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

The organization, general anatomy, and surface microanatomy of all regions of the gills of a representive bivalve mollusc, Placopecten magellanicus Gmelin, were studied using stereo-microscopic, histological, and scanning electron microscopic techniques. Individuals were collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada. In addition to correcting earlier accounts of this structure, a number of new observations are reported. The orientation of the ciliated spurs appears to be responsible for the sinusoidal arrangement of the gill filaments. Micrographs showing the structure of the dorsal respiratory expansion are presented. The entire abfrontal surface of the principal filament, including the dorsal respiratory expansion, is densely ciliated and mucosecretory. These characteristics may aid in the establishment of a respiratory current and in the prevention of gill damage during escape responses. All nonciliated regions of the gill filaments are covered with microvilli, thus greatly increasing the surface area of the gill. The feeding mechanism is discussed in relation to the dorsal and ventral ciliated tracts. Symbiotic ciliate protozoans are constantly dislodged from the gill filaments and transported via the ventral mucus string to the buccal region. The nutritional implications of these observations are discussed.     

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.     

Digestion in a subtidal population of Mercenaria mercenaria (Bivalvia)

The digestive processes of Mercenaria mercenaria (Linn.) were studied in a natural subtidal population. Hourly sampling of 4 to 5 quahogs was conducted over one 25 h period in August, 1978, at Woods Hole, Ma., USA. Crystalline style length did not vary significantly with time, tide, particulate C, N or suspended material in the water. Changes in secretory activity of the style sac epithelium were not evident from histological sections. Based on cellular morphological differences, digestive tubules were individually classified into one of 4 categories, indicative of the state of intracellular digestion within the digestive cells. All 4 tubule types were present in individuals from each sample hour, signifying that intracellular digestion occurs continually within the digestive gland. A 3 h time lag is evident between times of peak C, N or suspended materials and apparent peak levels of absorptive (Type II) tubules.Contribution No. 76 Marine Science Institute, Northeastern University; Nahant, MA 01908, USA     

Phenotypic variations in the gills of the symbiont-containing bivalve Lucinoma aequizonata

The marine bivalve Lucinoma aequizonata (Lucinidae) maintains a population of sulfide-oxidizing chemoautotrophic bacteria in its gill tissue. These are housed in large numbers intracellularly in specialized host cells, termed bacteriocytes. In a natural population of L. aequizonata, striking variations of the gill colors occur, ranging from yellow to grey, brown and black. The aim of the present study was to investigate how this phenomenon relates to the physiology and numbers of the symbiont population. Our results show that in aquarium-maintained animals, black gills contained fewer numbers of bacteria as well as lower concentrations of sulfur and total protein. Nitrate respiration was stimulated by sulfide (but not by thiosulfate) 33-fold in homogenates of black gills and threefold in yellow gill homogenates. The total rates of sulfide-stimulated nitrate respiration were the same. Oxygen respiration could be measured in animals with yellow gills but not in animals with black gills. The cumulative data suggest that black-gilled clams maintained in the aquarium represent a starvation state. When collected from their natural habitat black gills contain the same number of bacteria as yellow gills. Also, no significant difference in glycogen concentrations of the host tissues was observed. Therefore, starvation is unlikely the cause of black gill color in a natural population. Alternative sources of nutrition to sulfur-based metabolism are discussed. Denaturing gradient gel electrophoresis (DGGE) performed on the different gill tissues, as well as on isolated symbionts, resulted in a single gill symbiont amplification product, the sequence of which is identical to published data. These findings provide molecular evidence that one dominant phylotype is present in the morphologically different gill tissues. Nevertheless, the presence of other phylotypes cannot formally be excluded. The implications of this study are that the gill of L. aequizonata is a highly dynamic organ which lends itself to more detailed studies regarding the molecular and cellular processes underlying nutrient transfer, regulation of bacterial numbers and host–symbiont communication. Received: 1 September 1999 / Accepted: 1 February 2000     

Variability of the frequency of male neoteny in Ostrea puelchana (Mollusca: Bivalvia)

Ostrea puelchana (D'Orbigny), a larviparous oyster from the Atlantic coast of South America, comprises two types of individuals: (1) adults which live for several years and become sexually developed after one year, these show a rhythmic successive hermaphroditism with male, hermaphroditic and female phases; (2) neotenic individuals settled on the adults, which live for only about one year and function exclusively as males. Adult oysters were collected in February 1982 from a natural bed in the San Matias Gulf, Argentina, and were transferred to two sites 200 km to the south of their original habitat, where they were maintained under culture conditions. At the beginning of the study, 17% of adult oysters were observed bearing neotenic individuals. After two years, the number of adults in the female phase had decreased, and neotenic individuals had entirely disappeared. The presence of female-phase adults would seem to be necessary for the existence of neotenic males. Further analysis of the results revealed high adult density to be directly associated with greatly decreased numbers of adults in the female phase; this may play a role in the decrease of neotenic individuals.     

On the functional morphology of Hiatella solida (Hiatellidae: Bivalvia)

Hiatella solida (Sowerby, 1834) is the only species of the genus Hiatella represented in Southern Brazilian waters. It occurs on the coast of São Paulo and Rio Grande do Sul. The bivalves were found nestling or attached by byssus threads to specimens of the sea squirt Polyandrocarpa zorritensis or between the tubes of the polychaete Phragmatopoma lapidosa in the intertidal zone. The structure, adaptation and functional morphology were studied in detail, including the functioning of the stomach. H. solida were found with mature ova in August.     

The behaviour of giant clams (Bivalvia: Cardiidae: Tridacninae)

Giant clams, the largest living bivalves, live in close association with coral reefs throughout the Indo-Pacific. These iconic invertebrates perform numerous important ecological roles as well as serve as flagship species—drawing attention to the ongoing destruction of coral reefs and their associated biodiversity. To date, no review of giant clams has focussed on their behaviour, yet this component of their autecology is critical to their life history and hence conservation. Almost 100 articles published between 1865 and 2014 include behavioural observations, and these have been collated and synthesised into five sections: spawning, locomotion, feeding, anti-predation, and stress responses. Even though the exact cues for spawning in the wild have yet to be elucidated, giant clams appear to display diel and lunar periodicities in reproduction, and for some species, peak breeding seasons have been established. Perhaps surprisingly, giant clams have considerable mobility, ranging from swimming and gliding as larvae to crawling in juveniles and adults. Chemotaxis and geotaxis have been established, but giant clams are not phototactic. At least one species exhibits clumping behaviour, which may enhance physical stabilisation, facilitate reproduction, or provide protection from predators. Giant clams undergo several shifts in their mode of acquiring nutrition; starting with a lecithotrophic and planktotrophic diet as larvae, switching to pedal feeding after metamorphosis followed by the transition to a dual mode of filter feeding and phototrophy once symbiosis with zooxanthellae (Symbiodinium spp.) is established. Because of their shell weight and/or byssal attachment, adult giant clams are unable to escape rapidly from threats using locomotion. Instead, they exhibit a suite of visually mediated anti-predation behaviours that include sudden contraction of the mantle, valve adduction, and squirting of water. Knowledge on the behaviour of giant clams will benefit conservation and restocking efforts and help fine-tune mariculture techniques. Understanding the repertoire of giant clam behaviours will also facilitate the prediction of threshold levels for sustainable exploitation as well as recovery rates of depleted clam populations.     

An ultrastructural study of Crassostrea angulata (Mollusca,Bivalvia) spermatogenesis

The cytoplasm of Crassostrea angulata (collected from the Atlantic coast of Northern Spain, Galiza, between 1992 and 1993) spermatogonia contains a few proacrosomal dense vesicles and intermitochondrial round granulo-fibrillar dense bodies. In primary spermatocytes, large cytoplasmic spaces appear occupied by large round and more scattered granulo-fibrillar dense bodies which are intermingled with cord-like dense structures and are in close association with mitochondria and the rough endoplasmic reticulum. As a result of Golgian activity, the number of dense proacrosomal vesicles increases and a flagellum arises from the centrioles. During the second meiotic division, several round dense bodies surrounding cord-like dense structures as well as dense proacrosomal vesicles, were observed between chromosomes. During spermiogenesis, no Golgian activity was noticed, and only the rough endoplasmic reticulum was observed in close association with proacrosomal vesicles. Similar results were observed for Ostrea edulis (collected from the Atlantic coast of southern Portugal, Algarve, between 1992 and 1993). Spermatozoa of both species are of the aquasperm type, and although their ultrastructure is quite similar, silver and phosphotungstic acid-chromic acid stainings enabled us to clearly distinguish both species at the ultrastructural level. The different morphological aspects of spermatogenesis in the Pteriomorphia and the use of silver and PTA stainings in the Mollusca are reviewed and discussed.     

Protein kinase C and ion transport in the posterior gills of the Chinese crabEriocheir sinensis

The possible involvement of protein kinase C in control of ion transport was investigated on a preparation of isolated, perfused posterior gills of the Chinese crabsEriocheir sinensis (collected in 1989 from lakes near Emden, northern Germany) acclimated to fresh water. 1-oleyl-2-acetyl-sn-glycerol (OAG) and phorbol 12-myristate 13-acetate (PMA), two activators of protein kinase C, when added to the perfusion saline, induced depolarisation of the transepithelial potential difference (PD) and an increase in transepithelial Na⁺ influx. The observed increase was proportional to OAG concentration up to 250µM, with a 2.5× accelerated Na⁺ influx. OAG and PMA remained without effect on Cl^– fluxes. The observed effects were in agreement with an activation, via protein kinase C, of the Na⁺/K⁺ ATPase located on the serosal side of the epithelium.     

Functional and environmental anaerobiosis in the razor-clam Ensis directus (Mollusca: Bivalvia)

In Ensis directus (Conrad) metabolic changes accompanying muscular activity and experimental anoxia were investigated. Octopine dehydrogenase, strombine dehydrogenase, alanopine dehydrogenase and lactic dehydrogenase were shown to be present in different parts of the body, but the activity of ODH always exceeded that of the other enzymes by far. Muscular activity connected with escape digging or evoked by electrical stimulation resulted in a decrease of stored phosphagen and in an accumulation of octopine. During recovery the normal concentrations of energy-rich phosphates were restored quickly, whereas octopine was removed more slowly. In experimental anoxia only minor quantities of L-and D-alanine, succinate, propionate, and acetate were formed. Quantitatively the most important endproduct was strombine, which rose linearly with the duration of anoxic incubation.     

Endogenous circatidal rhythm in the Manila clam Ruditapes philippinarum (Bivalvia: Veneridae)

Manila clams, Ruditapes philippinarum, removed from their natural environment and maintained for 9 weeks in continuously immersed conditions exhibited a clear endogenous circatidal rhythm in oxygen consumption. The clams exhibited a semidiurnal rhythmicity in oxygen consumption after showing a diurnal pattern in the first few days (5 to 7 d) of the experiment. The results of the present study indicate that activity rhythms of clams are controlled not only by exogenous factors, but also by an endogenous circatidal periodicity. Received: 8 April 1998 / Accepted: 1 February 1999     

Integrating wetlands and riparian zones in river basin modelling

Wetlands, and in particular riparian wetlands, represent an interface between the catchment area and the aquatic environment. They control the exchange of water and related chemical fluxes from the upper catchment area to surface waters like streams and lakes. Their influence on water and nutrient balances has been investigated mainly at the patch scale. In this study an attempt was made (a) to integrate riparian zones and wetlands into eco-hydrological river basin modelling, and (b) to quantify the impacts of riparian wetland processes on water and nutrient fluxes in a meso-scale catchment located in the northeastern German lowland. The investigation was performed by analysing hydro-chemical field data and applying the eco-hydrological model SWIM (Soil and Water Integrated Model), which was extended to reproduce the relevant water and nutrient flows and retention processes at the catchment scale in general, and in riparian zones and wetlands in particular. The main extensions introduced in the model were: (1) implementation of daily groundwater table dynamics at the hydrotope level, (2) implementation of water and nutrient uptake by plants from groundwater in riparian zones and wetlands, and (3) assessment of nutrient retention in groundwater and interflow. The simulation results indicate that wetlands, though they represent relatively small parts of the total catchment area, may have a significant impact on the overall water and nutrient balances of the catchment. The uncertainty of the simulation results is considerably high, with the main sources of uncertainty being the model parameters representing the geo-hydrology and the input data for land use management.     

Labial palps of the blue mussel Mytilus edulis (Bivalvia: Mytilidae)

As a basis for understanding the functions of labial palps in the blue mussel Mytilus edulis, the structure and histology of palps were studied using light and scanning electron microscopy. Mussels used in the present study were collected in August 1993 and April 1994. The palp ridged surface is characterized by the presence of a smooth but densely ciliated dorsal fold, upon which rests the corresponding demibranch ventral region. The underside of the dorsal fold and the palp ridges fuse to form vestigial ciliated tracts. The dorsal fold is capable of contraction, allowing it to cover variable amounts of the ridged surface. Two different types of mucocyte are present on the palp ridged surface: subepithelial, glandular, acid-dominant secretion mucocytes and epithelial mucocytes characterized by neutral secretions. In histological section, these mucocytes appear to be concentrated on anatomical features known to intervene in particle handling. The anatomical and histological features of the smooth surface are typical of bivalve labial palps, except that the dense ciliation of the dorsal fold begins in the dorsal region of the smooth surface, indicating the possible origin of this feature. Previous studies on M. edulis point to the palps as the probable site for both ingestion volume control and particle selection; the anatomical basis of the present study should facilitate further research on these aspects.     

Effects of UV-B radiation on the gills of Catla catla during early development

Scanning electron microscopic (SEM) study of gills of Catla catla catla (17-day-old) exposed to UV-B radiation (145?µW?cm^?2 at the water surface) for three different exposure times: 5, 10 and 15?min was conducted. Fish without UV-B exposure served as control. UV-B radiation damaged both gill filaments and lamellae. The intensity of damage was minimal in 5?min exposed fish, followed by 10?min exposed fish and maximal in 15?min. The gill epithelium was severely damaged in 15?min irradiated fish compared to control. Pavement cells (PVCs) were damaged and the numbers of microridges within PVCs decreased. The deep boundary of PVC was not clear. In some area of gill epithelium, PVCs were destroyed and mitochondrion-rich cells (MRCs) were exposed. The 5?min exposure reduced the number of microridges in the PVCs, but the boundary of PVCs was still visible. MRCs in the gill epithelium were not exposed in 5?min exposed fish. The damage to PVCs and subsequent exposure of MRCs in UV-B irradiated fish may hamper respiratory functions and disturb osmoregulation in catla.     

Co-ordinated rhythms of digestion,absorption and excretion in Mytilus edulis (Bivalvia: Mollusca)

Sequential alternation of extracellular digestion in the stomach and intracellular digestion in the diverticula appears widespread among bivalves. The present study documents some physiological consequences of such processes in Mytilus edulis L. collected during 1981 from Whitsand Bay, Cornwall, England. Pronounced temporal fluctuations in faecal deposition are described that relate, in terms of amplitude and period, to both sinusoidal rhythmicity established for ammonia excretion and changes in the morphology of digestive tubules. Although at least partially synchronised among replicate groups of mussels, these cycles bore no consistent relationship with exogenous influences. Hourly fluctuation in the net absorption efficiency for nitrogen, as evidenced by the mean percentage ±2 SE, measured over 24 h sampling periods, was considerable (16.0±53.7, 49.3±10.9 and 52.8±6.6 for mussels acclimated in March, June and October, respectively). This variation in absorption derived from an inverse relationship between the percentage nitrogen within faeces and the rate of faecal egestion. Accordingly, peaks of faecal deposition presumably represented the pulsed remnants of intracellular digestion. Co-ordinated rhythms of digestion, absorption and excretion were thus evident in M. edulis. These processes displayed seasonally dependent periodicities of approximately 8, 3 and 4 h in March, June and October, respectively. It was concluded that, at least for M. edulis, this previously unquantified rhythmicity of physiological processes warrants careful consideration during assays commonly undertaken in the complication of nutrient and energy budgets.     

Functional morphology of Petricola (Rupellaria) typica (Bivalvia: Petricolidae)

The bivalve Petricola typica occurs from North Carolina (USA) to Brazil where, according to Rios (1970), its range extends to Rio Grande do Sul. The species can be found in coral blocks of Mussismilia hispida, inside colonies of Schizoporella unicornis (a calcarean bryozoan) and, more frequently, inside the sandy reefs built by the polychaete Phragmatopoma lapidosa (intertidal zone). Initial settlement probably occurs in a pre-existing cavity, which is enlarged to form a hole into which the bivalve fits exactly when its valves open. The shell, mantle, and siphons are described in detail. Special attention was paid to the ctenidia, labial palps, and stomach.     

Heteroplasmy in a deep-sea protobranch bivalve suggests an ancient origin of doubly uniparental inheritance of mitochondria in Bivalvia

Most metazoan species have strict maternal inheritance of the mitochondrial genome. In bivalves, a unique inheritance pattern called doubly uniparental inheritance (DUI) occurs in at least seven bivalve families. In this system of mitochondrial inheritance, males inherit and carry mtDNA from both parents, while females only carry mtDNA from the mother. Here, we present evidence of mitochondrial heteroplasmy in deep-sea protobranch bivalves. Divergent 16S rRNA and cytochrome b sequences were obtained within individuals of Ledella ultima. Ledella sublevis also exhibited divergent 16S sequences. Levels of divergence between 16S sequences within individuals were 27 and 15 % for each species, respectively. Ratios of homoplasmic to heteroplasmic individuals were not significantly different from 1:1, in agreement with sex ratios in protobranchs. The results provide the first evidence for mitochondrial heteroplasmy in the protobranchs and suggest DUI might have evolved much earlier in the evolution of the Bivalvia than previously thought.     

Population structure and growth of Donax trunculus (Bivalvia: Donacidae) in the western Mediterranean

A comparison of shell growth in Donax trunculus (collected between 1988 and 1990 of Cullera, Spain) has been carried out using an analysis of cohort progression in monthly length frequency distributions, hyaline surface shell growth rings and internal microgrowth bands. In the Mediterranean there are two periods of recruitment of D. trunculus, one in the summer (July to September) and the other in winter (December to February). Clams recruited to the population in winter display a clear cessation in shell growth during the following summer which may possibly be correlated with spawning, whereas individuals of the summer recruited cohort show no growth cessation the following summer and continue to deposit shell during this period. The normally opaque shell of D.trunculus reveals the presence of translucent hyaline growth rings when the shells are backlit by a strong light source, and these have been shown to be laid down in the shell during summer months. Formation of a hyaline ring is accompanied by a narrowing of the microgrowth patterns present in shell sections. Both the hyaline rings and the length frequency distributions have been used to determine the age and growth rate of D. trunculus.     

Na-K-ATPase generates an active transport potential in the gills of the hyperregulating shore crab Carcinus maenas

Perfusing and bathing isolated gills of shore crabs Carcinus maenas with artificial saline or sea (brackish) water enabled us to determine potential differences (PDs) between ambient bathing medium and perfusion solution. Establishment of diffusional PDs was avoided by employment of the same solution on the internal and external side. The PDs measured were therefore of an active nature. We compared the properties of the PDs with the well-known properties of the Na-K-ATPase: dependence on biological energy (ATP), on salinity and sodium concentration, susceptibility of PD to depletion of internal K and to the addition of 5 mM internal ouabain. Considering also the magnitude of the PDs measured, the results obtained indicate that it is the Na-K-ATPase that generates an active transport potential for Na in the gills of shore crabs. This PD represents the driving force for the active uptake of Na in crabs that hyperregulate their body fluids when they inhabit environments of reduced salinity regimes or fluctuating salinities in tidal estuaries. This process counteracts diffusional losses of Na in crabs exposed to dilute media.