2 is a suppressor of denticle phenotype of raw1

NOT Suppressor of
Statement
Reference

zip2 is a non-suppressor of embryonic midgut constriction phenotype of rib1

zip2 is a non-suppressor of embryo | dorsal closure stage phenotype of rib1

Other
Additional Comments
Genetic Interactions
Statement
Reference

Heterozygosity for zip2 partially suppresses the increased lethality and associated "head hole" phenotype of stepKG09493/stepk08110 transheterozygous.

A zip2/zipEbr mutant background (which reducing tension in cells) induces cell death at the anterior boundary when fweScer\UAS.P\T.Lose-A.C.T:Ivir\HA1 is expressed under the control of Scer\GAL4dpp.PU in the anterior compartment of the wing disc.

A zip2 heterozygous mutant background suppresses the actin remodelling and subsequent basolateral invasion of epithelial cells seen in flies expressing CskGD9345 in a stripe of cells at the anterior/posterior boundary of the larval wing disc under the control of Scer\GAL4ptc-559.1.

par-6Δ226, zip2 double mutant embryos display merged dorsal and head holes.

zip2 dominantly enhances the shortening of the wing which is seen in DrakKO and in Drakdel flies.

A zip2/+ background enhances the visual system degeneration seen in Scer\GAL4GMR.PF, Ppt1Scer\UAS.cKa flies.

Many adults expressing zipR1171C.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4en-e16E in a zip2/+ ; Rho1E3.10/+ background show both leg and wing abnormalities (26%), while others show wing abnormalities only (56%).

Some adults expressing zipD1430N.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4en-e16E in a zip2/+ ; Rho1E3.10/+ background show both leg and wing abnormalities (11%), while others show wing abnormalities only (5.4%).

Some adults expressing zipD1847K.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4en-e16E in a zip2/+ ; Rho1E3.10/+ background show both leg and wing abnormalities (8.2%).

Some adults expressing zipR1933X.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4en-e16E in a zip2/+ ; Rho1E3.10/+ background show both leg and wing abnormalities (14%), while others show wing abnormalities only (4.7%).

Partially suppresses the salivary gland phenotype of raw1 and rib1. Partially suppresses the failure of dorsal closure seen in raw1 embryos; a small amount of dorsal closure of the epidermis is seen. Has no effect on dorsal closure or midgut formation in rib1 embryos. Partially suppresses the mutant phenotype of cuticular denticles and hairs seen in rib1 and raw1 embryos.

Xenogenetic Interactions
Statement
Reference

Expression of two copies of Dpse\l(2)gl+t12.2 raises the penetrance of the zip1/zip2 failed dorsal closure phenotype to 100%. Ubiquitous overexpression of zipFL.Scer\UAS.T:Avic\GFP, under the control of the Scer\GAL4sqh.PW, rescues dorsal closure in more than 84% of zip1/zip2; Dpse\l(2)gl+t12.2/Dpse\l(2)gl+t12.2 embryos. 61% of these rescued flies survive to hatch as larvae, 41% survive to pupal stages and 2.8% eclose as adults. Expression of zipFL.Scer\UAS.T:Avic\GFP in the epidermis and a few amnioserosa cells, driven by Scer\GAL4e22c, rescues the zip1/zip2; Dpse\l(2)gl+t12.2/Dpse\l(2)gl+t12.2 dorsal closure phenotype in 88% of embryos. Rescued flies survive to larval stages in 60% of cases and to pupal stages in 21% of cases, although no flies eclose. Expression of zipFL.Scer\UAS.T:Avic\GFP in the amnioserosa, driven by Scer\GAL4c381, rescues the dorsal closure phenotype of Dpse\l(2)gl+t12.2/Dpse\l(2)gl+t12.2 mutants in 82% of embryos. However, none of the rescued embryos survive to larval stages. Similarly, when zipFL.Scer\UAS.T:Avic\GFP expression is driven in the leading edge by Scer\GAL4LE, dorsal closure is rescued in 87% of embryos, but only 1% of flies survive to larval stages. zipFL.Scer\UAS.T:Avic\GFP expression in epidermal stripes, driven by Scer\GAL4en-e16E, rescues dorsal closure in only half of embryos and does not rescue lethality.

The addition of four copies of Dpse\l(2)gl+t12.2 to zip2 animals results a failure of germband retraction, at approximately 8.2 to 9.2 hours after egg laying, but muscles remained attached to the epidermis.

Complementation and Rescue Data
Comments

Expression of zipFL.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW rescues the embryonic lethality of zip1/zip2 animals. 86 +/- 2% of the rescued animals survive to the pupal stage, but only 6 +/- 7% eclose as adults.

Expression of zipR1171C.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW rescues the embryonic lethality of zip1/zip2 animals. 62 +/- 10% of the rescued animals survive to the pupal stage, but only 18 +/- 12% eclose as adults.

Expression of zipD1430N.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW rescues the embryonic lethality of zip1/zip2 animals. 72 +/- 5% of the rescued animals survive to the pupal stage, but no animals survive to the adult stage.

Expression of zipD1847K.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW rescues the embryonic lethality of zip1/zip2 animals. 85 +/- 12% of the rescued animals survive to the pupal stage, but no animals survive to the adult stage.

Expression of zipR1933X.Scer\UAS.T:Avic\GFP under the control of Scer\GAL4sqh.PW rescues the embryonic lethality of zip1/zip2 animals. 85 +/- 4% of the rescued animals survive to the pupal stage, but no animals survive to the adult stage.

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Mutant
Wild-type
Stocks (1)
Notes on Origin
Discoverer
External Crossreferences and Linkouts ( 0 )
Synonyms and Secondary IDs (6)
References (36)