# The tilt ( tt ) mutation of Drosophila melanogaster maps to the cis-regulatory region of the Iroquois Complex (Iro-C) located in the sosondowah ( sowah) gene

**Authors:** Samuel Gruber, Arno Houtman, Hailey Reisert, Mina Amini, Caroline Fiore, Paula Gonzalez, Veronica Han, Aeva Jazic, Mie Kusupholnand, Max Miller, Jiung Nam, Ziqin Wang, Yang Yu, Peter Dong, Allen S. W. Oak, Arun Sharma, Eric P Spana

PMC · DOI: 10.17912/micropub.biology.001704 · microPublication Biology · 2025-07-16

## TL;DR

This paper identifies the genetic location of the tilt mutation in fruit flies and links it to regulatory elements of the Iroquois Complex genes.

## Contribution

The study maps the tilt mutation to the cis-regulatory region of the Iroquois Complex within the sosondowah gene.

## Key findings

- The tilt mutation is located in the sosondowah gene, which regulates the Iroquois Complex genes araucan and caupolican.
- A gtwin retrotransposon with su(Hw) insulator binding sites is associated with the tilt mutation.
- Mutations in su(Hw) suppress the tilt phenotype, suggesting a functional link.

## Abstract

The
tilt
(

tt

) mutation first described by Morgan and Bridges in 1915 has adult visible phenotypes in wing posture and vein formation. We have mapped

tt

to a genomic region within the
sosondowah
(

sowah

) gene that houses the cis-regulatory elements that control expression of the Iroquois Complex genes
araucan
(

ara

) and

caup
olican

(

caup

) in the wing hinge and wing veins. Sequence analysis of the

tt
1

allele has identified a gtwin retrotransposon containing su(Hw) insulator binding sites within this region of

sowah

. We find that mutations in
su(Hw)
suppress the
tilt
phenotype, providing a potential mechanism for the insertion to affect the function of the Iroquois Complex.

## Linked entities

- **Genes:** tt (tilt) [NCBI Gene 246931], sowah (sosondowah) [NCBI Gene 39438], ABCC6 (ATP binding cassette subfamily C member 6) [NCBI Gene 368], caup (caupolican) [NCBI Gene 39440], su(Hw) (suppressor of Hairy wing) [NCBI Gene 41740]
- **Species:** Drosophila melanogaster (taxon 7227)

## Full-text entities

- **Genes:** Egfr (Epidermal growth factor receptor) [NCBI Gene 37455] {aka C-erb, CG10079, D-EGFR, D-Egf, DEGFR, DER}, sowah (sosondowah) [NCBI Gene 39438] {aka BcDNA:GM09162, CG10632, Dmel\CG10632}, N (Notch) [NCBI Gene 31293] {aka 1.1, 16-178, 16-55, Ax, CG3936, CT13012}, hh (hedgehog) [NCBI Gene 42737] {aka CG4637, Dmel\CG4637, Dmhh, Hg, Mir, Mrt}, lncRNA:CR43612 (long non-coding RNA:CR43612) [NCBI Gene 12798224] {aka CR43612, Dmel\CR43612}, ac (achaete) [NCBI Gene 30981] {aka 990 E5 F1, AS-C, AS-C T5, AS-C T5ac, ASC, Achaete}, caup (caupolican) [NCBI Gene 39440] {aka CG10605, Dmel\CG10605, IRO-C, Iro, Iro-C, IroC}, ara (araucan) [NCBI Gene 39439] {aka CG10571, Dmel\CG10571, IRO-C, Iro, Iro-C, IroC}, RpS4 (Ribosomal protein S4) [NCBI Gene 39484] {aka CG11276, Dmel\CG11276, Rp S4, S4e, SX4, anon-EST:Posey270}, dpp (decapentaplegic) [NCBI Gene 33432] {aka BMP, Bmp, CG9885, DPP-C, Dm-DPP, DmDPP}, AdamTS-B (ADAM metallopeptidase with thrombospondin type 1 motif B) [NCBI Gene 31490] {aka CG4096, CT13580, Dmel\CG4096}
- **Diseases:** wing posture defect (MESH:D054972)
- **Chemicals:** LL05518 (-)
- **Species:** Drosophila melanogaster (fruit fly, species) [taxon 7227]
- **Mutations:** A 400X, A 1000X, A 200X, A 100X

## Full text

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## Figures

1 figure with captions in the complete paper: https://tomesphere.com/paper/PMC12311624/full.md

## References

25 references — full list in the complete paper: https://tomesphere.com/paper/PMC12311624/full.md

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Source: https://tomesphere.com/paper/PMC12311624