Adult stem cells often divide asymmetrically to produce one self-renewed stem cell and one differentiating cell, thus maintaining both populations. The asymmetric outcome of stem cell divisions can be specified by an oriented spindle and local self-renewal signals from the stem cell niche. Here we show that developmentally programmed asymmetric behavior and inheritance of mother and daughter centrosomes underlies the stereotyped spindle orientation and asymmetric outcome of stem cell divisions in the Drosophila male germ line. The mother centrosome remains anchored near the niche while the daughter centrosome migrates to the opposite side of the cell before spindle formation.Adult stem cells maintain populations of highly differentiated but short-lived cells throughout the life of the organism. To maintain the critical balance between stem cell and differentiating cell populations, stem cells have a potential to divide asymmetrically, producing one stem and one differentiating cell (1). The asymmetric outcome of stem cell divisions can be specified by regulated spindle orientation, such that the two daughter cells are placed in different microenvironments that either specify stem cell identity (stem cell niche) or allow differentiation (2,3).Drosophila male germline stem cells (GSCs) are maintained through attachment to somatic hub cells, which constitute the stem cell niche. Hub cells secrete the signaling ligand Upd, which activates the Janus kinase-signal transducer and activator of transcription (JAK-STAT) pathway in the neighboring germ cells to specify stem cell identity (4,5). Drosophila male GSCs normally divide asymmetrically, producing one stem cell, which remains attached to the hub, and one gonialblast, which initiates differentiation. This stereotyped asymmetric outcome is controlled by the orientation of the mitotic spindle in GSCs: The spindle lies perpendicular to the hub so that one daughter cell inherits the attachment to the hub, whereas the other is displaced away (6).

No abstract

Summary Basement membranes (BM) are specialized extracellular matrices that are essential for epithelial structure and morphogenesis. However, little is known about how BM proteins are delivered to the basal cell surface, or how this process is regulated during development. Here, we identify a mechanism for polarized BM secretion in the Drosophila follicle cells. BM proteins are synthesized in a basal ER compartment from localized mRNAs, and are then exported through Tango1-positive ER exit sites to basal Golgi clusters. Next, Crag targets Rab10 to structures in the basal cytoplasm where it restricts protein delivery to the basal surface. These events occur during egg chamber elongation, a morphogenetic process that depends on follicle cell planar polarity and BM remodeling. Significantly, Tango1 and Rab10 are also planar polarized at the basal epithelial surface. We propose that the spatial control of BM production along two tissue axes promotes exocytic efficiency, BM remodeling and organ morphogenesis.

In Drosophila melanogaster the primordial germ cells are normally formed at the posterior tip of the egg during the preblastoderm stage. In order to determine whether the posterior polar plasm is capable of inducing the formation of primordial germ cells in another region of the embryo, portions of this cytoplasm were transferred from wild-type embryos of the early cleavage stage to the anterior tip of mwh e embryos of the same age. At various times after the injection (15-200 min), embryos were fixed for histological analysis. Alternating thick and thin sections were examined for the presence of experimentally induced pole cells. In more than half of the embryos analyzed in this way, one to six cells were found containing the polar granules as well as round nuclear structures, both of which are characteristic of normal pole cells and are not present in blastoderm cells. In order to determine whether these "pole cells" function normally, i.e., develop further into germ cells, the cells induced at the anterior tip of mwh e blastoderm embryos were introduced into the posterior region of y w sns hosts of the same age. The flies resulting from these embryos were mated toy w sn3 partners. In addition to the expected y w sn3 progeny, wild-type flies heterozygous for mwh e and, therefore, descended from the experimentally induced pole cells were found in 4% of the crosses. Such flies did not appear in the control experiments after transfer of normal anterior cells from noninjected blastoderm embryos.These results demonstrate that the posterior polar plasm can be transferred to the anterior tip of the embryo and that in this presuniptive somatic region it still retains its capacity to determine the formation of the primordial germ cells.