Mortality of newly metamorphosed eastern oysters (Crassostrea virginica) in mesohaline Chesapeake Bay

 Stocks of eastern oysters, Crassostrea virginica (Gmelin), in mesohaline Chesapeake Bay, USA, exhibit a high degree of inter-annual and spatial variability in recruitment. We found that cumulative oyster spatfall on off-bottom collector plates, measured throughout the summer in 14 years over a span of three decades, was highly positively correlated (r ² = 0.8) with juvenile oyster recruitment on adjacent oyster bars. Total abundances of juvenile oysters on these bars were, however, generally 99.7% lower than predicted from cumulative seasonal larval settlement on collector plates. We propose that although the number of larvae metamorphosing was the key factor in determining the gross annual pattern of recruitment to these mesohaline oyster bars, the actual magnitude of recruitment was governed by post-settlement processes, such as competition for limited resources and predation. We tested the hypothesis that predation may be partly responsible for high post-settlement juvenile oyster mortality. We performed a series of 3-d field investigations over two summers (1989, 1990) at a mesohaline site, employing cages of various mesh sizes (400, 800, 1500 μm) to protect hatchery-reared spat of 0.5 to 4.0 mm shell height. Mortality rates for spat held for 3 d in the estuary (17.8%) were significantly higher (P = 0.0001) for the smallest spat (0.5 to 2.0 mm) compared with those of 2.01 to 4.0 mm (4.2%). In 1990, but not in 1989, enclosure within 400 and 800 μm mesh cages significantly (P = 0.004) increased survival during 3-d deployments (9.4 and 10.1%, respectively) compared with spat unprotected by mesh cages (21.9%). In a series of laboratory predation studies that used the entire community of invertebrates that could penetrate the cages, microscopic juvenile polyclad flatworms, Stylochus ellipticus, were the only organisms that we observed crawling into living oysters and feeding on oyster tissue. Large flatworms (50 to 200 mm²) are known to be important predators on oysters, but this ability of flatworms that were so small (<ca. 5 mm²) and translucent as to be almost invisible without magnification to feed on immediate post-metamorphic oysters has not been documented previously. Our results suggest that the rate of mortality due to predation in mesohaline Chesapeake Bay is much reduced once spat survive for 2 to 3 weeks post-metamorphosis. Thus, it is likely that predation in the 1 to 2 week period immediately after settlement may be a crucial factor in the structuring of eastern oyster populations. Received: 21 December 1998 / Accepted: 2 December 1999     

Mitochondrial DNA analysis of native and selectively inbred Chesapeake Bay oysters,Crassostrea virginica

Mitochondrial DNA (mtDNA) genotypes were examined in 1989 in threeCrassostrea virginica (Gmelin) populations native to Chesapeake Bay, USA, and in one population selectively inbred for rapid growth for ten generations. We wished to determine whether this character would be useful as a genetic marker for distinguishing between the inbred line and the native oysters and to determine whether detectable genetic differences exist among present-day native populations ofC. virginica. Thirty mtDNA haplotypes were identified. The average percentage nucleotide difference between native haplotypes was 1.8%. Inbred oysters were characterized by mtDNA haplotypes distinctly different from ancestral native oysters, indicating a high degree of genetic differentiation between the two groups (average percentage mtDNA nucleotide sequence divergence,=21.8%). The common native haplotype was not present in the selectively inbred sample, and six of the seven haplotypes detected in the inbred oysters were not found in the survey of native oysters. Chi-square tests on haplotype frequencies indicated that the native populations were not significantly different from one another. However, the distribution and relatedness of haplotypes suggest that significant change in the oyster gene pool may have occurred over the past few decades.     

Growth patterns in oysters, Crassostrea virginica, from different estuaries

We analyzed a data set collected over 15 yr, containing growth data from strains of eastern oysters, Crassostrea virginica (Gmelin, 1791), initiated from parent populations in Long Island Sound, Delaware Bay, and lower Chesapeake Bay. The long-term growth data proved to be a powerful tool for examining patterns of growth differentiation among separated populations of C. virginica. The oyster strains had been grown in a common environment in lower Delaware Bay for up to seven generations. We found that the oyster strains with origins in Long Island Sound were significantly larger over several generations than oyster strains from Delaware Bay and Chesapeake Bay. Chesapeake Bay oyster strains were larger than Delaware Bay oyster strains at 1.5 yr old, but Delaware Bay oysters were larger at 2.5 yr. Year-to-year variation in environmental conditions had a strong significant effect on absolute oyster size and the relative sizes of the oyster strains. Persistent differences between oyster strains from different origins over several generations support a hypothesis that these estuarine populations have experienced long-term genetically-based population differentiation. This result is consistent with hypotheses of population differentiation of oysters based on observations of local reproductive timing. Received: 12 August 1997 / Accepted: 26 May 1998     

Metabolism of palmitic,linoleic, and linolenic acids in adult oysters,Crassostrea virginica

This study investigated incorporation and metabolism of saturated [(1-¹⁴C) 16:0] and unsaturated [(1-¹⁴C) 18:26 and (1-¹⁴C) 18:33] fatty acids in adult eastern oysters,Crassostrea virginica Gmelin (spawned from parents obtained in 1986 from Mobjack Bay, Virginia, USA), and the influence of temperature on these processes. InC. virginica, incorporation of injected palmitic (16:0) and linolenic (18:33) acids was increased when oysters which had been grown in warm water (22 to 23°C) were transfered to cold water (5 to 7°C) for 8 to 18 d. Incorporation of linoleic acid (18:26) was unchanged under these conditions. The changes in concentration may have been linked to depression of metabolism in these oysters, in particular that of 16:0, which was reduced by 90%. Oxidation of incorporated fatty acids was much higher in warm than in cold water. Cold-temperature conditioning ofC. virginica altered the distribution of fatty acids among the neutral and polar lipid fractions. Long-term exposure to cold water increased the proportion of fatty acids in the polar fraction, which may be related to maintenance of membrane fluidity. Short-term exposure to cold water had the opposite effect, which may be due to increased energy requirements as the oyster adapts to new conditions. Reutilization of¹⁴C-acyl groups demonstrated de novo synthesis of 16:0 and 18:0 fatty acids. Only limited elongation and no desaturation of the administered fatty acids was observed.     

Depuration of twelve trace metals in tissues of the oysters Crassostrea gigas and C. virginica

The depuration of 12 trace metals in the mantle, gill, digestive gland, and kidney of Crassostrea gigas and C. virginica was investigated under natural field conditions; oysters from a relatively contaminated environment (Redwood Creek in south San Francisco Bay) were transplanted to a relatively clean environment (Tomales Bay). In the transplanted oysters, the digestive gland and kidney depurated Cd, Cu, Hg, Ag, and Zn more readily than the mantle and gill. Other trace metals As, Fe, Mn, Ni and Se showed varying depuration patterns. The results for Cr and Pb were inconclusive, since initial concentrations were too low to follow any losses. Interspecific differences in trace metal depuration were observed. Biological half-lives for most trace metals were on the order of 23 to 60 d for C. gigas and on the order of 70 to 180 d for C. virginica.     

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.     

Ultrastructure of the gonad and gametogenesis in the eastern oyster,Crassostrea virginica. II. Testis and spermatogenesis

The pelagic harpacticoid copepod, Macrosetella gracilis (A. Scott), is found in association with colonies of the nitrogen-fixing (diazotrophic), bloomforming cyanobacterium Trichodesmium spp. in tropical and subtropical waters. M. gracilis is one of the few direct grazers of these often toxic cyanobacteria. Experiments investigating NH ₊ ⁴ regeneration by M. gracilis were conducted in the Caribbean in September 1992 and the Coral Sea, Australia in November 1994. Rates of M. gracilis ingestion of Trichodesmium thiebautii labelled with ¹⁵N₂ measured in the eastern Caribbean indicated that M. gracilis could consume 33 to 45% of total T. thiebautii colony N d^-1 and >100% of new N fixed d^-1. We also measured the release of NH ₊ ⁴ by M. gracilis feeding on T. thiebautii, as well as by non-feeding copepods, using ¹⁵N isotope dilution methods. In non-feeding copepods, rates of NH ₊ ⁴ release increased as numbers of copepods were increased as both copepod numbers and food availability increased. In the presence of T. thiebautii colonies, M. gracilis had an average rate of NH ₊ ⁴ regeneration of 7.7±1.5 nmol N copepod^-1 h^-1 (±SE), which was significantly higher than when food was absent (1.9±0.7 nmol N copepod^-1 h^-1). Rates of M. gracilis excretion were relatively high based on excretion: ingestion ratios, which could be due to having a high-N food source readily available, to sloppy-feeding effects, or as a response to toxins in the cyanobacterium. Incubations of M. gracilis with and without T. erythraeum resulted in significant increases in [NH ₊ ⁴ ] as a function of copepod density only. Ammonium leakage from the cyanobacterium and/or microheterotroph associates was relatively low. M. gracilis, through excretion and possible mechanical breakage of cells while grazing, appears to provide a direct link between atmospherically derived new nitrogen and regenerated NH ₊ ⁴ in the oligotrophic systems where Trichodesmium spp. are abundant.     

胶州湾某水域蛤蜊(Ruditapes philippinarum)、牡蛎(Crassostrea ariakensis)中的雌激素含量

为了探讨海洋贝类对环境中雌激素的富集情况,采用液-液萃取和固相萃取方法提取了胶州湾李村河入海口蛤蜊、牡蛎和底泥中的雌酮(E1)、雌二醇(E2)、雌三醇(E3)和雌炔醇(EE2),并用GC-MS方法测定了上述4种物质的含量,此外还采用暴露实验方法研究了蛤蜊吸收底泥中雌激素物质的影响因素.测定结果显示,该水域中蛤蜊全组织中E1含量为0.45～5.20ng·g-1,E2含量为nd～4.34ng·g-1,E3和EE2未检出;牡蛎体内的E1、E2和E3含量分别为4.64～29.66、2.16～7.20和0.96～7.96ng·g-1,EE2未检出;底泥中E1、E2和E3含量分别为0.43～2.36、nd～0.59、0.52～1.13ng·g-1,EE2未检出.蛤蜊和牡蛎体内的雌激素物质含量高于其栖息的底泥.暴露实验结果显示,随着底泥中雌激素物质含量的增加或暴露时间增加,蛤蜊体内雌激素物质的含量增加,贝类可吸收并富集底泥中的雌激素物质.     

Different salinity tolerance mechanisms in Atlantic and Chesapeake Bay conspecific oysters: glycine betaine and amino acid pool variations

Crassostrea virginica (Gmelin) collected in 1989 from several sites within the Chesapeake Bay have narrower salinity tolerances than conspecific oysters collected in 1989 from several Atlantic coast sites (Georgia to Cape Cod). The basis of this physiological difference appears to be the biochemical mechanisms that control cellular osmolality following salinity stress. When adapted to the same salinity, the amino acid pools of both gill and adductor muscles of Atlantic oysters are larger than those of Bay oysters and different in composition. The Atlantic oyster tissues rely primarily on taurine for salinity tolerance, while the Bay oyster tissues have relatively less taurine, depending instead upon alanine, glycine and proline to adapt to high salinity. In addition, Atlantic oyster gill and adductor have 10 to 25 times the glycine betaine concentrations of these tissues from Bay oysters, depending upor the salinity of acclimation. The betaine concentration varies with salinity in Atlantic oysters, but does not change in Bay oysters. The results suggest that these biochemical differences are the basis of the narrower salinity tolerance in Bay oysters. The biochemical differences may reflect genetic differences between Bay and Atlantic oysters.     

Toxicity of ammonia,nitrite ion,nitrate ion,and orthophosphate to Mercenaria mercenaria and Crassostrea virginica

The 96-h lethal tolerance limits of the hard clam Mercenaria mercenaria (Linné) and the oyster Crassostrea virginica (Gmelin) to ammonia, nitrite ion, nitrate ion, and orthophosphate were defined. Sublethal effects of the chemicals upon the rates at which the shellfish removed algal cells from suspension were also studied. In comparison with other marine and aquatic species which have been studied, hard clams and oysters are extremely tolerant.     

Latitudinal patterns of shell thickness and metabolism in the eastern oyster Crassostrea virginica along the east coast of North America

The goal of this study was to quantify growth and metabolic responses of oysters to increased temperatures like those that will occur due to global warming. Impact of temperature on eastern oyster (Crassostrea virginica) shell growth and metabolism was investigated by sampling 24 sites along the eastern North American seaboard ranging from New Brunswick, Canada, to Florida, USA, in March and August 2013. There was a positive correlation between oyster shell thickness and site temperature. At southern sites, shells were up to 65 % thicker than at the northernmost site, likely due to higher precipitation of CaCO₃ in warmer water. This was supported by laboratory experiments showing that thicker shells were produced in response to temperatures 2, 4, and 6 °C above ambient seawater temperatures (8–14 °C) in Connecticut, USA. Field experiments with oyster respiration were conducted during winter and summer at 13 sites to compare responses to thermal stress with latitude. Respiration rates were much higher during summer than winter, but the combination of summer and winter data fell along the same exponential curve with respect to temperature. At all sites, temperature-specific metabolic rates at elevated temperatures were lower than predicted, indicating significant seasonal acclimatization by C. virginica.     

Accumulation and depuration of mercury in the American oyster Crassostrea virginica

To study the kinetics of mercury uptake in oysters, adult Crassostrea virginica (Gmelin) were held in seawater containing 10 g mercury/l (ppb) or 100 g mercury/l (ppb), added in the form of mercuric acetate, for 60 days. Mercury concentration in tissues was determined by analysis of individually homogenized oyster meats, using wet digestion and flameless absorption spectrophotometry. After 45 days, average mercury tissue concentration was 140,000 g mercury/kg tissue (ppb) and 28,000 g mercury/kg tissue (ppb) in the 100 ppb and 10 ppb experimental groups, respectively. After this time, concentrations dropped sharply, probably due to spawning. Clearance of mercury from tissue was studied by exposing treated adults to estuarine water (with no additions) for 30 days (100 ppb group) and 160 days (10 ppb group). Tissue concentrations in the 100 ppb mercury environment group declined from 115,000 to 65,000 ppb, and those of the 10 ppb group declined from 18,000 to 15,000 ppb, in 18 days; there-after, no further decline occurred in either group. Oysters accumulated mercury 1,400 times and 2,800 times above the environmental concentrations of 100 and 10 ppb mercury, respectively. Total self-purification was not achieved over a 6 month cleansing period.     

Accumulation,release and retention of petroleum hydrocarbons by the oyster Crassostrea virginica

Two Crassostrea virginica populations, differing in fat content, were experimentally exposed to a complex petroleum-hydrocarbon fraction. The hydrocarbons in this mixture were accumulated by both groups of oysters, and their lipid content, as well as the concentration of hydrocarbon in the water, were found to affect the rate and extent of accumulation. Hydrocarbons accumulated were rapidly, although incompletely, discharged when the oysters were transferred to an uncontaminated system. Amounts of hydrocarbons discharged and amounts retained after discharge are probably related to the level of contamination. The data can be interpreted as indicating that equilibration and the occurrence of multiple compartments where hydrocarbons can reside are factors involved in the uptake and retention of nonbiogenic hydrocarbons by oysters. The petroleum hydrocarbons contained in the oysters differed from the contaminating oil by displaying a greater aromatic content. In addition, gas-liquid chromatograms of aliphatic fractions of the hydrocarbons in the oysters rapidly showed a degraded appearance; the possibility that the oysters themselves are modifying the oil cannot be excluded.Contribution No. 3098 from the Woods Hole Oceanographic Institution.     

An exploratory study with the proton microprobe of the ontogenetic distribution of 16 elements in the shell of living oysters (Crassostrea virginica)

Study of the chemical composition of shell of exoskeletonous organisms in the past has required the sacrifice of the organism. Because the beam of the proton microprobe is relatively nondestructive and analyzes the surface layer of the shell, organisms do not have to be killed. The present paper presents results of a preliminary experiment in which distribution of elements (Na to Sr) in shell of living juvenile oysters, Crassostrea virginica (Gmelin), was studied in situ with a proton microprobe at monthly intervals for four months. The relative concentration of 16 elements was measured in the newly deposited prismatic edge of the right valve of three oysters reared in controlled laboratory conditions. Na, Mg, Al, Si, S, Cl, K, Ca, Ti, Cr, Mn, Fe, Cu, Zn, Br, and Sr were detected in concentrations as low as a few parts per million relative to the concentration of standards added to pure CaCO₃. Concentration of elements varied nominally among shells of the three individual oysters and in their successive ontogenetic stages. Fluctuations in concentration of Na, Mg, S, Cl, Ca, Mn, Fe, Cu, and Zn were generally similar in the two normally growing oysters, but differed from those in the oyster that stopped growing. Trends in concentration of Al, Si, and Sr were similar in the three oysters: those of Br were variable. Relative concentrations of Na, Cl, S, Mn, Fe, and Zn increased slightly with age of oysters, that of the other elements stayed relatively constant. Concentration of most elements was higher in shell than in seawater. Variable concentrations, especially of Na, Cl, and Si in valve edges, tend to support the hypothesis of earlier workers that separate mineral phases are present as impurities entrapped within the shell during calcification.     

Fluxes of dissolved and particulate calcium in selected tissues of Crassostrea virginica

Rates of calcium incorporation by selected tissues of Crassostrea virginica increased in a step-wise fashion from lowest values among organisms exposed to ambient total calcium concentrations of 45 mg l^-1, intermediate values among oysters exposed to 135, 225, and 315 mg l^-1, to highest rates for oysters exposed to 360 mg l^-1. Although excised visceral mass tissue had highest rates of calcium incorporation relative to mantle, muscle, and both organic and inorganic portions of the shell, mantle tissue appeared to have the most dynamic response to changes in ambient calcium concentrations. Rates of dissolved calcium incorporation from ambient water were approximately two to three orders of magnitude higher than comparable rates from ingested algal food. Behavioral response to concentrations of selected inoic, species in the ambient environment may have been responsible for observed differences in rates of calcium incorporation.     

Interaction of genotype and salinity in larvae of the oyster Crassostrea virginica

Adult Crassostrea virginica were obtained from 4 populations and spawned in the laboratory. The larvae from the within-population crosses and the hybrid crosses were raised at 4 salinities. There were no significant differences in survival of the larvae between the populations. However, one set of hybrids did show overdominance in survival. There were genetic differences between the populations in growth rate, but the expression of the differences depended upon the salinity, i.e, there was significant genotype-environment interaction. There were expressions of nonadditive genetic effects in the hybrid crosses, but the direction and magnitude was dependent upon the salinity. There was as much difference between populations from the same estuary as there was between populations from geographically isolated populations.     

Environmental stress,competition and dominance of Crassostrea virginica near Beaufort,North Carolina,USA

Observations and experiments were made at 2 intertidal areas near Beaufort, North Carolina, USA from May 1977 to July 1978 to determine why the oyster Crassostrea virginica dominated the community in areas protected from wave action but not in areas directly exposed to waves. Barnacles, oysters, the green alga Enteromorpha sp. and the mussel Brachiodontes exustus were the main occupiers of primary space at the mid and low intertidal levels of exposed areas. The intertidal community at the protected site consisted of a mid intertidal occupied by the barnacles Balanus amphitrite and Chthamalus fragilis, and the oyster C. virginica, and a low intertidal dominated by C. virginica. The exposed area was highly variable with high colonization and mortality for all species producing large seasonal changes in structure. The protected site remained constant throughout the year; there was no evidence of further colonization of either barnacles or oysters and mortality was very low. Recolonization experiments, selective removal of species, and growth and survival data demonstrated that C. virginica does not become dominant at exposed locations because (1) the constant wave shock at the ocean site reduces growth and increases mortality of young and adult oysters and (2) oysters are outcompeted by the mussel B. exustus. The monopolization of space by C. virginica at protected sites contrasts with studies north of Cape Hatteras where the abundance of predators produces a more diverse and heterogeneous community. Predation was unimportant in Beaufort because predators were absent at the exposed areas and the oyster drill Urosalpinx cinerea was restricted to the subtidal zone at the protected site. This absence of predators indicates a higher level of environmental stress near Beaufort compared to areas farther north.     

Genetics of larvae and spat growth rate in the oyster Crassostrea virginica

The heritability of oyster (Crassostrea virginica) larval growth rate was estimated to be in the range of 0.25 to 0.50 and a significant part of this genetic variation is of the additive type. Larval growth rate and spat growth rate were found to be highly correlated. These results suggest that a selection program for faster growing larvae and spat would be successful.     

The ecological energies of growth,respiration and assimilation in the intertidal American oyster Crassostrea virginica

Seasonal variations in the growth, respiration and assimilation of the intertidal oyster Crassostrea virginica (Gmelin) of different sizes were determined. The instantaneous growth rates for intertidal oysters decreased with increasing size and with lower temperatures. Q₁₀ values computed from instantaneous growth rates were approximately 2 during the warm growing season, but were higher in the colder months. Oxygen consumption increased with temperature and body size. A model was developed to predict oxygen consumption at any environmental temperature from 10° to 30°C for oysters ranging in weight from 0.1 to 100.0 g. Q₁₀ values computed from oxygen-consumption rates decreased with increasing temperature and increasing body size. Intertidal oysters utilize a large proportion of their assimilated energy in growth.Supported by a Belle W. Baruch Fellowship in Marine Ecology.