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

The metabolic rates (oxygen uptake, ammonia excretion, phosphate excretion) of epipelagic marine zooplankton have been expressed as a function of body mass (dry, carbon, nitrogen and phosphorus weights) and habitat temperature, using the multiple-regression method. Zooplankton data used for this analysis are from phylogenetically mixed groups (56 to 143 species, representing 7 to 8 phyla, body mass range: 6 orders of magnitude) from various latitudes (habitat temperature range:-1.4° to 30°C). The results revealed that 84 to 96% of variation in metabolic rates is due to body mass and habitat temperature. Among the various body-mass units, the best correlation was provided by carbon and nitrogen units for all three metabolic rates. Oxygen uptake, ammonia excretion and phosphate excretion are all similar in terms of body-mass effect, but differ in terms of temperature effect. With carbon or nitrogen body-mass units, calculated Q₁₀ values are 1.82 to 1.89 for oxygen uptake, 1.91 to 1.93 for ammonia excretion and 1.55 for phosphate excretion. The effects of body mass and habitat temperature on the metabolic quotients (O:N, N:P, O:P) are insignificant. The present results for oxygen-uptake rate vs body mass do not differ significantly from those reported for general poikilotherms by Hemmingsen and for crustaceans by Ivleva at a comparable temperature (20°C). The importance of a body-mass measure for meaningful comparison is suggested by the evaluation of the habitat-temperature effect between mixed taxonomic groups and selected ones. Considering the dominant effects of body mass and temperature on zooplankton metabolic rates, the latitudinal gradient of community metabolic rate for net zooplankton in the ocean is estimated, emphasizing the non-parallelism between community metabolic rates and the standing stock of net zooplankton.     

Influence of temperature changes on oxygen uptake and ammonia and phosphate excretion,in relation to body size and weight,in Oikopleura dioica (Appendicularia)

The relationship between the rates of oxygen consumption, ammonia and phosphate excretion of a pelagic tunicate, the larvacean Oikopleura dioica Fol, 1872 were assessed as a function of size, dry weight and ash-free dry weight at 15°, 20° and 24°C. O. dioica has higher respiration and excretion rates than copepods of similar weight, but the weight exponent of the allometric power function: Y=aX ^b is similar to that of other poikilotherms. Temperatures above 20°C have a depressing effect on respiration and ammonia excretion. 90% of the variance in metabolic rates is explainable by body mass and temperatures Q₁₀ values for oxygen consumption, ammonia and phosphate excretion, respectively, are 2.45, 1.86 and 1.75 between 15° and 20°C, and 3.75, 2.90 and 3.60 between 20° and 24°C. Metabolic quotients (O:N, O:P, N:P) indicate a protein-oriented diet. The results of this study suggest weak metabolic regulation in O. dioica, an energetic strategy which allows an immediate response to favourable changes in feeding conditions.     

Temperature effects on the metabolism of larvae of the Antarctic starfish Odontaster validus, using a novel micro-respirometry method

Oxygen consumption of individual larvae of the Antarctic sea-star Odontaster validus was measured during the 50-day period following fertilisation. Values ranged from 0.76 pmol O₂ h^-1 for one specimen at the coeloblastula stage to 77.6 pmol O₂ h^-1 for one bipinnaria larva. At 0°C the mean oxygen consumption rate of an individual larva increased from 10.9 pmol O₂ h^-1 (standard error of the mean, SEM, 0.13) for a gastrula larva, 13 days post-fertilisation, to 25.4 pmol O₂ h^-1 (SEM 3.5) at the bipinnaria stage (50 days post-fertilisation). Gastrulae reared at -0.5°C did not have significantly different oxygen consumption rates between days 13 and 45 post-fertilisation (mean=11.4 pmol O₂ h^-1). Individual metabolic rates were highly variable, covering more than a 40-fold range. At 2°C gastrula oxygen consumption was on average 45% higher (17.35 pmol O₂ h^-1), giving a Q₁₀ temperature effect of 4.4. For bipinnaria, mean oxygen consumption in 2°C larvae (31.4 pmol O₂ h^-1) was not significantly different from that in larvae at -0.5°C, suggesting bipinnaria metabolism may be less sensitive to temperature change than earlier stages. At 2°C the bipinnaria stage was reached at 30-35 days compared with 45-50 days at 0°C, giving a Q₁₀ of 4.5 for temperature effects on development. The method here used a new, highly sensitive micro-respirometry method that is inexpensive and straightforward in design. Individual larvae of O. validus were held in 35- to 50-µl respirometers. These larvae have very low metabolic rates, and published work on such organisms have utilised at least 25 individuals per chamber. The oxygen content of the respirometers was measured using a 25-µl sample injected into a couloximeter. Oxygen consumption rates down to -1 pmol h^-1 can be detected. Under optimum conditions oxygen consumption of a single larva of -4 pmol O₂ h^-1 was measured with an accuracy of ᆨ%. Values of ~15 pmol h^-1 could routinely be measured with this accuracy. This method would allow oxygen consumption to be evaluated in individual field-caught larvae of most marine ectotherms.     

An experimental and theoretical analysis of the effect of added weight on the energetics and hydrostatic function of the swimbladder of European sea bass (Dicentrarchus labrax)

We conducted experiments to determine the effect of the increasing ultrasonic/radio transmitter weight on the routine metabolic rate of sea bass. We measured the oxygen consumption (MO₂) of fish tagged externally with a dummy transmitter made of a hollow pipe, the weight of which was adjusted with lead to represent in water 0, 1 and 4% (R_tf) of the animal weight. We then developed a theoretical model to estimate, for a given fish size, the range of added weight that fish can compensate for through swimbladder regulation. When R_tfБ%, MO₂ of untagged and tagged fish did not differ significantly. However, when R_tf reached 4%, fish that carried a tag incurred a significant elevation of oxygen consumption, which represented 28% of their total useable power (or metabolic scope). This result strongly supports the view that a high R_tf ratio contributes to a decrease in available metabolic energy by diverting energy from, e.g., growth or swimming performance. A comparison between the tagged fish and the theoretical model reinforced the hypothesis that, when R_tf attained 4%, the increase in metabolic rate reflected a supplementary and costly swimming effort necessary to maintain vertical position. In this condition, the swimbladder cannot regulate the buoyancy of tagged fish.     

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).     

Respiration and ammonia excretion by marine metazooplankton taxa: synthesis toward a global-bathymetric model

For thirteen representative taxa of metazooplankton from various depth horizons (<4,200 m) of the world’s oceans, respiration rate (681 datasets on 390 species) and ammonia excretion rate (266 datasets on 190 species) are compiled and analyzed as a function of body mass (dry mass, carbon or nitrogen), habitat temperature, habitat depth and taxon. Stepwise-regression analyses reveal that body mass is the most important parameter, followed by habitat temperature and habitat depth, whereas taxon is of lesser importance for both rates. The resultant multiple regression equations show that both respiration rate and ammonia excretion rate (per individual) increase with increase in body mass and habitat temperature, but decrease with habitat depth. Some taxa are characterized by significantly higher or lower rates of respiration or ammonia excretion than the others. Overall, the global-bathymetric models explain 93.4–94.2 % of the variance of respiration data and 80.8–89.7 % of the variance of ammonia excretion data. The atomic O:N ratios (respiration/ammonia excretion) are largely independent of body mass, habitat temperature, habitat depth and taxon, with a median of 17.8. The present results are discussed in light of the methodological constraints and the standing hypotheses for the relationship between metabolic rate and temperature. Perspectives for model improvement and possible application of it to plankton-imaging systems for rapid assessment of the role of metazooplankton in C or N cycles in the pelagic ecosystem are briefly discussed.     

Metabolic rates of juvenile scalloped hammerhead sharks (Sphyrna lewini)

Oxygen consumption of juvenile scalloped hammerhead sharks, Sphyrna lewini, was measured in a Brett-type flume (volume=635 l) to quantify metabolic rates over a range of aerobic swimming speeds and water temperatures. Oxygen consumption (log transformed) increased at a linear rate with increases in tailbeat frequency and swimming speed. Estimates of standard metabolic rate ranged between 161 mg O₂ kg^-1 h^-1 at 21°C and 203 mg O₂ kg^-1 h^-1 at 29°C (mean-SD: 189ᆣ mg O₂ kg^-1 h^-1 at 26°C). Total metabolic rates ranged from 275 mg O₂ kg^-1 h^-1 at swimming speeds of 0.5 body lengths per second (L s^-1) to a maximum aerobic metabolic rate of 501 mg O₂ kg^-1 h^-1 at 1.4 L s^-1. Net cost of transport was highest at slower swimming speeds (0.5-0.6 L s^-1) and was lowest between 0.75 and 0.9 L s^-1. Therefore, these sharks are most energy efficient at swimming speeds between 0.75 and 0.9 L s^-1. These data indicate that tailbeat frequency and swimming speed can be used as predictors of metabolic rate of free-swimming juvenile hammerhead sharks.     

How size relates to oxygen consumption,ammonia excretion,and ingestion rates in cold (<Emphasis Type="Italic">Enteroctopus megalocyathus</Emphasis>) and tropical (<Emphasis Type="Italic">Octopus maya</Emphasis>) octopus species

The scaling of metabolic rates with body mass is one of the best known and most studied characteristics of aquatic animals. Herein, we studied how size is related to oxygen consumption, ammonia excretion, and ingestion rates in tropical (Octopus maya) and cold-water (Enteroctopus megalocyathus) cephalopod species in an attempt to understand how size affects their metabolism. We also looked at how cephalopod metabolisms are modulated by temperature by constructing the relationship between metabolism and temperature for some benthic octopod species. Finally, we estimated the energy balance for O. maya and E. megalocyathus in order to validate the use of this information for aquaculture or fisheries management. In both species, oxygen consumption and ammonia excretion increased allometrically with increasing body weight (BW) expressed as Y = aBW ^b . For oxygen consumption, b was 0.71 and 0.69 for E. megalocyathus and O. maya, respectively, and for ammonia excretion it was 0.37 and 0.43. Both species had low O/N ratios, indicating an apparent dependence on protein energy. The mean ingestion rates for E. megalocyathus (3.1 ± 0.2% its BW day⁻¹) and O. maya (2.9 ± 0.5% its BW day⁻¹) indicate that voracity, which is characteristic of cephalopods, could be independent of species. The scope for growth (P = I − (H + U + R) estimated for E. megalocyathus was 28% higher than that observed in O. maya (320 vs. 249 kJ day⁻¹ kg⁻¹). Thus, cold-water cephalopod species could be more efficient than tropical species. The protein and respiratory metabolisms of O. maya, E. megalocyathus, and other octopod species are directly dependent on temperature. Our results offer complementary evidence that, as Clarke (2004) stated, the metabolic response (R and U) cannot be determined mechanistically by temperature, as previously proposed (Gillooly et al. 2002).     

Experimental evaluation of the energy balance in Octopus vulgaris, fed ad libitum on a high-lipid diet

A complete energy balance equation was estimated for the common octopus Octopus vulgaris at a constant temperature of 20°C, fed ad libitum on anchovy fillet (Engraulis encrasicolus). Energy used for growth and respiration or lost with faeces and excreted ammonia was estimated, along with total energy consumption through food, for six specimens of O. vulgaris (with masses between 114 and 662 g). The energy balance equation was estimated for the specimens at 10-day intervals. During each 10-day interval, food consumed, body mass increase and quantity of faeces voided were measured. The calorific values of octopus flesh, anchovy flesh and faeces were measured by bomb calorimetry. Oxygen consumption and ammonia excretion rates were monitored for each specimen during three 24-h experiments and daily oxygen consumption and ammonia excretion were estimated. It was found that 58% of the energy consumed was used for respiration. The amount of energy invested in somatic and gonadal growth represented 26% of the total energy budget. The energy discarded through faeces was 13% of consumed energy. The estimated assimilation efficiency (AE) values of O. vulgaris feeding on anchovy (80.9–90.7%) were lower than the AE values estimated for other cephalopod species with different diets of lower lipid content such as crabs or mussels. Specific growth rates (SGR) ranged 0.43–0.95 and were similar to those reported for other high-lipid diets (bogue, sardine) and lower than SGR values found for low-lipid, high-protein diets (squid, crab, natural diet). Ammonia excretion peak (6 h after feeding) followed the one of oxygen consumption (1 h after feeding). The values of atomic oxygen-to-nitrogen (O:N) ratio indicated a protein-dominated metabolism for O. vulgaris.     

Oxygen-dependent asynchrony of embryonic development in embryo masses of brachyuran crabs

Among brooding species, passive and active means to provide oxygen to embryos can be observed. Among passive oxygen providers, lower oxygen availability in the center than at the periphery of embryo masses seems to delay development of inner embryos. We investigated the differences in patterns of oxygen supply to the periphery and the center of embryo masses in two active oxygen providers, the brachyuran crabs Cancer setosus and Homalaspis plana, and evaluated the consequences on: (1) the proportion of time that early- and late-stage embryos were exposed to low or high oxygen partial pressure (PO₂), (2) oxygen consumption of the embryos from the center (inner) and the periphery (outer) of the embryo mass at those PO₂ levels that the embryos experience throughout development, and (3) development of inner and outer embryos. We found that oxygen availability in the embryo masses of brachyuran crabs exhibited dramatic contrasts between the periphery and the center during early development and that these differences decreased throughout embryonic development. These dissimilar patterns of oxygen availability produced differences in the proportion of the time that the embryos were exposed to high and low PO₂ levels throughout development. PO₂ affected oxygen consumption of the inner and outer embryos in the same fashion, but the oxygen demand of inner embryos was lower. Furthermore, development of inner embryos was delayed, in comparison to outer embryos of the same female. We suggest that the asynchrony in the development of inner embryos, in comparison to outer embryos, is due to oxygen limitation, since oxygen availability affects embryonic oxygen consumption. The differences between development of inner and outer embryos is relatively small, when compared to other marine invertebrates, probably because female crabs are able to adjust oxygen supply to the embryos according their needs, while passive oxygen providers are not. However, active oxygen provision may affect investment in reproduction. Our results could have important implications both on studies of larval development and survival and in understanding the life-history tradeoffs of aquatic invertebrates.     

Respiration and ammonia excretion of euphausiid crustaceans: synthesis toward a global-bathymetric model

Respiration and ammonia excretion rates of 19–24 euphausiids from the epipelagic through bathypelagic zones of the world’s oceans were compiled. Body mass (expressed in terms of dry mass, carbon or nitrogen), habitat temperature and sampling depth were designated as parameters in multiple regression analysis. Results suggested that the three parameters were highly significant, contributing 71–89 % of the variance in respiration rates and 69–81 % of the variance in ammonia excretion rates. Atomic O:N ratios derived from simultaneous measurements of respiration and ammonia excretion rates ranged from 11 to 90 (median: 27), and no appreciable effects of the three parameters on O:N ratios were detected. If global-bathymetric models for the metabolism and chemical composition of copepods and chaetognaths are compared with those of euphausiids, it becomes evident that euphausiids are unique in that they maintain high metabolic rates and accumulate moderate amounts of energy reserves (lipids).     

The effects of feeding or addition of dissolved inorganic nutrients in maintaining the symbiosis between dinoflagellates and a tropical marine cnidarian

The availability of solid food (Artemia nauplii) and dissolved inorganic nutrients (ammonium, nitrate, phosphate) to the shallow-water marine hydroid Myrionema amboinense was manipulated for 1-8 days in order to investigate their role in the growth of intracellular symbiotic dinoflagellates (zooxanthellae) of the genus Symbiodinium. Symbionts from hydroids collected from the field or maintained under laboratory conditions (25°C, 12 h:12 h light:dark cycle, 80 µE m^-2 s^-1 fluorescent lighting) always exhibited a single peak in mitotic index (MI) at dawn. Symbionts in freshly collected field animals had an MI peak of about 15%. Symbiotic dinoflagellates in hydroids fed Artemia nauplii twice daily in the laboratory maintained this dawn peak of MI between 10% and 15%, but in the absence of feeding or added inorganic nutrients, this peak declined to less than 1% within 2-4 days. In contrast, when hydroids were placed in solutions containing ammonium (20 µM NH₄Cl), nitrate (10 µM NaNO₃), and a combination of ammonium and phosphate (2 µM Na₂HPO₄) immediately after collection, the algal MI remained between 5% and 15% for 4-7 days; the addition of 2 µM phosphate did not increase MI relative to unfed rates. When unfed animals were placed in dissolved nitrogen or fed Artemia, the symbiont MI increased from <1% to 10-17% within 2-3 days; P alone had no effect. However, the increase resulting from added inorganic nutrients was temporary, lasting only 5-7 days. These observations suggest that algal division in the host is maintained indefinitely in the field or by feeding particulate foods twice daily in the laboratory, but the addition of inorganic nutrients alone (ammonium, nitrate and ammonium/phosphate) appeared to support the completion of a maximum of one additional round of cell division. Nutrients required for continued growth and division of symbiotic dinoflagellates are linked to host feeding and host growth; without external food, neither host nor symbiont continue to grow. The same phenomenon is seen in zooxanthellate anemones, clams and corals, where total numbers of symbionts appear to be linked to changes in host-tissue biomass (protein), achieving relatively stable densities in M. amboinense, corals and other cnidarian symbioses, depending on their local environmental conditions. The results of the present study help explain the cellular responses of algal symbionts in reef-dwelling invertebrates to additions of dissolved inorganic nutrients to coral-reef ecosystems.     

Effects of emersion and elevated haemolymph ammonia on haemocyanin–oxygen affinity of Cancer pagurus

The subtidal crab Cancer pagurus (L.) experiences involuntary periods of emersion associated with practices used in their marketing and distribution. During 24 h emersion, impaired gill function caused an increase of circulating total ammonia (TA=NH₃+NH₄⁺) of 0.35 mmol TA l^-1 (167%). The oxygen-binding characteristics of the haemocyanin of C. pagurus were examined at 10°C in the presence of total ammonia (0.2-1.0 mmol TA l^-1). The haemocyanin-oxygen affinity was decreased in the presence of TA ((logP₅₀/(log[TA]=0.16). Emersion induced significant acidosis and elevated circulating levels of haemolymph TA, lactate and urate, but all had returned to normal levels within 24 h of re-immersion. The accumulation of haemocyanin-modulating substances during 24 h emersion compensated partially (40%) for the effect of the acidosis, but the net effect of the emersion period was a significant decrease in oxygen affinity, corresponding to an increase of P₅₀ (10°C ) from 1.24 kPa (immersed) to 1.96 kPa (24 h emersion). The implications of the findings are considered in terms of the effects and adaptations to emersion.     

Patterns of nitrogenous waste excretion and gill urea transporter mRNA expression in several species of marine fish

Many prior studies of nitrogenous waste excretion in marine fish have examined excretion patterns for short time periods, and with relatively coarse sampling schemes (e.g., an initial and a final sample point). Recent studies of a ureotelic marine fish (the gulf toadfish, Opsanus beta) have demonstrated that urea excretion in this species occurs in brief but massive bursts, lasting from 0.5 to 3 h, and often only once per day. The present study sought to determine if prior sampling protocols may have underestimated the amount of urea being excreted by marine fish. A survey of 16 marine species (the teleosts: Myoxocephalus octodecemspinosus, Scophthalamus aquosa, Cyclopterus lumpus, Lophius americanus, Aprodon cortezianus, Cymatogaster aggregatus, Parophrys vetulis, Microstomus pacificus, Hippoglossoides elassodon, Bathyagonus nigripinnus, Ophiodon elongatus, Hemilepidatus spinosus, Icelinus terrius; the elasmobranch: Raja rhina; and the hagfish: Eptatretus stoutii) was undertaken for ammonia-N and urea-N excretion using a long sampling period (48 h) and hourly sample collection. Apart from the obvious exception of an elasmobranch, ammonia excretion was confirmed to be predominant in marine fish, with urea excretion constituting between 1.4 and 23.8% of the total of ammonia plus urea excreted. Notably, no pulses of urea excretion were detected. Despite the relatively low level of urea excretion, expression of urea transporter-like mRNA (detected using the toadfish urea transporter, tUT, cDNA as a probe) was discovered in gills of many of the species surveyed for nitrogen excretion patterns, although no signal was detected in the hagfish. These results suggest that urea excretion takes place through a specific transport pathway. Finally, more detailed analysis of nitrogen excretion in one of the surveyed species, the plainfin midshipman (Porichthys notatus) demonstrates that "total" nitrogen excretion estimated by summing ammonia and urea excretion underestimates true total nitrogen excretion by 37-51%.     

Characterization of urease activity in three marine phytoplankton species,<Emphasis Type="Italic"> Aureococcus anophagefferens</Emphasis>,<Emphasis Type="Italic"> Prorocentrum minimum</Emphasis>, and<Emphasis Type="Italic"> Thalassiosira weissflogii</Emphasis>

The availability of different forms of nitrogen in coastal and estuarine waters may be important in determining the abundance and productivity of different phytoplankton species. Although urea has been shown to contribute as much as 50% of the nitrogen for phytoplankton nutrition, relatively little is known of the activity and expression of urease in phytoplankton. Using an in vitro enzyme assay, urease activities were examined in laboratory cultures of three species: Aureococcus anophagefferens Hargraves et Sieburth, Prorocentrum minimum (Pavillard) Schiller, and Thalassiosira weissflogii (Grunow) Fryxell et Hasle. Cultures of P. minimum and T. weissflogii were grown on three nitrogen sources (NO₃^m, NH₄⁺, and urea), while A. anophagefferens was grown only on NO₃^m and urea. Urease was found to be constitutive in all cultures, but activity varied with growth rate and assay temperature for the different cultures. For A. anophagefferens, urease activity varied positively with growth rate regardless of the N source, while for P. minimum, urease activity varied positively with growth rate only for cultures grown on urea and NH₄⁺. In contrast, for T. weissflogii, activity did not vary with growth rate for any of the N sources. For all species, urease activity increased with assay temperature, but with different apparent temperature optima. For A. anophagefferens, in vitro activity increased from near 0-30°C, and remained stable to 50°C, while for P. minimum, increased in vitro activity was noted from near 0-20°C, but constant activity was observed between 20°C and 50°C. For T. weissfloggii, while activity also increased from 0°C to 20°C, subsequent decreases were noted when temperature was elevated above 20°C. Urease activity had a half-saturation constant of 120-165 wg atom N l^у in all three species. On both an hourly and daily basis, urease activity in A. anophagefferens exceeded nitrogen demand for growth. In P. minimum, urease activity on an hourly basis matched the nitrogen demand, but was less than the demand on a daily basis. For T. weissflogii, urease activity was always less than the nitrogen demand. These patterns in urease activity in three different species demonstrate that while apparently constitutive, the regulation of activity was substantially different in the diatom. These differences in the physiological regulation of urease activity, as well as other enzymes, may play a role in their ecological success in different environments.     

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.     

Metabolic properties of Northern krill, Meganyctiphanes norvegica, from different climatic zones. I. Respiration and excretion

Adaptive processes linked to overall metabolism were studied in terms of oxygen consumption and ammonia excretion in each of three self-contained krill populations along a climatic gradient. In the Danish Kattegat, krill were exposed to temperatures which ranged from 4°C to 16°C between seasons and a vertical temperature gradient of up to 10°C during summer. In the Scottish Clyde Sea, water temperatures varied less between seasons and the vertical temperature gradient in summer was only 3°C. Temperatures in the Ligurian Sea, off Nice, were relatively constant around 12-13°C throughout the year, with a thin surface layer (20-30 m) of warm water developing during summer. The trophic conditions were rich in the Kattegat and, particularly, in the Clyde, but comparatively poor in the Ligurian Sea. Oxygen consumption increased exponentially with increasing experimental temperature, which ranged from 4°C to 16°C. Overall respiration rates were between 19.9 and 89.9 µmol O₂ g^-1 dry wt h^-1. Krill from the Kattegat, the Clyde Sea, and the Ligurian Sea all exhibited approximately the same level of oxygen consumption (30-35 µmol O₂ g^-1 dry wt h^-1) when incubated at the ambient temperatures found in their respective environments (9°C, 5°C, and 12°C). This indicates that krill adjust their overall metabolic rates to the prevailing thermal conditions. The exception to this were the respiration rates of Ligurian krill from winter/spring, which were about twice as high as the rates from summer krill despite the fact that the thermal conditions were the same. This effect appears to result from enhanced somatic activity during a short period of increased food availability and reproduction. Accordingly, krill appears to be capable of adapting to both changing thermal and trophic conditions, especially when nutrition is a limiting factor in physiological processes.     

Leptocephalus energetics: assembly of the energetics equation

The principles of energetics were used to examine the energetic requirements of leptocephali. Respiration and excretion rates and daily growth rates combined with proximate composition were used to examine the allocation of energy into each of the three main components of energetics: metabolism, excretion and growth. The daily energetic requirements for leptocephali, referred to as type 2 larvae based upon their unique developmental strategy, were compared to the requirements of non-leptocephalus larvae, known as type 1. Leptocephalus daily energetic requirements were also compared to the energy available from the leptocephalus' proposed food sources. The four species of eel larvae selected were all from the order Anguilliformes: Paraconger caudilimbatus (Poey), Ariosoma balearicum (Delaroche), Gymnothorax saxicola Jordan and Davis, and Ophichthus gomesii (Castelnau). The allocation of energy to each of the components of energetics as well as the total energetic requirements for the leptocephali proved to be very different from those of type 1 larvae. Metabolism received the majority, 60-92%, of the energy required per day. Growth and excretion were allocated 4-39% and <1-21%, respectively, of the total energy needed per day. Leptocephali required <50% of the energy needed by type 1 larvae of equal dry mass. The unique growth strategy used by leptocephali allows them to increase rapidly in size while allocating the majority of their energy, not to growth as in most larval fish, but to metabolism.     

Effects of prolonged starvation on O2 consumption,NH 4 + excretion,and chemical composition of the bathypelagic mysid Gnathophausia ingens

The large bathypelagic mysid Gnathophausia ingens was collected in January 1980 at 400 to 700 m depth from the San Clemente Basin off southern California. Instars 7-8 and Instars 10-12 were starved in the laboratory for up to 19 wk. Oxygen consumption and ammonia excretion rates, and water, protein, lipid, and ash contents were determined periodically during starvation. Protein and lipid were metabolized in approximately equal amounts by starved individuals after the initial weeks of food deprivation. Unidentified components (probably non-protein nitrogenous compounds) apparently were oxidized within the first 7 wk of starvation. Oxygen consumption and ammonia excretion by Instars 7-8 decreased steadily during 19 wk of starvation. In contrast, stable or increasing respiration and excretion rates were observed for fed mysids. The mean respiration rate of Instars 10-12 did not change significantly during 13 wk of starvation, although ammonia excretion rates decreased. Low metabolic rates and large lipid reserves probably help G. ingens to withstand long periods of starvation in the mesopelagic environment. Calculations based on the laboratory data demonstrate that small, infrequent meals could account for the rates of metabolism and growth observed for G. ingens in the field.     

Biomass, photosynthesis and growth of Laminaria saccharina in a high-arctic fjord, NE Greenland

Biomass, photosynthesis and growth of the large, perennial brown alga Laminaria saccharina (L.) Lamour. were examined along a depth gradient in a high-arctic fjord, Young Sound, NE Greenland (74°18'N; 20°14'W), in order to evaluate how well the species is adapted to the extreme climatic conditions. The area is covered by up to 1.6-m-thick ice during 10 months of the year, and bottom water temperature is <0°C all year round. L. saccharina occurred from 2.5 m depth to a lower depth limit of about 20 m receiving 0.7% of surface irradiance. Specimen density and biomass were low, likely, because of heavy ice scouring in shallow water and intensive feeding activity from walruses in deeper areas. The largest specimens were >4 m long and older than 4 years. In contrast to temperate stands of L. saccharina, old leaf blades (2-3 years old) remained attached to the new blades. The old tissues maintained their photosynthetic capacity thereby contributing importantly to algal carbon balance. The photosynthetic characteristics of new tissues reflected a high capacity for adaptation to different light regimes. At low light under ice, or in deep water, the chlorophyll a content and photosynthetic efficiency (!) were high, while light compensation (E_c) and saturation (E_k) points were low. An E_c of 2.0 µmol photons m^-2 s^-1 under ice allowed photosynthesis to almost balance, and sometimes exceed, respiratory costs during the period with thick ice cover but high surface irradiance, from April through July. Rates of respiration were lower than usually found for macroalgae. Annual elongation rates of leaf blades (70-90 cm) were only slightly lower than for temperate L. saccharina, but specific growth rates (0.48-0.58 year^-1) were substantially lower, because the old blades remained attached. L. saccharina comprised between 5% and 10% of total macroalgal biomass in the area, and the annual contribution to primary production was only between 0.1 and 1.6 g C m^-2 year^-1.