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First published online September 30, 2004
doi: 10.1242/10.1242/dev.01373


Development 131, 4895-4905 (2004)
Published by The Company of Biologists 2004


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How different is Venus from Mars? The genetics of germ-line stem cells in Drosophila females and males

Lilach Gilboa and Ruth Lehmann*

Developmental Genetics Program, Skirball Institute and Department of Cell Biology at NYU School of Medicine and Howard Hughes Medical Institute, 540 First Avenue, New York, NY 10016, USA



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Fig. 1. Schematic of the anterior part of a female ovariole and male testis in Drosophila melanogaster. The description of the developmental stages of the germ cells is to the left of each structure and the somatic cells are to the right. Anterior is up. In the female ovariole (A) and male testis (B), germ-line stem cells (GSCs) are located at the anterior tip of the gonad. Upon division, the posterior daughter cell differentiates to a cystoblast (Cb, females) or a gonialblast (males). The differentiation may be gradual, as depicted by increasingly stronger shades of green. The daughter cell divides further and forms a cyst. GSCs and their immediate daughter cells harbor a spherical fusome (here marked in black). This organelle grows and extends into every cell of the cyst (black marking within the cyst). In females, the niche includes terminal filament (TF) and cap cells, which are located most anteriorly. Inner sheath (IS) cells may also be part of the niche and may perform similar functions to those of early somatic cyst cells in males. Somatic stem cells (SSCs) are located `midway' down the germarium and they give rise to the follicle cells that envelope the cyst. In males, SSCs are attached to the hub, and their descendents (cyst cells) encapsulate the gonialblast. Color-coding is used to mark cells that have a similar function in males and females. Shades of red have been used for TF and cap cells as these two populations, although similar, are not identical in their gene expression profiles (Forbes et al., 1996aGo).

 


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Fig. 2. The anterior part of a female ovariole and male testis in Drosophila melanogaster. Anterior is up. (A) An ovariole from a female Drosophila carrying a ß-galactosidase (ß-gal) enhancer trap in the dad gene, stained with antibodies against ß-gal (green), and with the monoclonal antibody 1B1 (red), which stains the membrane cytoskeleton in somatic cells and the fusome within germ cells. The terminal filament and cap cells are at the anterior and are marked with brackets. GSCs are located just posterior to cap cells, and an arrowhead marks a spherical fusome in one GSC that is abutting cap cells. An arrow marks a branched fusome within a cyst in a more posterior location within the germarium. ß-gal staining is strong in GSCs, but can also be observed in their immediate daughters and early cysts. This may be due to the produrence of the ß-gal protein. (B) A testis from a male Drosophila carrying a ß-gal enhancer trap in the escargot gene, stained with anti-ß-gal (green) and 1B1 (red) antibodies. Hub cells express ß-gal most strongly and are circled. Arranged around the hub are GSCs, which also express ß-gal. Staining of ß-gal in gonialblasts and early cysts may be due to the produrence of the protein. Posterior to the hub are the dividing cysts; a fusome in one cyst is marked by an arrow. (B inset) The same genotype as in B, stained with anti-ß-gal (green) and with anti-Fasciclin III (red), which outlines the cells of the hub, showing its compact structure. Scale bar: 20 µm.

 


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Fig. 3. Major participants in GSC maintenance and differentiation in (A) female and (B) male gonads. Germ-line stem cells (GSCs) and their differentiating daughter cells (Cb, cystoblast in females and gonialblast in males) are shown in shades of green. The somatic cells of the niche (cap cells in females and hub cells in males) are colored pink. Other somatic cells [inner sheath (IS) cells in females, and somatic stem cells (SSCs) and cyst cells in males) may affect either GSCs or their differentiating daughter cells, or both, and are shown in blue. Proteins and organelles that act within each cell type are noted. Major and minor signaling pathways emanating from the niche are marked with large and small arrows, respectively.

 





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