Ultrastructural analysis of the ovary and oogenesis in Spinicaudata and Laevicaudata (Branchiopoda) and its phylogenetic implications

Recent molecular studies have indicated a close relationship between Crustacea and Hexapoda and postulated their unification into the Pancrustacea/Tetraconata clade. Certain molecular analyses have also suggested that the crustacean lineage, which includes the Branchiopoda, might be the sister group of Hexapoda. We test this hypothesis by analyzing the structure of the ovary and the ultrastructural features of oogenesis in two branchiopod species, Cyzicus tetracerus and Lynceus brachyurus, representing two separate orders, Spinicaudata and Laevicaudata, respectively. The female gonads of these species have not been investigated before. Here, we demonstrate that in both studied species the ovarian follicles develop inside characteristic ovarian protrusions and comprise a germline cyst surrounded by a simple somatic (follicular) epithelium, supported by a thin basal lamina. Each germline cyst consists of one oocyte and three supporting nurse cells, and the oocyte differentiates relatively late during ovarian follicle development. The synthesis of oocyte reserve materials involves rough endoplasmic reticulum and Golgi complexes. The follicular cells are penetrated by a complex canal system and there is no external epithelial sheath covering the ovarian follicles. The structure of the ovary and the ultrastructural characteristics of oogenesis are not only remarkably similar in both Cyzicus and Lynceus, but also share morphological similarities with Notostraca as well as the basal hexapods Campodeina and Collembola. Possible phylogenetic implications of these findings are discussed.     

The structure of the ovary and oogenesis in the earthworm,Dendrobaena veneta (Annelida,Clitellata)

The structure of the ovary and the type of oogenesis were determined in the earthworm Dendrobaena veneta (Oligochaeta, Haplotaxida, Lumbricidae) with histological, electron-microscopic and immunocytochemical methods. In this species the ovary is of the alimentary, nutrimentary type because it contains oocytes and the nurse cells (trophocytes). The ovarian stroma is built by somatic cells, the processes of which are connected to each other via numerous desmosomes. The somatic cells and their processes envelop the germ cells tightly and play a supportive role. Oogonia, oocytes and trophocytes are arranged in distinct zones in the ovary. Trophocytes form chains of cells, which are interconnected by intercellular bridges. Numerous microtubules are located within the latter. The oocytes are distally arranged in the ovary. Vitellogenesis involves both auto- and heterosyntheses. The results obtained were compared with the reports on oogenesis in other representatives of Annelida.     

Germline stem cells and neo-oogenesis in the adult human ovary

It remains unclear whether neo-oogenesis occurs in postnatal ovaries of mammals, based on studies in mice. We thought to test whether adult human ovaries contain germline stem cells (GSCs) and undergo neo-oogenesis. Rather than using genetic manipulation which is unethical in humans, we took the approach of analyzing the expression of meiotic marker genes and genes for germ cell proliferation, which are required for neo-oogenesis, in adult human ovaries covering an age range from 28 to 53 years old, compared to testis and fetal ovaries served as positive controls. We show that active meiosis, neo-oogenesis and GSCs are unlikely to exist in normal, adult, human ovaries. No early meiotic-specific or oogenesis-associated mRNAs for SPO11, PRDM9, SCP1, TERT and NOBOX were detectable in adult human ovaries using RT-PCR, compared to fetal ovary and adult testis controls. These findings are further corroborated by the absence of early meiocytes and proliferating germ cells in adult human ovarian cortex probed with markers for meiosis (SCP3), oogonium (OCT3/4, c-KIT), and cell cycle progression (Ki-67, PCNA), in contrast to fetal ovary controls. If postnatal oogenesis is confirmed in mice, then this species would represent an exception to the rule that neo-oogenesis does not occur in adults.     

Taxonomic significance of the inner structure of the ovary in the genusImpatiens (Balsaminaceae)

The inner structure of the ovary inImpatiens was examined from the viewpoint of taxonomy, based on the materials obtained during the 1979 Botanical Expedition to Thailand. The species studied areI. kerriae, I. larsenii, I. hongsonensis, I. psittacina, I. salaengensis, I. charanii, I. kanburiensis andI. macrosepala. It is worthy of special mention that they are all characterized by 4 lateral sepals, large round outer sepals, a nipple-like short spur of the lip and connate wing petals. Contrary to the prevailing information about the ovary ofImpatiens, these Thai species were found to have 4-locular and 4-carpellate ovaries. The arrangement and the number of ovules in a loculus show a tendency of reduction from biseriate to uniseriate and from several to single or none. The species of more advanced type have smaller numbers of ovules and often sterile loculi of the ovary, as represented byI. macrosepala. The types of ovule development were proved to correspond to the degree of specialization of floral characters. Supported by a Grant in Aid of Scientific Research of Ministry of Education, No. 404126 and No. 504352, and partly reported at the Meeting of the Japan Society of Plant Taxonomists in March 1981.     

A visual screen for diet-regulated proteins in the Drosophila ovary using GFP protein trap lines

The effect of diet on reproduction is well documented in a large number of organisms; however, much remains to be learned about the molecular mechanisms underlying this connection. The Drosophila ovary has a well described, fast and largely reversible response to diet. Ovarian stem cells and their progeny proliferate and grow faster on a yeast-rich diet than on a yeast-free (poor) diet, and death of early germline cysts, degeneration of early vitellogenic follicles and partial block in ovulation further contribute to the ∼60-fold decrease in egg laying observed on a poor diet. Multiple diet-dependent factors, including insulin-like peptides, the steroid ecdysone, the nutrient sensor Target of Rapamycin, AMP-dependent kinase, and adipocyte factors mediate this complex response. Here, we describe the results of a visual screen using a collection of green fluorescent protein (GFP) protein trap lines to identify additional factors potentially involved in this response. In each GFP protein trap line, an artificial GFP exon is fused in frame to an endogenous protein, such that the GFP fusion pattern parallels the levels and subcellular localization of the corresponding native protein. We identified 53 GFP-tagged proteins that exhibit changes in levels and/or subcellular localization in the ovary at 12–16 hours after switching females from rich to poor diets, suggesting them as potential candidates for future functional studies.     

Amblyomma triste (Koch, 1844) (Acari: Ixodidae): morphological description of the ovary and of vitellogenesis

The present study presents the morphology, histology, and the dynamics of vitellogenesis in females of the tick Amblyomma triste. The ovary in this species is of the panoistic type, therefore it lacks nurse cells. It is composed of a layer of epithelial cells that outwardly form the wall of the ovary, but also originate the pedicel, the structure that attaches the oocytes to its external margin, as well the oocytes themselves. In Amblyomma triste, the oocytes develop in four synchronic stages, which differs from the process in other tick species. The classification of the stages of the oocytes was carried out based on the presence of four morphologic characteristics: cytoplasm appearance; site of the germ vesicle; presence, quantity, and constitution of the yolk granules and presence of chorium.     

Germline cyst formation and incomplete cytokinesis during Drosophila melanogaster oogenesis

Germline cyst formation via incomplete cytokinesis (IC) is necessary to generate functional eggs and sperm in various organisms. Drosophila melanogaster oogenesis is an ideal system for studying IC. 29 stages of germline cyst formation can be identified in D. melanogaster oogenesis. We have defined necessary terminology to describe IC and have developed a method to measure the sizes of contractile rings and ring canals. Time course study of germline cyst formation demonstrates that contractile ring constriction proceeds to a defined end point unique for each mitotic division. Contractile rings constrict to a greater degree, resulting in smaller ring diameters, for each subsequent round of mitotic division. Contrary to conventional wisdom, ring canal growth is not initiated until well after the fourth mitotic division. Ring canals grow, in an orderly manner, with ring canals derived from the first mitotic division enlarging first followed by those from the second, then those from the third, and finally those from the fourth mitotic division. This work establishes a foundation for identifying genes specific for IC and for elucidating the molecular mechanism underlying this aspect of germline cyst formation.     

Morphological characterization of the ovary and oocytes vitellogenesis of the tick Rhipicephalus sanguineus (Latreille, 1806) (Acari: Ixodidae)

This study presents the morphology of the ovary, as well as the process of the vitellogenesis in oocytes of the tick Rhipicephalus sanguineus. The ovary of these individuals is of the panoistic type; therefore, it lacks nurse cells. This organ consists of a single tubular structure, continuous, and composed of a wall formed by small epithelial cells with rounded nuclei which delimit the lumen. The oocytes in the different developmental stages in this tick species were classified into five stages (I-V). They remain attached to the ovary during vitellogenesis by a cellular pedicel and afterwards the mature oocytes (stage V) are released into the ovary lumen.     

The structure and phylogenetic significance of the flagellar transition region in the chlorophyll c-containing algae.

An electron-dense helix is the most conspicuous structure in the flagellar transition region of members of the algal class Chrysophyceae. This “transitional helix” (TH) lies immediately distal to a partition across the flagellar axoneme which occurs exactly at the level at which the flagellum enters the cell body. The helix surrounds the central axonemal pair and lies at a distance of 10 nm from the 9 peripheral doublets. From the new data presented and a survey of published observations on the structure of the transition region of all the chlorophyll c-containing classes of algae, it is shown that a TH characteristic of the Chrysophyceae, Xanthophyceae and Eustigmatophyceae. The number of TH gyres varies from 3 to 6 in the Xanthophyceae and from 1 to 8 in the Chrysophyceae. In any one species, however, the TH is the same size in both the long flagellum which bears tubular mastigonemes and in the short smooth flagellum, though in some chrysophytes where the short flagellum is vestigial the number is fewer than in the normal flagellum. A TH appears to be absent from the Rhaphidophyceae and zoids of the Bacillariophyceae and Phaeophyceae though the structure of the transition region in these groups otherwise resembles that of the Chrysophyceae, Xanthophyceae and Eustigmatophyceae.The value of transition region variation in determining evolutionary relationships among the chlorophyll c-containing algal classes is assessed against a background of current ideas on their taxonomy and phylogeny. The relevant structural and biochemical features are tabled, and a phylogenetic scheme is presented which appears most logically to interpret these data. It is suggested that the line leading to the Eustigmatophyceae probably diverged from that leading to the strictly heterokont classes Xanthophyceae, Chrysophyceae, Phaeophyceae and Bacillariophyceae before evolution of a girdle lamella in the chloroplast and a photoreceptor apparatus involving a swelling at the proximal end of the short flagellum and an intraplastidial eyespot. The possession of a TH by both the Chrysophyceae and Xanthophyceae adds further support to the concept of their close relationship based on a range of other features. The exceptional absence of a TH from the chrysophycean genera Pedinella and Pseudopedinella reinforces the idea that these taxa are remote from the main chrysophycean line. Absence of a TH from the Phaeophyceae and Bacillariophyceae which otherwise share many important features with the Chrysophyceae and Xanthophyceae is probably a result of loss owing to the functional and morphological specialization of the zoids of these two groups. Transition region structure does not clarify the possible relationships of the Rhaphidophyceae, Prymnesiophyceae, Cryptophyceae or Dinophyceae.The proposed phylogeny supports the idea of a mutually related “heterokont” protist assemblage comprising the Chrysophyceae, Xanthophyceae, Phaeophyceae, Bacillariophyceae and possibly Rhaphidophyceae and the Oomycetes (water moulds) though in the latter the TH is replaced by a dense cylinder with a corrugated wall which may or may not be homologous with it. Structures resembling a TH have been described in a wide variety of other flagellated cells including the prasinophyte Pyraminonas orientalis, one species of the colourless flagellate genus Bicosoeca and the proteromonads Karotomorpha and Proteromonas. Only in the latter genera does homology with a TH seem likely on present evidence, suggesting that flagellates of this type may have evolved from chrysomonad-like ancestors.     

Development of the calyx and lateral oviduct during oogenesis in Aedes aegypti

Summary The lateral oviduct and calyx of nulliparous Aedes aegypti on a sucrose diet are both flattened sacs, lacking a well defined lumen. Both are formed of an inner epithelial and an outer muscular layer, each one cell thick. The lateral oviduct is surrounded by a circular muscle sheath which is continuous with the ovarian sheath. Each ovariolar sheath is continuous with the outer layer of the calyx. The structure of both the lateral oviduct and the calyx is greatly modified after the initial blood meal. A distinct lumen develops; there is an extensive development of the outer muscular layers, and the inner epithelial layers become invaginated forming deep crypts lined with extensive microvilli. The follicular stem, which joins the primary follicle to the calyx in each ovariole, is not hollow and does not mark the opening into the calyx through which the mature egg can pass. The eggs gain access to the oviductal system after the calyx extends around the follicular epithelium of the primary follicle, when breaks appear in the calyx wall opposed to the follicular epithelium, until the mature eggs, eventually lie in a highly distended thin-walled sac of calyx from which they have direct and easy access to the lateral oviduct. After oviposition, this sac contracts to occupy once more a compact axial position in the ovary. Remnants of the follicular epithelium, containing many lysosomes are attached to the calyx at this time.     

Eggs over easy: cell death in the Drosophila ovary

Programmed cell death is the most common fate of female germ cells in Drosophila and many animals. In Drosophila, oocytes form in individual egg chambers that are supported by germline nurse cells and surrounded by somatic follicle cells. As oogenesis proceeds, 15 nurse cells die for every oocyte that is produced. In addition to this developmentally regulated cell death, groups of germ cells or entire egg chambers may be induced to undergo apoptosis in response to starvation or other insults. Recent findings suggest that these different types of cell death involve distinct genetic pathways. This review focuses on progress towards elucidating the molecular mechanisms acting during programmed cell death in Drosophila oogenesis.     

Phylogenetic significance of the structure of adult ovary and oogenesis in a primitive chilognathan diplopod,Hyleoglomeris japonica Verhoeff (Glomerida,Diplopoda)

Some histological details of the adult ovary of Hyleoglomeris japonica are described for the first time in the glomerid diplopods. The ovary is a single, long sac-like organ extending from the 4th to the 12th body segment along the median body axis, lying between the alimentary canal and the ventral nerve cord. The ovarian wall consists of a layer of thin ovarian epithelium which surrounds a wide ovarian lumen. A pair of longitudinal “germ zones,” including female germ cells, runs in the lateral ovarian wall. Each germ zone consists of two types of oogenetic areas: 1) 8–12 narrow patch-shaped areas for oogonial proliferation, arranged metamerically in a row along each of the dorsal and ventral peripheries, and 2) the remaining wide area for oocyte growth. Oogonial proliferation areas include oogonia, very early previtellogenic oocytes, and young somatic interstitial cells, among the ovarian epithelial cells. The larger early previtellogenic oocytes in the oogonial proliferation areas are located nearer to the oocyte growth area, and migrate to the oocyte growth area. They are surrounded by a layer of follicle cells and are connected with the ovarian epithelium of the oocyte growth area by a portion of their follicles. They grow into the ovarian lumen, but their follicles are still connected with the oocyte growth area. Various sizes of the previtellogenic and vitellogenic oocytes in the ovarian lumen are connected with the oocyte growth area; the smaller oocytes are connected nearer to the dorsal and ventral oogonial proliferation areas, while the larger ones are connected nearer to the longitudinal middle line of the oocyte growth area. Following the completion of vitellogenesis and egg membrane formation in the largest primary oocytes, the germinal vesicles break down. Ripe oocytes are released from their follicles directly into the ovarian lumen to be transported into the oviducts. Ovarian structure and oogenesis of H. japonica are very similar to those of other chilognathan diplopods. At the same time, however, some characteristic features of the ovary of H. japonica are helpful for understanding the structure and evolution of the diplopod ovaries. Some aspects of the phylogenetic significance in the paired germ zones of H. japonica are discussed. J. Morphol 231:277–285, 1997. © 1997 Wiley-Liss, Inc.     

Fine structure of the pharyngeal bulb in Trilobodrilus and its phylogenetic significance within archiannelida.

Transmission electron microscopical studies of the pharyngeal bulb in the archiannelid Trilobodrilus sp. proved the presence of two opposing myofilament systems in the plate muscle cells forming this bulb. These muscles are obliquely striated. The significance as well as the origin of this specialized plate-muscle cells in the pharyngeal bulb of archiannelids is discussed in the light of their controversial phylogenetic and systematic position. In addition a hypothesis is suggested to explain the origin of the special arrangement of two opposing systems of myofilaments in one single muscle cell.     

The initial stages of oogenesis and their relation to differential fertility in the honey bee (Apis mellifera) castes

Neither the overall differences in ovariole number nor the caste-specifically modulated expression of vitellogenin can fully explain the striking caste differences in honey bee reproduction, in particular the mechanisms that block oogenesis in virgin queens and in workers kept in the presence of a queen. For this reason we investigated the initial stages of oogenesis in queens in relation to mating status and in workers exposed to different social conditions. A striking feature in ovarioles of both castes was a considerably elongated terminal filament which consisted not only of normal terminal filament cells but also contained apparently undifferentiated cells that were tentatively considered as stem cells. BrdU incorporation was detected in the upper germarium, as well as in the terminal filament. Cytoskeleton analysis by TRITC-phalloidin labeling for F-actin, and immunofluorescence detection for β-tubulin did not reveal structural differences in the early oogenesis steps between queens and queenless workers. In contrast, queenright workers showed signs of a disorganized microtubule and microfilament system that could explain the histological evidence for progressive cell death observed in the germaria. In addition to cytoplasmic tubulin we also detected marked intranuclear foci indicating the presence of nuclear β_II-tubulin.     

The rhabdom structure in the ommatidia of the Heteroptera (Insecta), and its phylogenetic significance

In its plesiomorphic state the insect ommatidium consists of eight retinula cells forming a fused rhabdom. It has long been observed that, in contrast to this pattern, Heteroptera have open rhabdoms. However, there has so far been no comprehensive and comparative study of heteropteran ommatidia. For this reason, we investigated the rhabdom structure in 36 species from all higher groups of Heteroptera, as well as from Coleorrhyncha and Auchenorrhyncha as outgroup representatives. In addition we surveyed the data of earlier authors, which brings the number of examined species to a total of more than 70. All examined Heteroptera do have open rhabdoms, with a system of six peripheral and two central rhabdomeres. Outgroup comparison shows that the open rhabdom is an autapomorphy of the Heteroptera. As for the rhabdom structure within the Heteroptera, we found further autapomorphic patterns in Corixidae (Nepomorpha), Gerromorpha, and Leptopodomorpha. Finally, the Cimicomorpha and Pentatomomorpha share a special pattern of the two central rhabdomeres, which we call V-pattern. This is a new synapomorphy of these two taxa. Accepted: 22 November 1999     

Morphological study of the ovary in Hanseniella caldaria (Myriapoda; Symphyla): The position of oocyte-growth and evolution of ovarian structure in Arthropoda

In Arthropoda, the ovary is classified into Chelicerata-type and Mandibulata-type, based on the oocyte-growth position within the ovary. By contrast, oocytes of Diplopoda and Chilopoda grow within the hemocoelic space. However, as the position of oocyte-growth in Symphyla and Pauropoda has not been confirmed, whether the hemocoelic nature of oocyte-growth is common among myriapods remains ambiguous. This study described the ovarian structure of Hanseniella caldaria to reveal the oocyte-growth position in Symphyla. The oocyte is surrounded by the follicle epithelium, and the inner surface of the follicle epithelium, i.e., the space between follicle cells and oocytes, is lined with a basement membrane. The follicle epithelial layer continues to the ovarian epithelium via the follicle extension with a continuous layer of basement membrane. Data on the architecture of the follicle suggest that the follicle pouch opens to the hemocoel. Hence, the oocyte of H. caldaria grows within the hemocoelic space. Based on our findings in H. caldaria and previous studies in a millipede and in centipedes, the hemocoelic nature of oocyte-growth is considered as a common feature among myriapods and a synapomorphy of the Myriapoda for which morphological synapomorphies have been ambiguous.     

物种谱系不确定性对群落谱系格局指标的影响

物种谱系关系常被用于衡量群落谱系格局及推断格局背后的生态过程,但多数研究往往忽视谱系关系的不确定性及其可能对群落谱系格局造成的影响.为此,本文以浙江天童20 hm^2样地内150个树种为研究对象,采用这些物种叶绿体DNA的rbcL和matK碱基序列构建1棵一致系统发育树和反映谱系不确定性的999棵系统发育树,然后结合样地物种分布数据计算标准化净亲缘指数(NRI)和最近亲缘指数(NTI),最后运用独立置换零模型衡量样地群落谱系格局.结果表明:物种系统发育树在拓扑结构和物种谱系分支节点年龄上均存在较大的不确定性,谱系不确定性随着谱系分支节点年龄的减小而增大,也随物种间平均谱系距离的增加而增加;在样方尺度上,物种谱系的不确定性增加了标准化NRI和NTI指数的变异,但对两个指数的影响几乎独立;其对两指数的空间分布影响不同,且程度不一,其中标准化NRI受到的影响相对更大;在群落尺度上,物种谱系的不确定性增加了标准化NRI和NTI的变异,平均变异系数分别为0.37和0.077,表明群落水平的标准化NRI更易受到谱系不确定性的影响.这说明物种谱系不确定性会传递到常用的群落谱系格局指标中,且不同指标受影响的程度不同,进而影响对群落谱系格局的衡量及相关生态过程的推断.该结论也暗示以往不考虑谱系不确定性的研究中,非随机的群落谱系格局比例可能被高估.