Doublesex controls specification and maintenance of the gonad stem cell niches in Drosophila.

Sex-specific development of the gonads is a key aspect of sexual dimorphism that is regulated by Doublesex/Mab3-related transcription factors (DMRTs) in diverse animal species. We find that in mutants for Drosophila dsx, important components of the male and female gonad stem cell niches (hubs and terminal filaments/cap cells, respectively) still form. Initially, gonads in all dsx mutants (both XX and XY) initiate the male program of development, but later half of these gonads switch to form female stem cell niche structures. One individual can have both male-type and female-type gonad niches; however, male and female niches are usually not observed in the same gonad, indicating that cells make a 'group decision' about which program to follow. We conclude that dsx does not act in an instructive manner to regulate male versus female niche formation, as these structures form in the absence of dsx function. Instead, dsx acts to 'tip the balance' between the male or female programs, which are then executed independently of dsx We show that bric a brac acts downstream of dsx to control the male versus female niche decision. These results indicate that, in both flies and mammals, the sexual fate of the somatic gonad is remarkably plastic and is controlled by a combination of autonomous and non-autonomous cues.

Targeted disruption of an EH-domain protein endocytic complex, Pan1-End3.

Pan1 is a multi-domain scaffold that enables dynamic interactions with both structural and regulatory components of the endocytic pathway. Pan1 is composed of Eps15 Homology (EH) domains which interact with adaptor proteins, a central region that is responsible for its oligomerization and C-terminal binding sites for Arp2/3, F-actin, and type-I myosin motors. In this study, we have characterized the binding sites between Pan1 and its constitutive binding partner End3, another EH domain containing endocytic protein. The C-terminal End3 Repeats of End3 associate with the N-terminal part of Pan1's central coiled-coil region. These repeats appear to act independently of one another as tandem, redundant binding sites for Pan1. The end3-1 allele was sequenced, and corresponds to a C-terminal truncation lacking the End3 Repeats. Mutations of the End3 Repeats highlight that those residues which are identical between these repeats serve as contact sites for the interaction with Pan1.

Nonautonomous sex determination controls sexually dimorphic development of the Drosophila gonad.

Sex determination in Drosophila is commonly thought to be a cell-autonomous process, where each cell decides its own sexual fate based on its sex chromosome constitution (XX versus XY). This is in contrast to sex determination in mammals, which largely acts nonautonomously through cell-cell signaling. Here we examine how sexual dimorphism is created in the Drosophila gonad by investigating the formation of the pigment cell precursors, a male-specific cell type in the embryonic gonad. Surprisingly, we find that sex determination in the pigment cell precursors, as well as the male-specific somatic gonadal precursors, is non-cell autonomous. Male-specific expression of Wnt2 within the somatic gonad triggers pigment cell precursor formation from surrounding cells. Our results indicate that nonautonomous sex determination is important for creating sexual dimorphism in the Drosophila gonad, similar to the manner in which sex-specific gonad formation is controlled in mammals.

The creation of sexual dimorphism in the Drosophila soma.

Animals have evolved a fascinating array of mechanisms for conducting sexual reproduction. These include producing the sex-specific gametes, as well as mechanisms for attracting a mate, courting a mate, and getting the gametes together. These processes require that males and females take on dramatically different forms (sexual dimorphism). Here, we will explore the problem of how sex is determined in Drosophila, and pay particular attention to how information about sexual identity is used to instruct males and females to develop differently. Along the way, we will highlight new work that challenges some of the traditional views about sex determination. In Drosophila, it is commonly thought that every cell decides its own sex based on its sex chromosome constitution (XX vs. XY). However, we now know that many cell types undergo nonautonomous sex determination, where they are told what sex to be through signals from surrounding cells, independent of their own chromosomal content. Further, it now appears that not all cells even "know" their sex, since key members of the sex determination pathway are not expressed in all cells. Thus, our understanding of how sex is determined, and how sexual identity is used to create sexual dimorphism, has changed considerably.