Spermatogenesis最新文献

Meiosis entails sorting and separating both homologous and sister chromatids. The mechanisms for connecting sister chromatids and homologs during meiosis are highly conserved and include specialized forms of the cohesin complex and a tightly regulated homolog synapsis/recombination pathway designed to yield regular crossovers between homologous chromatids. Drosophila male meiosis is of special interest because it dispenses with large segments of the standard meiotic script, particularly recombination, synapsis and the associated structures. Instead, Drosophila relies on a unique protein complex composed of at least two novel proteins, SNM and MNM, to provide stable connections between homologs during meiosis I. Sister chromatid cohesion in Drosophila is mediated by cohesins, ring-shaped complexes that entrap sister chromatids. However, unlike other eukaryotes Drosophila does not rely on the highly conserved Rec8 cohesin in meiosis, but instead utilizes two novel cohesion proteins, ORD and SOLO, which interact with the SMC1/3 cohesin components in providing meiotic cohesion.

The ability of stem cells to divide asymmetrically to produce both self-renewing and differentiating daughter cells sustains many adult tissues, but germline stem cells (GSCs) are unique among stem cells as they perpetuate the genome of the species. The cellular and molecular mechanisms regulating most mammalian stem cells in their endogenous local microenvironments, or niches, are quite challenging to study. However, studies of stem cell niches such as those found in the Drosophila gonads have proven very useful. In these tissues, GSCs are housed in a readily identifiable niche, and the ability to genetically manipulate these cells and their neighbors has uncovered several fundamental mechanisms that are relevant to stem cells more generally. Here, we summarize recent work on the regulation of GSCs in the Drosophila testis niche by intercellular signals, and on the intracellular mechanisms that cooperate with these signals to ensure the survival of the germline. This review focuses on GSCs within the adult Drosophila testis; somatic stem cells in this tissue are reviewed by Zoller and Schulz in this issue.(1) For a review of the testis niche as a whole, see de Cuevas and Matunis,(2) and for more comprehensive reviews of the Drosophila testis, refer to Fuller(3) and Davies and Fuller.(4).

Among most animals with internal fertilization, females store sperm in specific regions of their reproductive tract for later use. Sperm storage enables prolonged fertility, physical and temporal separation of mating from fertilization and, when females mate with multiple males, opportunities for differential use of the various males' sperm. Thus, stored sperm move within the female reproductive tract as well as to several potential fates - fertilization, displacement by other sperm or ejection by the female. Drosophila melanogaster is a leading model system for elucidating both the mechanisms and evolutionary consequences of female sperm storage and differential male fertilization success. The prominence of Drosophila is due, in part, to the ability to examine processes influencing sperm movement and fate at several biological levels, from molecules to organ systems. In this review, we describe male and female factors, as well as their interactions, involved in female sperm storage and differential male fertilization success.

The creation of sexual dimorphism in the gonads is essential for producing the male and female gametes required for sexual reproduction. Sexual development of the gonads involves both somatic cells and germ cells, which often undergo sex determination by different mechanisms. While many sex-specific characteristics evolve rapidly and are very different between animal species, gonad function and the formation of sperm and eggs appear more similar and may be more conserved. Consistent with this, the doublesex/mab3 Related Transcription factors (DMRTs) are important for gonad sexual dimorphism in a wide range of animals, including flies, worms and mammals. Here we explore how sexual dimorphism is regulated in the Drosophila gonad, focusing on recent discoveries relating to testis development. We will discuss how sex determination in both the germline and the soma are utilized to create a testis, including the role of the key somatic sex determination factor doublesex.

In all animals, germline cells differentiate in intimate contact with somatic cells and interactions between germline and soma are particularly important for germline development and function. In the male gonad of Drosophila melanogaster, the developing germline cells are enclosed by somatic cyst cells. The cyst cells are derived from cyst stem cells (CySCs) of somatic origin and codifferentiate with the germline cells. The fast generation cycle and the genetic tractability of Drosophila has made the Drosophila testis an excellent model for studying both the roles of somatic cells in guiding germline development and the interdependence of two separate stem cell lineages. This review focuses on our current understanding of CySC specification, CySC self-renewing divisions, cyst cell differentiation, and soma-germline interactions. Many of the mechanisms guiding these processes in Drosophila testes are similarly essential for the development and function of tissues in other organisms, most importantly for gametogenesis in mammals.

Drosophila spermatogenesis has become a paradigmatic system for the study of mechanisms that regulate adult stem cell maintenance, proliferation and differentiation. The dramatic cellular differentiation process from germline stem cell (GSC) to mature sperm is accompanied by dynamic changes in gene expression, which are regulated at transcriptional, post-transcriptional (including translational) and post-translational levels. Post-transcriptional regulation has been proposed as a unique feature of germ cells. However, recent studies have provided new insights into transcriptional regulation during Drosophila spermatogenesis. Both signaling pathways and epigenetic mechanisms act to orchestrate the transcriptional regulation of distinct genes at different germ cell differentiation stages. Many of the regulatory pathways that control male gamete differentiation in Drosophila are conserved in mammals. Therefore, studies using Drosophila spermatogenesis will provide insight into the molecular mechanisms that regulate mammalian germ cell differentiation pathways.

Cytokinesis separates the cytoplasm and the duplicated genome into two daughter cells at the end of cell division. This process must be finely regulated to maintain ploidy and prevent tumor formation. Drosophila male meiosis provides an excellent cell system for investigating cytokinesis. Mutants affecting this process can be easily identified and spermatocytes are large cells particularly suitable for cytological analysis of cytokinetic structures. Over the past decade, the powerful tools of Drosophila genetics and the unique characteristics of this cell system have led researchers to identify molecular players of the cell cleavage machinery and to address important open questions. Although spermatocyte cytokinesis is incomplete, resulting in formation of stable intercellular bridges, the molecular mechanisms are largely conserved in somatic cells. Thus, studies of Drosophila male meiosis will shed new light on the complex cell circuits regulating furrow ingression and substantially further our knowledge of cancer and other human diseases.

Despite their conserved functional role in sexually reproducing organisms, spermatozoa are a diverse and rapidly evolving cell type. This phenomenon is largely attributed to sexual selection in polygamous species where sperm from multiple males compete to fertilize a limited number of oocytes. Drosophila have proven to be a particularly informative model system for the study of spermatogenesis and in this review we discuss how the characterization of the Drosophila melanogaster sperm proteome has advanced our understanding of the evolutionary genomics of sperm form and function. We summarize the molecular evolutionary characteristics of sperm genes and highlight recent evidence demonstrating the importance of novel gene creation in the evolution of sperm function and competitive ability. Comparative proteomic evidence is also provided, supporting an overall functional conservation between the Drosophila and mouse sperm proteomes. This analysis reveals a diverse repertoire of proteins functioning in proteolytic pathways, as well as the presence of proteins of the complement and innate immunity systems. We propose that these pathways may have functional relevance to post-mating female immunological responses as well as coevolved interactions with pathways expressed in the female reproductive tract, including those involved in sperm-oocyte recognition and fertilization.

Drosophila melanogaster spermatids undergo dramatic morphological changes as they differentiate from small round cells approximately 12 μm in diameter into highly polarized, 1.8 mm long, motile sperm capable of participating in fertilization. During spermiogenesis, syncytial cysts of 64 haploid spermatids undergo synchronous differentiation. Numerous changes occur at a subcellular level, including remodeling of existing organelles (mitochondria, nuclei), formation of new organelles (flagellar axonemes, acrosomes), polarization of elongating cysts and plasma membrane addition. At the end of spermatid morphogenesis, organelles, mitochondrial DNA and cytoplasmic components not needed in mature sperm are stripped away in a caspase-dependent process called individualization that results in formation of individual sperm. Here, we review the stages of Drosophila spermiogenesis and examine our current understanding of the cellular and molecular mechanisms involved in shaping male germ cell-specific organelles and forming mature, fertile sperm.