Ultrastructure of the ovary and oogenesis in the jellyfish Linuche unguiculata and Stomolophus meleagris,with a review of ovarian structure in the Scyphozoa

Ovarian structure and oogenesis has been examined in six scyphozoan species including the semaeostome Diplumularis antarctica Maas, 1908 (collected in 1987 in McMurdo Sound, Antarctic), the rhizostomes Cassiopea xamachana Bigelow, 1892 (collected in Belize in 1988), and Stomolophus meleagris L. Agassiz, 1862 (collected in Ft. Pierce Inlet in 1988), and the coronates Periphylla periphylla (Peron and Lesueur, 1810), Nausithoe atlantica Broch, 1914 (both collected in the Bahamas in 1988), and Linuche unguiculata (Schwartz, 1788) (collected in Nassau Harbor, Bahama Islands in 1989). Based on these findings and information on five other scyphozoan species from additional literature sources, at least two fundamentally different types of ovaries exist in the Scyphozoa. In semaeosotome and rhizostome species, oocytes develop in close association with specialized gastrodermal cells called trophocytes which may serve a nutritive function. However, coronate species lack trophocytes and oocytes develop freely in the mesoglea. The ovaries of S. meleagris and L. unguiculata are used as models to represent the ultrastructural events occurring during oogenesis in species having trophocytes and those lacking them, respectively. In both L. unguiculata and S. meleagris, the ovaries arise as evaginations of the gastrodermis in the floor of interradial pouches. Germ cells appear to originate from endodermally-derived gastrodermal cells and migrate into the mesoglea prior to vitellogenesis. In L. unguiculata, the oocytes develop freely within the mesoglea throughout vitellogenesis, while in S. meleagris each oocyte maintains contact with specialized gastrodermal cells called trophocytes. In the vitellogenic oocytes of both species, numerous invaginations of the oolemma result in the formation of intraooplasmic channels throughout the ooplasm. These channels are intimately associated with cisternae of rough endoplasmic reticulum and may play some role in yolk precursor uptake by substantially increasing the surface area of the oocyte. Vitellogenesis is similar in both species and involves the autosynthetic activity of the Golge complex and rough endoplasmic reticulum, and the heterosynthetic incorporation of yolk precursors through receptor-mediated endocytosis. However, in the oocytes of S. meleagris, the trophocytes probably play a role in the transfer of nutrients from the gastrovascular cavity to the oocyte. The present study suggests that scyphozoans were among the first metazoans to develop ovarian accessory cells during their reproductive evolution. The trophocyte-oocyte association observed in some scyphozoans is similar to but structurally less complex than the trophonema-oocyte association described from anthozoans. Scyphozoan ovarian morphology helps support the view that the Scyphozoa share a closer phylogenetic relationship with the Anthozoa than with the Hydrozoa.     

Oogenesis in the marine mussel Mytilus edulis: an ultrastructural study

Ultrastructural changes occurring during the course of development in oocytes of Mytilus edulis are described for mussels collected at monthly intervals over a period of one year (September 1981 to October 1982) from a site in Cornwall, England. During early stages of oogenesis the oocyte is surrounded by a small number of follicle cells but, as development proceeds, the follicle cells are restricted to the stalk region which attaches the oocyte to the acinar wall. Contact between the follicle cells and the developing oocyte is maintained by means of desmosomelike gap junctions. Organelles and inclusion bodies present in the ooplasm during oogenesis include rough endoplasmic reticulum (RER), Golgi bodies, mitochondria, free ribosomes, Balbiani's vitelline body, annulate lamellae and yolk and cortical granules. The RER, in particular, varies considerably throughout the course of development. Evidence for uptake of exogenous macromolecules into oocytes by pinocytosis is presented; it occurs in the basal region of previtellogenic oocytes prior to the formation of the vittelline coat. Lipid-yolk granules invariably have mitochondria in close association and, during the winter months, develop in close proximity to small, apparently glycogen-rich vesicles possibly suggesting that conversion of glycogen to lipid takes place in developing oocytes. Oocyte degeneration was commonly observed and involves initial breakdown of the plasma membrane followed by rupture of the vitelline coat. The oocyte contents once released into the acinar lumen are resorbed by the epithelial cells of the gonoducts, which are prevalent throughout the mantle of ripe individuals.     

Ultrastructure of oogenesis in the holopelagic polychaetes Rhynchonerella angelini and Alciopa reynaudii (Polychaeta: Alciopidae)

Ultrastructural features of oogenesis were examined in the pelagic polychaetes Rhynchonerella angelini and Alciopa reynaudii which were collected from Bahamian waters by a manned submersible during 1979 and 1980. No definitive ovary was detected in either species. Oogonia are released into the coelom as packets of cells, where they undergo mitotic division while surrounded by an envelope of sheath cells. Cytokinesis is incomplete, resulting in intercellular bridges between oogonia. Oocytes undergo early stages of meiosis characterized by the presence of synapsed chromosomes, followed by a period of rapid cytoplasmic and nuclear growth. Oocytes are released from the packets in the early vitellogenic phase into the coelom, where they undergo yolk synthesis as solitary coelomic cells. Vitellogenesis includes both autosynthetic and heterosynthetic processes. Autosynthesis involves the fusion of secretory vesicles formed by the combined activity of the rough endoplasmic reticulum and Golgi complex, with convoluted electron-dense tubular bodies of unknown origin. Heterosynthesis involves the intense uptake of exogenous precursors through endocytosis and their fusion into nascent yolk bodies which, in turn, are presumed to fuse with autosynthetically-derived yolk bodies. No nutrient stores were detected in somatic tissues. Early and middle stages of vitellogenic oocytes were absent from the coelom. This absence combined with the high level of endocytotic activity suggests that vitellogenesis occurs rapidly. These features, in combination with the presence of an exceptionally thin body wall and gut, might serve as related adaptations for predator avoidance by the maintenance of relatively low tissue-density. Alciopid, phyllodocid, and nereid polychaetes share some common reproductive features including the presence of dispersed ovaries, clusters of syncytial germ cells which undergo meiosis while enveloped by somatic cells and the release of oocytes from the clusters prior to vitellogenesis.     

Ovary structure and oogenesis in Cirratulus cirratus (Polychaeta: Cirratulidae)

Oogenesis in polychaetes has, in most species, two main phases: a coelomic phase during which vitellogenesis occurs, and an ovarian phase. Details of the latter are known in very few species. Efficient reproduction requires coordination between these two phases of oocyte differentiation. The ovaries of Cirratulus cirratus (O. F. Müller) occur 2/segment throughout the fertile region of the body; 5 main stages of oocyte differentiation can be recognised on them: stem cells, oogonia, premeiotic oocytes, early oocytes and terminal oocytes. A quantitative study of the composition of the ovary throughout the reproductive cycle shows that the ovary size is approximately constant, but that the percentage of cells in the terminal oocyte stage is reduced to zero prior to spawning. This stage in oocyte differentiation is therefore a critical one, at which oogenesis may be arrested. The ovary alternates between a proliferative phase when the terminal oocytes are present, and a non-proliferative phase when they are absent. The overall rate of germ cell production for each ovary is low, less than 1 cell/ovary/day, and the, ovary tissue turnover time is greater than 2 years. A qualitative model describing the production of oocytes is presented, and the implications of the low rate of germ cell production are discussed. A low rate of germ cell proliferation, as in Cirratulus cirratus, implies that the rate of oocyte output into the coelom will be controlled by factors influencing the later stages of ovarian oocyte differentiation.     

Oocyte development in the kuruma prawn Penaeus japonicus

Female kuruma prawns (Penaeus japonicus Bate) with undeveloped, early developing, developing, nearly ripe and ripe ovaries, were collected from Ise Bay, Japan, in 1984. Oocyte development of the kuruma prawn was classified into ten stages according to morphological characters, namely: (1) synapsis stage, (2) chromatin nucleolus stage, (3) early perinucleolus stage, (4) late perinucleolus stage, (5) oil globule Stage I, (6) oil globule Stage II, (7) yolkless stage, (8) yolk granule stage, (9) prematuration stage, and (10) maturation stage. The synapsis stage is a multiplication stage. The chromatin nucleolus stage, early and late perinucleolus stages are previtellogenesis and primary growth stages. Oil globule Stage I is an initial stage of primary vitellogenesis and secondary growth. Follicle cells on the oil globule Stage I oocytes expand rapidly and reach maximum size during oogenesis. Yolk granule stage oocytes are in the initial stages of secondary vitellogenesis. Strongly acidophilic yolk granules accumulate within basophilic vesicles of the cytoplasm. The yolk granules are first concentrated in the inner part of the cytoplasm, then gradually spread to the periphery. Cortical crypts, which are separated from the oocyte cytoplasm by the cytoplasmic membrane, are situated outside of oocyte cytoplasm. Germinal vesicle breakdown (GVBD) is initiated in the late phase of prematuration and continues until the late phase of maturation immediately prior to spawning. At the beginning of the maturation stage, the oocytes are ovulated, after which the nuclei further shrink and migrate out-wards. After ovulation, meiotic division of the ovarian oocyte progressed up to the metaphase of primary maturation division. Finally, the meiotic metaphase is visible just beneath the cytoplasmic membrane in the mature oocyte. Though ovulation is synchronous within the same ovary, GVBD is not completely synchronous. Ovulated mature oocytes have many club-shaped cortical crypts in the peripheral part of the cytoplasm and contain extensive accumulations of yolk granules dispersed throughout the cytoplasm. The apical end of the club-shaped cortical crypts and cytoplasmic membrane are coated by the vitellin envelope in the mature oocyte.     

Reproductive cycle of two populations of Ophionereis schayeri (Ophiuroidea) in New South Wales

Reproduction of two populations of Ophionereis schayeri (Müller and Troschel) in New South Wales was investigated from November 1991 through January 1993. The reproductive cycle was documented by histological examination of the gonads and by image analysis to determine the oocyte size-frequency distributions. Both populations of O. schayeri were mature in December and January and had a major summer spawning period between January and February. Thereafter, the condition of the gonads exhibited interindividual variability, with asynchronous low-intensity gamete release through August. O. schayeri has the potential to spawn for up to 8 mo of the year. This breeding pattern with synchronous spawning in summer and asynchronous gamete release through winter was similar in both populations. During autumn and winter, the gonads contained developing gametes and advanced gametes ready for spawning. The eggs spawned by O. schayeri during these seasons started their growth during the previous spring vitellogenic period, and continued to grow through summer. Upon reaching full size, they were stored for spawning outside the main breeding period. Spring is marked by increased spermatogenic and vitellogenic activity followed by maturation and spawning. Throughout their development, the oocytes of O. schayeri are surrounded by a follicle consisting of periodic acid-Schiff-positive (PAS+) haemal fluid and PAS+ yolk by the oocytes. In the testes, the haemal sinus projects into the centre of the spermatocyte columns, thus appearing to play a nutritive role in support of both oogenesis and spermatogenesis. Late vitellogenic oocytes of O. schayeri are firmly anchored to the germinal epithelium by an attachment complex consisting of specialised attachment cells and basophilic strands that radiate between the oolemma and the nucleus. Spawning is associated with rupture of the follicles, which remain as prominent, empty U-shaped profiles in the ovaries. O. schayeri produces copious numbers of 200 m-diam oocytes, suggesting that this species is a broadcast spawner and that it has a modified ophiopluteus or vitellaria larva.     

Embryogenesis and acquisition of algal symbionts by planulae of Xenia umbellata (Octocorallia: Alcyonacea)

Early embryogenesis of the internally brooding soft coral Xenia umbellata and acquisition of algal symbionts in the course of its planular ontogenesis have been examined by scanning and transmission electron microscopy and by light microscopy. The endoderm of adult X. umbellata harbours symbionts mainly in the tentacles and in the peripheral solenia system. The colonies are gonochoric brooders. Algal symbionts were never found in the sperm sacs, and were only rarely found in the follicular tissue enclosing the oocytes. Fertilized eggs pass into endodermal brood pouches where embryogenesis occurs. Cleavage is holoblastic and leads to formation of a solid blastula. Algal symbionts are conspicuously embedded in the parental mesoglea that coats the young embryo, most probably transmitted by surface adherence. At a further stage, this integument disappears and the algae reside extracellularly among the cells of the newly-formed blastula. After subsequent cell proliferation developing planulae possess an inner mass of yolk-laden cells that contain numerous symbiotic algae. Gradually the yolk disintegrates, leaving a cavity enclosed by ectoderm, a thin mesoglea and an inner endoderm with intracellular symbionts. The mature planulae have already been provided with numerous intracellular symbionts by the time they are expelled from the brood pouches. The markedly early symbiont acquisition by the embryos of X. umbellata may help support their developmental requirements in the course of planular ontogenesis.     

Route of egg yolk protein uptake in the oocytes of kuruma prawn,Penaeus japonicus

The route of egg yolk protein uptake into the oocytes of kuruma prawn, Penaeus japonicus, was studied using immunohistochemical and electron microscopical methods. Although a significant immunofluorescence with anti-vitellin-immunoglobulin was observed in the enlarged follicle cells surrounding oil globule stage oocytes of the early vitellogenic ovary, no fluorescence was detected in shrunken follicle cells surrounding oocytes in the yolk granule stage. Electron microscopically, yolk granule stage oocytes have an irregular surface with numerous well-developed microvilli. In contrast, the surface of follicle cells is relatively smooth. The irregular surface of yolk granule stage oocytes was covered with a layer of electron dense material. Similar dense material was found in the spaces between the neighboring follicle cells on the yolk granule stage oocytes. The outer surface of the follicle cells on yolk granule stage oocytes was covered by dense materials which were similar to those found on the irregular surface of oocytes. Micropinocytotic vesicles containing dense material were found in the ooplasm near the irregular surface with numerous well-developed microvilli. Dense material was concentrated in the peripheral part of the small forming yolk bodies of yolk granule stage oocytes. This suggests that the electron dense material, probably egg yolk protein, transferred to the surface of yolk granule stage oocytes from the spaces between the neighboring follicle cells may be incorporated into the ooplasm by pinocytosis through the microvilli and subsequently aggregate to form yolk bodies.     

Ovarian maturation of the multi-spawning spider crab <Emphasis Type="Italic">Maja brachydactyla</Emphasis> (Decapoda: Majidae) with special reference to yolk formation

In the study of the reproductive biology of the spider crab Maja brachydactyla, the morphology of the female reproductive system and yolk formation have long been overlooked. Females spawn two or three times during their annual reproductive cycle in northern Spain (Galicia). The ovaries consist of two lobes. The right and left lobes are connected by a small cross-lobe at the level of the heart and merge at the posterior edge. Before merging, the ovaries descend to the ventral part of the body, joining the spermathecae in the vagina, which opens through a chitin tube to the gonopore, located in the sternite, at the level of the third walking leg. No morphological changes have been observed between either the different parts of the ovaries or the different annual spawning periods. At the start of vitellogenesis, the oocyte of M. brachydactyla is characterized by a large number of vesicles in the cytoplasm. These vesicles are surrounded by a unit membrane whose size increases as the oocyte matures and contain fine granular material including a variable number of ovoid, electron-dense granules. The vesicles are of diverse origin, although most of them develop directly from the mitochondria and the Golgi complex (endogenous phase of vitellogenesis). In a subsequent phase, a series of substances (principally lipoproteins) are incorporated into the ooplasma by means of micropinocytosis. These substances are also involved in yolk formation (exogenous phase of vitellogenesis). During vitellogenesis in M. brachydactyla, mitochondria play the most important role since they are not only the energetic centre of the cellule, but they also act as containers of high-energy reserve substances: the yolk granules.     

Feeding rates of the jellyfish Aurelia aurita on fish larvae

We quantified feeding rates of field caught Aurelia aurita feeding on yolk sac cod (Gadus morhua) larvae in a series of incubation experiments. A short-time (~1 h) functional response experiment with a wide range of prey concentrations (0.5–16 prey l⁻¹, initial concentration) revealed that ingestion rates increased linearly over this range, such that clearance rates were similar between the different prey concentrations. This suggests that A. aurita is capable of efficiently utilizing dense prey patches. This indication was further supported by a linear increase of prey captured by A. aurita during 2.5 h of feeding at extremely high prey concentration (>200 prey l⁻¹). Clearance rate in darkness scaled with jellyfish diameter to a power of ~1.7 for jellyfish 3.9–9.5 cm in diameter. The jellyfish did not alter their umbrella pulse frequency in response to presence of fish larvae. There were no significant differences between A. aurita feeding rates in light and darkness for yolk sac prey ages 0–7 days (at 7.5°C). Although prey vision and escape abilities of fish may develop rapidly during early larval ontogeny, these factors apparently have little impact on interactions with predators such as A. aurita during the yolk sac stage.     

Skeleton and sclerite formation in the precious red coral Corallium rubrum

The carbonate skeleton of the gorgonian coral Corallium rubrum (L.) is composed of both a skeletal axis and numerous sclerites scattered in the mesoglea. Studies carried out on these skeletal elements and their associated tisues using microscopy and X-ray microanalysis, suggest a close relationship between the process of sclerite formation and skeletogenesis. The skeleton is surrounded by an axial epithelium composed of a single cell type. These cells associate intimately with mesogleal sclerites and scleroblasts, incorporating them into a nascent skeleton at the branch tip. Subsequent (sub-apical) growth appears to occur solely through the agency of the axis epithelial cells that serve to physically separate mesogleal sclerites and scleroblasts from contact with the axis. The epithelium is associated with the production of layered calcite crystals and irregular protuberances that constitute the mature, calcareous skeleton. Free sclerites in the mesoglea appear to be the product of multiple cells that are cytologically indistinguishable from those in the axis epithelium. Like the axis, sclerites are produced as layers of calcite crystals with irregular protuberances. The protuberances differ only slightly from those of the axis, and the skeleton is mineralogically indistinguishable from the sclerites. Thus, the skeleton of red coral is not primarily the product of fused sclerites. Instead, we suggest that the axis epithelium treats the incipient skeleton as if it were the core of a single sclerite, and conversely, that the mesogleal scleroblasts of C. rubrum constitute a fragmented axis epithelium.     

Sun-compass migration by Aurelia aurita (Scyphozoa): population retention and reproduction in Saanich Inlet,British Columbia

The scyphomedusa Aurelia aurita in Saanich Inlet, a north-south oriented fjord in British Columbia, uses a sun compass to migrate in a southeasterly direction during the day. When the sky is overcast and at night, A. aurita orients randomly and is dispersed passively by gentle tidal currents. The net result is daily reaggregation of medusae into enormous swarms along the southeastern shore of the fjord. Observations of spawning A. aurita in these swarms suggest that sun-compass migration and aggregative behavior may have evolved to facilitate reproduction and to maintain the population within this fjord throughout the year.     

Role of ovarian follicle cells in vitellogenesis and oocyte resorption inCapitella sp. I (Polychaeta)

Capitella sp. I has a short generation time, is polytelic and produces broods of relatively large, yolky eggs at intervals of as short as 5–7 d at 20°C. The follicle cells surrounding vitellogenic oocytes in the medial portion of the ovary contain large nuclei, extensive arrays of rough endoplasmic reticulum and Golgi complexes. The coelomic surfaces of follicle cells in the outer layer are covered by blade-like microvilli, which project into the coelom. In the dorsal and lateral portions of the ovary, where mature oocytes are found, the follicle cells are covered by flat, stellate-shaped, peritoneal cells that resemble the podocytes of many invertebrate excretory organs and of the mammalian glomerulus. Changes occurring in the ovary during resorption of oocytes in the middle to advanced stages of vitellogenesis are described. Oogonia and previtellogenic oocytes are not resorbed and perhaps subsequently undergo vitellogenesis. The ovary in this highly opportunistic polychaete species may play a significant regulatory role in the control of reproduction by producing yolk precursors, regulating their uptake from the coelomic fluid via the activity of the follicle cells and by resorbing mature oocytes in response to stress.     

Responses of ovaries and testes of Lytechinus variegatus (Echinodermata: Echinoidea) to dietary administration of estradiol, progesterone and testosterone

 In many vertebrates, environmental factors influence gamete differentiation and growth of the mature gonad through alteration of sex steroid production or action; however, it is unclear how gamete differentiation and gonadal growth are regulated in echinoids. The purpose of this study was to examine the influence of dietary administration of estradiol (E2), progesterone (P4), testosterone (T) and finasteride (F, a 5α-reductase inhibitor) on the ovaries and testes of mature Lytechinus variegatus (Lamarck) during gonadal growth. Echinoids were fed a formulated diet supplemented with steroids or steroids in combination with finasteride for 36 d. The effects of dietary administration of steroids on L. variegatus were both steroid- and sex-specific. The mean ovary index was 54% greater in individuals fed E2 than from individuals fed the control (C) diet (10.0 ± 1.1 vs 6.5 ± 0.7, respectively; P < 0.05). Individuals fed E2, P4, E2/P4, and P4 in combination with F had significantly smaller oocytes (P < 0.005) than individuals fed C. The volume fraction occupied by nutritive phagocytes in ovarian tubules from individuals fed E2/P4 or P4 were significantly larger than the volume fraction occupied by nutritive phagocytes from individuals fed C, indicating that E2 and 5α-reduced progestins may promote nutrient allocation to nutritive phagocytes. Although oocytes from the individuals fed T alone were significantly smaller than those fed T in combination with F, oocytes from individuals in both treatments were significantly larger compared to oocytes from individuals fed C. These data suggest that upon removal of 5α-reduced androgens, T is able to promote an increase in oocyte diameters. In contrast, the mean testis index was 56% greater in individuals fed P4 than in individuals fed C (8.9 ± 0.6 vs 5.7 ± 0.9, respectively; P < 0.05); the testis index did not increase in individuals fed P4/F, suggesting that 5α-reduced progestins stimulate testicular growth. Testes growth in the presence of 5α-reduced progestins was accomplished by significant increases in the volume fraction occupied by nutritive phagocytes and by a significant reduction in the volume fraction occupied by spermatogenic columns in testicular tubules. These data further indicate that 5α-reduced progestins (or 5α-reduced androgens) may inhibit spermatogenic column formation. In conclusion, E2 stimulated ovarian growth but inhibited oocyte growth, whereas T had no affect on ovarian growth but promoted oocyte growth in L. variegatus. We hypothesize that the E2 (or E2 metabolites) and/or 5α-reduced androgens in combination with T regulate oocyte growth in the echinoid L. variegatus. In addition, 5α-reduced progestins promoted nutrient accumulation in nutritive phagocytes within the ovaries and the testes. Furthermore, 5α-reduced progestins stimulated growth of the testes and inhibited spermatogenic column formation, suggesting that 5α-reduced progestins regulate nutrient accumulation into nutritive phagocytes and spermatogenic column formation in L. variegatus. The differences in estrogen effects between echinoids and asteroids may be related to differences in gonad morphology and, ultimately, the differences in cellular signalling pathways (paracrine vs endocrine). Received: 22 May 1999 / Accepted: 24 May 2000     

Ultrastructure of the gonad and gametogenesis in the eastern oyster,Crassostrea virginica. I. Ovary and oogenesis

The ultrastructural features of the ovary and oogenesis are described in the eastern oyster,Crassostrea virginica (Gmelin, 1791). The ovary is a diffuse organ consisting of highly branching acini in which oocytes develop. The acini are surrounded by a matrix of vesicular connective tissue (VCT tells) which serves a nutrient storage function. Each acinus is bathed by fluid within a surrounding connective tissue compartment, the hemocoel, which likely serves as a means of transporting nutrients to the oocytes. Oocytes begin growth while positioned near to the inner acinus wall. As differentiation proceeds and they enter the late stages of vitellogenesis, they become stalle-shaped and project into the acinus lumen. Follicle tells are closely associated with oocytes during the early and middle stages of vitellogenesis but they are largely confined to the basal, stalked region of late-stage oocytes. Vitellogenesis occurs through a process of autosynthesis, involving the combined activity of the Golgi complex and rough endoplasmic reticulum, and heterosynthesis in which extraovarian precursors are incorporated into oocytes via receptor-mediated endocytosis involving the basal surface of the oocytes. It is suggested that the follicle tells play some important role during oogenesis but probably are not the major source of yolk precursors. The VCT celas are probably the main source of nutrients for vitellogenesis.     

Functional morphology of the gonads of the articulate brachiopodTerebratulina retusa

The reproductive anatomy, and ultrastructural features of the gonads of the articulate brachiopodTerebratulina retusa (Linnaeus), are documented based on collections made between October 1985 and October 1986 from the Firth of Lorn, west coast of Scotland. This species is dioecious, and maturity is achieved in both sexes at shell lengths greater than ~5.5 mm. There is no obvious external sexual dimorphism except for slight differences in the coloration of the gonads; testes are white/cream, ovaries are yellow/orange. The gonads occur as four palmate lobes, a pair in each valve. Gonads are formed within a mantle sinus (vascula genitalia), which is an anterior extension of the coelom, that opens posteriorly into the visceral cavity and to the exterior via a pair of metanephridia. The latter serve as gonoducts during spawning. Gametes are borne on genital lamellae formed from a reticulate lattice of connective tissue. The lamellae are an extension of the ileoparietal band and are fused along one margin to the inner mantle membrane. Developing oocytes are closely affixed to the genital lamellae and originate from a pool of proliferating germ cells at its base. Vitellogenic oocytes that are at an advanced stage are released from the genital lamellae, but are retained within thevascula genitalia. Liberated oocytes continue to accumulate yolk and eventually occlude thevascula genitalia, before being spawned. Coelomocytes were identified within the gonads. In spent females these cells appear to be phagocytic and involved in the resorption of necrotic material, while in the male they may serve as nutritive cells.     

Gonadal morphology and gametogenesis in the sea pen Pennatula aculeata (Anthozoa: Pennatulacea) from the Gulf of Maine

The ultrastructural features of gametogenesis are described in male and female colonies of the sea pen Pennatula aculeata. Specimens were collected for observation and fixation at 113 to 231 m depth in the Gulf of Maine, USA, in August 1993. The species is gonochoric, and all stages of gametogenesis are observed in both male and female colonies >45 mm in height. Gametogenesis shows several features that differ from sea anemones. The developing oocytes and sperm cysts are completely encompassed by gastrodermally derived follicle cells, and they are released from the mesenteries into the coelenteron before they are fully differentiated. Following maturation in the coelenteron, the eggs and intact sperm cysts are expelled through the mouths of the autozoids during spawning. The expulsion of sperm cysts suggests that they function as primitive spermatophores, perhaps as a way of reducing sperm dilution. Vitellogenesis results in the biosynthesis of lipid droplets which are the sole nutrient reserves in the egg. Heterosynthetic vitellogenesis is characterized by the importation of lipid precursors into the oocyte, and there is some indirect evidence that hypertrophic follicle cells play a role in production, transport, and/or mediation of these precursors. Spermatogenesis is similar to that of other anthozoans. The spermatozoon has a cone-shaped head, a posterior nuclear fossa, a ring of lipid-like bodies in the midpiece, a prominent cytoplasmic collar surrounding the proximal flagellum, and a single mitochondrion, but the posterior region of the sperm also contains previously undescribed concentric rings of cisternae resembling smooth endoplasmic reticulum. Received: 23 March 1998 / Accepted: 31 July 1998     

Quantitative and ultrastructural studies on the reproductive biology of the polychaete Pholoe minuta in Galway Bay

Previous works by the present authors have identified, in a qualitative fashion, the reproductive cycle of Pholoe minuta (Fabricius) in inner Galway (53°13.5N; 9°7W). This quantitative, histological study, carried out over the period late May 1981 to late April 1982, shows that P. minuta is polytelic (two to three annual spawnings in an individual's lifespan). Individuals become sexually mature for the first time when approaching 3 yr of age. Estimates were made of the size of the reproductively active population (47 to 49%), sex ratio (1:1), duration (10 to 11 mo) and pattern (unimodal) of gametogenesis and fecundity levels (210 198 eggs m^-2). Maturation (mean oocyte size=85 m) and spawning (for 1 wk during late March through early April) were shown to be synchronised. Throughout oogenesis, follicle cells play an important role in binding together oocyte clusters, and desmosomal contacts are evident between oocyte and follicle cell plasmalemmas. No evidence for the synthesis of yolk precursors in the follicle cell cytoplasm was discovered. Oocytes are rich in Golgi apparatuses (yolk synthesising) and nuclear pores are numerous. Rough endoplasmic reticulum is distended in nature. Egg envelope development does not appear to commence until after the onset of vitellogenesis. Mature spermatozoa are of the primitive bullet-shaped type, with an unmodified acrosome. A possible method of elevation membrane formation is described.     

Oogenesis and spawn-formation in the pigmy lion fish Dendrochirus brachypterus (Pteroidae)

Dendrochirus brachypterus possesses a special type of ovary in which the germinative epithelium surrounds a spongy stroma hanging in the lumen. The inner side of the ovarian wall is covered by a special type of epithelium that produces a gelatinous envelope around the gametes (spawn) during reproduction. In general, oogenesis in the lion fish resembles that of other teleosts, the characteristic difference being in the formation of ovigerous peduncles-protrusions of the spongy stroma, each of which bears a single oocyte. During oogenesis the oocytes rise above the germinal epithelium, enveloped in their follicular epithelia. The larger the oocyte the greater the length of its peduncles and the further its position from the germinal layer (Fig. 3). During this phase of development, a marked multiplication of yolk granules occurs; they increase their diameter and finally homogenize at ovulation. Under favorable thermal conditions, oogenesis continues all year round. The peduncular growth of oocytes enables the growing egg cells to move away from the egg-producing surface, thus making space available for the ovary to produce more eggs. This ability should have a survival value for these fish if we take into consideration that the most vulnerable stage in their life history is that of planktonic larvae (Fishelson, 1975).     

Spawning pattern and type of fecundity in relation to ovarian allometry in the round herring <Emphasis Type="Italic">Etrumeus teres</Emphasis>

Spawning pattern (assessed by seasonal changes in ovarian developmental stages) and type of fecundity (assessed by analysis of oocyte-size frequency distributions) of the round herring Etrumeus teres were studied in relation to ovarian growth and seasonal changes in the gonadosomatic (GSI), hepatosomatic (HSI) and liposomatic (LSI) index as well as the somatic condition of spawners (CS) in a spawning ground of southern Japan. Except for summer, mature and recently spawned ovaries occurred all year round. Oogonia and primary oocytes were present in all ovaries, and cortical alveoli stage (CA) oocytes occurred in all mature, hydrated and partially spent (PS) females (PS: females containing post-ovulatory follicles). Before hydration, a clutch of larger yolked oocytes, undergoing synchronous growth (range 0.7–1.1 mm), was present in mature ovaries which was completely separated from a more heterogeneous clutch of oogonia, primary and secondary oocytes (<0.150 mm) and oocytes in the CA stage (range 0.15–0.60 mm). As vitellogenesis progressed, the yolked clutch increased in size but the CA oocytes remained arrested. The latter entered into the secondary growth phase when hydration started in the advanced batch. Ovarian growth was isometric in all developmental stages, validating the use of GSI, which showed a consistent monthly evolution among years. Spawning stopped in summer (July and August) and peaked in winter and spring. HSI correlated positively with GSI on both a monthly mean basis (r = 0.76) and individual fish basis (liver weight explained 67–83% of the variability in ovary weight when females were grouped into 1-unit GSI intervals) suggesting a significant role of liver in vitellogenesis. LSI and CS also showed marked seasonal changes peaking from summer to middle autumn. Overall results suggest that E. teres is a multiple spawner with a group-synchronous ovarian development and indeterminate annual fecundity, with the three processes linked to an isometric growth of the ovary. We propose that such a reproductive pattern is an adaptation to produce batches of large pelagic eggs through a protracted spawning season.