Mortality and growth of juvenile hard clams Mercenaria mercenaria during brown tide

Blooms of the picoplankton-sized alga Aureococcus anophagefferens Hargraves et Sieburth, also known as "brown tide," frequently recur in Great South Bay, New York, USA. A field study compared mortality and growth rates of juvenile hard clams, Mercenaria mercenaria (L.), raised at a site that experienced a bloom of A. anophagefferens (>1᎒⁶ cells ml^-1) to clams grown in a nonbloom site from May to September 2000 at two sites on the north and south shores of Long Island. High (67%) cumulative clam mortality was observed for clams raised at the brown tide site (south shore) during the bloom (June-July), and individuals exhibited no measurable growth during that period. After the bloom subsided (mid-July), clams that survived the brown tide suffered low (<5%) cumulative mortality over the remaining sampling period (July-September). These survivors exhibited rapid growth for 4 weeks shortly after the bloom's decline. Subsequently, their growth slowed to rates comparable with those observed at the nonbloom site (north shore). By contrast, a low (<5%) cumulative mortality was observed for M. mercenaria raised at the site with no brown tide, and positive growth occurred throughout the study. These data suggest that some juvenile clams not only survive brown tide but also recover following the bloom.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

Effects of exposure to toxic and non-toxic dinoflagellates on oxygen consumption and locomotion in stage 1 larvae of the crabs Cancer oregonensis and C. magister

Sublethal effects on larval crabs upon exposure to toxic dinoflagellates were examined in the laboratory in early 1999. Specifically, oxygen consumption rates and geotaxis responses were determined for stage 1 larvae of the crabs Cancer oregonensis (Dana) and C. magister Dana that were exposed to non-toxic (Alexandrium tamarense, strain 115) or toxic (A. fundyense, strain 1719) dinoflagellates or to freshly hatched nauplii of the brine shrimp Artemia sp. In C. oregonensis, larvae exposed to the toxic dinoflagellate showed reduced rates of oxygen consumption compared to those exposed to non-toxic dinoflagellates or brine shrimp nauplii. Larvae exposed to a filtrate of the non-toxic dinoflagellate showed no change in oxygen consumption, but a reduced rate when exposed to filtrate from the toxic alga at densities >5᎒² cells ml^-1. In C. magister, larvae exposed to the non-toxic A. tamarense or the toxic A. fundyense had reduced oxygen consumption rates. Larvae exposed to filtrates of non-toxic and toxic dinoflagellates had no change in oxygen consumption. In geotaxis tests, C. oregonensis larvae exposed for 1 day to the toxic A. fundyense reduced their level of locomotion compared to those exposed to non-toxic A. tamarense or to brine shrimp nauplii. C. magister larvae showed no change in activity after a 1-day exposure to the toxic A. fundyense. After a 4-day exposure to A. fundyense, C. magister larvae had much reduced locomotion. Reduced locomotory activity in larvae exposed to toxic algae is consistent with the changes in oxygen consumption rates. Responding to exposure to toxic algae by reducing locomotion may affect vertical migration in these negatively buoyant crab larvae, resulting in sinking below a toxic alga bloom, at least temporarily.     

Environmental controls on shell growth rates and δ<Superscript>18</Superscript>O of the shallow-marine bivalve mollusk <Emphasis Type="Italic">Phacosoma</Emphasis><Emphasis Type="Italic">japonicum</Emphasis> in Japan

Microgrowth patterns and the oxygen isotope composition of juvenile, shallow-marine bivalve mollusk shells of Phacosoma japonicum (Reeve) in Japan were analyzed and cross-calibrated with environmental parameters. Mark-and-recovery experiments indicate that a pair of two microgrowth lines and two microgrowth increments is produced every lunar day. This finding makes it possible to assign exact calendar dates to each portion of the shell. Average daily growth rates decrease by 61% from age two to three and 55% from age three to four. The length of the growing season and the growth rate are mainly controlled by temperature: shell growth ceases below 14.2°C (age two) and 16.8°C (age four) and is most rapid between 24.6°C and 27.2°C. Based on local temperature cycles, the growing season is longest in Seto Inland Sea, central Honshu (from May to November) and shortest at Hakodate Bay, North Japan (from June to October). The annual oxygen isotope profiles of the shells reflect the temperature cycle and the varying amounts of freshwater added to the seawater by precipitation. The most negative '¹⁸O values of -3.15‰ occur during the rainy season, i.e. during the monsoon and typhoon seasons. Growth rates are only slightly affected by salinity changes. Strongly reduced growth rates during the second half of the year at Seto Inland Sea and to a lesser extent at Tokyo Bay are explained by nutrient deprivation. Our study provides the basis for the use of P. japonicum in high-resolution ecological studies and environmental reconstructions.     

Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus

The chemical composition of kelps (e.g. polyphenolics) deters grazing by herbivores, but kelp detritus is potentially a source of nutrition for suspension feeders. The effects of kelp detritus derived from two species [Agarum fimbriatum Harvey and Costaria costata (Turner) Saunders] on feeding of oysters, Crassostrea gigas Thunberg, and mussels, Mytilus trossulus Gould, were examined in feeding experiments. Fresh and aged kelp particles were sequentially presented in combination with the microalga Rhodomonas lens at an initial total concentration of 5᎒^-4 ml^-1. Aging of kelp particles for 4 days in seawater significantly reduced the concentration of polyphenolics without changing the total carbon or nitrogen content. Clearance rates of both mussels and oysters were significantly lower in the presence of fresh versus aged kelp particles, and clearance rates declined overall with declining polyphenolic concentrations. Video endoscopy was used to examine feeding selectivity at the level of the gill in oysters in the same food treatments used in the clearance rate experiments. Comparison of particle composition in the water versus the pseudofeces in both oysters and mussels was also used as a measure of feeding selectivity. When presented with R. lens in combination with fresh and aged kelp particles selectivity for R. lens tended to be greater against fresh than aged particles, and there was some indication that this was stronger for A. fimbriatum than for C. costata particles. The ability to select was lower at very high polyphenolic concentrations, which may reflect poisoning of sensory binding sites. These data suggest that bivalves distinguish among particles of varying chemical composition and respond by changing their clearance rates and their selectivity.     

Diel movement patterns of the Hawaiian stingray,<Emphasis Type="Italic"> Dasyatis lata</Emphasis>: implications for ecological interactions between sympatric elasmobranch species

The Hawaiian stingray, Dasyatis lata, is a common benthic elasmobranch in nearshore Hawaiian waters. Acoustic telemetry was used to track the movements of seven rays in Kaneohe Bay, Oahu, Hawaii. Rays were tracked continuously over 31-74 h periods. Geographical movements were analyzed to determine space utilization and rate of movement. Rays were found to utilize significantly larger activity spaces at night (0.83ǂ.70 km²) (mean-SD) than during the day (0.12ǂ.15 km²). Mean total activity space for rays tracked was 1.32ǂ.75 km². Rates of movement were also significantly higher at night (0.34ǂ.30 km h^-1) than during the day (0.15ǂ.22 km h^-1). Average straight-line swimming speed was 0.64ǂ.16 km h^-1, with a maximum observed swimming speed of 1.9 km h^-1. Tidal stage had no effect on rate of movement. Comparison with previously published data on juvenile scalloped hammerhead sharks, Sphyrna lewini, in Kaneohe Bay revealed a high degree of overlap in habitat use and time of activity, suggesting possible ecological interactions between these two species.     

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.     

Ultrastructural,biochemical, and immunological characterization of two populations of the mytilid mussel <Emphasis Type="Italic">Bathymodiolus</Emphasis><Emphasis Type="Italic">azoricus</Emphasis> from the Mid-Atlantic Ridge: evidence for a dual symbiosis

A recently described species of mytilid mussel, Bathymodiolus azoricus Von Cosel et al., 1999, was observed to be the dominant organism at the hydrothermal vents off the Azores, at both the Lucky Strike and Menez Gwen sites. Evidence suggests this species of Bathymodiolus represents yet another example of the intriguing dual symbiosis known in three other species of deep-sea mytilid mussels. Transmission electron micrographs (TEM) show the majority of gill bacteriocytes in mussels sampled from both populations to contain two distinct symbiont morphotypes. One morphotype is characterized by large size (mean diameter, 1.25 µm), coccoid shape, and stacked intracytoplasmic membranes that are consistent with the morphology of type I methanotrophs. The second morphotype is smaller (mean diameter, 0.35 µm) and was observed in coccoid or rod shapes. Immunoblots revealed the presence of ribulose 1,5-bisphosphate carboxylase/oxygenase (RubisCO) and methanol dehydrogenase (MeDH) in both populations of mussels. Activities of these enzymes, as well as sulfate adenylyl transferase (ATP sulfurylase) and adenylyl sulfate reductase (APS reductase), were detected in gill extracts. The activities measured for the two populations were highly variable, though the population sampled from Lucky Strike showed higher RubisCO activity. Stable carbon isotope values (Lucky Strike, '¹³C=-32.6ǂ.3‰; Menez Gwen, '¹³C=-22.8ǂ.4‰) are in the range of previously reported stable carbon isotope measurements for mytilid mussels hosting a dual symbiosis. Collectively, these results provide evidence for the activity of both sulfur-oxidizing and methane-oxidizing metabolic pathways in B. azoricus. Furthermore, evidence for a greater dependence on methanotrophy in the Menez Gwen mussel population is offered by analysis of cell counts from TEMs. Higher methanotroph numbers, and putatively activity, in this population of mussels are further supported by published geochemical data indicating higher methane concentrations in the vent fluids at Menez Gwen. This finding suggests that environmental conditions may regulate a balance between the physiological activities of different symbiont populations associated with these mussels. The existence of a dual symbiosis could thus confer greater environmental tolerance and increased niche space to the mytilid host in the stochastic hydrothermal vent habitat.     

Combined effect of temperature and food concentration on the grazing rate of the rotifer Brachionus plicatilis

We developed a predictive relationship to determine the grazing rate of Brachionus plicatilis at given temperatures and food concentrations; this function could be applied to experimental culturing and aquaculture practices. Grazing experiments were conducted at temperatures between 5°C and 40°C and at food concentrations, of the flagellate Isochrysis galbana, ranging between ~0 and 10⁶ ml^-1. In total, 136 grazing rates were determined, using the prey depletion method, for rotifers acclimated to treatments for 0.5 or 4 h. The response of grazing rate to temperature and food concentration was described using a model that combined a rectangular hyperbolic function for food concentration and a sigmoidal function for temperature. Using non-linear curve-fitting methods an equation was obtained: G=(452F)/(159000+F)Ǵ.94/(1+2190002T^-4.35) , where G is the grazing rate (flagellates rotifer^-1 min^-1), F is the food concentration (flagellates ml^-1), and T is temperature (°C). The equation indicates a maximum grazing rate of ~35 prey rotifer^-1 min^-1, above ~4᎒⁵ prey ml^-1 and 25°C.     

Trace metal ingestion and assimilation by the green mussel <Emphasis Type="Italic">Perna viridis </Emphasis>in a phytoplankton and sediment mixture

We examined trace metal ingestion and assimilation (Cd, Se, Zn) by the green mussel Perna viridis in a mixture of diatoms and sediment at concentrations below and above the pseudofeces production levels. Dual gamma radiotracers (¹⁰⁹Cd, ⁶⁵Zn) were used to investigate particle selection on marine diatoms and sediments. The diatom (Thalassiosira weissflogii) was radiolabeled with ¹⁰⁹Cd, and the natural sediment was radiolabeled with ⁶⁵Zn. By comparing the ratios of ¹⁰⁹Cd:⁶⁵Zn in the particle mixture, mussel tissues and pseudofeces within a short-term exposure period (35 min), the results demonstrated that the green mussels were able to selectively ingest the diatom particles at a high particle load. Efficiency of selection for nutritious particles (e.g. diatoms) increased with increasing ratio of sediments in the particle mixture. Pseudofeces contained a higher ratio of sediments relative to that in the feeding suspension. No major particle selection was observed at concentrations below the level for pseudofeces production. The assimilation of Cd, Se and Zn by the green mussels was quantified using a pulse-chase feeding technique. The assimilation of Se and Zn by the green mussels from ingested diatoms was reduced with the presence of sediment within the mussel gut, presumably due to the resorption of metals onto the sediment, leading to a quicker passage of metals through the digestive tract and a lower proportion of metals subjected to intensive digestion. In contrast, the presence of diatoms did not significantly affect metal assimilation from ingested sediment. A significant correlation between metal assimilation efficiency and metal gut passage time was also observed. Metal assimilation by the green mussels appeared to be little dependent on the particle concentration in seawater. Our study suggests that particle selection may potentially alter metal influx from ingested food sources, particularly at high particle concentrations. Selective feeding of nutritious particles, coupled with a high assimilation efficiency from these ingested particles, may increase metal influx into mussels from the dietary phase.     

Large viruses and infected microeukaryotes in Ross Sea summer pack ice habitats

A variety of Ross Sea summer pack ice habitats between 66 and 75°S were examined for viruses 𔒦 nm capsid diameter. Maximum abundances of these viruses likely to infect eukaryotes were 10⁶-10⁷ ml^-1 brine in surface, interior, and bottom habitats and constituted up to 18% of the total (all sizes) viruses. There is abundant ultrastructural evidence for infection of a variety of microheterotrophs and some autotrophs. One station exhibited the classical characteristics of a lower latitude algal bloom with potential viral control. The blooming alga, Pyramimonas tychotreta Daugbjerg 2000, was infected, as were two abundant heterotrophs, Cryothecomonas spp. and an unidentified flagellate, that fed on P. tychotreta. Infections were observed in only one life history stage (multiflagellate cells) of P. tychotreta, suggesting a relationship among virus-induced lysis, life-history stages, physiology, and environmental factors regulating the life cycle. There is good evidence that diatoms are not a likely source of the large viruses, and viruses in general are not a major food source for ice microheterotrophs in summer.     

Foraging behavior of juvenile Carcinus maenas (L.) and Cancer pagurus L.

Information concerning the way juvenile crabs choose their diet from a variety of prey types can be useful for a better understanding of community dynamics, as well as for the adequate management of natural resources. Prey size and species selection by juvenile Carcinus maenas (15-35 mm carapace width, CW) and Cancer pagurus (20-40 mm CW) feeding on four bivalves of contrasting shell morphology were investigated. When offered a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas, and Cerastoderma edule presented individually, crabs generally showed evidence of size-selective predation. Cancer pagurus selected larger mussels relative to the size of their chelae (relative prey size, RPS) than did Carcinus maenas of similar and even larger carapace width. However, the RPS of selected O. edulis and Cerastoderma edule were similar for all crabs, suggesting that certain prey features constitute effective barriers even to the powerful chelae of Cancer pagurus. When offered a wide size range of mussels and oysters simultaneously, all crabs consistently selected mussels. When offered O. edulis and Crassostrea gigas, crabs consumed both these oyster species in similar numbers. Carcinus maenas consumed similar numbers of mussels and cockles; Cancer pagurus, however, showed no preference for either prey in the smaller size classes but selected more mussels than cockles as prey increased in size. Although previous studies report that adult Carcinus maenas select prey species according to their profitability (amount of food ingested per unit of handling time, milligrams per second), consumption rates of the size classes of prey selected by juvenile shore crabs did not always parallel prey value. Although variations in crab strength can account for many of the differences between the foraging strategy of juvenile and adult C. maenas, our results suggest that juvenile crabs are less species selective than adults as a result of the restrictions imposed on small individuals that have limited access to larger prey.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.     

Significance of food type for growth of ephyrae Aurelia aurita (Scyphozoa)

We studied growth of newly released Aurelia aurita ephyra larvae fed five different food types, including a large-sized copepod, a phytoflagellate, and suspended POM (particulate organic matter) made from bivalve meat. Experiments were run at saturated food concentration in two different temperatures over 10 days. The effect of small differences in temperature was inconsistent and interacted with the effect of food type, which, in turn, was highly significant. A low average growth rate (4-9% day^-1) was shown when feeding on the large-sized copepod Calanus finmarchicus (80 µg AFDW individual^-1), in spite of an extremely high daily ration of up to 1500% of body AFDW. When feeding on the cryptophyte Rhodomonas baltica (ca. 8 µm cell diameter), the ephyrae showed an average growth rate over the 10 day experiment of 7-11%, but with a considerably higher growth rate during the first days. Suspended POM generated an average growth rate of 7-9% day^-1, whereas fresh bivalve meat, manually placed into the stomach of the ephyra, gave an average growth rate of 12-14% day^-1. Artemia nauplii (ca. 3 µg AFDW individual^-1), used as a general reference, resulted in higher growth rates than any of the other food types (17-31% day^-1). We conclude that A. aurita ephyrae can capture and feed on phytoplankton, large copepods, and POM; that phytoplankton might be of nutritive significance early in development; and that the high quantity of large-sized copepods ingested is inefficiently converted to growth during early development. POM is a potential food source because of the ability of the ephyrae to encounter and ingest it, although concentration, size distribution, and nutritional composition of natural POM probably constrain its effect on growth.     

Population structure of two Antarctic ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, in winter sea ice

The spatial distribution and population structure of two dominant ice-associated copepods, Drescheriella glacialis and Paralabidocera antarctica, were studied during winter at nine locations in east Antarctic fast ice. These species accounted for at least 90% of the total metazoan abundance at each location. Abundances were high, reaching 175 individuals l^-1 (190,000 m^-2) for D. glacialis and 660 l^-1 (901,000 m^-2) for P. antarctica. These abundances were probably partly supported by the high biomass of ice-algae (Pearson correlation coefficient, r=0.75), as indicated by chlorophyll-a concentrations (1.7-10.1 µg l^-1). The population structures of each species suggested very different life-history strategies. All developmental stages of D. glacialis were isolated from the ice cores, including females with egg sacs, supporting the hypothesis that this species reproduces in the sea ice during winter. This strategy might assist D. glacialis in leading a continually colonising existence, whereby it responds opportunistically to the availability of favourable habitat patches. The populations of P. antarctica were composed primarily of nauplii (>99%), consistent with past observations of a synchronised life cycle for this species. The strong coupling of the developmental cycle of P. antarctica to the growth and decay of sea ice suggests that local extinctions might occur in areas where ice break-out is unpredictable.     

Deleterious effects of a nonPST bioactive compound(s) from <Emphasis Type="Italic">Alexandrium tamarense</Emphasis> on bivalve hemocytes

The known negative effects of shellfish toxin-producing dinoflagellates on feeding, burrowing and survival of some bivalve mollusks has prompted questions concerning whether they might also impair the internal defense system of affected bivalves and make them more susceptible to disease agents. The primary components of the cellular defense system are hemocytes. Many toxic dinoflagellates are too large to be ingested whole by hemocytes and would most likely be exposed to intracellular toxins only after the algae are consumed, broken down, and the water-soluble toxins, released. Therefore, we conducted a series of experiments in which hemocytes from two suspension-feeding bivalves—the Manila clam, Ruditapes philippinarum, and the softshell clam, Mya arenaria—were exposed in vitro to filtered extracts of one highly toxic paralytic shellfish toxin (PST)-producing and one nonPST-producing strain of Alexandrium tamarense (isolates PR18b, 76 ± 6 STXeq cell⁻¹ and CCMP115, with undetectable PST, respectively). We measured adherence and phagocytosis, two hemocyte attributes known to be inhibited by bacterial pathogens and other stressors. We found no measurable effect of a cell-free extract from a highly concentrated suspension of the PST-producing strain on hemocytes of either bivalve species. Instead, extract from the nonPST-producing strain had a consistent negative effect on both clams, resulting in significantly lower adherence and phagocytosis compared to strain PR18b and filtered seawater controls. The bioactive compound produced by strain CCMP115, which has yet to be characterized, may be similar to the PST-independent allelopathic compounds described for Alexandrium spp., which act on other plankters. These compounds and those produced by other harmful algae are known to cause immobilization, cellular deformation and lysis of co-occurring target organisms. Thus, nonPST producing Alexandrium spp., which do not cause paralysis and burrowing incapacitation of clams, may still produce a compound(s) that has negative effects not only on hemocytes, but on other molluscan cell types and their functions, as well.     

Settlement patterns of newly settled plaice (Pleuronectes platessa) in a non-tidal Swedish fjord in relation to larval supply and benthic predators

In demersal fish species with a pelagic larval stage, settlement patterns may be a consequence of variations in larval supply, habitat selection at settlement, and processes acting between the time of settlement and the time of benthic sampling. This study describes temporal (1994-1998) and spatial variation in plaice (Pleuronectes platessa L.) settlement densities in four semi-isolated nursery areas with similar habitat characteristics, in the non-tidal Gullmarsfjord on the west coast of Sweden. Juvenile abundance varied by a factor of ten, both among years and among nursery grounds. For the 3 years when larval sampling was undertaken (1994-1996) and all nursery areas, there was a significant positive relationship between larval supply and juvenile abundance (linear regression: r²=0.45, n=24, P<0.001). On the southern side of the fjord, a significant positive relationship between larval and juvenile abundance was found in one area (r²=0.62, n=6, P<0.05). The absolute mortality rate of plaice after settlement was related to the initial settlement density (r²=0.95, n=20, P<0.001), and to the abundance of predatory shrimps Crangon crangon (r²=0.44, n=20, P<0.01). Plaice otoliths were found in 6% of the shrimp stomachs analysed from an area with high density (13.3 m^-2) of newly settled plaice. The present study suggests that the density of juvenile plaice was limited by larval supply to the nursery grounds. Consistency in the relative abundance of juveniles among nursery grounds between years also suggested that some nursery areas may be in the settlement shadow of others. The irregular nature of the coastline in combination with larval depletion could thereby cause small-scale (10³-10⁴ m) variation in settlement densities of the same order of magnitude as the inter-annual variability in recruitment to individual nursery grounds.     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.     

Doubly uniparental transmission of mitochondrial DNA length variants in the mussel <Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">trossulus</Emphasis>

Doubly uniparental inheritance of mitochondrial DNA (mtDNA) is an unusual feature found in marine mussels and a few other bivalve species. In addition to a mitochondrial genome (F) inherited through the female line, heteroplasmic individuals (males) contain a second highly diverged genome (M) that is inherited through the paternal line. The Baltic mussel Mytilus trossulus is characterized by the presence of two phylogenetically close female and male genomes. We examined M. trossulus sampled from the Gulf of Gdansk, southern Baltic. Somatic tissues and gametes were surveyed for the presence of different types of mtDNA. Length variants were identified using PCR (polymerase chain reaction) amplification of the mtDNA fragment containing the major noncoding region. Seventeen length variants present in homo- and heteroplasmic individuals were found in a sample of 343 individuals analyzed. Two length variants of the major noncoding region (PCR product lengths: 1520 and 1370 bp) were the most frequent. Length heteroplasmy was found in 46.8% of males, and was mostly caused by the presence of the short length variant (PCR product length: 1370 bp) of the major noncoding region. MtDNA variants were also detected by restriction analysis of a 1.3 kb segment of coding region (ND2-COIII) amplified by PCR. This study provides evidence that the short length variant is always transmitted to sperm and has taken over the role of the M genome. The remaining length variants were present in both males and females. Longer length variants were transmitted mainly through the female line. A possibility of much higher incidence of genome role reversals in Baltic M. trossulus, in comparison with other populations of mussels is discussed.     

Estimating the influence of prawn stocking density and seagrass type on the growth of juvenile tiger prawns (Penaeus semisulcatus): results from field experiments in small enclosures

Fine mesh enclosures (0.9 m² in basal area, 1 m high, with 100 µm mesh) and a jet-net retrieval system were developed to test the influence of juvenile prawn stocking density on growth rates in (1) different months (April and October/November) and (2) different types of intertidal seagrass beds in the Embley River estuary of tropical Australia. Small juvenile tiger prawns (3-6 mm in carapace length, CL) were stocked in enclosures at densities of 4-32 prawns per enclosure (4.4-35.5 prawns m^-2) on a high biomass seagrass bed (about 70 g m^-2 of mostly Enhalus acoroides) and one with low biomass (about 10 g m^-2 of mostly Halodule uninervis). After 2-3 weeks in the enclosures, recovery rates, and hence possibly survival, were greater on the high biomass Enhalus than on the low biomass Halodule. However, not all fish and crustaceans could be excluded from the enclosures. Growth rates were twice as fast on the high biomass Enhalus than on the low biomass Halodule. It is likely that the high biomass Enhalus, with its greater surface area, supported more epiphytic flora and fauna and reduced the potential for interference competition between prawns, compared with the low biomass Halodule. Growth rates on Enhalus were significantly faster at a stocking density of 4 prawns per enclosure (1.3 mm CL week^-1) than at a stocking density of 16 and 32 prawns per enclosure (both 0.8 mm CL week^-1), and did not differ significantly between April and October/November (temperatures were about 30°C at both times). The mean growth rate at 8 prawns per enclosure (1.1 mm CL week^-1) did not differ significantly from those at 4, 16 and 32 prawns per enclosure. These results from two seagrass beds suggest that the carrying capacity for juvenile tiger prawns was greater in the high biomass Enhalus than the low biomass Halodule bed.