Observations on shell deformities,ultrastructure, and increment formation in the bay scallop Argopecten irradians

Shell morphology and ultrastructure were examined in the bay scallop Argopecten irradians, cultured in recirculating seawater systems under various conditions of feeding, lighting, and handling. On a unialgal diet of Thalassiosira pseudonana, scallop growth ranged from 120 to 183 m d^-1 at 20°C in the laboratory, about two-thirds of the growth rate found in the field. However, shell deposited in the laboratory differed from that in the field in several ways. In the field, scallops formed costae as an unpigmented, corrugated marginal shell layer; shell deposited in the laboratory lacked this layer and was therefore darker. Also, microstructure of the exterior shell surface of field scallops was coarsely granular, while that of cultured scallops was relatively smooth. Excessive handling of scallops in the laboratory resulted in marginal thickening of valves, a deformity which was completely arrested by a change from daily to weekly handling. Scallops cultured in the same tank with oysters developed shell-thickening on the interior of the valves. It is postulated that shell abnormalities in bivalves result from disruption of complex behavioral processes associated with shell deposition and may be elicited by a variety of natural and experimental irritants. Under natural lighting regimes and optimal conditions for growth, scallops deposited exactly one shell increment per day, but under continuous lighting, deposition of growth increments often became aphasic. In one 28-d experiment, there was a strong correlation between number of growth increments formed and increase in shell height, suggesting that shell ridge formation occurred intermittently, rather than daily, when shell growth rates fell below approximately 150 m d^-1.     

Gpi genotypic effect on quantitative traits in the northern bay scallop,Argopecten irradians irradians

The relationships between glucose-6-phosphate isomerase (Gpi) genotype and quantitative variation related to reproduction and growth were explored over the lifespan of a single cohort of northern bay scallops, Argopecten irradians irradians (Lamarck), from September 1986 to April 1988 in the Niantic River estuary, Connecticut, USA. Analyses revealed that Gpi genotype explained an increasingly significant proportion of variation in scallop size (up to 15%) as the cohort aged. The genotype-specific effects were consistent across sampling dates and among measured traits. There was no evidence for heterosis with respect to size at this locus; however, rare genotypes contributed substantially to the relationship and showed some tendency to fall on the extremes of the phenotypic distribution. The strength and consistency of the genotypic effect on scallop size suggest that genetic variation for Gpi, or some locus in linkage disequilibrium with Gpi, may translate into biochemical and/or physiological variation and affect fitness in this species.     

Cadmium accumulation and toxicity in the kidney of the bay scallop Argopecten irradians

Specimes of bay scallops, Argopecten irradians, collected from shallow-water eel-grass beds near Beaufort, North Carolina (USA) in September 1979, were exposed to 0.7 ppm cadmium in flowing seawater for 5 d. This exposure resulted in a massive extrusion of the calcified concretions of most of the kidney epithelial cells, although marked morphological damage consisting of cytoplasmc degeneration was apparent in only a few focal areas of the kidney. In addition, unique cytoplasmic membrane-bound bodies were observed in epithelial cells of cadmium-treated but not control scallop kidneys. In some cells, these bodies appeared to fuse with the main concretion vacuole. Kidneys of cadmium-treated scallops accumulated cadmium to 200 ppm on a wet wt basis; of this 60% was associated with concretions (2 000 ppm dry wt) and 38% with the membranous pellet obtained after ultracentrifugation at 105 000 g for 1 h. Approximately 2% of total kidney cadmium was associated with the cytosolic fractions but, unlike zinc or manganese, which were bound to either high or low molecular weight species, a large component of the cadmium in this fraction was bound to a protein peak of approximately 21 000 daltons. Results of this study indicate that kidney concretions of A. irradians play a major role in the control of renal cadmium accumulation and excretion and hence the toxicity of cadmium to this organism.     

Combined effects of temperature and salinity on embryos and larvae of the northern bay scallop Argopecten irradians irradians

The combined effects of temperature and salinity on embryonic development and on larval survival and growth to setting size of the northerm bay scallop Argopecten irradians irradians (Lamarck) were studied in the laboratory. A 6x6 complete factorial design was used; temperatures ranged from 10° to 35°C, at 5C° intervals, and salinities ranged from 10 to 35S, at 5S intervals. Response-surface contour diagrams were generated to provide estimates of conditions for optimal responses. Normal development of embryos occurred over a very narrow range of temperature and salinity. Survival of larvae occurred over a wider range of temperature and salinity than did embryonic development or growth of larvae. Satisfactory growth (>70% of the maximum observed value) occurred only at high temperature-high salinity conditions; optimal conditions for survival occurred at similar salinities, but at slightly lower temperatures. Temperatures of 35°C or greater and/or salinities of 10S or less were lethal for all life stages studied. Both salinity and temperature exerted significant effects on development and survival, but temperature was clearly the dominant factor influencing growth. It is suggested that northern bay scallop embryos and larvae be reared at their respective optimal temperature-salinity levels so as to increase efficiency of aquaculture operations.This paper is adapted from a thesis submitted to the College of Fisheries, University of Washington, in partial fulfillment of the requirements for the MS degree. This study was conducted at the NMFS Laboratory in Milford, Connecticut, USA     

Resource allocation and population genetics of the bay scallop,Argopecten irradians irradians: effects of age and allozyme heterozygosity on reproductive output

The relationships among multiple-locus heterozygosity, age, reproduction and growth were examined over the ca. 2-yr lifespan of a cohort of northern bay scallops, Argopecten irradians irradians (Lamarck), from the Niantic River estuary, Connecticut, USA. Electrophoretic analyses revealed a relatively low proportion of polymorphic loci (=0.35) and low level of heterozygote deficiency ( D. across 6 loci=-0.05) in this population. Allele frequency distributions showed a high degree of temporal stability within and among cohorts. No significant correlation was found between multiple-locus heterozygosity and either somatic or reproductive growth. Our results lend support to the postulated linkage between low heterozygote deficit and lack of a heterozygosity/growth association in marine bivalves, and in the pectinid group in particular. Partial reproductive senility occurs in these scallops in their second year of life. Somatic tissue weight increased exponentially with increasing shell height, whereas pre-spawning gonad mass attained asymptotic values in 2-yr-old scallops. Thus, weight-specific reproductive effort was significantly lower in second- than first-year scallops (24 and 33% respectively). This pattern could be generated if high individual reproductive effort in the first year were correlated with accelerated (early) post-reproductive senescence. This kind of resource allocation, in which somatic growth is given precedence over reproductive growth in older individuals, has been previously reported in only one other bivalve, a pectinid (Chlamys islandica), in which the decline in reproductive effort occurs only after the scallops reach an age of ca. 20 yr.     

Substrate catabolism related to reproduction in the bay scallop Argopecten irradians concentricus,as determined by O/N and RQ physiological indexes

Weight specific rates of oxygen consumption carbon dioxide production, and ammonia-N excretion, measured for a Florida population of the bay scallop Argopecten irradians concentricus between May and September, 1982 and October and November, 1983 were significantly correlated (P<0.0005) to environmental factors that co-varied seasonally with metabolic shifts related to reproduction. Mean O/N and CO₂/O₂ (RQ) molar ratios indicates that scallop energy metabolism varied over the course of the reproductive cycle. Resting stage individuals (May-early June) had RQ values close to 0.7, indicative of a predominantly lipid-based metabolism. During the initial stages of gametogenesis (late June-early July) scallops catabolized primarily carbohydrate, as evidenced by maximum O/N (>22) values and RQ values close to 1.0 RQ values >1.0 indicated a possible carbohydrate to lipid conversion during the period of cytoplasmic growth (late July-early September). As gametes matured and spawning commenced (late September-November), metabolism became primarily protein based, as indicated by O/N and RQ values around 9.0 and 0.8, respectively. This pattern of substrate catabolism supports existing data on the storage and utilization of specific energy reserves with respect to reproduction in this species.     

Mercury stress on juvenile bay scallops,Argopecten irradians,under various salinity-temperature regimes

Survival of juvenile bay scallops, Argopecten irradians, after 96 h of exposure to mercury at various salinity and temperature regimes was studied. Scallop survival was significantly affected by mercury concentration and by salinity, as well as by the interaction between temperature and concentration and between temperature and salinity. Toxicity of mercury at low concentrations was enhanced by high temperature and low salinity, whereas at high mercury concentrations this effect diminished.     

Geographic variation in a predator-induced defense and its genetic basis

Predator-induced morphological defenses are a well-known form of phenotypic plasticity, but we continue to have a limited understanding of geographic variation in these responses and its genetic basis. Here we examine genetic variation and geographic differentiation in the inducible defenses of tadpoles (Rana pirica) in response to predatory salamander larvae (Hynobius retardatus). To do so, we crossed male and female frogs from a "mainland" Japanese island having predaceous salamanders and a more isolated island not having predaceous salamanders and raised resulting offspring in the presence and absence of H. retardatus. Mainland tadpoles exhibited a higher capacity to express the inducible morphology (a more bulgy body) than those from the predator-free island, and expression of the bulgy morph in mainland-island hybrids produced phenotypes that were intermediate to those produced by pure crosses. In addition, parental sex had no effect on expression of the bulgy morph. Our results support the hypothesis that geographic variation in inducible defenses is linked to the additive effects of autosomal alleles that are shaped by differences in historical exposure to the inducing predator.     

Geographic variation,niche breadth and genetic differentiation at different geographic scales in the mussels Mytilus californianus and M. edulis

Genetic differentiation was investigated in the marine mussels Mytilus californianus Conrad and M. edulis Linn. from the west coast of North America. In allopatry with M. californianus, M. edulis occurs throughout the intertidal zone; however, in microgeographic sympatry its ecological range is restricted to above the M. californianus mussel bed and to patches of substratum opened by natural disturbances within the bed. Over the same geographic scale, the broader-niched M. edulis shows greater among-locality genetic difference and greater levels of polymorphism than M. californianus at two enzyme loci. Genetic differentiation on a geographic basis was investigated in M. californianus at a single rock (on a scale of meters), on an island (on a scale of kilometers), throughout a strait (on a scale of 10² kilometers), and along the west coast of North America (on a scale of 10³ km). Differentiation was minimal over the west coast, and could be explained by microhabitat differences in a local area. The minimal differentiation in west coast mussels relative to strong geographic differentiation of M. edulis on the east coast of North America may be related to the steeper latitudinal thermal gradient on the east coast. Local ecologically related microgeographic variation can result in biased and misleading estimates of genetic heterogeneity but microgeographic variation at enzyme loci may not be due to selection at the loci investigated or may even be due to the breeding structure of the mussel populations, as suggested by Tracey et al. (1975).     

Microgeographical shell variation in Littorina striata, a planktonic developing periwinkle

Littorina striata is a strictly Macaronesian, intertidal periwinkle with planktonic development. The species produces both nodulose and smooth shells, which co-occur at Ilheu de Vila Franca do Campo, a drowned crater situated about 1000?m off the south coast of São Miguel, Azores. The present work describes and analyzes the shell variation, temporal change and ecological distribution of the two shell types at this crater over a 3-year period. Nodulose shells were more common in the sheltered lagoon inside the crater, while smooth specimens dominated the outside of the wave-exposed crater. Moreover, nodulose specimens were smaller and weighed less compared to smooth ones. However, regardless of morphotype, shells from the lagoon had a smaller aperture and were less globose than those from the outside. Within an exposure regime, smooth specimens had a larger aperture. These patterns remained constant over time. They are tentatively interpreted as functional adaptations to thermal stress and wave exposure. The mechanisms that maintain these patterns are still unknown. Yet, aperture height of specimens transplanted from the lagoon to the outside increased markedly over a period of 5 months, whereas no comparable changes were observed in other experimental groups (i.e. transplanted from the outside into the lagoon, transplanted within the outside and transplanted within the lagoon; the latter two transplant groups being the blancos). This suggests that at least the aperture size in L. striata may be an ecophenotypically plastic trait.     

Temporal variation of recruits as a basis of ephemeral genetic heterogeneity in the western rock lobster Panulirus cygnus

Allozymes were examined in quantitative lunar monthly collections of larval recruits of the western rock lobster Panulirus cygnus George over three recruitment seasons at two sites nearly 350 km apart in Western Australia. At Alkimos, the southern site, recruitment occurs in a relatively narrow peak early in the spring, whereas at the northern Houtman Abrolhos Islands, recruitment extends into the summer months. In the 1995/1996 recruitment season, the frequency of the GPI ^* 100 allele increased from early to late in the season, but the frequencies were indistinguishable at the two sites in each monthly collection. The combination of this temporal variation in allelic frequencies with the contrasting patterns of recruitment at the Abrolhos Islands and Alkimos resulted in genetically different cohorts at the two sites. This pattern was ephemeral, as it was not repeated in the subsequent two years. Thus, ephemeral genetic patchiness in P. cygnus can be generated by the locally-specific genetic mix of recruits obtained from a common larval pool. This mechanism is the probable explanation of previously observed temporal and possible spatial genetic variation in adult P. cygnus, and highlights the importance of studying recruitment in order to understand the genetic structure of marine species. Received: 22 February 1999 / Accepted: 8 June 1999     

Origin,morphology and fate of the nonmineralized shell of Coryphella salmonacea,an opisthobranch gastropod

Little is known about initial shell formation in molluscs or about the possible functions of the embryonic shell field invagination (SFI). The present paper describes formation and loss of an unusual shell in embryos of the nudibranch Coryphella salmonacea (Couthouy, 1838), collected in February–March, 1981–1984. The first organic shell material was attached to shell field cells just outside the SFI at two weeks after fertilization (5°C). As viewed with transmission electron microscopy, the organic shell in section consisted of two electron dense layers having a total thickness of 16 nm. The shell continued to grow through at least the fifth week of encapsulated development. In the eighth week, the shell still covered the visceral mass; there was no evidence that mantle tissue reflexed and covered the shell or that shell materials were resorbed by the mantle. The shell and sometimes the operculum were discarded within the embryonic capsule prior to hatching at the end of Week 8. The cup-like shell is unusual in that it was shed as a wrinkled organic sheet which apparently lacked mineralization. Fully-developed shells were non-birefringent and did not produce calcium peaks when examined by electron microprobe. Both the location of the first organic shell material outside the SFI and the apparent absence of shell mineralization in the presence of an SFI are consistent with the hypothesis that the SFI is nonsecretory. C. salmonacea is probably in the evolutionary process of losing its shell, a structure of little or no function during encapsulated, nonplanktonic development.Contribution No. 130 of the Marine Science Laboratory, Northeastern University     

Physiological correlates of genetic variation for rate of behavioral development in the honeybee, Apis mellifera

Two factors that influence age at onset of foraging in honeybees are juvenile hormone (JH) and colony age demography (older bees inhibit behavioral development of younger bees). We tested the hypothesis that genetic variation among bees for these factors influences genetic variation in behavioral development. Pairs of colonies showing genetic differences in rates of behavioral development were identified in a screening experiment and bees from these colonies were used for physiological and behavioral assays. Six pairs were assayed, three with European bees only and three with both European and Africanized bees. There was genetic variation for the following four components: (1) production of JH in four pairs (experiment 1); (2) sensitivity to JH in three pairs (experiment 2); (3) sensitivity to social inhibition in three pairs (experiment 3), and (4) potency of social inhibition in four pairs (experiment 4). Cross-fostering assays (experiment 5), which allowed all four components to be evaluated simultaneously, revealed genetic variation for production of JH, sensitivity to JH, or sensitivity to social inhibition in five of six pairs, and potency of social inhibition in five of six pairs. There was often evidence for genotypic differences in more than one component, and no consistent pattern of association among any of the components. Africanized bees had faster rates of behavioral development than European bees, but there were no racial differences in patterns of variation among the four components. These results indicate that there are at least several, apparently distinct, physiological processes associated with JH and colony age demography upon which natural selection can act to alter the rate of behavioral development in honeybees. Received: 8 December 1998 / Received in revised form: 29 July 1999 / Accepted: 8 August 1999     

A biochemical genetic study of population structure in queen scallop (Chlamys opercularis) stocks in the Northern Irish Sea

Electrophoresis was used to survey a number of enzymes for loci of potential value as stock markers in Chlamys opercularis (L.). Two loci, Pgi and Pgd, were then studied in larger samples from each of two localities around the Isle of Man. One Pgi allele was found to be restricted to a particular year-class in both populations. This was confirmed by subsequent studies of three further populations, in each of which the same allele occurred only in the one year-class. Otherwise, no significant differences in allele frequencies were found between year-classes or between populations in any of the samples used. From the results it is concluded that there is temporal variation in the sources of recruitment of C. opercularis stocks around the Isle of Man. Possible origins of larvae contributing to these stocks are discussed in the light of what is known of current circulation patterns in the northern Irish Sea.     

Burrow morphology,tube formation,and microarchitecture of shell dissolution by the spionid polychaete Polydora websteri

Burrow morphology, detrital tube formation within the burrow, and ultrastructure of the surface of the burrow of Polydora websteri Hartman, 1943, are described in the shells of Crassostrea virginica (Gmelin) and of Mytilus edulis Linné. Observations were made on live worms living normally in artificial preparations of polished shell covered with transparent plastic film. P. websteri is capable of settling wherever there are crevices in shell surfaces, and slowly penetrates the shell forming a U- or flask-shaped cavity. A detrital tube is formed within the burrow from detritus collected and pressed in place by the worm. During this process, the modified setae of the 5th setiger are thrust against the wall of the inner tube and help to keep the diameter of the inner tube constant throught its length. Individuals of P. websteri burrow into the shell when placed on preparations of polished shell of Mytilus edulis under plastic film. As they do so, they secrete a viscous fluid which dissolves interprismatic and interlamellar organic matrices and then dissolves the exposed crystals. Etched prisms and lamellae reveal a complex pattern of internal dissolution. Enlargement of the burrow by the worm thus takes place by chemical dissolution of shell and also probably by flushing of loosened partially dissolved prisms and lamellae from the tube. Some shell fragments become incorporated in the walls of the detrital tube. There is no evidence of setal abrasion on the shell walls of the burrow even at the ultrastructural levelUniversity of Delaware College of Marine Studies Contribution No. 99.     

Postingestive selection in the sea scallop (Placopecten magellanicus) on the basis of chemical properties of particles

Suspension-feeding bivalves are adapted to use a highly variable mixture of particles by sorting them before and after ingestion. Postingestive sorting in bivalves has been confirmed for several bivalve species, but few studies have attempted to isolate the factors influencing postingestive selection among different particles presented simultaneously. The ability of the sea scallop, Placopecten magellanicus (Gmelin), to sort a mixture of organic (¹⁴C-labeled dinoflagellates, Prorocentrum minimum) and inorganic (⁵¹Cr-labeled beads, diameter 16-18 µm) particles of similar size and shape within the stomach was examined. The study was carried out in August 1997 using scallops collected near Deer Island, New Brunswick, Canada. Sorting of the two particle types was measured by dissecting the scallops at intervals after feeding and comparing the ¹⁴C:⁵¹Cr ratios within the stomach, digestive gland, and feces. The ¹⁴C:⁵¹Cr ratio in the stomach decreased over time, indicating that sea scallops were sorting organic from inorganic particles. The ability of P. magellanicus to sort particles solely on the basis of chemical properties was tested in a second study by presenting them with a mixture of protein-coated and uncoated beads of two different colors. This study was carried out in February 2000 using scallops collected near Maces Bay, New Brunswick, Canada. Gut retention times of the two types of beads were measured using flow cytometry. Scallops retained protein-coated beads in the gut longer than uncoated beads, indicating postingestive selection by chemical properties. This study provides the first evidence for simultaneous postingestive sorting by bivalves of particles based solely on chemical properties. This would potentially enable P. magellanicus to preferentially retain particles of higher food quality longer than those of poor quality, thereby enhancing digestive efficiency.     

Vocal dialect and genetic subdivisions along a geographic gradient in the orange-tufted sunbird

At least four hypotheses have been suggested to explain the formation and maintenance of song dialects among birds: historic processes (epiphenomenon), genetic or local adaptation, acoustic adaptation, and social adaptation. We studied spatial and temporal distribution of dialect in the orange-tufted sunbird (Nectarinia osea), a small nectarivorous bird that expanded its breeding range in Israel during the past 100 years from the southern part of Rift Valley to the entire country. Sunbird range expansion was concurrent with the establishment of many small settlements with an ethos of gardening, which introduced many ornithophilous plants. We recorded songs and genetically screened individual sunbirds in 29 settlements distributed across a 380 km north–south gradient along the Rift Valley. We show that dialects cluster together into geographical regions in 70% of cases, a moderate concurrence to geography. Settlement establishment date, geographical position, and genetic distance between local populations (i.e., settlements) were all poor predictors for the variance among song dialects. The specific effect of habitat was not tested because all sampled localities were similar in their physical and acoustic properties. Using a network analysis, we show that dialects seem to aggregate into several network communities, which clustered settlement populations from several regions. Our results are best explained by either the epiphenomenon hypothesis or the social adaptation hypothesis, but at present our data cannot state unequivocally which of these hypotheses is better supported. Last, we discovered a negative association between network centrality and genetic diversity, a pattern that requires further examination in other systems.     

Seasonal variation in the properties of muscle mitochondria from the tropical scallop Euvola (Pecten) ziczac

In scallops, gametogenesis leads to considerable transfer of energetic reserves from the adductor muscle to the gonads. During an annual cycle, the scallops are exposed to changes in temperature and food availability. As these changes may affect muscle metabolic capacities, we examined whether the properties of the mitochondria in the phasic adductor muscle were modified during the annual cycle of the scallop Euvola (Pecten) ziczac (L. 1758). During our study, temperature and chlorophyll a levels generally showed an inverse relationship: high temperatures and low chlorophyll a levels occurred from mid-April to early June. Lower temperatures and higher chlorophyll a levels were found from January to late March and from mid-June to mid-September. Throughout the annual cycle, the substrate preferences and the pH sensitivity of the isolated muscle mitochondria changed little, whereas the maximal oxidative capacities and respiratory control ratios (RCR) varied considerably. Consistently, the maximal capacities for substrate oxidation were 30 to 80% lower in mitochondria isolated in May than at other times in the year. The RCR values of mitochondrial oxidation of glutamate, glutamine and succinate varied throughout the year with lower values characterizing the mitochondria from scallops harvested in May and in certain cases in August. In May, adductor muscles had lower protein levels than at other times. These data suggest that the requirements of gametogenesis, coupled with␣the high temperatures and low food availability occurring during April and May, led to a mobilization of muscle proteins which concomitantly decreased the oxidative capacity of isolated mitochondria. Received: 29 November 1996 / Accepted: 5 December 1996     

Clinal variation in anuran advertisement calls: basis for acoustic isolation?

Summary The calls of male treefrogs (Eleutherodactylus coqui) were studied along an altitudinal gradient in the Luquillo Mountains in eastern Puerto Rico. The fundamental frequency of each note in the advertisement call was highly negatively correlated with altitude. Although males responded vigorously to playbacks of calls of other males recorded at their own altitude, males at high altitudes responded less frequently to recorded calls of males from low altitudes, and males at low altitudes responded less frequently to recorded calls of males from high altitudes. These results are discussed in relation to potential isolation in contiguous populations of E. coqui.