Time-course of digestive-enzyme acclimation in the cockle Cerastoderma edule

We determined the temporal evolution of amylase, cellulase, laminarinase and protease in the digestive gland and crystalline style of cockles Cerastoderma edule held over 9 to 12 d in the presence and absence of food. Cockles were fed a constant diet of 1.5 mm³ l⁻¹ of Tetraselmis suecica for 9 to 12 d and were then starved for 6 to 8 d in late summer (September 1992) and in winter (January 1993). Feeding increased the dry weight and total cellulase, laminarinase and protease activities of the digestive gland irrespective of season, whereas amylase activity remained unchanged. In winter (i.e. when cockles are metabolically weak) the response was faster and stronger, especially for protease. An additional experiment in September starved cockles for 20 d before resuming feeding. In agreement with the seasonal differences, the presence of food after prolonged starvation induced a rapid and marked increase in protease in the digestive gland of the cockles. In winter, the possible effects of the biochemical composition of food on their enzymatic response were tested by feeding two groups of cockles with the same ration of T. suecica but harvested at different growth phases. A compensatory induction of cellulases occurred in cockles fed on T. suecica with a lower carbohydrate content. In the crystalline style, the protein level and carbohydrase fell during the first day of feeding and increased during the first day of subsequent starvation. These results indicate that the release of enzymes from the style prevails over the incorporation of enzymes during the early stages of feeding, whereas the opposite occurs during starvation. Received: 15 February 1998 / Accepted: 22 February 1999     

Accumulation of polychlorinated biphenyls in the cockle Cerastoderma edule and the tellin Macoma balthica

Two species of marine bivalve, Cerastoderma edule (Linnaeus) and Macoma balthica (Linnaeus), were exposed to particles contaminated with three commercial polychlorinated biphenyl (PCB) mixtures, Aroclors^® 1242, 1254 and 1260. The accumulation of PCB in bivalve tissue was monitored for periods of up to 40 days. In addition to measuring total PCB concentrations, which reached 60 ppm of Aroclor 1242 in M. balthica, the fate of individual PCB homologues of different chlorine content, ranging from 2 to 8 chlorine atoms per biphenyl molecule, was also determined. Selective accumulation of PCBs with 5 chlorine atoms per biphenyl molecule occurred, whilst isomers of very low and very high molecular weights were accumulated at much slower rates.     

Anaerobic metabolism of the common cockle Cardium edule

Changes in the level of the adenylates (ATP, ADP, AMP), arginine phosphate, arginine, octopine, D-lactate, succinate, and aspartate were determined in the foot of Cardium edule, after 12 h of anoxia and after aerobic recovery for up to 12 h. Control levels of the adenylates, the calculated energy charge and the arginine phosphate content were restored after about 1 h of recovery. On the other hand, the recovery of the D-lactate, succinate, and aspartate pools was a slow process, taking place over about 12h.     

Interannual variation in the reproductive cycle and biochemical composition of the cockle Cerastoderma edule from Mundaca Estuary (Biscay,North Spain)

Seasonal determinants of body weight, biochemical composition and reproductive condition in the cockle Cerastoderma edule L. from the Mundaca Estuary, Spain, were performed from December 1983 to July 1985. Interannual differences in timing of gametogenesis and spawning between 1984 and 1985 were correlated with temporal variations in the cycles of storage and utilisation of carbohydrate reserves. Young individuals exhibited larger annual fluctuations in soft-body weight, displaying higher growth rates in spring and greater diminishment of energy reserves during autumn and winter than older individuals. Carbohydrate content displayed a similar tendency, levels being lower in young individuals throughout the year.     

Seasonal changes in the physiology,reproductive condition and carbohydrate content of the cockle Cardium (=Cerastoderma) edule (Bivalvia: Cardiidae)

Pronounced seasonal cycles in the rates of oxygen consumption and feeding were found for Cardium (=Cerastoderma) edule L. measured in the field under ambient conditions. The cockles had a maximum rate of oxygen consumption (0.89 ml O₂ g^-1 h^-1) in April which declined to a minimum of 0.35 ml O₂ g^-1 h^-1 in March. Their feeding rate was variable but had a maximum value (3.91 l g^-1 h^-1) in April and a minimum value (0.73 l g^-1 h^-1) in October. There was no apparent seasonal variation in absorption efficiency, with a mean value of 67.6%. Gametogenesis was initiated in January and the population reached a peak in reproductive condition in April/May, followed by a 3 month spawning period. Carbohydrate reserves were synthesised during spawning, and were then utilised during the winter and early spring. An adaptive function for a reduction in time spent feeding is postulated, and correlations between the rates of certain physiological processes and some exogenous and endogenous variables are discussed.     

Potential of bivalves' secondary settlement differs with species: a comparison between cockle (Cerastoderma edule) and clam (Ruditapes philippinarum) juvenile resuspension

Juvenile bivalves may be dispersed by entering a bysso-pelagic phase where they drift through the water mass aided by a long thread. The ability to resuspend and control the specific weight in two bivalve species, the cockle Cerastoderma edule (L.) and the Japanese clam Ruditapes philippinarum (Adams and Reeves), was documented with juveniles through flume and still-water experiments. Cockle juveniles initially placed on an unsuitable substratum were exposed to two shear velocities (u _*). At the end of the experiment, 42 (±15)% (for u _*=0.51 cm s⁻¹) and 79 (±9)% (for u _*=0.99 cm s⁻¹) of individuals were retrieved from the sand area which represents only 8% of the total flume surface. Most juveniles (70.5%) with shell lengths <2.5 mm migrated from the unsuitable Plexiglas substratum to the sand array by resuspension in the water column. The percentage was lower (21.5%) for larger individuals. The same experimental design was applied to clams, which immediately adhered to the Plexiglas substratum and remained attach to it. Sinking rates of live and dead specimens of both species were measured in a 1 m long transparent PVC tube. Cockle fall velocities showed severe deceleration, probably due to byssus secretion (up to 15-fold slower than dead cockles), sometimes interrupted by brutal acceleration probably due to byssal rupture. Cockles were able to reduce their sinking rate for shell lengths up to 4.25 mm. By contrast, clam sinking rates were constant, and similar to dead clam sinking rates. Specific weights of all experimental juveniles were calculated in relation to their lengths, and their passive motion into the boundary layer was theoretically assessed with Shields curve. In short, C. edule and R. philippinarum can both exhibit dense populations in the field with a good capacity to colonize, although juveniles display different abilities to resuspend in the water column. Received: 27 January 1997 / Accepted: 13 February 1997     

Effect of tidal submersion on lysozyme activity in Mytilus edulis and Tellina tenuis

Changes in mean style weight and in lysozyme activity of the style, digestive gland, gill and mantle of Mytilus edulis and Tellina tenuis from the Clyde Sea area, Scotland, were investigated over tidal cycles in March and August, 1981. For M. edulis, significant changes occurred in the style weight, style lysozyme activity and digestive gland lysozyme activity during a 22 h period. These appear to be related to a diurnal cycle rather than a tidal cycle. Changes in the weight of the style of M. edulis may be caused by dissolution during feeding, and style lysozyme may be secreted independently of the style matrix. The activity of lysozyme in T. tenuis is unaffected by the tide, suggesting that this intertidal bivalve can feed throughout the tidal cycle.     

Does displacement of spawning time occur in the sibling species Cerastoderma edule and C. lamarcki?

The spawning cycles of the sibling species Cerastoderma edule (L.) and C. lamarcki (Reeve) were studied in allopatric and sympatric Danish populations from 1978–1980. The results do not support earlier findings of reproductive character displacement in sympatric populations of the two species. New records of hybrid formation and intermediate forms are presented and discussed.     

Energetics of growth and reproduction in cockles (Cerastoderma edule): Seasonal and age-dependent variations

Seasonal patterns of growth were compared interannually between 1983 and 1985 for different ageclasses of an intertidal population of cockles,Cerastoderma edule L., in the Mundaka Estuary (Basque Country, North Spain). Growth data and simultaneous determinations of respiratory metabolism were integrated over defined periods of time in order to compute assimilation and net growth efficiency. Consistent differences between the exponents that scale both assimilation and metabolism to body size led to growth efficiency values that were a decreasing function of age. Total growth was partitioned into three components: somatic growth, reproductive output and reserve storage, and the corresponding effort indexes were calculated. Despite important differences in total growth, reproductive output was constant between years, indicating a higher reproductive effort under poorer growth conditions. Increased costs of reproduction that appeared to be associated with this higher reproductive effort did not affect somatic maintenance, since previously stored carbohydrate reserves were primarily used to fuel gametogenesis.     

Evolution of Cardium (Cerastoderma) edule,C. lamarcki and C. glaucum: studies of DNA-variation

Genetic relations between populations of Cardium (Cerastoderma) edule (L.) and C. lamarcki (Reeve) from Danish waters and C. glaucum (Bruguiére) from the Mediterranean Sea were studied using DNA analysis. Repetitive sequences of nuclear DNA were compared by Southern blotting of enzyme digested DNA fragments, separated on agarose gel by electrophoresis. Fragments were visualized by autoradiography using ³²P labelled probes. These were nick translated, randomly sheared fragments of the whole chromosomal genomes of two species of distant taxa — Mytilus edulis and Mya arenaria, respectively. The number of detected DNA bands was 18 for C. edule, 32 for C. lamarcki, and 24 for C. glaucum. Measurements of genetic distances, based on comparisons of these repetitive sequences using a simple dissimilarity coefficients, show the greatest distance, i.e., 0.84 between C. edule and C. lamarcki, 0.69 between C. edule and C. glaucum, and the smallest, 0.52 between C. lamarcki and C. glaucum. Comparisons of the three Cardium types, using gene spliced and cloned C. edule nuclear DNA (nDNA) fragments as probes, accorded with these findings. Also, analysis of mitochondrial DNA (mtDNA) from eggs and hepatopancreas, although not in itself conclusive, accorded with the results of the analyses of nDNA. MtDNA was measured by electron micrograph to 15 000 base pairs (bp), and sizes of restriction enzyme digested fragments were compared by gel electrophoresis. The results of the comparisons based on repetitive sequences supported by the two other studies suggest that C. edule resembles the ancestral form more than the two others, and that C. lamarcki is the most different and recently evolved form. This hypothesis is discussed in relation to the geographical distribution of the species complex.     

Genetic relations between the bivalves Cardium (Cerastoderma) edule,Cardium lamarcki and Cardium glaucum,studied by means of crossed immunoelectrophoresis

Rabbit antisera were raised against Cardium edule (L.) from the Waddensea locality, Skallingen, against Cardium lamarcki (Reeve) from the Danish locality, Dybsø Fjord, and against Cardium glaucum (Bruguière) from the French Mediterranean coast at Ile des Embiez. Allopatric and sympatric populations of C. lamarcki and C. edule from Swedish and Danish localities, and allopatric populations of C. glaucum from the Mediterranean Sea, Red Sea and Black Sea were studied by means of the described method and by starch gel electrophoresis of malate dehydrogenase (MDH). By means of crossed immuno-electrophoresis, using intermediary gels containing antibodies against one of the Cardium types and receiver gels containing antibodies against one of the others, type-specific and type-non-specific reactions were recorded. The results are compared with a similar investigation of Mytilus edulis, M. galloprovincialis and M. californianus. Based on immunological properties, it is shown that C. edule is a valid species different from the C. lamarcki/C. glaucum complex. It is further suggested that C. lamarcki and C. glaucum should still be referred to by these two names as significant genetic differences between them is revealed by this immunological study.     

Seasonal variations in biochemical constituents during the reproductive cycle of the female dog cockle Glycymeris glycymeris

In an attempt to describe the biochemical events associated with the main stages of the annual and reproductive cycles of the female dog cockle Glycymeris glycymeris L., we studied seasonal variations in the various stages of oocyte development of the ovaries, and the glycogen, total protein and total lipid content of five body tissues – adductor muscle, foot, tunic coat, visceral mass and mantle. From November 1991 to November 1994, microscopic examination of the ovaries and measurement of the tissue concentrations of glycogen, total proteins and total lipids in these five body tissues were made monthly on ten female dog cockles originating from the sea area around Douarnenez (south Brittany, France). Morphological studies revealed that in the population investigated the annual cycle is characterised by three major periods: a first period of vitellogenesis extending from February/March to April/May and preceding a spawning in spring; a second period of vitellogenesis extending from May/June to September/October and leading to either no spawning, a single autumnal spawning event, or to two spawning events in summer and autumn; and a third period extending from October/November to February/March and characterised by a high level of oocyte lysis. In the muscular body tissues of the dog cockle, i.e. the adductor muscles, the foot and the tunic coat (the muscular envelope containing the visceral mass), the concentrations of glycogen, total proteins and total lipids underwent very similar variations during the annual cycle. During each stage of vitellogenesis, a typical glycogen–protein–lipid sequence was observed in the muscular tissues that was characterised firstly by a peak of glycogen concentration 2 to 3 mo before spawning, followed by a peak in total proteins 1 mo before spawning, and finally by a peak in lipid content just before spawning. A similar glycogen–protein–lipid sequence was also recorded in the first half of the winter period. However, these events were followed by general atresia affecting all oocytes in the gonad. Maximum energetic value of biochemical constituents in females coincided with peaks in lipid content in the visceral mass and mantle. These biochemical events occurred principally immediately before and at the end of oocyte lysis (December/January). A drop in the total energetic value, affecting mainly the visceral mass and the mantle, was recorded each year during the period January to March, coinciding with the period of shell growth in this species. Our data clearly indicate that in female G. glycymeris all muscular tissues contribute to the storage of glycogen and proteins, and suggest that glycogen may be the source of energy triggering vitellogenesis. Biochemical and microscopic observations revealed that oocyte development takes place during the first half of winter, but that these oocytes undergo atresia in December/January. The metabolites produced from oocyte lysis could contribute to somatic growth, which occurs in late winter. Received: 3 March 1997 / Accepted: 23 July 1997     

Feeding current characteristics of three morphologically different bivalve suspension feeders, Crassostrea gigas, Mytilus edulis and Cerastoderma edule, in relation to food competition

Introduced Pacific oysters (Crassostrea gigas) have shown rapid expansion in the Oosterschelde estuary, while stocks of native bivalves declined slightly or remained stable. This indicates that they might have an advantage over native bivalve filter feeders. Hence, at the scale of individual bivalves, we studied whether this advantage occurs in optimizing food intake over native bivalves. We investigated feeding current characteristics, in which potential differences may ultimately lead to a differential food intake. We compared feeding currents of the invasive epibenthic non-siphonate Pacific oyster to those of two native bivalve suspension feeders: the epibenthic siphonate blue mussel Mytilus edulis and the endobenthic siphonate common cockle Cerastoderma edule. Inhalant flow fields were studied empirically using digital particle image velocimetry and particle tracking velocimetry. Exhalant jet speeds were modelled for a range of exhalant-aperture cross-sectional areas as determined in the laboratory and a range of filtration rates derived from literature. Significant differences were found in inhalant and exhalant current velocities and properties of the inhalant flow field (acceleration and distance of influence). At comparable body weight, inhalant current velocities were lower in C. gigas than in the other species. Modelled exhalant jets were higher in C. gigas, but oriented horizontally instead of vertically as in the other species. Despite these significant differences and apparent morphological differences between the three species, absolute differences in feeding current characteristics were small and are not expected to lead to significant differences in feeding efficiency.     

The effect of microbial activity upon the sedimentary sulphur cycle

The sulphate content of an intertidal sediment at Anglesey (UK) was shown to vary during the year, the sulphate-reducing activity of the microbial population being limited by low temperature during winter, and by the low numbers of sulphate-reducing bacteria during May, 1969. A corresponding annual variation of the sedimentary sulphide could not be demonstrated, and the sulphide which was formed biologically during the course of the year was almost entirely lost from the sediment. This loss of sulphide was probably due to subsurface oxidation; and flushing, by water entering at the base of the sediment. It is suggested that sulphide was only precipitated within the sediment during summer when sulphate reduction was active, and even then only during part of the tidal cycle when flushing did not occur.     

Effect of the presence of the shore crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, on burrowing behaviour and clearance rate of the common cockle, <Emphasis Type="Italic">Cerastoderma edule</Emphasis>

Bivalves demonstrate various morphological and behavioural adaptations to reduce the risk of being attacked by predators. This paper examines how the presence of the crab Carcinus maenas (L.), a natural predator of the cockle Cerastoderma edule (L.), affects its burrowing depth and clearance or feeding rate. Cockles were placed in experimental tanks and treated with three levels of predatory disturbance: (1) unfed crab loose inside the tank, (2) unfed crab inside a cage suspended in the water column and (3) no crab present. Cockles’ burrowing depth was measured in two sediment types: mud and sand. Cockles burrowed more deeply in treatments with no crabs. Burrowing depth in sand was significantly greater than in mud. Two factors may contribute to the reduction in burial depth of C. edule in the presence of C. maenas: the change in the vertical orientation of the cockle and the ‘cough response’. No significant difference was found in the cockles’ clearance rate among the different levels of predator threat.     

Dynamics of heavy-metal transfer between biotic (Cytheria chione and Cerastoderma edule) and abiotic (water and sediment) components in marine environment (Bay of Martil, Moroccan Mediterranean coast)

This study deals with the transfer of heavy metals (Hg, Cd, Pb, Cr, Ni, Ti, Zn, and Cu) between abiotic and biotic components (Cytheria chione and Cerastoderma edule) in the inshore intertidal zones of Tetouan/Martil over 2 yr of investigation (1992-1994). Analysis of the heavy-metal accumulation kinetics in Cytheria chione and Cerastoderma edule shows that their highest contents occur during the spring-summer period when an important proliferation of the plankton biomass occurs. However, the accumulation of Cd is higher during the period between winter and spring. In general, heavy-metal concentrations are higher at Cytheria chione than at Cerastoderma edule. These results, according to previous studies, suggest the presence of significant correlations and dynamic reciprocal transfer of heavy metals among seawater, sediment, and molluscs. They also suggest that the level of contamination of decreasing heavy metals follows the sequence: sediment, organisms, and water. Among the molluscs investigated, the variability of the thallium (Tl) contents is difficult to detect due to the very weak presence of this metal in the analysed tissues.     

Studies on the reproductive cycles of Cardium edule and C. glaucum

Two allopatric populations of Cardium edule and two of C. glaucum were sampled over a period of 16 months and their gonad maturity cycles compared by histological methods. The onset of both spermatogenesis and oogenesis in C. edule took place between September and October; in C. glaucum it was slightly later, between October and November. Development of the gonads continued slowly throughout the winter, C. glaucum developing marginally faster than C. edule. Maturation accelerated in April and May, with the gonads of both species becoming fully ripe in late May and early June. Spawning of the two species approximately coincided at all the localities investigated. Spawning in C. edule was better coordinated than in C. glaucum. Transplanting cockles of one species to the typical habitat of the other did not appreciably alter the spawning period. The separation of spawning periods in sympatric associations of the two cockles is discussed.     

Egg release and settlement patterns of dioecious and hermaphroditic fucoid algae during the tidal cycle

The timing of gamete release by fucoid algae, although known to be restricted to calm days is not clearly understood within a circadian time scale. The need for externally fertilizing species to avoid gamete dilution suggests that in wave-exposed areas spawning may occur during particular tidal phases. However, this may differ between mating systems, as selfing species may be less affected by gamete dilution. In this study, two different approaches were used to determine when egg release occurs during the tidal cycle in two sister species with different mating systems. First, egg settlement of Fucus vesiculosus (dioecious) and Fucus spiralis (selfing hermaphrodite) was quantified on removable substrates (egg settlement disks) every day for 2 months and settlement patterns were used to statistically estimate the radius of a circle that would encompass 99% of each patch of settled eggs (the egg dispersal radius). Also, egg release was quantified every 2 h during the tidal cycle. A significantly larger egg dispersal radius (P < 0.02) was found for F. spiralis than F. vesiculosus, and this difference was somewhat site dependent with a greater difference between species in exposed sites. The egg dispersal radius was negatively correlated with significant wave height and positively correlated with sea surface temperature for both the species (P < 0.05), with a greater effect of both the factors for F. spiralis than for F. vesiculosus. Egg release during the tidal cycle was variable between species and experiments, with F. vesiculosus releasing more eggs, later in the day, and at a lower tide, than F. spiralis, which released fewer eggs, throughout the day and at all tides. The dioecious species, F. vesiculosus, may have developed a specific adaptation for timing the egg release to periods when emersed in exposed habitats to avoid rapid dilution of gametes that require outcrossing for fertilization. On the other hand, egg release for F. spiralis, which can self-fertilize, occurred both when emersed and immersed, suggesting this species has developed less synchrony with specific environmental factors. Site dependence also suggests local wave conditions can modulate timing of release. The data are consistent with the relaxation of the selective constraints of water motion on fertilization success in a selfing hermaphrodite, relative to an obligate outcrossing species. Results support the idea that species with different mating systems evolve different sensitivities to environmental cues for gamete release with specific implications for inbreeding and successful external fertilization in the ocean.     

除草剂对土壤氮素循环的影响

土壤微生物参与土壤氮素循环的生物学与生物化学过程,对氮素形态转化与去向产生很大影响。在现代农作物生产上农田普遍施用除草剂,除草剂进入土壤生态环境中影响土壤微生物种群数量、活性和土壤氮素循环过程,在一定程度上改变氮素各去向的比例。因此,除草剂的施用对植物氮素吸收利用和土壤氮的环境释放具有一定效应。文章综述了除草剂对生物固氮、土壤氮矿化与转化、氨挥发、硝化反硝化、温室气体N2O排放、植物的氮吸收利用、土壤氮损失等方面的影响,并提出了今后进一步研究的方向,为减少氮素损失和温室气体排放以及除草剂使用的安全性评价提供参考。