Steroidogenesis and apoptosis in the mammalian ovary

Ovarian cell death is an essential process for the homeostasis of ovarian function in human and other mammalian species. It ensures the selection of the dominant follicle and the demise of excess follicles. In turn, this process minimizes the possibility of multiple embryo development during pregnancy and assures the development of few, but healthy embryos. Degeneration of the old corpora lutea in each estrous/menstrual cycle by programmed cell death is essential to maintain the normal cyclicity of ovarian steroidogenesis. Although there are multiple pathways that can determine cell death or survival, crosstalk among endocrine, paracrine and autocrine factors, as well as among protooncogenes, tumor suppressor genes, survival genes and death genes, plays an important role in determining the fate of ovarian somatic and germ cells. The establishment of immortalized rat and human steroidogenic granulosa cell lines and the investigation of pure populations of primary granulosa cells allows systematic studies of the mechanisms that control steroidogenesis and apoptosis in granulosa cells. We have discovered that during initial stages of granulosa cell apoptosis progesterone production does not decrease. In contrast, we found that it is elevated up to 24h following the onset of the apoptotic stimuli exerted by starvation, cAMP, p53 or TNF-alpha stimulation, before total cell collapse. These observations raise the possibility for an alternative unique apoptotic pathway, one not involving mitochondrial Cyt C release associated with the destruction of mitochondrial structure and steroidogenic function. Using mRNA from apoptotic cells and affymetrix DNA microarray technology we discovered that granzyme B, a protease that normally resides in T cytotoxic lymphocytes and natural killer cells of the immune system is expressed and activated in granulosa cells. Thus, the apoptotic signals could bypass mitochondrial signals for apoptosis, which can preserve their steroidogenic activity until complete cell destruction. This unique apoptotic pathway assures cyclicity of estradiol and progesterone release in the estrous/menstruous cycle even during the initial stages of apoptosis.     

The mammalian gonocytes fate: oogenesis or spermatogenesis

Germ line cells become gonocytes after the completion of migration and colonization of gonadal anlages. After the contact with the somatic cells of genital ridges the reprogramming of gonocytes takes place. Upon entry the embryonic gonads germ cells undergo the most complete demethylation during germ line development, their chromatin tends to have an open conformation for a short period. This event promotes the susceptibility to meiosis-inducing factors signaling. The choice of the further path of gonocytes development just after the gonadal sex differentiation, mitotic arrest and meiotic entry are discussed. Analisis of the action mechanisms of meiosis-inducing and meiosis-preventing factors, especially retinoic acid and enzymes of its degradation and synthesis, was performed.     

A test of the production line hypothesis of mammalian oogenesis

Summary Germ cells in female mammals become committed to meiosis and enter its prophase sequentially in fetal life and, according to the Production Line Hypothesis, the oocytes thus generated are released after puberty as mature ova in the same sequence as that of meiotic entry in fetu. This hypothesis in its original and complete form has a subordinate proposition that concerns chiasma (Xma) frequency; it postulates that Xma number would decrease with fetal age. Consequently, univalents would increase, leading to errors of chromosome disjunction at the first meiotic division (MI), and thus to maternal age-dependent numerical chromosome anomalies. By using an in vitro/in vivo approach, we radioactively labelled the DNA of germ cells at premeiotic synthesis as they sequentially entered meiosis, while the fetal ovaries were in culture. At the end of this in vitro phase, pachytene/diplotene (P/D) stages were studied to determine their labelled fraction. The ovaries were then transplanted to spayed females and, after the in vivo phase, mature ova were harvested and the proportion of labelled first and second meiotic metaphases (MI/MII) determined. By marking the germ cells with label while in vitro during periods equivalent to early and late gestation, and by comparing the observed proportions of labelled MI/MII with those of oocytes labelled at P/D, we concluded that, in the mouse, ova do not mature at random for release, but are formed according to a production line system in which the time of release after puberty is related to the time of entry into meiosis in fetu.     

Structure of the ovary and mode of oogenesis in a freshwater crab Potamon dehaani

Structure of the adult ovary and oogenetic mode were examined in the freshwater crab Potamon dehaani. An H‐shaped ovary consisting of a pair of long ovarian sacs connected by a narrow bridge tube is located in the cephalothorax on the dorsal side of the stomach. A short oviduct with a seminal receptacle is connected with the posterior end of each ovarian sac, and a genital pore opens on the sternum of the sixth thoracic segment. The ovarian wall consists of a layer of ovarian epithelium that infolds to form a number of oogenetic pouches of various sizes. These are present mainly in the anterior regions of the ovarian sacs, are scarce in the posterior regions of the ovarian sacs, and are absent from the bridge tube. Each oogenetic pouch contains an egg or a relative large oocyte in its lumen. Germaria containing oogonia, very early previtellogenic oocytes, and somatic interstitial cells are located in the ovarian epithelium near the necks of the oogenetic pouches in the anterior regions of the ovarian sacs and are randomly scattered throughout the ovarian epithelium in the posterior regions of the ovarian sacs. In cross section, the germaria appear to be concentrated into a central germarial cluster in the ovarian sac. In the posterior regions of the ovarian sacs, however, the germaria are randomly scattered throughout the ovarian epithelium. An early previtellogenic oocyte leaves its germarium and raises the ovarian epithelium infolds to form a new oogenetic pouch in which it grows to maturity. Mature eggs are ovulated from the oogenetic pouches into the ovarian lumen, transferred from the ovarian lumen into the oviducts, fertilized there by sperm stored in the seminal receptacles, and then oviposited through the genital pores. The female reproductive system is surrounded wholly and tightly by a thin, cellular, membranous sheath, which has often been mistaken as the ovarian epithelium in some decapod crustaceans. J. Morphol. 239:107–114, 1999. © 1999 Wiley‐Liss, Inc.     

不同食料对野蚕黑卵蜂卵巢发育和卵子发生的影响

以20%浓度蜂蜜、蔗糖、葡萄糖、果糖、甘露糖为食料,研究了它们对野蚕黑卵卵巢发育和卵子发生的影响。结果表明,与对照相比(蒸馏水),蜂蜜、蔗糖、葡萄糖和果糖能促进野蚕黑卵蜂雌蜂卵巢发育和卵子发生,可使其卵巢管中较高的成熟卵量维持较长时间,即可延缓该蜂的卵子重吸收。甘露糖对野蚕黑卵蜂的卵子形成有一定的促进作用,但其所起的作用显然不如蜂蜜及其它3种糖类,并且也不能延缓该蜂的卵子重吸收。     

Interactions between somatic cells and germ cells throughout mammalian oogenesis

Oocytes and their companion somatic cells maintain a close association throughout oogenesis and this association is essential for normal oocyte and follicular development. This review summarizes current concepts of the role of the somatic cells in the regulation of mammalian oocyte growth, the maintenance of meiotic arrest, the induction of oocyte maturation, and the acquisition of full embryonic developmental competence during oocyte maturation in vitro. Gap junctions appear to mediate these regulatory processes. The regulatory interaction of oocytes and somatic cells, however, is not unidirectional; the oocyte participates in the proliferation, development, and function of the follicular somatic cells. The oocyte secretes factors that enable the cumulus cells to synthesize hyaluronic acid and undergo cumulus expansion in response to hormonal stimulation. In addition, the oocyte produces factors that promote the proliferation of granulosa cells. These interactions in vitro do not appear to require the mediation of gap junctions. The oocyte also promotes the differentiation of granulosa cells into functional cumulus cells, but this function of the oocyte appears to require the continued presence and close association of the oocyte and granulosa cells. Therefore, oocytes and follicular somatic cells are interdependent for development and function.     

The ovary structure and oogenesis in the basal crustaceans and hexapods. Possible phylogenetic significance

Recent large-scale phylogenetic analyses of exclusively molecular or combined molecular and morphological characters support a close relationship between Crustacea and Hexapoda. The growing consensus on this phylogenetic link is reflected in uniting both taxa under the name Pancrustacea or Tetraconata. Several recent molecular phylogenies have also indicated that the monophyletic hexapods should be nested within paraphyletic crustaceans. However, it is still contentious exactly which crustacean taxon is the sister group to Hexapoda. Among the favored candidates are Branchiopoda, Malacostraca, Remipedia and Xenocarida (Remipedia + Cephalocarida). In this context, we review morphological and ultrastructural features of the ovary architecture and oogenesis in these crustacean groups in search of traits potentially suitable for phylogenetic considerations. We have identified a suite of morphological characters which may prove useful in further comparative studies.     

Continuities between mitochondria and endoplasmic reticulum in the mammalian ovary

Summary An electron microscope study of developing mouse oocytes has revealed a close morphological relationship between mitochondria and endoplasmic reticulum. In many instances, it was noted that the outer mitochondrial membrane was continuous with the reticular membranes. These cytoplasmic membranes are smooth or studded with ribosomes. These continuities establish an open channel between the endoplasmic reticulum and mitochondria. Similar connections are also found in isolated preparations of mitochondria from the adult guinea pig ovary. The functional significance of these observations are discussed in relation to biochemical studies which demonstrate a transfer of protein from endoplasmic reticulum to mitochondria.     

Ultrastructure of the ovary and oogenesis in the polychaete Capitella jonesi (Hartman, 1959)

Ultrastructural features of the ovary and oogenesis in the polychaete Capitella jonesi (Hartman, '59) have been described. The ovaries are paired, sac-like follicles suspended by mesenteries in the ventral coelom throughout the midbody region of the mature worm. Oogenesis is unsynchronized and occurs entirely within the ovary, where developing gametogenic stages are segregated spatially within a germinal and a growth zone. Multiplication of oogonia and differentiation of oocytes into the late stages of vitellogenesis occur in the germinal region of the ovary, whereas late-stage vitellogenic oocytes and mature eggs are located in a growth zone. Follicle cells envelop the oocytes in the germinal zone of the ovary and undergo hypertrophy and ultrastructural changes that correlate with the onset of vitellogenesis. These changes include the development of extensive arrays of rough ER and numerous Golgi complexes, formation of microvilli along the surface of the ovary, and the initiation of extensive endocytotic activity. Oocytes undergo similar, concomitant changes such as the differentiation of surface microvilli, the formation of abundant endocytotic pits and vesicles along the oolemma, and the appearance of numerous Golgi complexes, cisternae of rough ER, and yolk bodies. Yolk synthesis appears to occur by both autosynthetic and heterosynthetic processes involving the conjoined efforts of the Golgi complex and rough ER of the oocyte and the probable addition of extraovarian (heterosynthetic) yolk precursors. Evidence is presented that implicates the follicle cells in the synthesis of yolk precursors for transport to the oocytes. At ovulation, mature oocytes are released from the overy after the overlying follicle cells apparently withdraw. Bundles of microfilaments within the follicle cells may play a role in this withdrawal process.