The tut genomic region has been introduced back into the attP site present in tut3. As a result of this manipulation, a 91bp fragment (attR site and vector sequence) is present upstream of the tut start codon, and a 68bp fragment (loxP and vector sequence) is present downstream of the tut stop codon. The presence of these fragments affects tut function.
tut4 homozygotes display relatively mild tumorigenic phenotype (due to spermatogonial overgrowth) in adult testes with large tumors (more than 1/6 of overall testis length) being very rare.
Homozygous testes contain mostly spermatocytes, mixed with a few spermatogonial tumours.
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Pcf11UAS.C.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Pcf11UAS.B/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Pcf11UAS.D/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Wdr33UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Scer\GAL4VP16.bam/SymUAS.A
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Scer\GAL4VP16.bam/IntS11UAS.A.Tag:HA,Tag:FLAG
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf73UAS.A
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Scer\GAL4VP16.bam/IntS9UAS.A.Tag:HA,Tag:FLAG
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Cpsf5UAS.B.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by CG4612UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by cbcUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by CstF50UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf100UAS.A
tut4 has neoplasia | adult stage | male phenotype, enhanceable by CstF64UAS.A/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, enhanceable by Not1MI07631/Not1[+]
tut4 has neoplasia | adult stage | male phenotype, enhanceable by pAbp[+]/pAbpk10109
tut4 has neoplasia | adult stage | male phenotype, enhanceable by pAbpEY11561/pAbpEY11561
tut4 has increased cell number | recessive phenotype, enhanceable by bam[+]/bamΔ86
tut4 has increased cell number | recessive phenotype, enhanceable by bgcn[+]/bgcnQS2
tut4 has increased cell number | recessive phenotype, enhanceable by bgcn[+]/bgcn20093
tut4 has neoplasia | adult stage | male phenotype, non-enhanceable by Scer\GAL4VP16.bam/Pabp2UAS.A.Tag:HA,Tag:FLAG
tut4 has neoplasia | adult stage | male phenotype, non-enhanceable by Scer\GAL4VP16.bam/su(f)UAS.E.Tag:HA,Tag:FLAG
tut4 has neoplasia | adult stage | male phenotype, non-enhanceable by hrgUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has neoplasia | adult stage | male phenotype, non-enhanceable by Not3KG10496/Not3KG10496
tut4 has neoplasia | adult stage | male phenotype, non-enhanceable by PAN2f00130/PAN2[+]
tut4 is an enhancer of increased cell number | dominant phenotype of bamΔ86
tut4 has testis phenotype, enhanceable by Pcf11UAS.C.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by Pcf11UAS.C.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Pcf11UAS.B/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by Pcf11UAS.B/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Pcf11UAS.D/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by Pcf11UAS.D/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Wdr33UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by Wdr33UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Scer\GAL4VP16.bam/SymUAS.A
tut4 has spermatogonium phenotype, enhanceable by Scer\GAL4VP16.bam/SymUAS.A
tut4 has testis phenotype, enhanceable by Scer\GAL4VP16.bam/IntS11UAS.A.Tag:HA,Tag:FLAG
tut4 has spermatogonium phenotype, enhanceable by Scer\GAL4VP16.bam/IntS11UAS.A.Tag:HA,Tag:FLAG
tut4 has testis phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf73UAS.A
tut4 has spermatogonium phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf73UAS.A
tut4 has testis phenotype, enhanceable by Scer\GAL4VP16.bam/IntS9UAS.A.Tag:HA,Tag:FLAG
tut4 has spermatogonium phenotype, enhanceable by Scer\GAL4VP16.bam/IntS9UAS.A.Tag:HA,Tag:FLAG
tut4 has testis phenotype, enhanceable by Cpsf5UAS.B.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by Cpsf5UAS.B.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by CG4612UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by CG4612UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by cbcUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by cbcUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by CstF50UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by CstF50UAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf100UAS.A
tut4 has spermatogonium phenotype, enhanceable by Scer\GAL4VP16.bam/Cpsf100UAS.A
tut4 has testis phenotype, enhanceable by CstF64UAS.A/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, enhanceable by CstF64UAS.A/Scer\GAL4VP16.bam
tut4 has testis phenotype, enhanceable by Not1MI07631/Not1[+]
tut4 has spermatogonium phenotype, enhanceable by Not1MI07631/Not1[+]
tut4 has testis phenotype, enhanceable by pAbp[+]/pAbpk10109
tut4 has spermatogonium phenotype, enhanceable by pAbp[+]/pAbpk10109
tut4 has testis phenotype, enhanceable by pAbpEY11561/pAbpEY11561
tut4 has spermatogonium phenotype, enhanceable by pAbpEY11561/pAbpEY11561
tut4 has spermatogonium phenotype, enhanceable by bam[+]/bamΔ86
tut4 has spermatogonium phenotype, enhanceable by bgcn[+]/bgcnQS2
tut4 has spermatogonium phenotype, enhanceable by bgcn[+]/bgcn20093
tut4 has testis phenotype, non-enhanceable by Scer\GAL4VP16.bam/Pabp2UAS.A.Tag:HA,Tag:FLAG
tut4 has spermatogonium phenotype, non-enhanceable by Scer\GAL4VP16.bam/Pabp2UAS.A.Tag:HA,Tag:FLAG
tut4 has testis phenotype, non-enhanceable by Scer\GAL4VP16.bam/su(f)UAS.E.Tag:HA,Tag:FLAG
tut4 has spermatogonium phenotype, non-enhanceable by Scer\GAL4VP16.bam/su(f)UAS.E.Tag:HA,Tag:FLAG
tut4 has testis phenotype, non-enhanceable by hrgUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has spermatogonium phenotype, non-enhanceable by hrgUAS.A.Tag:HA,Tag:FLAG/Scer\GAL4VP16.bam
tut4 has testis phenotype, non-enhanceable by Not3KG10496/Not3KG10496
tut4 has spermatogonium phenotype, non-enhanceable by Not3KG10496/Not3KG10496
tut4 has testis phenotype, non-enhanceable by PAN2f00130/PAN2[+]
tut4 has spermatogonium phenotype, non-enhanceable by PAN2f00130/PAN2[+]
The moderate spermatogonial tumor phenotype (with large tumors, >1/6 of overall testis length, being very rare) observed in testes of tut4 homozygous mutant males is exacerbated (to a varying degree) by Scer\GAL4bam.T:Hsim\VP16-driven expression of any of the following: Pcf11Scer\UAS.C.T:Ivir\HA1,T:Zzzz\FLAG, Pcf11Scer\UAS.B, Pcf11Scer\UAS.D, Wdr33Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, SymScer\UAS.A, IntS11Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, Cpsf73Scer\UAS.A, IntS9Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, CG3689Scer\UAS.B.T:Ivir\HA1,T:Zzzz\FLAG, CG4612Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, generally weaker enhancement is observed upon expression of cbcScer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, CstF50Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, Cpsf100Scer\UAS.A or (very weak) CstF64Scer\UAS.A and none at all upon expression of Pabp2Scer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG, su(f)Scer\UAS.E.T:Ivir\HA1,T:Zzzz\FLAG and hrgScer\UAS.A.T:Ivir\HA1,T:Zzzz\FLAG in the mutant background (the latter two appear to have a somewhat suppressive effect as no large tumors are observed in these flies).
The severity of the tut4 mutant tumorigenic phenotype is also strongly enhanced by combination with a single copy of either Not1MI07631 or pAbpk10109 or by pAbpEY11561 homozygosity, no enhancement but rather an amelioration of the tumor phenotype is observed upon combination with Not3KG10496 in homozygous or PAN2f00130 in heterozygous state. None of these genetic perturbations cause a significant tumor phenotype if combined with just one copy of tut4.