Germ Cell Cysts Research Articles

Analysis of LHX8 mutation in premature ovarian failure

The LHX8 (LIM homeobox 8) gene encodes a LIM homeodomain transcriptional regulator that is preferentially expressed in germ cells and critical for mammalian oogenesis. The authors investigated whether nucleotide changes were present in the LHX8 gene of Caucasian women with premature ovarian failure (POF), as compared with control women. When the authors sequenced 95 Caucasian women with POF, they discovered two novel single-nucleotide polymorphisms (SNPs) in intron 3 (c.769+10G>T) and 3' untranslated region (c.1787A>G) of the LHX8 gene. These polymorphisms also were found in controls (n = 94), at frequencies that were not statistically different from those in POF women. Mutations in the LHX8 exons are uncommon in Caucasian women with POF.

Estradiol, Progesterone, and Genistein Inhibit Oocyte Nest Breakdown and Primordial Follicle Assembly in the Neonatal Mouse Ovary in Vitro and in Vivo

In developing mouse ovaries, oocytes develop as clusters of cells called nests or germ cell cysts. Shortly after birth, oocyte nests dissociate and granulosa cells surround individual oocytes forming primordial follicles. At the same time, two thirds of the oocytes die by apoptosis, but the link between oocyte nest breakdown and oocyte death is unclear. Although mechanisms controlling breakdown of nests into individual oocytes and selection of oocytes for survival are currently unknown, steroid hormones may play a role. Treatment of neonatal mice with natural or synthetic estrogens results in abnormal multiple oocyte follicles in adult ovaries. Neonatal genistein treatment inhibits nest breakdown suggesting multiple oocyte follicles are nests that did not break down. Here we investigated the role of estrogen signaling in nest breakdown and oocyte survival. We characterized an ovary organ culture system that recapitulates nest breakdown, reduction in oocyte number, primordial follicle assembly, and follicle growth in vitro. We found that estradiol, progesterone, and genistein inhibit nest breakdown and primordial follicle assembly but have no effect on oocyte number both in organ culture and in vivo. Fetal ovaries, removed from their normal environment of high levels of pregnancy hormones, underwent premature nest breakdown and oocyte loss that was rescued by addition of estradiol or progesterone. Our results implicate hormone signaling in ovarian differentiation with decreased estrogen and progesterone at birth as the primary signal to initiate oocyte nest breakdown and follicle assembly. These findings also provide insight into the mechanism of multiple oocyte follicle formation.

Mouse oocytes within germ cell cysts and primordial follicles contain a Balbiani body

The Balbiani body or mitochondrial cloud is a large distinctive organelle aggregate found in developing oocytes of many species, but its presence in the mouse has been controversial. Using confocal and electron microscopy, we report that a Balbiani body does arise in mouse neonatal germline cysts and oocytes of primordial follicles but disperses as follicles begin to grow. The mouse Balbiani body contains a core of Golgi elements surrounded by mitochondria and associated endoplasmic reticulum. Because of their stage specificity and perinuclear rather than spherical distribution, these clustered Balbiani body mitochondria may have been missed previously. The Balbiani body also contains Trailer hitch, a widely conserved member of a protein complex that associates with endoplasmic reticulum/Golgi-like vesicles and transports specific RNAs during Drosophila oogenesis. Our results provide evidence that mouse oocytes develop using molecular and developmental mechanisms widely conserved throughout the animal kingdom.

Characterization of NOBOX DNA Binding Specificity and Its Regulation of Gdf9 and Pou5f1 Promoters

Nobox (newborn ovary homeobox gene) deficiency disrupts early folliculogenesis and the expression of oocyte-specific genes in mice. Here, we identified several cis-acting sites, TAATTG, TAGTTG, and TAATTA as NOBOX DNA binding elements (NBEs) using a library of randomly generated oligonucleotides by cyclic amplification of sequence target assay and mutation analyses. We show that NOBOX preferentially binds to the NOBOX binding elements with high affinity. In addition, we found that promoter regions of mouse Pou5f1 and Gdf9 contain one (-426) and three NOBOX binding elements (-786, -967, and -1259), respectively. NOBOX binds to these putative NOBOX binding elements with high affinity and augmented transcriptional activity of luciferase reporter driven by mouse Pou5f1 and Gdf9 promoters containing the NOBOX binding elements. In chromatin immunoprecipitation assays, DNA sequences from Pou5f1 and Gdf9 promoters co-precipitated with anti-NOBOX antibody. These results suggest that NOBOX directly regulates the transcription of Pou5f1 and Gdf9 in oocytes during early folliculogenesis.

Tumor Necrosis Factor (TNF) Receptor Type 2 Is an Important Mediator of TNF alpha Function in the Mouse Ovary1

It is believed that a finite pool of primordial follicles is established during embryonic and neonatal life. At birth, the mouse ovary consists of clusters of interconnected oocytes surrounded by pregranulosa cells. Shortly after birth these structures, termed germ cell cysts or nests (GCN), break down to facilitate primordial follicle formation. Tumor necrosis factor alpha (TNF) is a widely expressed protein with myriad functions. TNF is expressed in the ovary and may regulate GCN breakdown in rats. We investigated whether it participates in GCN breakdown and follicle formation in mice by using an in vitro ovary culture system as well as mutant animal models. We found that TNF and both receptors (TNFRSF1A and TNFRSF1B) are expressed in neonatal mouse ovaries and that TNF promotes oocyte death in neonatal ovaries in vitro. However, deletion of either receptor did not affect follicle endowment, suggesting that TNF does not regulate GCN breakdown in vivo. Tnfrsf1b deletion led to an apparent acceleration of follicular growth and a concomitant expansion of the primordial follicle population. This expansion of the primordial follicle population does not appear to be due to decreased primordial follicle atresia, although this cannot be ruled out completely. This study demonstrates that mouse oocytes express both TNF receptors and are sensitive to TNF-induced death. Additionally, TNFRSF1B is demonstrated to be an important mediator of TNF function in the mouse ovary and an important regulator of folliculogenesis.

Irx3 is differentially up-regulated in female gonads during sex determination

Irx3 is a member of the Iroquois homeobox gene family that encodes a protein known for its essential role in spinal cord development. Transcript screening of male and female gonads during the critical period of sex determination (E12–13.5) revealed a sexually dimorphic expression pattern for Irx3 with female gonads exhibiting a sixfold increase in expression over time. Whole mount in situ hybridization confirmed the sexually dimorphic nature of Irx3 expression and immunohistochemical analysis of gonads at E13.5 determined that IRX3 and GATA4 proteins co-localized to somatic cells of XX gonads. The Irx3 signal persisted in germ cell-depleted XX gonads resulting from Busulfan treatment suggesting that its expression was independent of germ cell regulation. Quantitative real-time PCR analysis over an extended time course determined that Irx3 message was low initially and then increased in XX gonads until E13.5, remained elevated until birth, diminished shortly after birth, and remained low in the adult ovary. In contrast, Irx3 message was 50% lower in male compared to female gonads at the initial time point, and continued to decrease over time. Further analysis of adult ovaries suggested that IRX3 expression is not present in any subpopulations of cells of the differentiated ovary. Together, these results suggest that the Irx3 signal is restricted to the somatic cell component of XX gonads and is present at a discreet period of ovarian development that ends abruptly at birth. This timing coincides with the transition of female primordial germ cells from mitotic proliferation to meiotic division, and the organization of germ cell cysts prior to primordial follicle development at birth.

Differentiating germ cells can revert into functional stem cells in Drosophila melanogaster ovaries.

Many tissues including blood, skin, gut and germ cells are continuously maintained by tissue stem cells. Under certain conditions, however, other organs can undergo repair using stem-cell-like progenitors generated by cell de-differentiation. Cell fates have been broadened experimentally, but mechanisms allowing de-differentiation to a stem cell state are poorly known. Germline stem cells begin to differentiate by forming interconnected germ cell cysts (cystocytes), and under certain conditions male mouse cystocytes have been postulated to revert into functional progenitors. Here we report that four- and eight-cell Drosophila germline cystocytes generated either in second instar larval ovaries or in adults over-producing the BMP4-like stem cell signal Decapentaplegic efficiently convert into single stem-like cells. These de-differentiated cells can develop into functional germline stem cells and support normal fertility. Our results show that cystocytes represent a relatively abundant source of regenerative precursors that might help replenish germ cells after depletion by genotoxic chemicals, radiation or normal ageing. More generally, Drosophila cystocytes now provide a system for studying de-differentiation and its potential as a source of functional stem cells.

TBTCl이 구피(Poecilia reticulata)의 행동과 번식에 미치는 영향

TBTCl(0.1, 0.32, 1, 3.2, 10, 25, 32, 50, 75 및 100 <TEX>$\mug/l$</TEX>)을 구피에게 144시간 동안 노출시켜 TBTCl의 구피 생체 내 축적정도 및 구피의 번식과 행동에 미치는 영향에 대해서 조사하였다. 0.1 <TEX>$\mug/l$</TEX> 이상의 모든 TBTCl 실험군에서 <TEX>$\SigmaBTs\; (TBT:\SigmaBTs)$</TEX>가 67% 이상으로 매우 높았고, 고농도 실험군일수록 이 비율이 높아져, 고농도의 TBTCl에 노출될수록 구피의 대사능력이 저해됨을 알 수 있었다. 또한 TBTCl은 구피의 번식력을 약화시키고 행동이상을 유발함이 밝혀졌다. 즉, 0.1<TEX>$\mug/l$</TEX>의 실험군부터 수컷의 구애행동과 수출관 내 정포의 수가 감소되었고, 0.32∼10 <TEX>$\mug/l$</TEX>의 실험군에서는 정소 내 전체 생식세포낭에 대한 정포낭의 수도 점차 감소하여 TBTCl이 정자변태 과정에 영향을 미치는 내분비계 장애물질로서 작용하고 있음이 시사되었다. 더욱이 25 <TEX>$\mug/l$</TEX>이상의 고농도군에서는 단기간 내에 치사율이 증가하고, 생식세포가 괴사되고, 정소의 지지세포와 수출관 내의 상피세포가 상해되고, 유영행동 이상이 급증하고, 섭식이 중단되는 등 급성독성이 관찰되어, TBTCl이 구피의 번식 뿐 아니라 생존을 저해하고 있음이 시사되었다. Sexually matured guppies (Poeiria reticulata) were exposed to TBTCI (0.1, 0.32, 1, 3.2, 10, 25, 32, 50, 75 and 100 <TEX>$\mug/l$</TEX>) for 144 hours to determine the bioaccumulation rate and effects on the reproduction and behavior. The ratio of TBT residues to <TEX>$\SigmaBTs\; (TBT:\SigmaBTs)$</TEX> was 67% or higher in all the guppies exposed to TBTCl, and the higher the level of TBTCl exposed, the higher the ratio of TBT:∑BTs, suggesting that the higher the level of TBTCl exposed, the lower the metabolism rate of the fish. TBTCl exposure led to a poor reproductivity and an abnormal sexual behavior in the fish, i.e. a reduced number of the male sexual sigmoid display and of spermatophore in the efferent duct was observed in the fish exposed to 0.1 <TEX>$\mug/l$</TEX> and higher levels of TBTCl, and a decreasing ratio of the testicular spermatophore cyst to the whole germ cell cysts was observed in the fish exposed to 0.32∼10 <TEX>$\mug/l$</TEX>)of TBTCl. The reduced ratio of the spermatophore cyst seems to be an effect of the endocrine disrupter inhibiting spermiogenesis. In the fish exposed to 25 <TEX>$\mug/l$</TEX> and higher levels of TBTCl, more serious effects, such as a rapid increase of mortality, the necrosis of most of the germ cells, great damages in Sertoli cells and epithelial cells of the efferent duct, a significant increase of abnormal swimming behavior, and a cessation of feeding were observed, which suggest the acute toxicity of TBTCl inhibiting not only the reproduction and behavior but also the survival of the fish itself.

Mouse Ovarian Germ Cell Cysts Undergo Programmed Breakdown to Form Primordial Follicles

In many organisms, early germline development takes place within cysts of interconnected cells that form by incomplete cytokinesis and later undergo programmed breakdown. We recently identified similar cell clusters within the fetal mouse ovary, but the fate and functional significance of these germ cell cysts remained unclear. Here, we show that mouse cysts undergo programmed breakdown between 20.5–22.5 dpc, during which approximately 33% of the oocytes survive to form primordial follicles. This process accounts for most of the perinatal reduction in germ cell numbers and germ cell apoptosis reported by previous authors, and suggests that perinatal germ cell loss is a developmentally regulated process that is distinct from the follicular atresia that occurs during adult life. Our observations also suggest a novel function for a transient cyst stage of germ cell development. Prior to breakdown, mitochondria and ER reorganize into perinuclear aggregates, and can be seen within the ring canals joining adjacent germ cells. Cysts may ensure that oocytes destined to form primordial follicles acquire populations of functional mitochondria, through an active process that has been evolutionarily conserved.

Biopsy of Mediastinal Tumors: Needle Biopsy versus Mediastinoscopy

Biopsy of Mediastinal Tumors: Needle Biopsy versus Mediastinoscopy

Cyclin A Associates with the Fusome during Germline Cyst Formation in the Drosophila Ovary

Regulated changes in the cell cycle underlie many aspects of growth and differentiation. Prior to meiosis, germ cell cycles in many organisms become accelerated, synchronized, and modified to lack cytokinesis. These changes cause cysts of interconnected germ cells to form that typically contain 2n cells. InDrosophila, developing germ cells during this period contain a distinctive organelle, the fusome, that is required for normal cyst formation. We find that the cell cycle regulator Cyclin A transiently associates with the fusome during the cystocyte cell cycles, suggesting that fusome-associated Cyclin A drives the interconnected cells within each cyst synchronously into mitosis. In the presence of a normal fusome, overexpression of Cyclin A forces cysts through an extra round of cell division to produce cysts with 32 germline cells. Female sterile mutations in UbcD1, encoding an E2 ubiquitin-conjugating enzyme, have a similar effect. Our observations suggest that programmed changes in the expression and cytoplasmic localization of key cell cycle regulatory proteins control germline cyst production.

Testis structure and symplastic spermatid formation during spermatogenesis of pipefishes

The pipefishes Syngnathus abaster and S. acus have paired testes of atypical organization. Each testis is a hollow tube consisting of a single germinal compartment of the tubular type. During the reproductive period, the germinal epithelium consists of small spermatocysts containing spermatogonia or primary spermatocytes. Cysts of older germ cells, such as secondary spermatocytes and spermatids were never observed. Developing symplastic spermatids were found in the lumen of the tubule together with mature sperm and large droplet‐containing cells. Most of the spermatids were giant cells with four nuclei at the same developmental stage. Symplastic spermatids, which presumably form by nuclear division not followed by cytokinesis, are a stage of spermatogenesis in pipefishes.

Temperature effects on sex differentiation of two South American atherinids, Odontesthes argentinensis and Patagonina hatcheri

The present study investigated the effects of water temperature (18, 21, and 25 °C) on the histological process of gonadal sex differentiation of two commercially important atherinid fishes from South America, Odontesthes argentinensis (sea pejerrey) and Patagonina hatcheri (Patagonian freshwater pejerrey). In both species, female gonadal sex differentiation began with the formation of lateral stromal cell outgrowths and the appearance of meiotic oocytes. The male gonads remained quiescent for about twice as long as the female gonads, with differentiation becoming evident by the formation of the main sperm duct and of cysts of germ cells at the periphery of the gonads. Meiosis in males occurred relatively long after somatic differentiation of the testis. The ovaries of O. argentinensis differentiated at 28 days (20.3 mm) at 25 °C, 42 days (24.0 mm) at 21 °C, and 56 days (23.8 mm) at 18 °C. In the males, differentiation was observed at 98 days at 25 and 21 °C (39.4 mm and 40.4 mm, respectively), but at 112 days under 18 °C (40.7 mm). In P. hatcheri, differentiation of females occurred at 21 days (17.8 mm) at 25 °C, 28 days (20.8 mm) at 21 °C, and 35 days (23.2 mm) at 18 °C. Male differentiation became evident at 56 days under 25 and 21 °C (30.8 and 32.7 mm, respectively), and at 70 days (37.7 mm) at 18 °C. The sex-ratios of O. argentinensis reared at 18 or 21 °C were female-biased whereas those at 25 °C were not; groups reared at 18 °C had significantly more females than groups from the same progeny reared at 25 °C. In contrast, the sex-ratios in all groups of P. hatcheri did not differ significantly from 1:1 and no significant differences were found between groups of the same progeny reared at different temperatures. These results suggest the occurrence of thermolabile sex determination (TSD) in O. argentinensis whereas in P. hatcheri gonadal sex appears to be strongly genetically determined.

Innervation of testes in the tilapia Oreochromis niloticus

The innervation of the testes of Oreochromis niloticus was examined immunohistochemically and ultrastructurally. Many thick nerve bundles, which penetrate the testes via the mesorchium, are distributed in the wall surrounding the efferent duct. The neurons appear to terminate at the surface of smooth muscle cells in the efferent duct wall. Thin nerve bundles penetrate the interstices among lobules containing cysts of spermatogenic germ cells. Terminal ends of these nerve fibres also make contact with smooth muscle cells enclosing lobules and with the surface of steroid-producing Leydig cells (SPC). These observations suggest that innervation of the testis may not only control the contracting and relaxing of smooth muscle cells but also control steroid secretion. Some fibres containing many electron-dense granules were observed in the interstices near the cluster of SPC. Such granules suggest that these fibres may be peptidergic.

Intermediate filaments in the testis of the teleost mosquito fish Gambusia affinis holbrooki: a light and electron microscope immunocytochemical study and Western blotting analysis

A light and electron microscope immunocytochemical study and Western blotting analysis has been performed on intermediate filaments (vimentin, desmin and cytokeratins) in the testis of the teleost fish Gambusia affinis holbrooki. An immunoreaction to vimentin was observed in the epithelium of the efferent ducts, testicular canal and their surrounding peritubular cells. Positive vimentin immunostaining was also observed in the cells located around seminiferous tubules (boundary cells), Leydig cells, interstitial fibroblasts, chromatophores, and blood vessel endothelial cells. In contrast to mammals, no vimentin immunoreactivity was found in the Sertoli cells. Immunoreactivity to desmin was weak in the epithelial cells of the efferent ducts and testicular canal and intense in the peritubular cells that surrounded these ducts. Desmin immunoreactivity was also observed in the seminiferous tubule boundary cells. The immunoreactivity was weak in the boundary cells that surrounded germ cell cysts containing spermatogonia or spermatocytes and intense in the boundary cells around cysts with elongated or mature spermatids. Immunoreactivity towards cytokeratins was observed only in testicular blood vessels. Cytokeratin immunolabelling was intense in the endothelium and weak in the vascular smooth muscle cells. No cytokeratin immunoreactivity was found in the Sertoli cells, germ cells, interstitial cells or in the efferent duct epithelium. The absence of intermediate filaments in the Sertoli cells, the absence of cytokeratins in the epithelium of the sperm excretory ducts, and the presence of desmin filaments in these epithelial cells are the most important differences with regards to the intermediate filament phenotype in mammalian testes.

Cytoskeleton in Sertoli cells of the mosquito fish (Gambusia affinis holbrooki).

There is little information about the distribution of cytoskeletal components in the testes of teleost fish. The aim of this paper was to know the distribution of some major cytoskeletal proteins (tubulin, actin, vimentin, desmin, and cytokeratins) in the Sertoli cells of Gambusia affinis holbrooki and in their efferent duct epithelial cells which are possibly originated from the Sertoli cells. Light and electron microscopic immunocytochemical studies and Western blotting analysis were performed in G. affinis testis. Actin immunoreaction was observed in the Sertoli cells at all spermatogenic stages, although the intensity of this reaction varied from one stage to another. Sertoli cells that support spermatogonia or spermatocytes showed a weak immunoreaction which was uniformly distributed throughout the cytoplasm and somewhat more concentrated at the level of the inter-Sertoli specialized junctions. Immunoreaction to actin increased during the first stages of spermiogenesis and was mainly localized beneath the plasma membrane. This immunoreaction was more intense in the basal than in the apical cytoplasm of Sertoli cells. In a more advanced stage of spermiogenesis, actin immunoreaction become stronger in the apical cytoplasm where Sertoli cells displayed cytoplasmic projections around each spermatid. After sperm release, the apical Sertoli cell cytoplasm still showed an intense actin immunoreaction. Intense immunoreaction to actin was also observed in the epithelial cells lining the efferent ducts. Immunoreaction to tubulin was diffuse throughout the Sertoli cell cytoplasm. No immunoreaction to vimentin or desmin was observed in the Sertoli cells during the spermatogenic process. Immunoreaction to both vimentin and desmin was observed in the efferent duct cells. Desmin immunoreaction was also observed in the seminiferous tubule boundary cells, mainly in the sections showing germ cell cysts at the last stages of spermiogenesis and in the peritubular cells that surrounded the efferent duct epithelium. Immunoreaction to cytokeratins was found in the endothelium of testicular blood vessels but not in the Sertoli cells or in the efferent duct epithelium. Immunoreaction pattern to cytoskeletal proteins in the Sertoli cells of G. affinis differs from that reported in mammalian Sertoli cells. These differences include the distribution of actin filaments and the absence of detectable vimentin immunoreaction in G. affinis Sertoli cells.

Spermatogenesis in the urodele Amhystoma dumerilii.

The male reproductive cycle of this paedomorphic species that occurs only in Lake Pátzcuaro, Michoacán, México, was investigated by documenting changes in germinal cells during the spermatogenic cycle. Cysts of germ cells divide synchronously to complete spermatogenesis during September through December, with the proportion of evacuated cysts or cysts containing spermatozoa increasing during this period. The chromatin changes during prophase I of meiosis reveal the usual leptotene, zygotene, pachytene, and diplotene stages. A basal body at the caudal end of the spermatozoan head connects to the flagellum. After spermiation, empty cysts contain a granular substance. Spermatogenesis in this species follows an annual cycle like other north temperate salamanders, rather than the continuous spermatogenesis of some tropical salamanders. © 1994 Wiley-Liss, Inc.

The blood-testis barrier and its breakdown following immunization to testis material in the Nile tilapia, Oreochromis niloticus

The blood-testis barrier and its changes following immunization to testis material, were investigated by light- and electron microscopy in a teleost fish, the Nile tilapia Oreochromis niloticus, using horseradish peroxidase and bovine serum albumin as tracers. In the normal testis, histochemistry using horseradish peroxidase revealed that a barrier composed of junctional complexes connecting adjacent Sertoli cells existed around the central lumina of the seminal lobules, and also around the germ-cell cysts containing spermatids at the middle or late phase of chromatin condensation. By contrast, bovine serum albumin was prevented from passing through the basement membrane and could not penetrate any of the spermatogenetic cysts, indicating that the basement membrane may be an ion-selective barrier. In tilapia immunized with allogeneic testis homogenate emulsified in Freund's complete adjuvant, bovine serum albumin could penetrate the spermatogenetic cysts, and horseradish peroxidase was able to pass through the intercellular spaces between Sertoli cells to the region nearer the seminal lobule lumen, due to the junctional complexes becoming loosened. The results suggest that the blood-testis barrier, both junctional complexes and the basement membrane, are broken down during immune responses.

Effects of hypophysectomy on the testis of the European eel ( Anguilla anguilla L.)

The pituitary control of testis activity in the freshwater silver European eel ( Anguilla anguilla L.) was investigated by study of some variations of gonad structure and particularly changes in plasma levels of testosterone and 11-oxotestosterone (radioimmunoassay) following hypophysectomy. Germ cells comprised only spermatogonia, which were associated with Sertoli cells to form cysts. No mitotic activity of spermatogonia was observed. Gonadosomatic index (GSI) was low. Plasma levels of the main androgens (testosterone, 11-oxotestosterone) were under 1 ng ml −1. The male gonad appeared immature. After hypophysectomy, GSI decreased rapidly and cysts of germ cells were rapidly dissociated. Plasma level of testosterone decreased significantly, whereas that of 11-oxotestosterone increased significantly. These changes were not observed in sham-operated eels. Thus pituitary control appears essential for the maintenance of structural organization and secretory activity of the immature testis.

Fine structure of spermatogenesis and mature spermatozoa in Cryptocellus boneti Bolivar y Pieltain, 1941 (Arachnida, Ricinulei)

The testes of Cryptocellus boneti are composed of a germinal and a secretory part. Epithelial cells of the latter are capable of phagocytosing mature spermatozoa besides secreting a proteinaceous secretory product delivered into the lumen of the testes. The germinal part comprises somatic cells surrounding cysts of germ cells. Mature spermatozoa are roundish cells. Their central cytoplasm is ensheathed by two electron-dense intracellular plates. An elongated nucleus and an axoneme of the usual 9 × 2 + 2 pattern are rolled up within the cell. An acrosomal complex comprising a small acrosomal vacuole and an acrosomal filament is present. Early spermatogenesis reveals that eight electron-dense plates are derived from Golgi vesicles and are distributed to the spermatids during meiotic divisions. Elongation of spermatids occurs during chromatin condensation. The nucleus is paralleled by the axonemal tubules which are attached via a specialized centriolar region to the anterior curved acrosomal vacuole.