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     

Physioecology of zooplankton. I. Effects of phytoplankton concentration,temperature, and body size on the growth rate of Calanus pacificus and Pseudocalanus sp.

Changes in dry weight and in weight-specific growth rates were measured for copepodite stages of Calanus pacificus Brodsky and Pseudocalanus sp. cultured under various combinations of phytoplankton concentration and temperature. Mean dry weight of early copepodites was relatively unaffected by either food concentration or temperature, but mean dry weight of late stages increased hyperbolically with food concentration and was inversely related to temperature. The food concentration at which maximum body weight was attained increased with increasing temperature and body size, and it was considerably higher for C. pacificus than for Pseudocalanus sp. This suggests that final body size of small species of copepods may be determined primarily by temperature, whereas final body size of large species may be more dependent on food concentration than on temperature. Individual body weight increased sigmoidally with age. The weight-specific growth rate increased hyperbolically with food concentration. The maximum growth rate decreased logarithmically with a linear increase in body weight, and the slope of the lines was proportional to temperature. The critical food concentration for growth increased with body size proportionally more at high than at low temperature, and it was considerably higher for C. pacificus than for Pseudocalanus sp. Because of these interactions, early copepodites optimized growth at high temperature, even at low food concentrations, but under similar food conditions late stages attained higher growth at low temperature. The same growth patterns were found for both species, but the rates were significantly higher for the larger species, C. pacificus, than for the smaller one, Pseudocalanus sp. On the basis of findings in this study and of analyses of relationships between the maximum growth rate, body size, and temperature from other studies it is postulated (1) that the extrapolation of growth rates from one species to another on the basis of similarity in body size is not justified, even for taxonomically related species; (2) that the allometric model is inadequate for describing the relationship between the maximum weight-specific growth rate and body size at the intraspecific level; (3) that the body-size dependence of this rate is strongly influenced by temperature; and (4) that species of zooplankton seem to be geographically and vertically distributed, in relation to body size and food availability, to optimize growth rates at various stages of their life cycles.Contribution No. 1127 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Short-term changes in feeding and digestion by the copepodCalanus pacificus

The planktonic marine copepodCalanus pacificus exhibits an enhanced feeding rate, or hunger response, when exposed to food following short periods of starvation. In a scries of laboratory experiments with copepods collected from the main basin of Puget Sound, Washington, during 1982 and 1984, we measured maximum ingestion rate, assimilation efficiency, and digestive enzyme activity to determine the time scales over which the feeding behavior ofC. pacificus responds to increases in food. These laboratory results were then compared to field studies of diel fluctuations in digestive enzymes and gut fluorescence ofC. pacificus in Dabod Bay, a fjord of Puget Sound, during September, 1980, and the closely relatedC. marshallae off the Washington coast, in August, 1981. Laboratory experiments demonstrated that the hunger response ofC. pacificus lasts approximately 6 h before ingestion rate returns to a steady state level of about one-half maximum. On the order of 12h of starvation were required to induce the maximum ingestion rate of the hunger response. Digestive enzyme activities did not change over these time scales. Assimilation efficiency peaked within a few hours of the onset of feeding, with low initial rates possibly related to the period of starvation prior to feeding. These results were consistent with diel patterns observed in the field. The hunger response ofC. pacificus appears to be controlled by processes within the gut, and our results are discussed in relation to recent studies of the digestive processes of calanoid copepods.Contribution No. 1772 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

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     

Effects of food-level acclimation on digestive enzyme activities and feeding behavior of Calanus pacificus

A recent hypothesis in the zooplankton literature states that zooplankton acclimate to ambient food concentrations such that higher digestive enzyme activities and, consequently, higher maximum ingestion rates are achieved at higher food levels. To test this hypothesis, adult female Calanus pacificus, collected from the main basin of Puget Sound, Washington, USA, in August 1979 and May 1982, were conditioned for 2 wk at different concentrations of the diatom Thalassiosira weissflogii (=fluviatilis). Ingestion rates and the activity of the digestive enzymes laminarinase, maltase, and cellobiase were measured periodically during acclimation and in a block-designed feeding experiment at the end of acclimation. Consistent with the hypothesis, maximum ingestion rate and digestive enzyme activity were positively correlated. However, in contrast to the hypothesized mechanism, this result arose because both maximum ingestion rate and digestive enzyme activity were negatively correlated with food concentration during acclimation. The enhanced ingestion of copepods following long-term (12 to 14 d) acclimation to low food is similar to that previously described for short-term (e.g. 1 d) starvation. It might be energetically optimal for copepods experiencing a patchy food environment to maintain higher levels of digestive enzymes at low food concentrations in order to exploit high concentrations of food when encountered.     

Feeding and bacteriolytic responses of the deposit-feeder Abarenicola pacifica (Polychaeta: Arenicolidae) to changes in temperature and sediment food concentration

Deposit-feeders can respond to seasonal fluctuations in food concentration both functionally (e.g. by adjusting feeding rates) and physiologically (e.g. by changing the concentration of bacteriolytic agents in gut fluids). Laboratory feeding experiments were carried out (11 to 21 July 1997) with the arenicolid polychaete worm Abarenicola pacifica (Healy and Wells). Objectives were to test for separate and interactive effects of sediment food concentration and temperature (6, 11, and 16 °C) on deposit-feeder functional (feeding rates) and physiological (bacteriolytic activity of gut fluids) responses. Food concentration was varied experimentally using sieved (1 mm) natural sediments (Md φ=2.00; 0.6% organic) mixed with combusted (500 °C, 8 h) sediments for final concentrations of 25, 50, and 100% natural sediment. Sediment food quality was measured as: (1) bioavailable amino acids (EHAA), (2) chlorophyll a (chl a), and (3) bacterial abundance. Feeding rates were inferred from egestion rates (ER, g h⁻¹) and analyzed with respect to worm size. Bacteriolytic activity of midgut fluids was assayed turbidimetrically against two bacterial isolates, after worms had fed on experimental sediments for 15 d. Temperature and food concentration both significantly affected feeding rates, with maxima occurring at 50 and 100% natural sediment mixtures, and at high (16 °C) temperature. ER was positively, but not significantly correlated with EHAA and chl a; a positive, significant correlation was detected between ER and sediment bacterial abundance. Overall, functional responses agreed with earlier compensatory intake models for deposit-feeders. However, the size and direction of these responses was temperature-sensitive, suggesting that these models need to be adjusted for changes in absorption rates. No effects of ambient temperature or food concentration on bacteriolytic rates were observed, possibly due to compensatory mechanisms or the presence of multiple bacteriolytic agents in gut fluids. Received: 28 June 1999 / Accepted: 14 March 2000     

Influence of flow speed on the functional response of a passive suspension feeder,the spionid polychaete <Emphasis Type="Italic">Polydora cornuta</Emphasis>

Passive suspension feeders rely on surrounding flow to deliver food particles to them. Therefore, the classic conception of functional response (feeding rate vs. food concentration) may require modification to account for flow speed as a second independent variable. I compared the functional response of Polydora cornuta at different velocities and determined whether food capture was proportional to particle flux (concentration × velocity). To understand feeding responses at a mechanistic level, I measured the functional responses in terms of contact and capture rates and determined particle retention efficiency. Experiments were run separately with two sizes of food particles, and with juvenile or adult worms. For both worm sizes and both particle sizes, capture rate in weak flow was directly related to concentration, but in strong flow it was constant. Worms were therefore unable to benefit from abundant food when in strong flow. The critical velocity at which the capture rate became constant was lower for adult worms than for juvenile worms, and it was lower for small particles than for large particles. Retention efficiency was constant among all treatments, and the results for contact rate were essentially the same as for capture rate. Therefore, the mechanics of particle contact must explain the effects of velocity on the functional response. Contact rate was not a constant proportion of particle flux; treatments with similar fluxes yielded different contact rates depending on the strength of flow. The results appeared to be caused by a velocity-induced behavioral change in appendage posture that affects contact rates: in moderate flow, worms form their feeding palps into helical coils, which they tighten as the velocity increases. I suggest this behavior constrains suspension feeding rates and the mechanical selection between particle sizes when worms are in strong flow, and that the effect changes with ontogeny. Because the results are consistent with patterns in measured growth rates of P. cornuta, I hypothesize that this influence of velocity on the functional response can constrain growth and population dynamics in this species.     

Diet and respiration of the small planktonic marine copepod Oithona nana

The functional responses of Oithona nana (Giesbr.) to various phytoplankton and zooplankton food species are described. The food species were divided into three size categories, the seasonal abundances of which were measured in Loch Turnaig, a Scottish sea loch in 1977. The seasonal variations in feeding rates in the sea for each size class were derived. The seasonal variation in respiration rate of O. nana was measured, and metabolic requirements were claculated as between 6 and 40% of the food material estimated as being eaten. O. nana differs from other common copepods in having a wide food-particle size spectrum and a low metabolic rate. It is suggested that these adaptations constitute the strategy whereby O. nana maintains its population levels throughout the year.     

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     

Switching between herbivory and carnivory by the planktonic marine copepod Calanus pacificus

Adult females of the omnivorous copepod Calanus pacificus, collected from the plankton off La Jolla, California, USA (June, 1978), fed disproportionately on the prey in greatest relative abundance when given mixtures of diatoms (Thalassiosira fluviatilis) and copepod (C. pacificus) nauplii as food. This switch from herbivorous to carnivorous behavior may be significant in nature during the decline of phytoplankton blooms. More generally, the widespread omnivorous habit among pelagic animals suggests a responsive and flexible trophic organization which contributes to the resiliency of planktonic communities in a dynamic physical environment.Contribution No. 1223 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Comparison of feeding and biodeposition of three bivalves at different food levels

Experiments on the edible mussel Mytilus edulis, the American oyster Crassostrea virginica, and the hard clam Mercenaria mercenaria, using flowing systems, showed that the feeding and biodeposition rates were affected by food concentration. At all levels of food concentration, the order of increasing feeding rate (both the percent of available particulate carbon and the actual amount of carbon removed) was: clam < oyster < mussel. All bivalves exhibited lower feeding rates (both precent and actual) at low food concentrations. However, the precent of available food removed quickly increased to a maximum at food concentrations typical for the natural environment. This maximum remained constant for the mussel and oyster, but declined with increasing food concentration for the clam. However, because this percentage was for increasing levels, the actual carbon removed continued to increase up to the highest food level for all three bivalves. In increasing order of biodeposition rate, the bivalves were: clam < oyster < mussel. The biodeposition rates of the three bivalves increased logarithmically with increased food concentration as a result of the production of pseudofeces. The feeding and biodeposition data were used to calculate assimilation rates, and this pointed out the higher efficiency of the oyster compared to the mussel and clam.Contribution No. 2993 from the Woods Hole Oceanographic Institution.     

Measuring feeding rates of pelagic herbivores: analysis of experimental design and methods

We have re-evaluated the experimental methods and statistical procedures used to determine the relationship between feeding rates of pelagic herbivores and food concentration. Analysis of our own experiments, on Calanus pacificus feeding on Gyrodinium resplendens, and of other published research on this subject suggests the need for improvements in experimental design and methodology. We show that the use of mean concentration is statistically erroneous. First, it produces an artificial increase in the degrees of freedom that may result in the acceptance of nonsignificant regression lines. Second, it negates the value of replication, which is required to estimate sources of error. We present an example of how replication may be used to improve control over sources of error. Furthermore, we recommend the use of initial concentration rather than mean concentration. Finally, we introduce alternative methods to determine clearance and ingestion rates that enable the investigator to use replication and thus to estimate experimental errors.     

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 of pico- and nanoplankton by the Mediterranean red coral Corallium rubrum

This study investigates the feeding behaviour of the precious red coral Corallium rubrum on bacterioplankton. The effects of flow rate, prey concentration, and seawater temperature were tested. The results obtained show that C. rubrum was able to prey on both pico- and nanoplankton cells. Flagellates constituted the major bacterioplankton food source in terms of carbon and nitrogen, representing from 43 to 70% of the C and N ingested. Flow speed (2, 6, and 11 cm s⁻¹) had no effect on grazing rates, maybe due to the small size of the ingested particles. Conversely, feeding rates increased with prey concentration and seawater temperature. There was a doubling of the picoplankton ingestion rate for a sixfold increase in its concentration. The ingestion of autotrophic flagellates, however, increased at the same time as their concentration, indicating a preference for this type of food. Considering the range of concentrations typically found in the Ligurian Sea, the ingestion of pico- and nanoplankton brings 148 ng C polyp⁻¹ day⁻¹ and 28 ng N polyp⁻¹ day⁻¹. This type of food represents only ca. 4.5% of the total carbon gained by C. rubrum from the different sources, but might be the most important in terms of nitrogen, phosphorus, and other essential elements.     

Blooms of surf-zone diatoms along the coast of the Olympic Peninsula,Washington. XI. Regeneration of ammonium in the surf environment by the Pacific razor clam Siliqua patula

A 20 month field study was conducted on ammonium excretion rates of the Pacific razor clam Siliqua patula Dixon along the beaches of Washington State, USA. Excretion rates of all those nutrients likely to be regenerated in sufficient quantity to affect surf diatom growth were measured; ammonium appeared to be the most important metabolite. Excretion of ammonium by razor clams far exceeded that by other beach fauna. Ammonium excretion rates of razor clams were positively correlated with shell length, but no correlation between ammonium excretion rate and water temperature was evident. This may be an artifact or may represent some degree of seasonal acclimation of the species to temperature. Weight-specific ammonium excretion rates were negatively related to clam size, indicating a possible large (and unknown) contribution of regenerated ammonium by smaller clams in their first year of growth; smaller clams were rarely captured during this study.Contribution No. 1048 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA.     

Feeding energetics and metabolism in demersal fish species from Antarctic,temperate and tropical environments

The energetics of feeding has been investigated in demersal fish with similar sedentary lifestyles from the Antarctic (Notothenia neglecta Nybelin), North Sea (Myoxocephalus scorpius L.) and Indian Ocean (Cirrhitichys bleekeri Bleeker). In general, the metabolic rates of fasting individuals were positively correlated with adaptation temperature: values for a standard 100 g fish (mg O₂/h) were 3.3 for N. neglecta at around 0 °C, 2.7 for winter-acclimatized M. scorpius at 5 °C, 4.3 for summer-acclimatized M. scorpius at 15 °C, and 7.0 for C. bleekeri at 25 °C. In all species, following a single satiating meal, oxygen consumption increased to a peak of 2 to 3.5 times the fasting values. Maximum rates of oxygen consumption after feeding were several-fold higher in the warm-than in the cold-water species. After controlling for the effects of body mass and energy intake by analysis of covariance, the duration of the increase in metabolic rate, referred to as specific dynamic action (SDA), was found to be 3 to 4 times shorter in the warm- than in the cold-water fish, ranging from 57 h in C. bleekeri to 208 h in N. neglecta. In contrast, the SDA was not significantly different in the various species, corresponding to 15 to 23% of the energy ingested. Seasonal influences on metabolism and feeding were also studied in N. neglecta acclimated to simulated winter (-1.0 to-0.5 °C; 3 h light:21 h dark) or summer (0 to 0.9 °C; 21 h light:3 h dark) conditions. The metabolic rates of fasting and fed individuals, and the characteristics of the SDA were found to be independent of acclimation conditions. This suggests that N. neglecta is capable of processing food at similar rates throughout the year. Energy stores and enzyme activities were measured in the swimming muscles and liver of fish fed ad libitum. Summer-acclimated fish had higher concentrations of liver triglyceride stores and elevated activities of some enzymes of intermediary metabolism relative to winter-acclimated fish. The observed changes in intermdiary metabolism are probably related to annual cycles of growth and reproduction. It is suggested that the low aerobic scope for physiological performance in Antarctic fish may necessitate the seasonal switching of energy allocation between growth and reproduction.     

Factors influencing particle selection and feeding rate in the polychaete Cistenides (Pectinaria) gouldii

Feeding behavior of the deposit feeding polychaete Cistenides (Pectinaria) gouldii was examined to determine factors affecting particle selection and feeding rate. Worms were found to select large particles preferentially and particle size selection increased with worm size. Particle selection behavior was unaffected by changes in sediment bacterial abundance. Feeding rates were affected by sediment size, bacterial density and worm size. Generally feeding rates increased in sediment containing more food, although the response was worm size specific. When viewed in a theoretical construct these results were inconsistent with predictions of deposit feeder optimal foraging models. Alternative explanations, such as morphological constraints placed upon the polychaete, may explain C. gouldii feeding behavior.     

Impact of ocean warming on the early ontogeny of cephalopods: a metabolic approach

The impact of a realistic warming scenario on the metabolic physiology of early cephalopod (squid Loligo vulgaris and cuttlefish Sepia officinalis) life stages was investigated. During exposure to the warming conditions (19 °C for the western coast of Portugal in 2100), the increase in oxygen consumption rates throughout embryogenesis was much steeper in squid (28-fold increase) than in cuttlefish (11-fold increase). The elevated catabolic activity–accelerated oxygen depletion within egg capsules, which exacerbated metabolic suppression toward the end of embryogenesis. Squid late-stage embryos appear to be more impacted by warming via metabolic suppression than cuttlefish embryos. At all temperature scenarios, the transition from encapsulated embryos to planktonic paralarvae implied metabolic increments higher than 100 %. Contrary to the nektobenthic strategy of cuttlefish newborns, the planktonic squid paralarvae rely predominantly on pulsed jet locomotion that dramatically increases their energy requirements. In the future, hatchlings will require more food per unit body size and, thus, feeding intake success will be crucial, especially for squid with high metabolic rates and low levels of metabolic reserves.     

Relationships between seston,available food and feeding activity in the common mussel Mytilus edulis

The feeding and metabolic rates of Mytilus edulis L. of different body sizes were measured in response to changes in particle concentrations ranging from 2 to 350 mg l^-1. Rates of oxygen consumption were not significantly affected by changes in seston concentration, whereas clearance rates gradually declined with increasing particle concentration. Pseudofaeces production was initiated at relatively low seston concentrations (<5 mg l^-1). Marked seasonal changes were recorded in the composition of suspended particulates (seston) in an estuary in south-west England. Total seston was sampled at frequent intervals throughout an annual cycle and analysed in terms of: particle size-frequency distributions, total dry weight (mg l^-1), inorganic content, chlorophyll a, carbohydrate, protein and lipid. The particulate carbohydrate, protein and lipid content provided an estimate of the food content of the seston. The results are discussed in terms of the food available to a nonselective suspension feeder, such as M. edulis, during a seasonal cycle. The effect of inorganic silt in suspension was mainly to limit by dilution the amount of food material ingested rather than to reduce the amount of material filtered by the mussel. In winter, the food content of the material ingested was 5%, and this increased to 25% during the spring and summer.