M. Le Pennec

Utrastructural study of oogenesis and oocytic degeneration in Pecten maximus from the Bay of St. Brieuc

An ultrastructural study of oocytic development enabled the identification of changes occurring during oogenesis in Pecten maximus collected from the Bay of St. Brieuc, France, in 1987. “Auxiliary cells”, closely associated with developing oocytes were observed. Each oocyte seems to be associated with only one secretory cell, which is characterised by an abundant rough endoplasmic reticulum at the onset of vitellogenesis. Contact between this cell and a developing oocyte is maintained by a desmosome-like junction which can be observed when the vitelline coat is formed. These “auxiliary cells” seem to play a trophic role in vitellogenesis, and may be involved in the formation of the vitelline coat of the oocytes. Oocytic degeneration is discussed in detail; in this species, it is a continuous phenomenon of varying intensity throughout the year. The ultrastructural changes resulting in lysis of the oocyte are described, and the evolution of atretic oocytes is examined.

Ultrastructure of the gill of the hydrothermal-vent mytilid Bathymodiolus sp.

Specimens of Bathymodiolus sp. were collected at 2 620 m depth during the “Biocyarise” Cruise (12°58′80″N; 13°56′60″W) in March 1984, and samples of the gill were fixed for ultrastructural observations. The gill of this hydrothermal-vent mytilid constitutes the main organ in the nutritional processes. The lamellae display abundant ciliation, normally comprised of frontal cilia, compound latero-frontal cirri and lateral cilia. At the ventral margin of each demi-branch, a longitudinal, ciliated, feeding groove is present. the lamellae are composed of numerous homorhabdic filaments connected by tufts of cilia. Each filament is made of a thin wall overlying a central lumen containing amoebocytes. Ultrastructural observations revealed the filament wall to be composed of four types of cells: (1) The ciliated cells of the frontal, latero-frontal and lateral ciliation, characterized by an abundance of mitochondria. (2) Mucous cells present to some degree among the ciliated cells, but more abundant on the distal edge and containing dense droplets of mucus. (3) Cells colonized at their apical pole by numerous bacteria enclosed in membrane-delimited clear spaces and composing the major part of the filament wall. (4) Thin ciliated cells separating the bacterial cells and characterised by a dense fringe of microvilli at their apical pole. The lumen of the filament contains amoebocytes of different morphological aspects which seem to accumulate electron-dense granules, possibly related to detoxification processes.

The pelagic life of the pectinid Pecten maximus—a review

Although marine bivalves go through a planktonic larval phase, knowledge of this phase is often poor because of the difficulty in identifying and monitoring the activity of these tiny organisms within the water mass. Some bivalves have been studied more than others, often because of their economic value. These species can serve as a model for improving our understanding of the circumstances of planktonic larval development, in particular by assessing its impact on recruitment. This review is aimed at summarizing the knowledge on the pelagic life of Pecten maximus acquired by research in France over the last 25 years. The comparison of these results with those obtained elsewhere for pectinid species, indicates certain characteristics that appear to be fairly common to all coastal infralittoral pectinids. Regardless of species, pelagic life of pectinids starts with the release of gametes, followed by fertilization and embryonic and larval stages until metamorphosis and recruitment to the benthic community. After consideration of the main characteristics of the pelagic stages and their sensitivity to certain environmental factors, a graphic synthesis is provided displaying their migratory behaviour and the possible consequences for recruitment.

Mode of particle ingestion in five species of suspension-feeding bivalve molluscs

In order to elucidate the mode of particle ingestion and the functional anatomy of the oesophagus in bivalves, a histological study was performed onMytilus edulis (Mytilidae),Crassostrea virginica (Ostreidae),Placopecten magellanicus, Chlamys varia, and juvenilePecten maximus (Pectinidae). Specimens were sampled from various sites in New Brunswick, Canada, and Brittany, France, from 1987 to 1989. The buccal, peribuccal, and oesophageal epithelia of all species contained a dense distribution of actively secreting mucocytes, although these were somewhat less abundant inCrassostrea virginica, which also has the shortest oesophagus. Mucocyte morphology, while constant within a family (Pectinidae), showed clear differences among families. Both acid and neutral mucopolysaccharides were secreted by the epithelial mucocytes of all species. Mucus and mucus-particle masses were observed in the peribuccal and buccal regions, as well as in the oesophageal lumina of all species, even in those specimens which had been maintained without feeding (Placopecten magellanicus) or held out of water for 48 h (C. virginica) prior to dissection and fixation. These results indicate that a basal level of mucus production and transport is continuous on the peribuccal, buccal, and oesophageal ciliated epithelia, regardless of the particle concentration in the external medium. Buccooesophageal glands, generally thought to be absent in the Bivalvia, were observed in one of the species examined (M. edulis). It is concluded that the mode of particle ingestion in these suspension-feeding bivalves is via ciliatransported mucus masses; the presence of buccooesophageal glands inM. edulis suggests a digestive role for the oesophagus in this species.

Chemoautotrophic symbionts and translocation of fixed carbon from bacteria to host tissues in the littoral bivalve Loripes lucinalis (Lucinidae)

Specimens of Loripes lucinalis (Lucinidae) living in reducing sediments were collected near a sewage outfall at low tide on the Moulin Blanc beach, Brest, France, from January to March 1987. Electron microscope studies revealed numerous Gram-negative-type bacteria in the gill cells. Ribulosebiphosphate carboxylase, a diagnostic enzyme of the Calvin-Benson cycle of CO2-fixation was measured only in the gill extracts. Various tissues of L. lucinalis were examined for activity of APS reductase, (EC 1.8.99.2), ATP sulphurylase (EC 2.7.7.4) and rhodanese (EC 2.8.1.1), enzymes involved in sulphide oxidation. APS reductase was only found in symbiont-containing tissues, i.e., gills. These enzymatic studies characterise the symbionts as chemoautotrophic sulphide-oxidizing bacteria. Histoautoradiography demonstrated that part of the carbon dioxide fixed by symbiotic bacteria in the gills is translocated to symbiont-free tissues of the bivalve. The ultrastructure of the gill is detailed and a nomenclature based on established and new terminology is proposed to describe the various cellular types comprising the gill filament.

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The internal anatomy and microanatomy of the gill of Placopecten magellanicus Gmelin collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada, was studied using thin-section light microscopy and transmission electron microscopy. Most of the spurs show no evidence of organic union, and hence do not participate in vascular exchange. However, the dorsal bend shows both ciliary and organic interfilamentar union. The internal structure and the hemocytes of the dorsal respiratory expansion are presented. The epithelium consists of three distinct cell types, bounded by apical microvilli. All regions of the gill contain an epithelial basal membrane, which is greatly convoluted in the interconnecting vessels of the dorsal respiratory expansion. The significance of these observations is discussed in relation to possible roles in respiration, transmembrane transport and nutrition. The apical surface of all ciliated cells is covered with an acellular matrix composed of clear spherical vesicles, which may serve a mechanical function for which mucus would be unsuited. The significance of the abundance of mucocytes on the abfrontal surface of the principal filaments is discussed in terms of the escape response of pectinid bivalves.

Early development of the gill and implications for feeding in Pecten maximus (Bivalvia: Pectinidae)

Despite the importance of understanding feeding in the early stages of bivalve development, little information is available concerning the organogenesis of the bivalve gill. The present study used histological and scanning electron microscopical techniques to present a detailed account of gill development in the early stages of the scallop Pecten maximus L. (Bivalvia: Pectinidae). Live specimens from larval cultures were observed daily using light microscopy, while five scallops were sampled for electron and light microscopy every 2 to 3 d from Day 18 to 35, then weekly to Day 56, with a final sampling on Day 58. Although development was continuous, four distinct stages were identified (1-primordia, 2-homorhabdic unreflected, 3-homorhabdic reflected, 4-heterorhabdic), partially recapitulating the presumed phylogenetic evolution of this character in the Pectinidae. The absence of a ventral grcove in all stages suggests that the particle transport mechanism of pectinids evolved independently of such a structure, which is found in other bivalve families. Similarly, the absence of latero-frontal cilia in all specimens up to the largest observed (4 mm) indicates that the single row found in adults is a later development, rather than a vestige of a more abundant ciliation in ancestral forms. The anatomical data, together with in vivo observations of feeding in postlarvae, suggest that the developmental stages of the P. maximus gill correspond to critical changes in gill function. The early life of P. maximus may thus be characterized by distinct functional changes in feeding.

Survival of Nucella lapillus in a tributyltin-polluted area in west Brittany : a further example of a male genital defect (Dumpton syndrome) favouring survival

A male genital defect was reported in a Nucella lapillus (L.) population at Dumpton Gap (England). This defect was termed “Dumpton syndrome” (DS) and appears to be a genetic feature. Its main characteristic is the absence of penis (aphally) in males and in females. In 1992, such a phenomenon was discovered in populations in the vicinity of Brest (Brittany, France). DS-affected females exhibit fewer tributyltin-induced imposex characteristics than expected in normal individuals. The percentage of female sterilization is thus lower, favouring population survival. In consequence, the DS is considered to be a “pollution-resistance” feature. Comparison with the Dumpton population revealed similarities and differences in the DS characteristics. It is thus hypothesized that the DS observed at the two locations is due to two different biological mechanisms. Indeed, aphallic males with a split prostate were observed but no underdevelopment of their vas deferens and testis was noted in the present study. In this gonochoristic gastropod species, the most DS-affected males in Brest possess an ovotestis and it is thus hypothesized that feminity is remnant in N. lapillus. Incidence of abnormality is ten times higher in females than in males. This suggested that a sex-difference operates in the Brest phenomenon.

Peribuccal organs ofPlacopecten magellanicus andChlamys varia (Mollusca: Bivalvia): Structure, ultrastructure and implications for feeding

In order to better understand the structure of bivalve peribuccal organs and relate this to existing functional paradigms of their role in feeding, the labial palps of two scallop species,Placopecten magellanicus from the Bay of Fundy, Canada (1985 and 1986), andChlamys varia from the Bay of Brest, France (1986), were examined using histological techniques and electron microscopy. The ridged palp surface displays a uniformly dense ciliation with relatively few mucocytes; these are essentially concentrated in the region of the secondary ledge and may, through their secretory activity, determine the fate of particle masses in this area. The mucus secretions of the ridged palp surface are qualitatively different from those of the smooth palp surface. Mucocytes are much more abundant on the smooth palp surface, where it is suggested that their homogeneous secretions attenuate the potentially adverse effects of anteriorly-directed cleansing and swimming currents. Two other cell types are found in the palp epithelia: ciliated cells, which are very numerous on the ridged surface and relatively rare on the smooth surface, and non-ciliated epithelial cells, which are very numerous on the smooth surface and rare on the ridged surface, where they are confined to the palp margin. In addition to the mechanical role of the ciliated cells and mucocytes, the ultrastructural characteristics of the ciliated and non-ciliated epithelial cells indicate a dichotomy of function between the ridged and smooth surfaces. The ridged surface epithelial cells present an ultrastructural specialization in the absorption of dissolved and colloidal matter, suggesting an accessory nutritive role, whereas the smooth surface simple epithelial cells show signs of active molecular synthesis. No specialised sensory cells were observed on the ridged surface; it is therefore not yet possible to conclude whether the labial palps are capable of selection based on individual particle characteristics.

Association between the mollusc bivalve Loripes lucinalis and a Chlamydia-like organism, with comments on its pathogenic impact, life cycle and possible mode of transmission

Loripes lucinalis is a littoral bivalve which has already been confirmed to harbour endo-cellular sulfur-oxidizing bacteria within its gills. Examination of the digestive gland of L. lucinalis collected from the Moulin Blanc Beach in the Bay of Brest (Brittany, France) revealed the existence of an additional association involving a Chlamydia-like organism. Three different forms of Chlamydia-like bacteria were observed: reticulate rod-shaped cells, electron-dense cells and enlarged cells. The reticulate rod-shaped cells and the electron-dense bodies are thought to represent the germinal initial body and infectious form of the bacteria, respectively. The enlarged cells were always associated with what are believed to be spherical or icosahedral phages. Initial infestation seems to occur by phagocytosis at the apical pole of the digestive cells of the tubule and duct epithelia. Within the host cell, the bacteria undergo binary fission and budding, forming an inclusion which gradually fills up the cell. Inclusions are generally between 15 and 30 μm in size, and > 85% of all individuals examined possessed inclusion bodies. The level of infestation varied between individuals, some being heavily colonized, but did not seem to be related to season. Histological and ultrastructural observations suggest that, once developed, the colony has three possible fates: (1) the cells will degenerate due to phage infection; (2) colony overcrowding will occur, causing the development of electron-dense bodies that will be released into the lumen; (3) the entire membrane-bound inclusion will be released into the lumen and subsequently into the pallial cavity. Inclusions within the pallial cavity may be ingested by the host or may even be phagocytized by bacteriocyte cells of the gill. It is proposed that this association could be a form of symbiosis and that L. lucinalis may, therefore, be a rare example of an organism adapted to harbour two very different symbioses.