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.     

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.     

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.     

Metabolism and elemental composition of zooplankton from the Barents Sea during early Arctic summer

Rates of oxygen consumption, ammonia excretion and phosphate excretion were measured on a hydromedusae (Aglantha digitale), pteropods (Limacia helicina, Clione limacina), copepods (Calanus finmarchicus, C. glacialis, C. hyperboreus, Metridia longa), an amphipod (Parathemisto libellula), a euphausiid (Thysanoessa inermis) and a chaetognath (Sagitta elegans), all of which were dominant species in the Barents Sea during early summer 1987. Water and ash contents and elemental composition (C and N) were also analysed on the specimens used in these metabolic experiments. Between species variations were 67.8% to 94.7% of wet weight in water content, 6.4% to 56.5% of dry weight in ash content, 16.7% to 61.0% of dry weight in carbon content, and 4.3% to 11.2% of dry weight in nitrogen content. Oxygen consumption rates ranged from 0.33 to 13.8 l O₂ individual^-1 h^-1, ammonia excretion rates, from 0.0072 to 0.885 gN individual^-1 h^-1 and phosphate excretion rates, from 0.0036 to 0.33 g P individual^-1 h^-1. In general, higher rates were associated with larger species, but considerable differences were also seen between species. The ratios between the rates (O : N, N : P, O : P) exhibited a wide species-specific variation, indicating differences in dominant metabolic substrates. Typical protein oriented metabolism was identified only in S. elegans. From the results of metabolic rate measurements and elemental analyses, daily losses of body carbon and nitrogen were estimated to be 0.50 to 4.15% and 0.084 to 1.87%, respectively, showing faster turnover rates of carbon than that of nitrogen. Comparison of daily loss of body carbon of the Barents Sea zooplankton with that of the Antarctic zooplankton indicated reduced rates of the former (63% on average).     

Oxygen uptake,ammonia excretion and phosphate excretion by krill and other Antarctic zooplankton in relation to their body size and chemical composition

Oxygen uptake, ammonia excretion and phosphate excretion were measured in 14 Antarctic zooplankton species, including various size classes of krill (Euphausia superba), during a cruise to the Antarctic Ocean adjacent to Wilkes Land in the summer of 1980. Elemental composition (C, N and P) was also determined on the specimens used in these metabolic rate measurements. The values obtained for C, N and P were 4.7 to 47.5%, 1.2 to 12.5% and 0.09 to 1.23% of dry weight, respectively. Regression analyses of metabolic rates on different measures of body weight (fresh, dry, C, N and P) were made on krill, salps and other zooplankton as arbitrarily defined groups and also on the combined groups to determine the best measure of body weight for intra- and interspecific comparison of metabolic rates. The correlations were highly significant in all regressions, although no common measure of body weight provided the best correlation for the three groups of animals. Except for the regression of ammonia excretion on C and N weight, all other regressions of metabolic rates and body weights were significantly different within these three groups. In the combined group, oxygen uptake and ammonia excretion were better correlated to C and N weights than to dry and P weights. For phosphate excretion in the combined groups, dry weight gave the best correlation. Despite these results, the choice of a particular measure of body weight was shown to be important in a comparison of the rates between krill and salps because of their widely different chemical compositions. Our results of rate measurements are compared with those of previous workers for some Antarctic zooplankton, particularly krill. Some of the previous data are in good agreement with ours, while others are not. Possible contributing factors are considered in the latter case. The ratios between the rates (O:N, N:P and O:P) fell within the general ranges reported for zooplankton from different seas. The O:N ratio was consistently low (7.0 to 19.8, by atoms) in all species, suggesting the importance of protein in their metabolic substrates. Protein-oriented metabolism was also supported by the results of C and N analyses which indicated no large deposition of lipid in these animals. From the results of metabolic rate measurements and elemental analyses, daily losses in bodily C, N and P for Antarctic zooplankton in summer were estimated as 0.4 to 2.8%, 0.6 to 2.5% and 1.3 to 19.4%, respectively. These values are approximately one order of magnitude lower than those reported for subtropical and tropical zooplankton.     

Metabolism and elemental composition of four oncaeid copepods in the western subarctic Pacific

Respiration rates and elemental composition (carbon and nitrogen) were determined for four dominant oncaeid copepods (Triconia borealis, Triconia canadensis, Oncaea grossa and Oncaea parila) from 0–1,000 m depth in the western subarctic Pacific. Across the four species of which dry weight (DW) varied from 2.0 to 32 μg, respiration rates measured at in situ temperature (3°C) increased with DW, ranging from 0.84 to 7.4 nl O₂ individual⁻¹ h⁻¹. Carbon (C) and nitrogen (N) composition of the four oncaeid species ranged from 49–57% of DW and 7.0–10.3% of DW, respectively, and the resultant C:N ratios were 4.8–8.3. The high C contents and C:N ratios were reflected by large accumulation of lipids in their body. Specific respiration rates (SR, a fraction of body C respired per day) ranged between 0.5 and 1.3% day⁻¹. Respiration rates adjusted to a body size of 1 mg body N (i.e. adjusted metabolic rates, AMR) of the four oncaeid species [0.6–1.1 μl O₂ (mg body N)^−0.8 h⁻¹ at 3°C] were significantly lower than those (1.7–5.1) reported in the literature for oithonid and calanoid copepods at the same temperature. The present results indicate that lower metabolic expenditure due to less active swimming (pseudopelagic life mode) together with rich energy reserve in the body (as lipids) are the characters of oncaeid copepods inhabiting in the epi- and mesopelagic zones of this region.     

Metabolism and elemental composition of a giant chaetognath Sagitta gazellae from the Southern Ocean

Oxygen consumption, ammonia excretion and phosphate excretion rates were measured on Sagitta gazellae Ritter-Zahony, in conjunction with body composition analyses (water, ash, carbon, hydrogen, nitrogen and phosphorus). Both water content (94.7% of wet weight) and ash content (53.0% of dry weight) recorded on S. gazellae were the highest and the lowest, respectively, among the chaetognath data being reported. Contents of carbon, hydrogen and nitrogen of S. gazellae were the lowest among published values of chaetognaths. Metabolic comparison with other chaetognaths living in similar subzero water temperature revealed reduced rates in S. gazellae, while no appreciable differences were seen in the metabolic quotients (O:N, N:P and O:P ratios). The O:N atomic ratios were 10.5 to 15.9 indicating protein oriented metabolism. Reduced metabolic activity of S. gazellae is not due to their body composition as calculated daily metabolic losses of body carbon (0.50%), body nitrogen (0.38) and body phosphorus (1.6%) were also found to be lower than respective values reported on other congeners and even those of other zooplankton living in the Antarctic waters.     

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.     

Linking abundance and diversity of sponge-associated microbial communities to metabolic differences in host sponges

Many sponge species contain large and diverse communities of microorganisms. Some of these microbes are suggested to be in a mutualistic interaction with their host sponges, but there is little evidence to support these hypotheses. Stable nitrogen isotope ratios of sponges in the Key Largo, Florida (USA) area grouped sponges into species with relatively low δ¹⁵N ratios and species with relatively high δ¹⁵N ratios. Using samples collected in June 2002 from Three Sisters Reef and Conch Reef in the Key Largo, Florida area, transmission electron microscopy (TEM) and denaturing gradient gel electrophoresis were performed on tissues of the sponges Ircinia felix and Aplysina cauliformis, which are in the low δ¹⁵N group, and on tissue of the sponge Niphates erecta, which is in the high δ¹⁵N group. Results showed that I. felix and A. cauliformis have large and diverse microbial communities, while N. erecta has a low biomass of one bacterial strain. As the low δ¹⁵N ratios indicated a microbial input of nitrogen, these results suggested that I. felix and A. cauliformis were receiving nitrogen from their associated microbial community, while N. erecta was obtaining nitrogen solely from external sources. Sequence analysis of the microbial communities showed a diversity of metabolic capabilities among the microbes of the low δ¹⁵N group, which are lacking in the high δ¹⁵N group, further supporting metabolic differences between the two groups. This research provides support for hypotheses of mutualisms between sponges and their associated microbial communities.     

Salps in the Lazarev Sea,Southern Ocean: I. Feeding dynamics

Feeding dynamics of the Antarctic salps Ihlea racovitzai and Salpa thompsoni were studied in the Lazarev Sea in fall 2004, summer 2005–2006 and winter 2006. Pigment concentrations in the guts of both species were positively correlated with ambient surface chlorophyll a (chl a). No evidence was found for salp clogging even at dense surface concentrations of up to 7 μg chl a L⁻¹. However, gut pigment concentrations had a lower range than ambient pigment concentrations, suggesting that salps increased retention times of ingested material in low-food environments. For medium-sized I. racovitzai and S. thompsoni, estimated individual daily rations reached 7–10 and >100% of body carbon in winter and summer, respectively. Daily respiratory needs of I. racovitzai and S. thompsoni accounted for 28 and 22% of daily carbon assimilation based on pigment ingestion rates in winter, and for 2 and 1% in summer, respectively. The grazing impact of the salp populations on the phytoplankton standing stock was negligible during all seasons due to generally low salp densities. Fatty acid trophic biomarkers in the salps suggest high year-round contributions of flagellates and modest contributions of diatoms to the salp’s diet. These markers showed low seasonal variability for I. racovitzai. The more pronounced seasonality of trophic markers in S. thompsoni were likely related to their generally deeper residence depth in winter linked to a seasonal alternation of sexual and asexual generations.     

Relation between respiration,excretion (ammonia and inorganic phosphorus) and activity of amylase and trypsin in different species of pelagic copepods from an Indian Ocean equatorial area

The relationship between respiration, nitrogen and phosphorus excretion and specific activity of amylase and trypsin was investigated from shipboard experiments with several species of copepods from the equatorial divergence area of the Indian Ocean, in June 1978. Statistical analysis of O:N ratios in 6 of the most common species revealed three main groups: a group displaying low O:N (Pontella fera, Candacia pachydactyla); a group with high O:N (Undinula darwini, Euchaeta marina, Temora discaudata); and a species displaying an intermediate O:N ratio: Scolecithrix bradyi. O:P and N:P did not differ significantly between species. There was a direct relationship between average O:N and the ratios of specific activity of the digestive enzymes amylase and trypsin. Species displaying low O:N and/or A:T ratios such as P. fera, C. pachydactyla and Oncaea venusta probably metabolize proteins more efficiently than they do plant carbohydrates (high nitrogen excretion and low amylase activity). Species displaying high O:N and A:T ratios, such as U. darwini, E. marina and T. dicaudata (low nitrogen excretion and high amylase activity) were assumed to use carbohydrates (starch) and proteins with equal efficiency. S. bradyi showed a large range of variations in trypsin activity and low amylase activity, resulting in a low average A:T ratio, but its O:N ratio was intermediate. Variations in O:N and O:P ratios were related to differences in the nutritional strategy of the different species, based on literature data concerning the anatomy of their mouth parts and their selectivity for animal material in mixed-food experiments.     

Defecation by Salpa thompsoni and its contribution to vertical flux in the Southern Ocean

Measurements of the defecation rate of Salpa thompsoni were made at several stations during two cruises west of the Antarctic Peninsula in 2004 and 2006. Rates were quantified in terms of number of pellets, pigment, carbon and nitrogen for a wide size range of both aggregate and solitary salps. Measured defecation rates were constant over several hours when salps were held at near-surface conditions from which they had been collected. The defecation rate per salp increased with both salp size and the ambient level of particulate organic matter (POM) in the upper water column. The weight-specific defecation rate ranged between 0.5 and 6% day⁻¹ of salp body carbon, depending on the concentration of available particulate matter in the water. Carbon defecation rates were applied to biomass estimates of S. thompsoni to calculate daily carbon defecation rates for the populations sampled during the two cruises. Dense salp populations of over 400 mg C m⁻² were calculated to produce about 20 mg C m⁻² day⁻¹, comparable to other major sources of vertical flux of organic material in the Southern Ocean. Measured sinking rates for salp fecal pellets indicated that the majority of this organic material could reach deep sediments within a few days, providing a fast and direct pathway for carbon to the deep ocean.     

Metabolism of epipelagic tropical ctenophores

Measurements of respiration and excretion at 25°C were made for five species of ctenophores collected during five cruises to the Bahamas (1982–1984). The mean element-specific respiration and ammonium excretion rates of freshly collected specimens of all species ranged from 4 to 16% d^-1, the mean atomic O:N ratios were 10 to 16, and ammonium averaged 60 to 90% of the total dissolved nitrogen excreted. For adult ctenophores, the carbon content ranged from 0.6% carbon (as percent of dry weight) for Bolinopsis vitrea to 3.7% carbon for Beroë ovata. There was a marked increase in the organic content (% carbon of dry weight) of small Bolinopsis vitrea with tentacles compared to fully lobate adults. B. vitrea had increasingly higher metabolic rates when held at food concentrations up to 100 copepods 1^-1 (about 250 g C 1^-1). The overall range between starved and well-fed B. vitrea was about two times for respiration and a factor of three for ammonium excretion. B. vitrea decreased from well-fed to a starved metabolic rate in about a day after removal from food. The metabolic rate of Eurhamphaea vexilligera was not measurably affected by short-term starvation or feeding (maximum 25 copepods 1^-1). In feeding experiments, E. vexilligera of 20 to 56 mm length fed at rates equivalent to clearance rates of 250 to 1 800 ml h^-1.     

Nitrogen cycling in microbial mats: rates and patterns of denitrification and nitrogen fixation

Spatial and temporal variations in nitrogen fixation and denitrification rates were examined between July 1991 and September 1992 in the intertidal regions of Tomales Bay (California, USA). Microbial mat communities inhabited exposed mudflat and vegetated marsh surface sediments. Mudflat and marsh sediments exhibited comparable rates of nitrogen fixation. Denitrification rates were higher in marsh sediments. Nitrogen fixation rates were lowest during January at both sites, whereas highest rates occurred during summer and fall. Denitrification rates were highest during fall and winter months in marsh sediments, while rates in mudflat sediments were highest during summer and fall. In mudflat sediments, nitrogen fixation and denitrification rates, integrated over 24 h, ranged from 6 to 79 mg N m^-1 d^-1 and 1 to 10 mg N m^-2 d^-1, respectively. Rates of denitrification represented between 6 and 20% of nitrogen fixation rates during the day, but exceeded or were equivalent to nitrogen fixation rates at night. The highest integrated rates of both nitrogen fixation and denitrification occurred during July, whereas, the highest percent loss occurred during spring when denitrification rates amounted to 20% of nitrogen fixation rates during the day. Over an annual cycle, inputs of fixed N to mudflat communities occurred exclusively during daylight. These results underscore the importance of determining integrated diel rates of both nitrogen fixation and denitrification when constructing N budgets. Using this approach, it was shown that microbial denitrification can represent a significant loss of combined nitrogen from mats on daily as well as monthly time scales.     

Elemental and biochemical composition of salps (Tunicata: Thaliacea)

Carbon and nitrogen content have been measured in the solitary and aggregate generations of 11 species of salps. Regression equations for each species and generation permit estimation of carbon or nitrogen content as a function of length of live individuals. Different species of the same length may have nearly tenfold differences in carbon content. Fractionation and biochemical analysis of some samples revealed that the organic content of salps is approximately 80% protein. Ash-free dry weights average 27% of dry weights; mean carbon content is 29% of ash-free dry weight. Excess ash-free dry weight not accountable as organic material is thought to be water of hydration.     

Metabolic rates of epipelagic marine copepods as a function of body mass and temperature

Metabolic rates (oxygen consumption, ammonia excretion, phosphate excretion) have been calculated as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using multiple regression. The metabolic data used for this analysis were species structured, collected from Arctic to Antarctic seas (temperature range: -1.7°C to 29.0°C). The data were further divided into geographical and/or seasonal groups (35 species and 43 data sets for oxygen consumption; 38 species and 58 data sets for ammonia excretion; 22 species and 31 data sets for phosphate excretion). The results revealed that the variance attributed to body mass and temperature was highest (93-96%) for oxygen consumption rates, followed by ammonia excretion rates (74-80%) and phosphate excretion rates (46-56%). Among the various body mass units, the best correlation was provided by the nitrogen unit, followed by the dry weight unit. The calculated Q₁₀ values varied slightly according to the choice of body mass units; overall ranges were 1.8-2.1 for oxygen consumption rates, 1.8-2.0 for ammonia excretion rates and 1.6-1.9 for phosphate excretion rates. The effects of body mass and temperature on the metabolic quotients (O:N, N:P, O:P) were insignificant in most cases. Although the copepod metabolic data used in the present analysis were for adult and pre-adult stages, possible applications of the resultant regression equations to predict the metabolic rates of naupliar and early copepodite stages are discussed. Finally, global patterns of net growth efficiency [growth (growth+metabolism)^-1] of copepods were deduced by combining the present metabolic equation with Hirst and Lampitt's global growth equation for epipelagic marine copepods.     

Effects of nutritional history on nitrogen assimilation in congeneric temperate and tropical scleractinian corals

The nutritional history of corals is known to affect metabolic processes such as inorganic nutrient uptake and photosynthesis, but little is known about how it affects assimilation efficiency of ingested prey items or the partitioning of prey nitrogen between the host and symbiont. The temperate scleractinian coral Oculina arbuscula and its tropical congener Oculina diffusa were acclimated to three nutritional regimes (fed twice weekly, starved, starved with an inorganic nutrient supplement), then fed Artemia nauplii labeled with the stable isotope tracer ¹⁵N. Fed corals of both species had the lowest assimilation efficiencies (36–51% for O. arbuscula, 38–57% for O. diffusa), but were not statistically different from the other nutritional regimes. Fed and starved corals also had similar NH₄⁺ excretion rates. This is inconsistent with decreased nitrogen excretion and reduced amino acid catabolism predicted by both the nitrogen recycling and conservation paradigms. In coral host tissue, ~90% of the ingested ¹⁵N was in the TCA-insoluble (protein and nucleic acids) and ethanol-soluble (amino acids/low molecular weight compounds) within 4 h of feeding. The TCA-insoluble pool was also the dominant repository of the label in zooxanthellae of both species (40–53% in O. arbuscula, 50–60% in O. diffusa). However, nutritional history had no effect on the distribution of prey ¹⁵N within the biochemical pools of the host or the zooxanthellae for either species. This result is consistent with the nitrogen conservation hypothesis, as preferential carbon metabolism would minimize the effects of starvation on nitrogen-containing biochemical pools.Communicated by P.W. Sammarco, Chauvin     

Nitrogen-fixation by cyanobacteria associated withCodium fragile (Chlorophyta): Environmental effects and transfer of fixed nitrogen

N₂-fixation associated with the green macroalgaCodium fragile subsp.tomentosoides (van Goor) Silva from Long Island, New York, USA, was attributable to several species of endophytic cyanobacteria. Rates of N₂-fixation ranged from 0.03 to 3.2µg N g^–1 dry wt h^–1 in freshly collected plants from several sites. Growth of the cyanobacteria appeared to be light-limited, due to the transmission of only 5 to 10% of incident light through the pigmented surface-layer of the macroalga. Daily irradiance was the most important factor determining both abundance of cyanobacterial cells and rate of N₂-fixation. The rate was also affected by instantaneous irradiance, and increased twofold from dark to ambient surface irradiance. Rates were reduced at low temperature (8°C) but showed no temperature effect between 12° and 26°C. External concentrations of dissolved inorganic nitrogen (DIN) up to 20µM did not influence N₂-fixation rate, but long-term exposure to 60µmol l^–1 d^–1 of NH ₄ ⁺ caused a reduction in the rate. InC. fragile grown under high daily irradiance and low external DIN concentration, ~50% of the assimilated-N was attributable to N₂-fixation. However, chlorophyllb extracted from plants grown with¹⁵N₂ showed an atom % excess¹⁵N of less than 0.1, suggesting that only a small proportion of the bacterially fixed-N was transferred to the seaweed. The association betweenC. fragile and its endophytic cyanobacteria appears to be based primarily on microhabitat suitability, rather than mutual metabolic dependence. It is doubtful that N₂-fixation by cyanobacteria is important to the ecological success of this seaweed species.     

Foraging ecology of leatherback sea turtles in the Western North Atlantic determined through multi-tissue stable isotope analyses

Leatherback turtles, Dermochelys coriacea, are highly migratory, spending most of their lives submerged or offshore where their feeding habits are difficult to observe. In order to elucidate the foraging ecology of leatherbacks off Massachusetts, USA, stable isotope analyses were performed on leatherback tissues and prey collected from 2005 to 2009. Stable isotope ratios of nitrogen and carbon were determined in whole blood, red blood cells, blood plasma, muscle, liver, and skin from adult male, female, and subadult leatherbacks. Isotopic values were analyzed by body size (curved carapace length) and grouped by sex, and groups were tested for dietary differences. Gelatinous zooplankton samples were collected from leatherback foraging grounds using surface dip nets and stratified net tows, and prey contribution to leatherback diet was estimated using a two-isotope Bayesian mixing model. Skin and whole blood δ¹³C values and red blood cell δ¹⁵N values were correlated with body size, while δ¹³C values of red blood cells, whole blood, and blood plasma differed by sex. Mixing model results suggest that leatherbacks foraging off Massachusetts primarily consume the scyphozoan jellyfishes, Cyanea capillata and Chrysaora quinquecirrha, and ctenophores, while a smaller proportion of their diet comes from holoplanktonic salps and sea butterflies (Cymbuliidae). Our results are consistent with historical observations of leatherback turtles feeding on scyphozoan prey in this region and offer new insight into size- and sex-related differences in leatherback diet.     

Metabolism and body composition of a copepod (<Emphasis Type="Italic">Neocalanus cristatus</Emphasis>: Crustacea) from the bathypelagic zone of the Oyashio region,western subarctic Pacific

Metabolism [respiratory oxygen consumption, electron-transfer-system (ETS) activity] and body composition [water, ash, carbon (C), nitrogen (N), carbon/nitrogen (C/N) ratio] of stage C5/C6 Neocalanus cristatus from 1000 to 2000 m depth of the Oyashio region, western subarctic Pacific, were determined during the period of July 2000 through June 2003. Compared with the C5 specimens from shallow depths (<250 m), those from 1000 to 2000 m were characterized by quiescent behavior, reduced respiration rates (30% of the rates at active feeding), very low water content (61–70% of wet weight), but high C content (56–64% of dry weight) and C/N ratios (7.2–10.6, by weight). Artifacts due to the recovery of live specimens from the bathypelagic zone appeared to be unlikely in this study, as judged by the consistent results between re-compression (100 atm) and non-compression (1 atm) respiration experiments, and between ETS activities and respiration rates directly measured. In addition, the respiration rates of C6 males and females of N. cristatus from the same 1000–2000 m depth were two to three times higher than the rates of C5 individuals, but were similar to the rates of a bathypelagic copepod, Paraeuchaeta rubra. Combining these results with literature data, C budgets of: (1) diapausing C5 specimens, weighing 6–10 mg dry weight; (2) molt to C6 females; and (3) the complete the life span were established, taking into account assorted losses in respiration during diapause at stages C5 and C6, molt production and egg production. Respiratory C losses by C5 and C6 specimens were estimated on the basis of body N as adjusted metabolic rates [AMR; µl O₂ (mg body N)^–0.843 h^–1], then N budgets were also computed subtracting N lost in the form of cast molts and eggs from the initial stock. Calculations revealed that allocation of the C stock was greatest to egg production (34–57%), followed by respiration (27%) and cast molts (3%), leaving residual C of 13–36% in spent C6 females. The present results for N. cristatus from the North Pacific are compared with those of Calanus spp. in the North Atlantic.Communicated by O. Kinne, Oldendorf/Luhe