Physioecology of zooplankton. II. Effects of phytoplankton concentration,temperature, and body size on the development and molting rates of Calanus pacificus and Pseudocalanus sp.

Developmental time and stage duration for Calanus pacificus Brodsky and Pseudocalanus sp. and the rate of loss of body carbon by molting for C. pacificus were estimated for copepodite stages cultured under various combinations of phytoplankton concentration and temperature. Mean development time and stage duration for C. pacificus decreased hyperbolically with increasing food concentration, and the minimum time required for reaching a given stage decreased logarithmically with a logarithmic increase in temperature. Low temperature retarded the development of early stages proportionally more than that of late stages, and stage duration increased logarithmically with increasing body weight. Therefore, copepodite development was not isochronal. The rate of loss of body carbon by molting was small, ranging from 0.2 to 2% day^-1. This rate increased hyperbolically with food concentration and was linearly related to the growth rate. The critical food concentration for the rates of development and molting increased with temperature and stage of development, but these rates were less dependent on food concentration than the growth rate. The development rate of Pseudocalanus sp. was higher than that of C. pacificus, and was less influenced by changes in food concentration and temperature. It is postulated that the inverse relationship between temperature and body size results from a differential effect of temperature and body size on the rates of growth and development. That is, with increasing body size the growth rate tends to become temperature-independent, but the development rate remains proportional to temperature. Thus, copepodites growing at low temperature can experience a greater weight increment between molting periods than individuals growing at high temperature, because the growth rate is similar at all temperatures but stage duration is longer at low temperature.Contribution No. 1128 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Physioecology of zooplankton. IV. Effects of phytoplankton concentration,temperature, and body size on the net production efficiency ofCalanus pacificus

Weight-specific rates of individual production, total metabolic expenditure and assimilation, and net production efficiencies were estimated forCalanus pacificus Brodsky of selected body weights cultured at various phytoplankton concentrations and temperatures. The weight-specific rate of individual production increased hyperbolically with food concentration, and the maximum rate of individual production decreased logarithmically with a linear increase in body weight propotionally more at high than at low temperature. The weight-specific rate of total metabolic expenditure decreased logarithmically with increasing body weight and was unaffected by changes in food concentration. The effects of food concentration and temperature on the weight-specific rate of assimilation were similar to those on the rate of individual production, but the effect of body size differed considerably. The diversity in the temperature and body-size dependence of the maximum weight-specific rates of various physiological processes suggest (1) that, except for the metabolic rate, the allometric model (log-log relation) is inadequate for describing relationships between maximum rates of physiological processes and body size within species, and (2) that the common assumption that temperature affects the rates of various physiological processes in similar ways is not justified. Net production efficiency increased hyperbolically with food concentration, and the maximum production efficiency first increased slightly and then decreased gradually with increasing body weight. Small copepods attained higher efficiency at high temperature but larger ones did so at low temperature. The critical food concentrations for production efficiency and for the rate of individual production increased with increasing temperature and body size. Because of the effects of interactions among critical food concentration, temperature, and body size on the rates of growth and individual production and on net production efficiency, early development stages ofC. pacificus optimized growth and food conversion efficiency at high temperature, but late stages, particularly at low food concentrations, grew best and transformed food more efficiently at low temperature.Contribution No. 1130 From the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Influence of cultivation and food concentration on body length of calanoid copepods

The cephalothorax length of the marine pelagic Copepoda Acartia clausi Giesbrecht, Temora longicornis (Müller), Centropages hamatus (Lilljeborg) and Pseudocalanus sp. was monitored at 15°C during prolonged cultivation through up to 55 filial generations and at different concentrations of food. The length of T. longicornis decreased considerably during the first 15 generations and remained rather constant thereafter. In the other species, body length increased slightly or remained almost constant. Genetic changes are probably involved. Food concentration influenced body size of all species, particularly C. hamatus, in which 80% of the natural size range may be explained by differences in food concentration. The idea that temperature is a dominant factor in determining the length of copepods should be reconsidered.     

Physioecology of zooplankton. III. Effects of phytoplankton concentration,temperature, and body size on the metabolic rate ofCalanus pacificus

Weight-specific rates of oxygen consumption of actively feeding copepodite stages ofCalanus pacificus Brodsky were measured under various combination of phytoplankton concentration and temperature. The rate decreased logarithmically with a logarithmic increase in dry body weight of copepods, and the relationship between these variables was described using a log-transformed allometric equation. The body-size dependence of the metabolic rate was independent of changes in food concentration and temperature, but the metabolic level increased linearly with a logarithmic increase in temperature and was not significantly affected by changes in food concentration. Respiration rates measured in this study forC. pacificus were about twice as high as rates reported for unfed closely related species of the same genus. An analysis of the metabolic cost of feeding processes suggests that metabolic models derived from feeding models may be of little ecological value at present.Contribution No. 1129 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Influence of food quantity and temperature on development and growth of the marine copepod Calanus chilensis from northern Chile

An experiment under laboratory conditions was conducted to test the hypothesis that development and growth of copepodite stages in Calanus chilensis are temperature-dependent and not subject to food shortage in the upwelling area of the Humboldt Current, northern Chile. Field data obtained from June 1994 to May 1995 in Bahía Mejillones (23°S) were used to define four combinations of temperature and food under which copepodites were reared from Stage CIII to adulthood. The high temperature was 18.1 °C and the low temperature 13.1 °C, whereas the high food level was in the range of 6.8 to 24.8 μg l⁻¹ chlorophyll a and the low level 1.0 to 6.8 μg l⁻¹ chlorophyll a. As food a mixture of three unknown species of phytoflagellates and the diatom Navicula cryptocephala was used. This phytoplankton was initially obtained from the same sampling sites as copepods and kept in f/2 media at stable levels and composition throughout the experiment. The development rate (1/t), estimated from the time (t) elapsing between Stage CIV and adult, was significantly affected by both temperature and food, although low-food effects were much more remarkable. Low-food conditions also significantly reduced body length and “structural” (lipid-discounted) body mass at adulthood, while temperature only affected body length. The weight-specific growth rate was also affected by food and temperature, but again food effects were much more drastic. The results indicate that C. chilensis is a highly sensitive species to lack of food, and is possibly subject to food shortage during its annual cycle in the coastal upwelling area of northern Chile. Food limitation may help explain the seasonal pattern of adult size reported by previous studies in the area and the lack of consistence between the number of generations predictable from a temperature-dependent model and that observed in the field during the annual cycle. Received: 10 September 1996 / Accepted: 29 October 1996     

Dynamics of zinc-65 specific activity and total zinc in benthic fishes on the outer continental shelf off Central Oregon

The dynamics of ⁶⁵Zn specific activity and total zinc in benthic fishes on the outer continental shelf off central Oregon (USA) were examined. A differential equation that relates specific activity of ⁶⁵Zn in fish to that in fish food was used to estimate 's (zinc uptake-rate coefficients) for 3 different size classes of the flounder Lyopsetta exillis, a small predator of pelagic Crustacea, and for 1 size class of the flounder Microstromus pacificus, a large predator of infauna. The 's obtained for L. exilis were very close to the obtained in the laboratory for the flounder Pleuronectes platessa. The estimated for M. pacificus was very much smaller than the 's estimated for the other two species. A model that related to predicted weight-specific feeding rates suggested that the smaller of M. pacificus was caused by a low absorption efficiency of zinc from its prey. Sensitivity studies indicated that time histories of specific activity in the fishes are not sensitive to moderate changes in . The negative correlation between specific activity in the diet and in the weight of L. exilis was the major cause of the negative correlative between specific activity and weight in this species. In M. pacificus, where composition of diet does not vary with size, specific activity was independent of weight. The time history of specific activity in M. pacificus was very much lower than those in the different size classes of L. exilis, a result caused mainly by the much lower specific activity of the prey of M. pacificus. Differences in specific activity among other benthic fishes were also correlated with differences in specific activity of their prey. The food-web dynamics responsible for these patterns are discussed. Variation in total zinc concentrations among species was small. Within species of flounder, zinc concentration varied only slightly or not at all with weight.     

Demographic and temporal structure of an allele frequency cline in the mussel Mytilus edulis

Feeding rates, patterns of prey selection, and starvation tolerance were investigated for adult males and females of the cyclopoid copepod Corycaeus anglicus collected from the waters of Friday Harbor, Washington, USA. Selection by C. anglicus was determined largely by prey body-size, but was also affected by species and developmental stage. Small developmental stages of all prey species were fed upon at relatively low rates. The small calanoid species Acartia clausii was increasingly vulnerable to predation by C. anglicus as it progressed through successive developmental stages. Larger prey species, Pseudocalanus sp. and Calanus pacificus, were more vulnerable in intermediate stages, the C3 and N6 stages, respectively. Larger and smaller prey were characteristically attacked at different sites on their bodies; however, attack sites fell within a similar range of body widths, 130 to 170 m. Males of Corycaeus anglicus killed a maximum of 1.4 prey d^-1 when feeding on the optimally-sized adult females of Acartia clausii, which are approximately equivalent to its own body length. Males fed at approximately double the rates of females. Despite its small size and apparent lack of metabolic stores, this cyclopoid is highly tolerant of starvation conditions. Median survival time without food is at least 2 wk for both males and females. In its predatory behavior, C. anglicus employs an ambush-type strategy and seems to be adapted for infrequent encounters with relatively large prey.Contribution No. 1412 from the School of Oceanography, University of Washington, Seattle     

Food selection by laboratory-reared larvae of the scaled sardine Harengula pensacolae (Pisces,Clupeidae) and the bay anchovy Anchoa mitchilli (Pisces,Engraulidae)

Food selection by laboratory-reared larvae of scaled sardines Harengula pensacolae, and bay anchovies Anchoa mitchilli, was compared. Natural plankton was fed to the larvae during the 22 days following hatching. Food levels in the rearing tanks were maintained at an average of 1,600 to 1,800 potential food organisms per liter. Larvae of both species selected as food copepod nauplii, copepodites, and copepods; initial feeding was on organisms of 50 to 75 body width. Larvae of H. pensacolae averaged 4.2 mm in total length at hatching and those of A. mitchilli about 2.0 mm. H. pensacolae larvae grew about 1.0 mm per day and A. mitchilli 0.70 mm per day. The mean number of food organisms in each digestive tract was greater in H. pensacolae than in A. mitchilli, and the difference in number increased as the larvae grew. Average size of food organisms eaten increased for both species with growth, because of selection by the larvae; the average size of copepodites and copepods in digestive tracts increased at a faste rate in H. pensacolae than A. mitchilli. A. mitchilli longer than 8 mm did not eat copepod nauplii.Contribution No. 170, Bureau of Commercial Fisheries Tropical Atlantic Biological laboratory, Miami, Florida 33149, USA.     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Photosynthate partitioning by phytoplankton in a New Zealand coastal upwelling system

A stimulation model of copepod population dynamics (development rate, fecundity, and mortality) was used to compute the predatory consumption necessary to control population growth in three dominant copepod species (Pseudocalanus sp., Paracalanus parvus, and Calanus finmarchicus) on Georges Bank, given observed seasonal cycles of copepod and predator populations. The model also calculated secondary production of each species. Copepod development rate and fecundity were functions of temperature while mortality was a function of predator abundance and consumption rate. Daily inputs of temperature and predator abundance (chaetognaths, ctenophores, and Centropages spp.) were derived from equations fit to field data. Model runs were made with various consumption rates until the model output matched observed copepod seasonal cycles. Computed consumption rates were low compared with published values from field and laboratory studies indicating that, even at conservative estimates of consumption, predators are able to control these copepod populations. Combined annual secondary production by the small copepod species, Pseudocalanus sp. and P. parvus, was nearly twice that of the larger C. finmarchicus with P. parvus having the highest total annual production.     

Annual population development and production by small copepods in Disko Bay,western Greenland

The population of small copepod species (approximately <1 mm) were investigated during an annual cycle in Disko Bay, western Greenland. The small species considered were Acartia longiremis, Pseudocalanus spp., Oithona spp., Oncaea spp., Microsetella spp., and Microcalanus spp. Most of the small species were present in the surface waters year round and numerically dominated the community, and in biomass from late summer and throughout winter. Oithona spp. was numerically the main contributor, while Pseudocalanus spp. dominated in terms of biomass. In the uppermost 50 m, maximum abundance, biomass and secondary production were observed in late September after the phytoplankton production practically had terminated and the winter initiated. The free spawning Acartia longiremis showed a strong seasonal fluctuation in biomass and egg production, in contrast to the egg carrying species Pseudocalanus spp. and Oithona spp. These had a long spawning season and maintained a more stable biomass year round. Secondary production was estimated by three different ways: (1) based on the obtained specific egg production rates, (2) a temperature dependent equation, and (3) a multilinear regression taking temperature, body weight and chlorophyll into consideration. The contribution of the small species was insignificant when compared to the large Calanus species during the spring- and post-bloom. However, during late summer and winter, where Calanus had left the upper water strata for hibernation, the small species played a crucial role in the pelagic carbon cycling.     

Depth zonation and metabolic adaptation in Dover sole,Microstomus pacificus,and other deep-living flatfishes: factors that affect the sole

Flatfishes of Monterey Bay, central California, undergo species replacements with increasing depth along a transect from 100 m on the continental shelf down to a depth of 1400 m on the continental slope. The Dover sole, Microstomus pacificus, differs from the other local flatfish species by undergoing an extensive ontogenetic vertical migration, occupying all depth zones at different life stages, and having its maximum spawning biomass in the oxygen minimum zone between 600 and 1000 m. Size-activity relationships and depth-activity relationships for the glycolytic enzyme lactate dehydrogenase (LDH) and for two enzymes associated with aerobic metabolism, malate dehydrogenase and citrate synthase (CS), were examined in white-muscle tissue of shallow-living, deep-living and ontogenetically-migrating species. Scaling coefficients (b) for weight-specific enzyme activity (log activity)=a+b (log wet weight), varied in sign as well as magnitude for fishes living at different depths. In the shallow-living California halibut Paralichthys californicus, LDH scaled positively (0.39) and CS scaled negatively (-0.15) with size, a pattern observed previously for most shallow-water fish species. The permanently deep-living species, the deepsea sole Embassichthys bathybius, differed in that both LDH and CS scaled strongly negative (-2.0 and-1.5, respectively). For the ontogenetically migrating Dover sole Microstomus pacificus, there was a shelf-slope transition. For the shelf specimens (200 m), LDH scaled positive (0.11) and CS negative (-0.29) and for the slope specimens (400 m), LDH scaled negative (-0.65) and CS strongly negative (-0.63). Rex sole, Glyptocephalus zachirus, showed a similar shelf-slope transition. Intraspecific depth-enzyme activity differences were not incremental, but changed abruptly between the continental shelf stations (100 to 200 m) and the continental slope (400 to 1400 m). Based on comparisons with laboratory-maintained individuals, the decline in the metabolic capacity of the white muscle of Dover sole is a phenotypic response to the low food and oxygen conditions of the continental slope. Contrary to expectation, anaerobic capacity (LDH activity) decreased in response to low oxygen conditions, suggesting that in a permanently hypoxic environment such as the oxygen minimum zone the metabolic strategy may be to not incur an oxygen debt that would be difficult to pay back.     

Towards a global model of in situ weight-specific growth in marine planktonic copepods

The dependency of in situ weight-specific fecundity of adult females (as egg production) and growth of juveniles (as somatic production) upon individual body weight in marine planktonic copepods was examined. A compilation was made of results where wild-caught individuals were incubated in natural seawater (often pre-screened to remove large organisms), at near in situ temperatures, over short periods of the order of 24 h. The results demonstrate that for the adult broadcast-spawning group weight-specific fecundity rates are dependent upon body weight, but independent of temperature. We postulate this may be the result of global patterns in available phytoplankton. Weight-specific growth rates are dependent upon individual temperature and body weight in juvenile broadcast-spawners, with rates declining as body weight increases. Sac-spawners have growth/fecundity rates that are independent of body weight in adults, juveniles, and both combined, but which are temperature-dependent. Globally applicable equations are derived which may be used to predict growth and production of marine copepods using easily quantifiable parameters, namely size-distributed biomass and temperature. Some of the variability in growth which remained unaccounted for is the result of variations in food quantity and quality in the natural environment. Comparisons of the rates compiled here over the temperature range 10 to 20 °C with previously compiled food-saturated rates over the same temperature interval, revealed that in situ rates are typically sub-optimal. Adults appear to be more food-limited than juveniles, adult rates in situ being 32 and 40% of those under food saturation in broadcasters and sac-spawners, respectively, while juvenile in situ rates are on average ∼70% of those at food saturation in both broadcasters and sac-spawners. Received: 18 September 1997 / Accepted: 13 May 1998     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.     

Zooplankton growth rates: the influence of size and resources in tropical marine copepodites

Growth rates were determined for copepodites of the genera: Acartia, Centropages, Corycaeus, Oithona, Paracalanus, Parvocalanus and Temora in nearshore waters of Jamaica from in situ microcosm incubations. At these high local temperatures (∼28 °C), total copepodite development time was as short as 4 to 5 d. Mean instantaneous growth rates (g) ranged from as high as 1.2 d⁻¹ to as low as 0.1 d⁻¹. In general, cyclopoid copepods appeared to grow more slowly than calanoids of the same size. Enhancement of resources by nutrient addition caused a 32% increase in growth rates in experiments from a mesotrophic site, but only a 17% increase at a more eutrophic site. Additionally, copepodites at both sites showed faster development and generally larger size at stage in response to nutrient addition. Growth rates were positively related to chlorophyll concentration in the >2 μm size-fraction. A significant relationship of growth rate to body size (r ² = 0.45) emerged across a wide range of trophic status, but it was confounded with resource availability. It appears that growth in tropical copepod copepodites may be frequently limited by resources in a size-dependent manner. Received: 30 May 1997 / Accepted: 13 May 1998     

Nitrogen balance in marine fish larvae: influence of developmental stage and prey density

The utilization and fate of nitrogen in larvae of plaice (Pleuronectes platessa), blenny (Blennius pavo) and herring (Clupea harengus), from the stage of first-feeding to metamorphosis, was examined under laboratory conditions. Rates of ammonia excretion, primary amine defaecation, and growth in terms of protein-nitrogen were monitored throughout larval life. Data were used to calculate daily ration, the coefficient of nitrogen utilization (absorption efficiency), and gross and net growth efficiencies. The developmental pattern of nitrogen balance was similar for plaice and blenny larvae. These species showed increasing growth efficiency (k₁: 55 to 80%) with decreasing weight-specific waste nitrogen losses with age. Absorption efficiencies. were high (83 to 98%) in plaice and blenny larvae, and tended to increase with development in the former species. Ration relative to body weight decreased with growth in both species. Herring larval development, although at a slower rate than blenny and plaice, appeared normal up to 33 d, after which high mortality occurred. Absorption efficiency in this species tended to decline (83 to 43%) with age, until metabolic costs exceeded the absorbed ration and growth ceased. Artemia sp. nauplii proved a suitable food source for the rearing of plaice and blenny larvae, but this diet may have long-term toxicity or deficiency effects on herring. Availability and density of food affected nitrogen balance in the larvae of all three species. Feeding stimulated the output of wastes in excretion and defaecation by a factor of up to ten times the 12-h non-feeding basal rates. Waste nitrogen output reached a peak some 2 to 3 h after commencement of feeding and returned slowly to the baseline in 5 to 10 h after cessation of feeding. There was an asymptotic increase in ration, ammonia output and growth of larvae as prey density increased. Ration saturated at a higher prey density (>4 prey ml^-1) than either growth or excretion rate (1 prey ml^-1). Thus the efficiency with which food is absorbed and utilized for growth must eventually decline in response to high prey density. The idea that larval fish are adapted to maximize ingestion and growth rate, rather than optimize growth efficiency and thus to respond to prey occurring in either low density or in occasional patches, is supported by these results.     

Über den Einfluß der Nahrungskonzentration und anderer Faktoren auf Filtrierleistung und Nahrungsausnutzung der Muscheln Arctica islandica und Modiolus modiolus

Filtration rates and the extent of phagocytosed food particles were determined in the offshore lamellibranchs Artica islandica and Modiolus modiolus in relation to particle concentration, body size and temperature. Pure cultures of the algae Chlamydomonas sp. and Dunaliella sp. were used as food. A new method for determining filtration rates was developed by modifying the classical indirect method. The concentration of the experimental medium (100%) was kept constant to ±1%. Whenever the bivalves removed algae from the medium, additional algae were added and the filtration rate of the bivalves expressed in terms of percentage amount of algae added per unit time. The concentration of the experimental medium was measured continuously by a flow colorimeter. By keeping the concentration constant, filtration rates could be determined even in relation to different definite concentrations and over long periods of time. The amount of phagocytosed food was measured by employing the biuret-method (algae cells ingested minus algae cells in faeces). Filtration rates vary continuously. As a rule, however, during a period of 24 h, two phases of high food consumption alternate with two phases of low food consumption during which the mussels' activities are almost exclusively occupied by food digestion. Filtration rate and amount of phagocytosed algae increase with increasing body size. Specimens of A. islandica with a body length of 33 to 83 mm filter between 0.7 to 71/h (30–280 mg dry weight of algae/24 h) and phagocytose 21 to 122 mg dry weight of algae during a period of 24 h. The extent of food utilization declines from 75 to 43% with increasing body size. In M. modiolus of 40 to 88 mm body length, the corresponding values of filtration rate and amount of phagocytosed algae range between 0.5 and 2.5 l/h (20–100 mg dry weight of algae) and 17 to 90 mg dry weight of algae, respectively; the percentage of food utilization does not vary much and lies near 87%. Filtration rate and amount of phagocytosed algae follow the allometric equation y=a·x ^b. In this equation, y represents the filtration rate (or the amount of phagocytosed algae), a the specific capacity of a mussel of 1 g soft parts (wet weight), x the wet weight of the bivalves' soft parts, and b the specific form of relationship between body size and filtration rate (or the amount of phagocytosed algae). The values obtained for b lie within a range which indicates that the filtration rate (or the amount of phagocytosed algae) is sometimes more or less proportional to body surface area, sometimes to body weight. Temperature coefficients for the filtration rate are in Arctica islandica Q₁₀ (4°–14°C)=2.05 and Q₁₀ (10°–20°C)=1.23, in Modiolus modiolus Q₁₀ (4°–14°C)=2.33 and Q₁₀ (10°–20°C)=1.63. In A. islandica, temperature coefficients for the amount of phagocytosed algae amount to Q₁₀ (4°–14°C)=2.15 and Q₁₀ (10°–20°C)=1.55, in M. modiolus to Q₁₀ (4°–14°C)=2.54 and Q₁₀ (10°–20°C)=1.92. Upon a temperature decrease from 12° to 4°C, filtration rate and amount of phagocytosed algae are reduced to 50%. At the increasing concentrations of 10×10⁶, 20×10⁶ and 40×10⁶ cells of Chlamydomonas/l offered, filtration rates of both mollusc species decrease at the ratios 3:2:1. At 12°C, pseudofaeces production occurs in both species in a suspension of 40×10⁶, at 20°C in 60×10⁶ cells of Chlamydomonas/l. At 12°C and 10–20×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is phagocytosed. At 40×10⁶ cells/l, the amount of phagocytosed cells is reduced by 26% as a consequence of low filtration rates and intensive production of pseudofaeces. At 20°C and 20–50×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is sieved out and phagocytosed; the concentration of 10×10⁶ cells/l is too low and cannot be compensated for by increased activity of the molluscs. With increasing temperatures, the amount of suspended matter, allowing higher rates of filtration and food utilization, shifts toward higher particle concentrations; but at each temperature a threshold exists, above which increase in particle density is not followed by increase in the amount of particles ingested. Based on theoretical considerations and facts known from literature, 7 different levels of food concentration are distinguishable. Experiments with Chlamydomonas sp. and Dunaliella sp. used as food, reveal the combined influence of particle concentration and particle size on filtration rate. Supplementary experiments with Mytilus edulis resulted in filtration rates similar to those obtained for M. modiolus, whereas, experiments with Cardium edule, Mya arenaria, Mya truncata and Venerupis pullastra revealed low filtration rates. These species, inhabiting waters with high seston contents, seem to be adapted to higher food concentrations, and unable to compensate for low concentrations by higher filtration activities. Adaptation to higher food concentrations makes it possible to ingest large amounts of particles even at low filtration rates. Suspension feeding bivalves are subdivided into four groups on the basis of their different food filtration behaviour.     

Ammonium cycling by Antarctic zooplankton in winter

Elemental composition and excretion rates of ammonium-nitrogen of zooplankton, ranging over more than five orders of magnitude in body size, were measured in mid-winter in coastal waters west of the Antarctic Peninsula. Excretion rates were constant for the initial 12 h of incubation in the four species tested, and experimental stocking densities of up to 126 mg dry wt l^-1 did not cause variability in the rate of ammonium production. Weight-specific excretion rates of freshly caught Euchaeta antarctica, Conchoecia sp., Thysanoessa macrura, Euphausia superba, and early stage copepodites of Metridia gerlachei were not significantly different from those reported in summer. However, adult copepods of M. gerlachei and Calanoides acutus appear to have reduced their nitrogen metabolism during winter. Turnover rates of body nitrogen increased with diminishing size, ranging from <0.5% body N d^-1 for large E. superba to >7% body N d^-1 for CII and CIII copepodites of M. gerlachei. Only the nitrogen turnover rates of C. acutus were sufficiently low as to suggest that it could survive the entire austral winter without feeding. Phytoplankton and bacterioplankton were virtually absent in both the water column and the sea-ice. We conclude that carnivory is the dominant trophic mode of the pelagic zooplankton community in Antarctica during winter. Production of ammonium-nitrogen by the zooplankton community probably accounts for M10% of the total ammonium regenerated prior to the annual spring bloom.     

Effects of prey concentration,prey size,predator life stage,predator starvation,and season on predation rates of the carnivorous copepod Euchaeta elongata

Adult females of the large carnivorous copepod Euchaeta elongata Esterly were collected from 1977 to 1980 in Port Susan, Washington, USA. Predation rates of the adult females increased with increasing prey abundance when fed the following 4 sizes of copepods: adult females of Calanus pacificus (average prosome length [PL] of 2 650 μm), adults of Aetideus divergens (PL of 1 560 μm), adult females of Pseudocalanus spp. (PL of 1 060 μm), and nauplii of C. pacificus (PL of 410 μm). Saturation feeding levels were reached when adult females of the predator were fed the small adult copepod, Pseudocalanus spp. Maximum biomass ingested of this small copepod was more than the maximum amount ingested of the larger copepods. Predation rates of the predatory copepodids at Stages IV and V also increased with increasing concentration of the 1 060 μm (PL) prey. High feeding rates exhibited by both adults and copepodids at Stage V of the predator indicate their importance as sources of mortality on populations of small copepods. Ingestion efficiency E _i (prey wholly consumed [prey attacked]^-1) varied as follows: adults of E. elongata were more efficient than copepodids of E. elongata; adults were more efficient than copepodids when ingesting smaller prey; starved adults were more efficient than fed ones; and both adults and copepodids were more efficient at low food concentrations. For adults of E. elongata, there were no marked seasonal variations in predation or respiratory rates that would represent acclimatory responses; however, small adults obtained during winter were more efficient at ingesting prey than were the larger adults gathered in summer. This seasonal variation in the efficiency of ingestion may be a useful indicator of physiological state: high E _i values could indicate that predators are starving in winter, and low E _i values could indicate that predators are satiated in summer.     

Ingestion-independent rates of ammonium excretion by the copepod Calanus pacificus

The relationship between food ingested and NH ₊ ⁴ excretion rate was investigated for female Calanus pacificus collected in August, 1982, from the San Juan Archipelago, Washington State, USA. The copepods were preconditioned to 6 densities of the diatom Thalassiosira weissflogii (0 to 10⁴ cells ml^–1) for 30 h before the experiment. The experiment was conducted with nutrients added in excess to maintain equal rates of NH ₊ ⁴ uptake by the diatoms at all densities. Although ingestion rates of C. pacificus varied from 0 to over 20% of body N d^–1 at the different food levels, excretion was a constant 6.6 nM NH ₊ ⁴ copepod^–1 h^–1 or about 10% of body N d^–1. This ingestion-excretion relationship, which is consistent with previous respiration and fecundity studies, suggests that the ecological dominance of C. pacificus only under conditions of high food abundance may be due to a dramatic increase in its growth efficiency as ingestion increases above the level supporting a constant metabolic rate. The maintenance of a constant level of metabolism during relatively short periods of low food abundance may be advantageous if it allows the copepod to exploit more effectively short-term variability in its food resulting from environmental heterogeneity or vertical migration.Contribution No. 1360 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA