FB2026_01 , released March 12, 2026
FB2026_01 , released March 12, 2026
Allele: Dmel\put10460
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General Information
Symbol
Dmel\put10460
Species
D. melanogaster
Name
FlyBase ID
FBal0009622
Feature type
allele
Associated gene
Dmel\put
Associated Insertion(s)
P{PZ}put10460
Carried in Construct
Also Known As
puntP, puntP1, putP1, putP, punt10460
Key Links
Genomic Maps

Allele class

amorphic allele - genetic evidence

Mutagen
Nature of the Allele
Allele class

amorphic allele - genetic evidence

(Ruberte et al., 1995)
Mutagen
P-element activity
(Letsou et al., 1995, Ruberte et al., 1995)
Progenitor genotype
Associated Insertion(s)
P{PZ}put10460
Cytology
Description

P-element insertion within the put locus.

(Letsou et al., 1995)

P-element insertion within the untranslated leader sequence 2bp from the 5' end.

(Letsou et al., 1995)

P-element insertion 399bp upstream of ATG initiator codon.

(Ruberte et al., 1995)
Allele components
Component
Use(s)
Inserted element
P{PZ}
Encoded product / tool
lacZ
Tagged with
Tag:NLS(P)
Mutations Mapped to the Genome
Associated Sequence Data
DDBJ / EMBL / GenBank
DNA sequence
Protein sequence
 
UniProtKB/Swiss-Prot
    UniProtKB/TrEMBL
      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 )
       
      Disease-implicated variant(s)
       
      Phenotypic Data
      Phenotypic Class
      Phenotype Manifest In

      antenna | pharate adult stage

      (Theisen et al., 1996)

      axon terminus | larval stage | somatic clone

      (Kaneko et al., 2024)

      dorsal tracheal branch primordium

      (Ribeiro et al., 2002)

      dorsocentral bristle

      (Phillips et al., 1999)

      embryonic/first instar larval cuticle

      (Perrimon et al., 1996)

      embryonic/first instar larval cuticle | dorsal

      (Perrimon et al., 1996)

      embryonic/first instar larval cuticle | dorsal | germline clone

      (Bartoszewski et al., 2004)

      embryonic/first instar larval cuticle | dorsal | heat sensitive

      (Simin et al., 1998)

      embryonic/first instar larval cuticle | heat sensitive

      (Letsou et al., 1995)

      embryonic/larval crystal cell

      (Milchanowski et al., 2004)

      embryonic/larval plasmatocyte

      (Milchanowski et al., 2004)

      embryonic abdominal segment 1 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 2 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 3 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 4 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 5 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 6 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 7 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 8 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 9 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 10 | heat sensitive

      (Simin et al., 1998)

      embryonic abdominal segment 11 | heat sensitive

      (Simin et al., 1998)

      embryonic dorsal epidermis

      (Riesgo-Escovar and Hafen, 1997)

      embryonic head | heat sensitive

      (Simin et al., 1998, Letsou et al., 1995)

      eye

      (Burke and Basler, 1996)

      eye | pharate adult stage

      (Theisen et al., 1996)

      eye disc | posterior | somatic clone

      (Hazelett et al., 1998)

      eye disc morphogenetic furrow

      (Burke and Basler, 1996)

      eye disc posterior compartment | somatic clone | third instar larval stage

      (Burke and Basler, 1996)

      female germline stem cell

      (Xie and Spradling, 1998)

      female germline stem cell (with put135)

      (Song et al., 2004)

      larval nociceptive neuropil | somatic clone

      (Kaneko et al., 2024)

      leg | dorsal | pharate adult stage

      (Theisen et al., 1996)

      leg | pharate adult stage | ventral

      (Theisen et al., 1996)

      macrochaeta

      (Phillips et al., 1999)

      male germline stem cell | germline clone

      (Kawase et al., 2004)

      male germline stem cell | heat sensitive (with put135)

      (Kawase et al., 2004, Shivdasani and Ingham, 2003)

      photoreceptor neuron | somatic clone

      (Hazelett et al., 1998)

      pigment granule

      (Burke and Basler, 1996)

      primary pigment cell | pupal stage P7

      (Cordero et al., 2007)

      sensory mother cell & dorsal mesothoracic disc

      (Tomoyasu et al., 1998)

      spermatogonium | heat sensitive (with put135)

      (Shivdasani and Ingham, 2003)

      tertiary pigment cell | pupal stage P7

      (Cordero et al., 2007)

      testis | heat sensitive (with put135)

      (Shivdasani and Ingham, 2003)

      wing

      (Phillips et al., 1999, Simin et al., 1998, Burke and Basler, 1996)

      wing vein

      (Simin et al., 1998, Burke and Basler, 1996)

      wing vein | ectopic

      (Letsou et al., 1995)
      Detailed Description
      Statement
      Reference

      put10460 third instar larval C4da neurons do not show any obvious defects in the dendritic arborization, as compared to controls.

      (Hoyer et al., 2018)

      put10460 somatic clones in the pupal retina show mild defects in the arrangement of inter-ommatidial pigment cells.

      (Cordero et al., 2007)

      put10460/+ flies display a wild-type bristle pattern.

      (Gibert et al., 2005)

      Embryos laid from put10460 germ-line clones and fertilised with wild-type sperm are weakly ventralised.

      (Bartoszewski et al., 2004)

      put135/put10460 males raised to adulthood at 18oC and then shifted to 22oC for 7 days have normal numbers of germline stems cells per testis. However, one week at 29oC is sufficient to almost completely eliminate germline stem cells in these animals. No increase in apoptosis occurs of germline stem cells occurs in these testes during culture at 29oC, suggesting that loss is due to differentiation of stem cells. Clones of male put10460 homozygous germline stem cells are still present in 15% of testes one week after clone induction and none are present two weeks after clone induction. This is in contrast to wild-type control clones, which are present in 82% of testes one week after clone induction and 64% two weeks after clone induction.

      (Kawase et al., 2004)

      The average number of crystal cells per embryo is reduced (to 13.5) in homozygous stage 13-14 embryos compared to wild type (36 +/- 2.2 cells on average). The number of plasmatocytes is also significantly reduced compared to wild-type embryos.

      (Milchanowski et al., 2004)

      At eclosion put10460/put135 adult females raised at 18oC have normal numbers of germ-line stem cells, but after 2 days at 29oC 53% of germaria in these flies have one or no germline stem cells.

      (Song et al., 2004)

      put135/put10460 males shifted to the restrictive temperature of 29oC after eclosion have smaller and thinner testes than control males after 7 days at 29oC. The testes also show an apparent reduction in the number of early germ cells, particularly spermatogonia and germ line stem cells.

      (Shivdasani and Ingham, 2003)

      The tracheal systems of put10460 mutant embryos fail to form dorsal branches.

      (Ribeiro et al., 2002)

      Clones of put135/put10460 induced in the pleura cause no mutant phenotype.

      (Kopp et al., 1999)

      put135/put10460 animals shifted from 16 to 28oC 6 days after egg deposition show loss of the dorsocentral macrochaetae, although lateral macrochaetae and the field of microchaetae remain. There is also loss of wing tissue. put135/put10460 animals shifted from 16 to 28oC during larval 7 days after egg deposition produce extra dorsocentral macrochaetae.

      (Phillips et al., 1999)

      Homozygous clones at the posterior margin of the eye disc do not show photoreceptor differentiation.

      (Hazelett et al., 1998)

      At 25oC, 18% of homozygous embryos have a strong dorsal-open phenotype (severe head defects and a large hole in the dorsal cuticle), 43% have an intermediate phenotype (the dorsal cuticle is noticeably reduced, and there is increased curvature in the abdominal segments, resulting in a "tail-up" phenotype) and 39% have a weak phenotype. put10/put10460 adults are fully viable at 25oC, but show defects in wing venation in 28% of cases.

      (Simin et al., 1998)

      put10460/put135 animals raised at the permissive temperature (18oC) and then shifted to the nonpermissive temperature (29oC) at about 48 hours before pupariation lack the dorsocentral sensory mother cells in the thoracic discs. put10460/put135 animals raised at the permissive temperature (18oC) and then shifted to 25oC have one or a few extra sensory mother cells in the thoracic discs.

      (Tomoyasu et al., 1998)

      Clonal analysis in the germarium reveals that mutant stem cells are lost more rapidly than wild type, though there is no effect on the formation of 16 cell cysts or their subsequent development. Stem cell half life is 0.9 weeks (wild type being 4.6 weeks). Stem cell division rate relative to control is 0.36. Cysts contain the normal 16 cells, including and oocyte.

      (Xie and Spradling, 1998)

      Embryos exhibit a partial dorsal closure phenotype.

      (Riesgo-Escovar and Hafen, 1997)

      Large mutant clones covering almost the entire adult eye are generally wild type (with a small amount of wild type tissue at the posterior border); ommatidia are normal, occasional vacuoles between the ommatidia are observed. The head of an adult fly with mutant clones has no eye structures, only head cuticle is present where the eye normally forms. Clones in the imaginal disc anterior to the morphogenetic furrow have no detectable phenotype. Clones posterior to the furrow are also normal demonstrating the furrow can progress normally through and beyond a mutant clone. Within clones traversing the furrow neuronal differentiation is retarded, slowing of the furrow is more marked in the middle of the clone suggesting partial rescue by surrounding wild type tissue. Clones at the posterior margin of the eye discs can cause local overproliferation and block neuronal differentiation. Neuronal differentiation does occur in lateral clones.

      (Burke and Basler, 1996)

      Wing clones induced in mid-to-late third instar larvae are small due to their late induction time but cause visible mutant phenotypes: gaps, splits, vein indentations and additional vein material abutting normal veins.

      (Burke and Basler, 1996)

      Homozygous embryos exhibit a dorsal open phenotype. Germline clones produce eggs with patterning defects: embryos are ventralised.

      (Perrimon et al., 1996)

      put135/put10460 transheterozygous pharate adults heat shocked during late first and early second instar have legs in which dorsal structures are absent and ventral structures are duplicated. On the distal tibia of the second leg there are two apical bristles which are indicators of the ventral-most cell fate in leg discs. Pharate adults exhibit reduced or missing eyes and duplicated antenna.

      (Theisen et al., 1996)

      Dorsal open phenotype and head defects become more severe at the higher temperatures. Heterozygotes with put51 and put88 are lethal at high temperatures, at low temperatures individuals survive and exhibit ectopic wing venation in the region of the posterior cross vein. Escaper females produce ventralised embryos.

      (Letsou et al., 1995)
      External Data
      Interactions
      Show genetic interaction network for Enhancers & Suppressors
      Phenotypic Class
      NOT Suppressor of
      Statement
      Reference

      put[+]/put10460 is a non-suppressor of abnormal neuroanatomy phenotype of spin10403

      (Yuva-Aydemir et al., 2011)
      Other
      Statement
      Reference

      Chc[+]/Chc1, put10460 has visible | dominant | heat sensitive phenotype

      (Gonzalez-Gaitan and Jackle, 1999)

      Chc1, put[+]/put10460 has visible | dominant | heat sensitive phenotype

      (Gonzalez-Gaitan and Jackle, 1999)
      Phenotype Manifest In
      NOT Enhanced by
      Statement
      Reference

      put10460 has embryonic/first instar larval cuticle | dorsal | germline clone phenotype, non-enhanceable by eca2

      (Bartoszewski et al., 2004)
      Enhancer of
      Suppressor of
      Statement
      Reference

      put[+]/put10460 is a suppressor of wing vein phenotype of Ctdnep1G0269

      (Liu et al., 2011)
      NOT Suppressor of
      Statement
      Reference

      put[+]/put10460 is a non-suppressor of eye disc phenotype of spin10403

      (Yuva-Aydemir et al., 2011)

      put[+]/put10460 is a non-suppressor of glial cell phenotype of spin10403

      (Yuva-Aydemir et al., 2011)

      put10460 is a non-suppressor of female germline stem cell | germline clone phenotype of bamΔ86

      (Jin et al., 2008)
      Other
      Additional Comments
      Genetic Interactions
      Statement
      Reference

      The aberrant wing vein phenotype seen in DdG0269/Y animals is suppressed by put10460/+.

      (Liu et al., 2011)

      The glial overmigration phenotype seen in spin10403 eye discs is not suppressed if the animals are also heterozygous for put10460.

      (Yuva-Aydemir et al., 2011)

      bamΔ86 put10460 double mutant germline stem cell (GSC) clones retain a similar ability to bamΔ86 single mutant GSCs to outcompete wild type GSCs in the same niches.

      (Jin et al., 2008)

      Transheterozygous put10460 pad1 flies have ectopic dorso-central bristles, while neither single heterozygote shows this phenotype.

      (Gibert et al., 2005)

      In btlLG19/+, put10460/+ embryos, most primary tracheal branches fail to migrate.

      (Myat et al., 2005)

      No ptcH84, put10460 animals exhibit head capsule defects.

      (Shyamala and Bhat, 2002)

      The addition of put10460 to dallygem/dally06464 flies, leads to enhancement of the anterior dorsocentral bristle phenotype seen in dallygem/dally06464 flies alone: 90% of flies have the phenotype, rather than 24%. The posterior supraalar bristle phenotype is also enhanced by the addition of put10460. All flies have the phenotype, rather than 86%.

      (Fujise et al., 2001)

      At 18oC and 25oC, Chc1/+, put10460/+ double heterozygotes have normal wings, while at 29oC they show a wing reduction and wing vein truncation phenotype, indicative of a problem with A/P pattern formation.

      (Gonzalez-Gaitan and Jackle, 1999)
      Xenogenetic Interactions
      Statement
      Reference
      Complementation and Rescue Data
      Images (0)
      Mutant
      Wild-type
      Stocks (1)
      Notes on Origin
      Discoverer

      A. Spradling.

      Comments
      Comments

      Three classes of put allele can be identified with respect to decreasing severity of wing venation phenotypes with put10 : Class I (put10460 = put62) > Class II (put51 = put88 = put135) > Class III (put10 = put24 = put97).

      (Simin et al., 1998)

      Complements: l(3)0347103471. Complements: l(3)0360803608. Complements: cv-c06951.

      (BDGP Project Members, 1994-1999)

      Mobilization of the P-element results in multiple independent excision lines that are homozygous viable and fertile.

      (Letsou et al., 1995)

      Mobilization of the P-element reverts mutant phenotype to wild type. Mutant phenotype can be rescued to pharate adult by P{hs-punt} without heat shock.

      (Ruberte et al., 1995)

      put135/put10460 transheterozygotes are referred to as "putts" embryos, non-permissive temperature 25oC.

      (Theisen et al., 1996)
      External Crossreferences and Linkouts ( 0 )
      Synonyms and Secondary IDs (13)
      References (44)