Sinking rates of fecal pellets from the marine copepod Pontella meadii

Sinking rates of fecal pellets produced by the marine copepod Pontella meadii, grazing on 4 different phytoplankton diets, ranged from 15 to 153 m/day, with a mean of 66 m/day. Sinking rates, in general, were directly related to fecal pellet volumes, but unrelated to the diets used to produce the fecal pellets. There were two-to-threefold variations in sinking rates between fecal pellets of the same volumes, often produced on the same diets. Twenty repetitions of timed sinking of a single fecal pellet revealed sinking rates varying from 33 to 79 m/day, as well as variations in sinking rates within the course of individual descents. It is suggested that copepod fecal pellets are of such small volumes and densities that their sinking rates are subject to microstructural variations in the most carefully controlled water columns. Scanning electron microscope observations revealed lack of structural damage to some of the diatom frustules in the fecal pellets, suggesting that superfluous feeding may have occurred. Thus, the accelerated sinking rates of copepod fecal pellets may provide a mechanism for nutritional enrichment of the deep-sea ecosystem with organic parcels containing incompletely-assimilated plant material.     

Sinking rates of fecal pellets from gelatinous zooplankton (Salps,Pteropods, Doliolids)

Sinking rates were determined for fecal pellets produced by gelatinous zooplankton (salps, Salpa fusiformis and Pegea socia; pteropods, Corolla spectabilis; and doliolids, Dolioletta gegenbaurii) feeding in surface waters of the California Current. Pellets from the salps and pteropods sank at rates up to 2 700 and 1 800 m d^-1, respectively; such speeds exceed any yet recorded for zooplankton fecal pellets. Fecal pellets of salps were rich in organic material, with C:N ratios from 5.4 to 6.2, close to values for living plankton. The relation between volume and sinking rate indicates that salp and pteropod pellets are slightly less dense than those of pelagic Crustacea; moreover, pellet density varied between different collection dates, probably because of differences in composition. In contrast, doliolid pellets sank at rates up to 208 m d^-1, a rate much lower than would be expected from pellet size. Thus, density and sinking rates of pellets are much more variable in zooplankton than would be expected from studies of crustaceans alone. Moreover, the extraordinarily high sinking rates of fecal pellets of salps indicates that these tunicates may be disproportionately important in the flux of biogenic materials during periods when they form dense population blooms.     

Morphology and sinking velocities of fecal pellets of copepod, molluscan, euphausiid, and salp taxa in the northeastern tropical Atlantic

Morphological characteristics and sinking velocities of naturally occurring fecal pellets of copepods, euphausiids, salps, and pelagic mollusks collected in the northeastern tropical Atlantic were investigated during the period of May-June 1992. The fecal pellets of copepods and euphausiids were cylindrical and distinguished only by their size. Those of salps were, in general, rectangular, and slight differences were noted according to the species. The fecal pellets of the molluscan pteropod Clio sp. were conical, while those of the molluscan heteropod Carinaria sp. were spiral. The sinking velocities ranged from 26.5 to 159.5 m day^-1 for copepod fecal pellets, from 16.1 to 341.1 m day^-1 for euphausiid pellets, from 43.5 to 1167.6 m day^-1 for salps' pellets (Cyclosalpa affinis, Salpa fusiformis, Iasis zonaria, and two unidentified species), from 65 to 205.7 m day^-1 for Clio sp. pellets, and from 120.3 to 646.4 m day^-1 for Carinaria sp. fecal pellets. The measured sinking velocities were compared with estimates predicted using the equations of Komar et al. (1981; Limnol Oceanogr 26:172-180), Stokes' law, and Newton's second law, using either a constant density of fecal pellets (1.22 g cm^-3) or densities estimated with the three different equations. When a constant density was used, the three equations overestimated the sinking velocities; Stokes' law resulted in the largest overestimation, and Newton's second law, the smallest. At the taxa level, the overestimation was greatest for euphausiid 1 fecal pellets and smallest for copepod fecal pellets. When the three equations were used to estimate fecal pellet density, the density estimated using the equation of Komar et al. was the greatest, and that using Stokes' law, the smallest, resulting in over- and underestimation of sinking velocities, respectively. Newton's second law resulted in an intermediate density and gave the closest estimate of sinking velocities. We propose that measurement of sinking velocities of a portion of the fecal pellets might guide in choosing an appropriate equation to be used for a reasonable interpretation of vertical mass flux.     

Production and fate of copepod fecal pellets across the Southern Indian Ocean

The vertical distribution of copepods, fecal pellets and the fecal pellet production of copepods were measured at seven stations across the Southern Indian Ocean from productive areas off South Africa to oligotrophic waters off Northern Australia during October/November 2006. We quantified export of copepod fecal pellet from surface waters and how much was retained. Furthermore, the potential impact of Oncaea spp. and harpacticoid copepods on fecal pellets degradation was evaluated and found to be regional substantial. The highest copepod abundance and fecal pellet production was found in the western nutrient-rich stations close to South Africa and the lowest at the central oligotrophic stations. The in situ copepod fecal pellet production varied between 1 and 1,000 μg C m⁻³ day⁻¹. At all stations, the retention of fecal pellets in the upper 400 m of the water column was more than 99% and the vertical export of fecal pellets was low (<0.02 mg m⁻² day⁻¹).     

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.     

Production,composition and sedimentation of salp fecal pellets in oceanic waters

Rates of fecal pellet production have been recorded from seven species of oceanic salps feeding on natural diets. Expressed as g C defecated per mg salp body C per hour, the values range between 3.7 and 27.7. Carbon: nitrogen ratios of the salp fecal pellets average 11.4; the organic matter of the pellets is mainly protein and carbohydrate. Sinking velocities of the pellets are very high, ranging from 320 to 2 238 m d^-1 for pellets from three species. However, the pellets sink slower than would be predicted from extrapolation of rates for crustacean pellets, probably due to the shape of the pellets and their density. The high rates of defecation, large size and rapid sedimentation of salp fecal pellets make them likely mechanisms for rapid transport of small particulate matter from surface waters to deep water and the benthos.     

Interactions of marine plankton with transuranic elements

In a series of laboratory experiments, the biokinetics of ²⁴¹Am, an important transuranium element, was studied in Meganyctiphanes norvegica, a euphausiid common in the northwestern Mediterranean. The euphausiids accumulated Am from water by passive adsorption onto exoskeletons, achieving wet weight concentration factors on the order of 10² after 1 wk exposure; concentration factors varied inversely with the size of the euphausiids and linearly with their surface area:wet weight ratios. Essentially all (96±10%) of the Am taken up from water was associated with the exoskeleton, so that negligible Am was retained by the euphausiids after molting. The retention half-time of Am in molts was 2.9 d. Euphausiids could also concentrate Am from feeding suspensions by ingesting Am-labelled diatom cells, although there was negligible Am assimilation (3±2% after 4 d feeding); after passage through the gut, virtually all (99%) of the ingested Am was defecated within 1 wk. The retention half-time of Am in fecal pellets was 41 and 51 d at 13° and 5°C, respectively. In oceanic waters, where the preponderance of ²⁴¹Am is in the dissolved phase, uptake of Am from water by euphausiids would be the dominant route of bioaccumulation. The results underscore the importance of sinking biogenic debris from zooplankters in mediating the vertical transport of Am in the sea. Given their retention half-times for ²⁴¹Am and their rapid sinking rates, fecal pellets and discarded molts have the potential to deliver most of their Am to the sediments.     

Differential feeding and fecal pellet composition of salps and pteropods,and the possible origin of the deep-water flora and olive-green “Cells”

Salps (mainly Salpa fusiformis and, to a lesser extent, Pegea socia) and a web-building pteropod (Corolla spectabilis) were studied in epipelagic waters of the central California Current. Although both kinds of gelatinous zooplankton trap phytoplankton in a mucus net, a fecal pellet analysis indicated that their diet differs significantly when they feed together, probably because of differences both in the pore sizes of their nets and in their feeding methods. Salps have a finemesh filter, on which they can retain even the smallest phytoplankton; thus, when small coccolithophores are abundant, as they were in our study, salp feces contain such cells and the coccoliths derived from them. In contrast, pteropods feeding in the same area produce fecal pellets consisting chiefly of larger phytoplankton, especially diatoms. Since fecal pellets transport most biogenic material to the deep sea, changes in herbivore species composition at a given geographic location can change the chemistry of materials entering deep water; at our study site, the more salps, the greater the calcite flux, and, the more pteropods, the greater the silica flux. In addition, fecal pellets of both salps and pteropods include partially digested residues of phytoplankton that appear as olive-green spheres, having an ultrastructure identical with that of the socalled olive-green cells. Presumably, fecal pellets, after sinking into deep water, ultimately disintegrate. releasing both the viable phytoplankton and the olive-green spheres into aphotic waters. Thus the feces of epipelagic herbivores are likely sources of much of the flora of the deep ocean.     

Role of sinking in diatom life-history cycles: ecological,evolutionary and geological significance

Rapid mass sinking of cells following diatom blooms, observed in lakes and the sea, is argued here to represent the transition from a growing to a resting stage in the life histories of these algae. Mass sinking is of survival value in those bloom diatoms that retain viability over long periods in cold, dark water but not in warm, nutrient-depleted surface water. Mechanisms for accelerating sinking speed of populations entering a resting or seeding mode are proposed. Previously unexplained features of diatom form and behaviour take on a new meaning in this context of diatom seeding strategies. Diatoms have physiological control over buoyancy as declining growth is accompanied by increasing sinking rates, where the frustule acts as ballast. Increased mucous secretion in conjunction with the cell protuberances characteristic of bloom diatoms leads to entanglement and aggregate formation during sinking; the sticky aggregates scavenge mineral and other particles during descent which further accelerates the sinking rate. Such diatom flocs will have sinking rates of 100 m d^-1 or more. This is corroborated by recent observations of mass phytoplankton sedimentation to the deep sea. This mechanism would explain the origin of marine snow flocs containing diatoms in high productivity areas and also the well-known presence of a viable deep sea flora. That mortality is high in such a seeding strategy is not surprising. A number of species-specific variables pertaining to size, morphology, physiology, spore formation and frustule dissolution rate will determine the sinking behaviour and thus control positioning of resting stages in the water column or on the bottom. It is argued that sinking behaviour patterns will be environmentally selected and that some baffling aspects of diatom form and distribution can be explained in this light. Rapid diatom sedimentation is currently believed to be mediated by zooplankton faecal pellets, particularly those of copepods. This view is not supported by recently published observations. I speculate that copepod grazing actually retards rather than accelerates vertical flux, because faecal pellets tend to be recycled within the surface layer by the common herbivorous copepods. Egestion of undigested food by copepods during blooms acts as a storage mechanism, as ungrazed cells are likely to initiate mass precipitation and depletion of the surface layer in essential elements. Unique features of diatoms are discussed in the light of their possible evolution from resting spores of other algae. An evolutionary ecology of pelagic bloom diatoms is deduced from behavioural and morphological characteristics of meroplanktonic and tychopelagic forms. Other shell-bearing protistan plankters share common features with diatoms. Similar life-history patterns are likely to be present in species from all these groups. The geological significance of mass diatom sinking in rapidly affecting transfer of biogenic and mineral particles to the sea floor is pointed out.     

Sinking rates and dissolution of midwater fish fecal matter

The sinking rates of fecal matter from 7 southern California midwater fish species were investigated. Feces were obtained from 162 specimens of Stenobrachius leucopsarus, Triphoturus mexicanus, Leuroglossus stilbius, Lampanyctus ritteri, Argyropelecus affinis and Parvilux ingens, which were collected in the Santa Barbara and San Clemente Basins between 1977 and 1979. In addition, feces obtained from 6 laboratory-maintained specimens of the midwater zoarcid Melanostigma pammelas were used for repeated sinking-rate measurements. The mean of the measured sinking rates for all species was 1.19 cm s^-1 (1 028 m d^-1), which is much higher than the known descent rates of euphausiid and copepod fecal pellets and of most other particulate organic detritus. Dissolution characteristics were also investigated for fecal matter from 4 species collected by the same series of net hauls: S. leucopsarus, T. mexicanus, A. affinis, and Sternoptyx obscura. The release of dissolved organic compounds from this material is low and does not represent a significant output during the relatively short time required to sink through the water column. These findings suggest that midwater fish fecal matter may represent a major source of organic transfer between the pelagic community and the benthos.     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Comparison of the chemical composition of particulate material and copepod faecal pellets at stations off the coast of Labrador and in the Gulf of St. Lawrence

Faecal pellets were collected in 1988 from copepods which had fed in situ or in laboratory experiments, using screened natural seawater as food, at two stations off the coast of Labrador and one in the Gulf of St. Lawrence. The chemical composition of the pellets and of particulate material in profiles and in laboratory food were measured in terms of particulate carbon, carbohydrate (soluble and insoluble), protein and lipid. Faecal pellet composition was somewhat similar in all experiments at the first two stations, where the compositions of particulate material in situ and copepod species assemblages were also similar. At the third station the compositions of faecal pellets and particulate material were slightly different from those at the other stations and the copepod species composition varied between sampling times. Faecal pellets at the first two stations had very low levels of soluble carbohydrate, while concentrations in the food were generally high, suggesting that it was efficiently metabolized by copepods, although it might have been absent because of sloppy feeding or release, after passage through the gut, in soluble form or from faecal pellets. Comparisons of POC: biogenic silica ratios in food and faecal pellets, calculated using data presented elsewhere (Head 1992; Mar. Biol. 112: 583–592), suggested that at these stations, where food concentrations were high (chlorophyll concentrations>8 gl^-1), copepods may have been assimilating carbon rather inefficiently.     

An approach for particle sinking velocity measurements in the 3–400?μm size range and considerations on the effect of temperature on sinking rates

The flux of organic particles below the mixed layer is one major pathway of carbon from the surface into the deep ocean. The magnitude of this export flux depends on two major processes—remineralization rates and sinking velocities. Here, we present an efficient method to measure sinking velocities of particles in the size range from approximately 3–400?μm by means of video microscopy (FlowCAM^?). The method allows rapid measurement and automated analysis of mixed samples and was tested with polystyrene beads, different phytoplankton species, and sediment trap material. Sinking velocities of polystyrene beads were close to theoretical values calculated from Stokes’ Law. Sinking velocities of the investigated phytoplankton species were in reasonable agreement with published literature values and sinking velocities of material collected in sediment trap increased with particle size. Temperature had a strong effect on sinking velocities due to its influence on seawater viscosity and density. An increase in 9?°C led to a measured increase in sinking velocities of ~40?%. According to this temperature effect, an average temperature increase in 2?°C as projected for the sea surface by the end of this century could increase sinking velocities by about 6?% which might have feedbacks on carbon export into the deep ocean.     

Origins and microenvironments of bacteria mediating fecal pellet decomposition in the sea

The fecal pellets of zooplankton are thought to be major carriers of organic matter from surface to deeper waters of the oceans. As the pellets descend, they release soluble components, partially due to breakdown by associated microorganisms. Previous laboratory work of other investigators has suggested that the surface of a fecal pellet rapidly acquires bacteria, which increase in abundance until they and their protozoan consumers disrupt the pellet membrane, spilling contents into the water. In contrast, our field collections of fecal pellets from free-floating particle interceptor traps (from the Vertex project off Central California in 1980 and off Mexico in 1981), suggest that microbial decomposition probably is initiated in the sea from inside the fecal pellets. Transmission and scanning electron microscopy indicate that bacterial populations are most abundant in the interior of fecal pellets obtained from the sea, but that the same pellets will acquire the surface bacterial lawn typically observed in laboratory studies if maintained aboard ship. If the fecal pellets are decomposed from the inside, then the principal agents are enteric bacteria or ingested, digestion-resistant bacteria, or both. Such bacteria may differ metabolically from those that colonize the fecal pellet surfaces. Further-more, the abundance of healthy-appearing bacteria inside the pellets suggests that their metabolic activities may produce microhabitats of reduced oxygen tension that could differ considerably from that of the pellet exteriors. Decomposition in these semi-enclosed microenvironments may proceed in a manner not yet predicted by models that attribute decomposition to well-aerated, surface-dwelling bacterial populations on fecal pellets in the sea.     

Comparison of the food of Triphoturus mexicanus and T. nigrescens,two lanternfishes of the Pacific Ocean

Prey (chiefly euphausiids and copepods) eaten by two myctophids (lanternfishes) are compared from incidence in fish stomachs and from abundance in the environment. One lanternfish species, Triphoturus mexicanus, lives in the California Current, and the other, T. nigrescens, lives in the central Pacific Ocean. Although these two environments are very different physically and biologically, the feeding habits of the two lanternfishes are surprisingly similar. Prey biomass is 94% euphausiids, 3% copepods, and 3% other organisms for T. mexicanus and 88% euphausiids, 4.5% copepods, and 7.5% other organisms for T. nigrescens; the difference between the fish species is not significant when tested statistically. The two fishes resemble one another in frequency distributions of ingested copepod individuals, copepod species, euphausiid individuals, and euphausiid species. During a single diurnal feeding period, both fishes eat a variety of copepod species but tend to eat only a single species of euphausiid. T. mexicanus grows to twice the length of T. nigrescens and eats proportionally larger euphausiids; however, both fishes eat copepods having the same median size. The frequencies of euphausiid species in the diets of both fishes differ from the frequencies in the environment. The chief differences between the feeding habits of the two lanternfishes are that T. nigrescens, in comparison to its congener, eats a greater variety of organisms during one diurnal feeding period and captures smaller euphausiids. The feeding patterns for each lanternfish species are consistent over distances of hundreds of kilometers and over many years of sampling.     

Experimental studies on elimination of zinc-65, cesium-137 and cerium-144 by euphausiids

The elimination of 3 radionuclides from Euphausia pacifica was measured over a 5 month period. The biological half-lives for ⁶⁵Zn, ¹³⁷Cs, and ¹⁴⁴Ce, calculated after the euphausiids had ingested radioactive Artemia nauplii, were found to be 140 days, 6 days, and 7.5 h, respectively. The percentages of body burdens lost in molts were greatest for the fission products, ¹⁴⁴Ce (21%) and ¹³⁷Cs (7%), and least for ⁶⁵Zn (1%). Elimination of the isotopes in the feces could not be followed because of the difficulty in collecting fecal material for analysis; however, 1 sample collected 2 months after the beginning of the elimination experiment had no measurable radioactivity. Loss of ⁶⁵Zn from molts and time to disintegration of the molts were found to be temperature dependent over a 5° to 15°C range, and the sinking rate of molts was both temperature and salinity dependent. Calculations showed that, in areas in the North Pacific outside the influence of upwelling, percentage ⁶⁵Zn loss from sinking molts (before disintegration of the molts) was likely to be the same throughout the year, since the molts would be exposed to about the same mean temperature in the water column in all seasons. Even though temperature structure in the upper layers changes with season, mean temperatures change very little when calculated over the sinking distance of intact molts. Intact molts would sink to slightly over 400 m in the absence of turbulence, and would lose 87% of their ⁶⁵Zn by the time they reached this depth. Sinking molts thus might contribute substantially to the vertical transport of ⁶⁵Zn in the sea. If loss of ⁶⁵Zn in fecal pellets is assumed to be small under our experimental conditions, and molting loss is only 1% of ⁶⁵Zn body burden, the major mechanism of ⁶⁵Zn loss from euphausiids feeding on non-radioactive food must be isotopic exchange with the water. Approximately 96% of the initial body burden was eliminated over a period of 5 months.Supported by USAEC research contract AT (45-1)1830, PHS grant ES00026, and a Richland Graduate Fellowship to S. W. Fowler.     

Cyclical contributions of the digestive epithelium to faecal pellet formation by the copepod Calanus helgolandicus

Faecal pellet formation within the gut of Stage V and adult females of the copepod Calanus helgolandicus Claus involves (1) cyclical processes of digestion and (2) the contribution of parts of the gut epithelium to the pellets. During an experimental regime in which dim lighting was restricted to day-time and feeding to night-time (17.00 to 09.00 hrs), the copepods responded with cyclical changes in both the quantity of pellets they produced and the fine structure of the contents. During the feeding period, the contents showed changes in the relative amounts of materials originating from disintegrated cells of the digestive epithelium and those derived directly from the ingested food. The vacuolar B-cells of the gut contribute to the content of the pellets and the distal, necrotic N-cells appear to be involved in forming the peritrophic membrane which encloses each pellet. Cells of the gut epithelium which are broken down during feeding are all replaced during the non-feeding period. Other individuals were taken directly from the sea and in these, also, the cells of the gut broke down during feeding and contributed to the faecal pellets. The supply of epithelial cells may limit the duration of the feeding period.     

Flux of cadmium through euphausiids

Flux of the heavy metal cadmium through the euphausiid Meganyctiphanes norvegica was examined. Radiotracer experiments showed that cadmium can be accumulated either directly from water or through the food chain. When comparing equilibrium cadmium concentration factors based on stable element measurements with those obtained from radiotracer experiments, it is evident that exchange between cadmium in the water and that in euphausiid tissue is a relatively slow process, indicating that, in the long term, ingestion of cadmium will probably be the more important route for the accumulation of this metal. Approximately 10% of cadmium ingested by euphausiids was incorporated into internal tissues when the food source was radioactive Artemia. After 1 month cadmium, accumulated directly from water, was found to be most concentrated in the viscera with lesser amounts in eyes, exoskeleton and muscle, respectively. Use of a simple model, based on the assumption that cadmium taken in by the organism must equal cadmium released plus that accumulated in tissue, allowed assessment of the relative importance of various metabolic parameters in controlling the cadmium flux through euphausiids. Fecal pellets, due to their relatively high rate of production and high cadmium content, accounted for 84% of the total cadmium flux through M. norvegica. Comparisons of stable cadmium concentrations in natural euphausiid food and the organism's resultant fecal pellets indicate that the cadmium concentration in ingested material was increased nearly 5-fold during its passage through the euphausiid. From comparisons of all routes by which cadmium can be released from M. norvegica to the water column, it is concluded that fecal pellet deposition represents the principal mechanism effecting the downward vertical transport of cadmium by this species.     

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.