Omnivorous feeding behavior of the Antarctic krill Euphausia superba

Feeding experiments were conducted at Palmer Station from December 1985 to February 1986 to examine the potential role of copepod prey as an alternative food source for Euphausia superba. Copepod concentration, copepod size, phytoplankton concentration, the duration of krill starvation and the volume of experimental vessels were altered to determine effects on ingestion and clearance rates. Krill allowed to feed on phytoplankton and copepods in 50-litre tubs showed greatly increased feeding rates relative to animals feeding in the much smaller volumes of water traditionally used for krill-feeding studies. Clearance rates on copepods remained constant over the range of concentrations offered, but clearance rates on phytoplankton increased linearly with phytoplankton concentration. Feeding rates increased when larger copepods were offered and when krill were starved for two weeks prior to experiments. Clearance rates of krill feeding on copepods were higher than, but not correlated with, their clearance rates on phytoplankton in the same vessel. E. superba may have a distinct mechanism for capturing copepods, perhaps through mechanoreception. Although our observed clearance rate of 1055 ml krill^-1 h^-1 indicates that krill can feed very efficiently on copepod prey, such feeding would meet less than 10% of their minimum metabolic requirements at the typical copepod concentrations reported for Antarctic waters. However, substantial energy could be gained if krill fed on the patches of high copepod concentrations occasionally reported during the austral summer, or if krill and copepods were concentrated beneath the sea ice during the winter or spring months. Our results, indicating efficient feeding on zooplankton and higher clearance rates on phytoplankton than previously believed, represent a step towards balancing the energy budget of E. superba in Antarctic waters.     

Respiration rate and cost of swimming for Antarctic krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in large groups in the laboratory

Constructing realistic energy budgets for Antarctic krill, Euphausia superba, is hampered by the lack of data on the metabolic costs associated with swimming. In this study respiration rates and pleopod beating rates were measured at six current speeds. Pleopod beating rates increased linearly with current speed, reaching a maximum of 6 beats s^–1 at 17 cm s^–1. There was a concomitant linear increase in respiration rate, from 1.8 mg O₂ g_D^–1 h^–1 at 3 cm s^–1 to 8.0 mg O₂ g_D^–1 h^–1 at 17 cm s^–1. The size of the group tested (50, 100 and 300 krill) did not have a significant effect on pleopod beating rates or oxygen consumption (ANCOVA, F=0.264; P>0.05). The cost of transport reached a maximum of 75 J g^–1 km^–1 at 5 cm s^–1, and then decreased with increasing current speed to 29 J g^–1 km^–1. When considered in light of energy budgets for E. superba, these data indicate that the cost of swimming could account for up to 73% of total daily metabolic expenditure during early summer.Communicated by G.F. Humphrey, Sydney     

Schooling behavior of Antarctic krill (Euphausia superba) in laboratory aquaria: Reactions to chemical and visual stimuli

Antarctic krill,Euphausia superba, often exhibit abnormal behavior in laboratory aquaria, usually hovering in a stationary position, unresponsive to most external stimuli. In the austral summer of 1985–1986 at Palmer Station on Anvers Island, Antarctica, we provided laboratory conditions which inducedE. superba to school in large aquaria. Captive krill swam horizontally and exhibited the full spectrum of behaviors normally displayed while schooling at sea. Schooling krill avoided visually contrasting stimuli, with avoidance distances correlated with stimulus size. Schools responded in qualitatively different ways to presentations of food, chemical compounds, and abrupt increases in light intensity. We describe the conditions necessary for aquarium schooling and discuss the importance of an appropriate social environment for displays of escape, avoidance, and feeding behaviors and of positional preference within the school.     

Contribution of heterotrophic material to diet and energy budget of Antarctic krill, Euphausia superba

A novel approach was used to estimate the heterotrophic carbon component in the diet of the Antarctic krill Euphausia superba. Over 200 specimens from seven samples collected in the Lazarev Sea (January 1993 and 1995), at the Antarctic Polar Front (January 1993), and around South Georgia (February/March 1994) were dissected, and the total carbon content of their stomachs was estimated with a CHN-analyser. Gut-pigment contents were also measured by the gut-fluorescence technique in specimens collected at the same time, and the equivalent amount of their gut carbon was then subtracted from the total organic carbon content of guts from the same samples. The remaining carbon was assumed to originate entirely from heterotrophic food sources. This heterotrophic component accounted for a substantial proportion of the total food consumed by Antarctic krill, ranging from 17.4 to 98.9% of the mass of the gut contents (mean = 78.8% ± 21.2 SD). The results make an important contribution to the elucidation of the energy budget of krill and its daily carbon ration. With a few exceptions, previous estimates were largely calculated from a solely autotrophic carbon source, and were unable to account for the metabolic requirements of E. superba. Krill plays an important role in Antarctic food webs, as it often constitutes ≃50% of the total biomass of the zooplankton, and produces fast-sinking, dense faecal pellets which are important in the vertical transport of organic carbon from the euphotic layer to the deep ocean. High consumption rates of smaller heterotrophic organisms by krill suggest that this large microphage may be more important than previously believed in re-packaging micro- and mesozooplankton into a longer-lasting and more easily sequestered carbon pool. Received: 26 October 1998 / Accepted: 14 October 1999     

Effect of temperature on the molting,growth and maturation of the antarctic krill Euphausia superba (Crustacea: Euphausiacea) under laboratory conditions

The effect of temperature on molting, growth, and maturation rates was studied on laboratory-maintained Euphausia superba. The length of intermolt periods (IMP's) was inversely proportional to temperature (20.10 d, SD=1.60, at 0.12°C; 16.87 d, SD=1.68, at 0.97°C; and 12.48 d, SD=0.90, at 4.48°C), and directly proportional to krill size at 0.12°C and 0.97°C. For individually maintained krill the maximum growth rate at 4.48°C (0.068 mm d^-1) was nearly twice that at 0.68°C (0.037 mm d^-1). There was no observable temperature effect on maturation rates. The maturation changes of juveniles at all temperatures indicated that more than two years are probably required to reach maturity. Mature males and females regressed to immature forms, suggesting that E. superba may reproduce in successive years. These results and previously reported field and laboratory data for E. superba and other euphausiid species suggest a 4+ year life span for this species.This work was supported by NSF grant DPP 76-23437     

Reproductive trade-off in male Antarctic krill,Euphausia superba

Freshly caught male and female Euphausia superba from the same swarm exhibited different rates of mortality subsequent to capture. Mortality was significantly higher for reproductive males (100%, n=68) than for females (3%, n=186) within the first 3 d of capture. Total lipid and triacylglycerol levels in male, female and juvenile Euphausia superba were analysed and compared. All reproductive male krill analysed from this swarm had low lipid levels (1 to 3% dry weight) with negligible triacylglycerol stores (0 to 2% of total lipid). Somatic lipid stores in female and juvenile krill ranged from 8 to 30% of which up to 40% was triacylglycerol. The levels of algal sterols in the digestive gland of males, females and juveniles indicate that all krill had been feeding recently. An analysis of the sex ratio of krill catches derived from data collected over seven summers from the Prydz Bay region showed a decrease in the proportion of males with increasing size. There was a sharp decline in numbers of male krill once they attained a length of 51 to 55 mm. Low lipid levels in redroductive male krill may be due to reproductive costs. The resulting low storage-lipid levels are accompanied by high mortality in male krill.     

Metabolism and body composition of two euphausiids (Euphausia superba and E. crystallorophias) collected from under the pack-ice off Enderby Land,Antarctica

Oxygen consumption, ammonia excretion and phosphate excretion rates were measured for Euphausia superba and E. crystallorophias captured under the pack-ice off Enderby Land, Antarctica, during November 1985. Water, ash, carbon, nitrogen and phosphorus composition were also determined. Compared with summer data, body composition of E. superba at this time of year was characterized by low carbon (40.2 to 40.7% of total dry wt), high nitrogen (11.8 to 12.0%), high phosphorus (1.2%) and high water (80.6 to 81.7% of total wet wt) content. Metabolic rates, expressed as percentage daily losses of body carbon, nitrogen and phosphorus were 1.3 to 1.6%, 0.38 to 0.42% and 1.9 to 2.2%, respectively; these fall within the ranges of summer data. E. superba fed on algae growing on the bottom of the ice, but body compositional features of this species suggest limited contribution of ice-algae to nutrition. Compared to E. superba, E. crystallorophias displayed greater metabolic rates and much higher body carbon content (40.9 to 45.0%), implying that feeding conditions under the ice were better suited to the requirements of E. crystallorophias. The only gravid females found belonged to E. crystallorophias.     

Growth of Antarctic krill Euphausia superba at South Georgia

 Antarctic krill Euphausia superba has a central role in the ecosystem of the Southern Ocean and knowledge of its growth rate is central to determining the factors influencing population dynamics. The length of Antarctic krill in the diet of Antarctic fur seals Arctocephalus gazella at South Georgia revealed a consistent increase in size between ca. 42 and ca. 54 mm over the period October–March, indicating growth rates much higher than predicted by existing models. Geographical variation in growth rate may result in 2-year-old krill at South Georgia attaining the same size as 3-year-old krill in the Antarctic Peninsula region. The effect of geographical variation in growth rate on the population structure of krill has important implications for comparing the fate of individual cohorts over large scales and in the interpretation of krill life-cycles. Received: 20 May 2000 / Accepted: 11 August 2000     

Zooplankton grazing on the coccolithophore Emiliania huxleyi and its role in inorganic carbon flux

Grazing and faecal pellet production by the copepods Calanus helgolandicus and Pseudocalanus elongatus, feeding on the coccolithophore Emiliania huxleyi, were measured under defined laboratory conditions, together with the chemical characteristics and sinking rates of the faecal pellets produced. Ingestion rates of both copepods were equivalent at comparable cell concentrations, the relationship between ingestion rate (I, cells copepod^-1 h^-1) and food concentration (C, cells ml^-1), being I=0.558C for both species. P. elongatus produced a larger number of smaller faecal pellets than C. helgolandicus, but egested a larger volume of material per individual. Only between 27 and 50% of the ingested coccolith calcite was egested in the faecal pellets, and it is possible that acid digestion in the copepod gut is responsible for these considerable losses. Average sinking rates of faecal pellets containing E. huxleyi coccoliths, produced by both species, were >100 m d^-1. The implications of the quantitative laboratory estimates for the vertical flux of inorganic carbon are considered using recently studied shelf-break and oceanic E. huxleyi blooms in the N. E. Atlantic as examples.     

Gut evacuation rates and pigment destruction in the Antarctic krill Euphausia superba

Krill grazing data collected during cruises in the region of the Antarctic Polar Front (S.A. Agulhas Voyage 70) and the South Georgia shelf (R.V. Africana Voyage 119) during the austral summer of 1993 were analyzed to estimate the variability of crucial parameters of the gut fluorescence technique in relation to food availability and krill feeding history. Gut evacuation rates (k) and passage or throughput times (1/k) varied in the ranges of 0.101 to 0.424 h^-1 and 2.3 to 9.9 h and were strongly correlated (p<0.001, r ²=0.98) to krill feeding activity (estimated as initial gut pigment content, G₀) but not to ambient chlorophyll a concentration. A significant differences was found when k values derived from incubations in filtered seawater and low charcoal particle concentrations (0.4 to 0.8 mgl^-1) were compared with values derived from krill fed high concentrations of charcoal (6 mgl^-1). The efficiency of gut pigment destruction was among the highest recorded for zooplankton organisms, 58.1 to 98.4%, and did not covary significantly (p>0.05) with ambient food concentration. However, the pigment lost per individual krill was strongly correlated with the total amount of pigment ingested (p<0.001, r ²=0.99). We suggest that both gut evacuation rates and pigment destruction efficiency may be realistically estimated only when krill is allowed to continue ingesting particles uninterruptedly. Charcoal particle concentration should be equivalent to the in situ wet weight of total seston per unit volume. An objective criterion for the standardization of the measurement and calculation of k values is also proposed.     

Seasonality of feeding and nutritional status during the austral winter in the Antarctic sea urchin Sterechinus neumayeri

The seasonal pattern of food intake and tissue energy status was measured over a 2 year period for two populations of the common Antarctic echinoid Sterechinus neumayeri living on contrasting substrata at Rothera Point, Adelaide Island, Antarctica. Food intake was estimated from faecal egestion, and both tissue mass and energy content assessed. Food availability was intensely seasonal, in that water column chlorophyll content and sediment pigment content varied markedly throughout the year. In response, both urchin populations showed an extremely seasonal cycle of faecal egestion, indicating a strong seasonality of feeding activity. Urchins from North Cove living on soft sediment fed at a considerably higher rate, and had a significantly larger Aristotle's lantern, than those in South Cove, 1 km away, living on hard substrata and taking a more cosmopolitan diet. Faecal egestion in both populations was zero for a 7 month period in the austral winter of 1997, and again for a 4 month period in 1998. During the austral summer of 1997/1998, 84% of the total annual energy intake took place in the period January to March. Significant decrease in gut tissue mass provided energy for maintenance in early winter, although progressive reduction of both gonad energy content (but without a detectable change in gonad mass) and body wall organic mass provided energy during the late winter period. The mass of reproductive tissue showed large differences between the two sites, but there was no marked decrease associated with spawning. Tissue proximate composition was assessed stoichiometrically from elemental composition and also checked by direct assay; gonad tissue was richer in lipid than gut tissue, though both were dominated by protein and contained only small amounts of carbohydrate. These data suggest that the very strong seasonality of food intake does not pose a significant energetic challenge for this species.     

Metabolic activity and elemental composition of krill and other zooplankton from Prydz Bay,Antarctica, during early summer (November–December)

Oxygen uptake and ammonia excretion rates, and body carbon and nitrogen contents were measured in krill (Euphausia superba) and eight other zooplankton species collected during November–December 1982 in the Prydz Bay, Antarctica. From these data, metabolic O:N ratios (by atoms), body C:N ratios (by weight) and daily metabolic losses of body carbon and nitrogen were calculated as a basis from which to evaluate seasonal differences in metabolism and nutritional condition. Comparison of the present data with mid-summer (January) data revealed that early-summer E. superba were characterized by higher metabolic O:N ratios (58.7 to 103, compared with 15.9 to 17.5 for mid-summer individuals). Higher O:N ratios of early-summer E. superba resulted largely from reduced ammonia excretion rates and, to a lesser degree, from slightly increased oxygen uptake rates. Body C:N ratios of E. superba were low in early-summer (3.8 to 4.2) compared with mid-summer krill (4.1 to 4.7) due to lowered body-carbon contents in the former (42.6 to 43.6% compared with 43.2 to 47.5% dry weight of midsummer individuals); gravid females formed an exception, since no seasonal differences in body elemental composition were detected for these. No significant changes in water content (75.3 to 81.4% wet wt) and nitrogen content (9.9 to 11.1% dry wt) in E. superba were evident between the two seasons. Seasonal differences in metabolic rates and elemental composition were less pronounced in a salp (Salpa thompsoni), but a higher metabolic O:N ratio occurred in early-summer individuals. Interspecific comparison of the seven remaining zooplankton species studied with twelve species from mid-summer zooplankton investigated in an earlier study indicated that higher metabolic O:N ratios in early-summer are characteristic not only of herbivore/filter-feeders, but also of some carnivores/omnivores. No relationship between metabolic O:N ratios and body C:N ratios was apparent either intraspecifically or interspecifically, within or between early-summer and mid-summer seasons.     

Moult in relation to some aspects of reproduction and growth in swarms of Antarctic krill,Euphausia superba

In a 14-d period 38 swarms of Antarctic krill,Euphausia superba Dana, were sampled in an area 55.5 x 55.5 km to the southwest of Elephant Island, at the north of the South Shetland Islands. Moult stage, maturity stage, and size of ca. 100 krill from each swarm were measured. Each of the characteristics varied greatly between swarms. Moulting krill were found in most swarms, but in one swarm all of the krill were just about to moult. Ways in which moulting may act as a possible sorting mechanism are discussed. Data for all the analysed krill (ca. 3000 specimens) were used to investigate the interdependence of moult rate, sexual maturation and growth. While all immature krill moulted at approximately the saine rate in the study, there were significant differences in the moulting rates of mature male and female krill. Gravid female krill continued to moult, although less frequently than mature males. As a consequence males had to attach spermatophores to females after each moult. It is likely that variation in moult rate of females and males was related to the energy expenditure required for ovary development in females and spermatophore production and searching behaviour in males. Spawning and moulting were only partly coupled in gravid females. Spawning appeared to take place predominantly during Moult Stage D₂. There was no evidence of intermoult growth by intersegmental dilation.     

Sampling biological characteristics of krill: Effect of heterogeneous nature of swarms

An effect of the very patchy distribution of Antarctic krill (Euphausia superba Dana) in various forms of aggregation is that a single swarm may not provide an unbiased estimate of population parameters such as mean length of krill in the local area. Here, we analyse the number of samples required to estimate the characteristics of a local population as precisely as if there were no differences between krill swarms in terms of their biological composition. Krill were intensively sampled over different spatial and temporal scales around South Georgia in 1981 and 1982, and in the Bransfield Strait in 1985. These varied from replicate hauls at a single station over 24 h and repeat sampling in restricted areas over periods of 6 to 14 d to regional surveys around South Georgia and in the Bransfield Strait. Various biological characteristics were measured such as length, maturity, moult stage and feeding state. Depending upon the biological characteristic examined and the area covered by the sampling programme, the number of samples needed to obtain the same degree of precision as would be found in the absence of heterogeneity varied from 3 to > 80 samples. This has important implications for the design of net-sampling programme for monitoring krill populations.     

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.     

Comparative lipid dynamics of euphausiids from the Antarctic and Northeast Pacific Oceans

To better understand the feeding and reproductive ecology of euphausiids (krill) in different ocean environments, lipid classes and individual lipid components of four different species of euphausiids from Northeast Pacific (temperate species) and Southern Ocean (Antarctic species) were analyzed in animals from multiple life stages and seasons. The dominant krill species in the Northeast Pacific Euphausia pacifica and Thysanoessa spinifera, were compared to the two major Antarctic species, Euphausia superba and E. crystallorophias. Analysis comprised total lipid and lipid classes together with individual fatty acid and sterol composition in adults, juveniles, and larvae. Antarctic krill had much higher lipid content than their temperate relatives (10–50 and 5–20% of dry mass for Antarctic and temperate species, respectively) with significant seasonal variations observed. Phospholipids were the dominant lipid class in both temperate krill species, while neutral storage lipids (wax esters and triacylglycerols for E. crystallorophias and E. superba, respectively) were the major lipid class in Antarctic krill and accounted for up to 40% of the total lipid content. Important fatty acids, specifically 16:0, 18:1ω9, 20:5ω3, and 22:6ω3, were detected in all four krill species, with minor differences between species and seasons. Detailed lipid profiles suggest that krill alter their lipid composition with life stage and season. In particular, larval Antarctic krill appear to utilize alternate food resources (i.e., sea-ice associated organisms) during austral winter in contrast to juveniles and adults (i.e., seston and copepods). Lipid dynamics in krill among krill in both systems appear closely linked to their life cycle and environmental conditions including food availability, and can provide a more complete comparative ecology of euphausiids in these environmentally distinct systems.     

Growth and grazing loss rates in single-celledPhaeocystis sp. (Prymnesiophyceae)

Growth and grazing loss rates of naturalPhaeocystis sp. single cells were measured using a seawater dilution technique. Measurements were performed during an intensePhaeocystis sp. bloom in the North Sea between 19 April and 5 May 1988. Experimental results yielded rapid carbon turnover rates. Population growth rates varied from 0.033 to 0.098 h^–1, grazing loss rates from 0.037 to 0.174 h^–1. From measured growth rates, average doubling rages of 1.3 doublings d^–1 were calculated. The growth rates would have resulted in maximum carbon production rates of 146 mg C m^–3 d^–1. Grazing rates increased in the course of the bloom and exceeded growth rates at the end. Grazing loss was caused primarily by microzooplankton feeding. Ciliates and heterotrophic dinoflagellates were identified as the major potential consumers of single cells ofPhaeocystis sp. at the beginning of the bloom. The grazing impact of larger microzooplankton species appeared to increase during the progressing bloom.     

The tunicate Salpa thompsoni ecology in the Southern Ocean. I. Distribution, biomass, demography and feeding ecophysiology

Distribution, density, and feeding dynamics of the pelagic tunicate Salpa thompsoni have been investigated during the expedition ANTARKTIS XVIII/5b to the Eastern Bellingshausen Sea on board RV Polarstern in April 2001. This expedition was the German contribution to the field campaign of the Southern Ocean Global Ocean Ecosystems Dynamics Study (SO-GLOBEC). Salps were found at 31% of all RMT-8 and Bongo stations. Their densities in the RMT-8 samples were low and did not exceed 4.8 ind m⁻² and 7.4 mg C m⁻². However, maximum salp densities sampled with the Bongo net reached 56 ind m⁻² and 341 mg C m⁻². A bimodal salp length frequency distribution was recorded over the shelf, and suggested two recent budding events. This was also confirmed by the developmental stage composition of solitary forms. Ingestion rates of aggregate forms increased from 2.8 to 13.9 μg (pig) ind⁻¹ day⁻¹ or from 0.25 to 2.38 mg C ind⁻¹ day⁻¹ in salps from 10 to 40 mm oral-atrial length, accounting for 25–75% of body carbon per day. Faecal pellet production rates were on average 0.08 pellet ind⁻¹ h⁻¹ with a pronounced diel pattern. Daily individual egestion rates in 13 and 30 mm aggregates ranged from 0.6 to 4.8 μg (pig) day⁻¹ or from 164 to 239 μg C day⁻¹. Assimilation efficiency ranged from 73 to 90% and from 65 to 76% in 13 and 30 mm aggregates, respectively. S. thompsoni exhibited similar ingestion and egestion rates previously estimated for low Antarctic (~50°S) habitats. It has been suggested that the salp population was able to develop in the Eastern Bellingshausen Sea due to an intrusion into the area of the warm Upper Circumpolar Deep Water     

Depth distribution of developing Euphausia superba embryos,predicted from sinking rates

Sinking rates of the embryos of Euphausia superba were measured during development in the laboratory at Palmer Station. Antarctic Peninsula, during austral summer 1982. Two densities of seawater were used in these experiments; the lower density (1.027 g cm^-3) is characteristic of the upper 300 m of the water column, the higher (1.029 g cm^-3) is characteristic of the deeper circumpolar deep water. During development, the eggs sank at an initial average rate of 175 m d^-1, declining to 51 m d^-1 during the gastrula to early limb bud stages and increasing before hatching to near initial rates. The larvae hatched within 6 d. We predict that the majority of the embryos of E. superba should hatch at a depth of about 850 m.     

Adjustment of the components of energy balance in the gastropod Crepidula fornicata in response to thermal acclimation

Synchronous measurements were made of the routine rate of oxygen consumption and the clearance rate of Phaeodactylum tricornutum at different exposure temperatures by specimens of the suspension-feeding gastropod Crepidula fornicata which had been acclimated to temperatures between 10° and 25°C. The results show that the cost of activity (l O₂ consumed h^-1/ml seawater cleared h^-1) increases dramatically in individuals exposed to short-term increases of temperature up to 30°C, especially in limpets acclimated to 10°C. The process of thermal acclimation, however, results in two compensatory adjustments in energy expenditure and uptake which profoundly affect the energetics of water transport. Firstly, the routine oxygen consumption shows lateral translation of the rate-temperature curve which results in the maintenance of a relatively uniform energy expenditure despite an increase in acclimation temperature from 10° to 25°C. Secondly, because of the form of the rate-temperature curve for filtration by C. fornicata, lateral translation in response to warm acclimation results in an increase in the maximal clearance rate. Lateral translation of the rate-temperature curves for feeding rates and for oxygen consumption in response to thermal acclimation may thus be linked to maintain a balance between energy gain and expenditure. In this way, the greatly increased cost of activity which would occur with increase of temperature in the absence of acclimation is evaded. The minimal maintenance energy requirement, and hence the greatest scope for growth and reproduction, is then adjusted to coincide with temperatures prevailing in the environment.