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Development 129, 4661-4675 (2002)
Copyright © 2002 The Company of Biologists Limited

intersex, a gene required for female sexual development in Drosophila, is expressed in both sexes and functions together with doublesex to regulate terminal differentiation

Carrie M. Garrett-Engele1,2,*,{ddagger}, Mark L. Siegal1,{ddagger},§, Devanand S. Manoli1, Byron C. Williams3, Hao Li1,{dagger} and Bruce S. Baker1

1 Department of Biological Sciences, Stanford University, Stanford, CA 94305-5020, USA
2 Department of Developmental Biology, Stanford University School of Medicine, Stanford, CA 94305, USA
3 Department of Molecular Biology and Genetics, Biotechnology Building, Cornell University, Ithaca, NY 14853-2703, USA
* Present address: Howard Hughes Medical Institute and Division of Basic Sciences, Fred Hutchinson Cancer Research Center, Seattle, WA 98109, USA
{dagger} Present address: Department of Functional Genomics, Novartis Pharmaceuticals, 556 Morris Avenue, Summit, NJ 07901, USA
{ddagger} These authors contributed equally to this work

§Author for correspondence (e-mail: mlsiegal{at}stanford.edu)

Accepted 10 July 2002

Previous genetic studies indicated intersex (ix) functions only in females and that it acts near the end of the sex determination hierarchy to control somatic sexual differentiation in Drosophila melanogaster. We have cloned ix and characterized its function genetically, molecularly and biochemically. The ix pre-mRNA is not spliced, and ix mRNA is produced in both sexes. The ix gene encodes a 188 amino acid protein, which has a sequence similar to mammalian proteins thought to function as transcriptional activators, and a Caenorhabditis elegans protein that is thought to function as a transcription factor. Bringing together the facts that (1) the ix phenotype is female-specific and (2) functions at the end of the sex determination hierarchy, yet (3) is expressed sex non-specifically and appears likely to encode a transcription factor with no known DNA-binding domain, leads to the inference that ix may require the female-specific protein product of the doublesex (dsx) gene in order to function. Consistent with this inference, we find that for all sexually dimorphic cuticular structures examined, ix and dsx are dependent on each other to promote female differentiation. This dependent relationship also holds for the only known direct target of dsx, the Yolk protein (Yp) genes. Using yeast 2-hybrid assay, immunoprecipitation of recombinant tagged IX and DSX proteins from Drosophila S2 cell extracts, and gel shifts with the tagged IX and DSXF proteins, we demonstrate that IX interacts with DSXF, but not DSXM. Taken together, the above findings strongly suggest that IX and DSXF function in a complex, in which IX acts as a transcriptional co-factor for the DNA-binding DSXF.

Key words: Drosophila, doublesex, hermaphrodite, intersex, Sex determination






© The Company of Biologists Ltd 2002