Enzyme activities of fish skeletal muscle and brain as influenced by depth of occurrence and habits of feeding and locomotion

Activities of lactate dehydrogenase (LDH), pyruvate kinase (PK), malate dehydrogenase (MDH) and citrate synthase (CS) were measured in the white skeletal muscle of marine fishes having different depths of occurrence and different feeding and locomotory strategies. There were significant depth-related differences in the two glycolytic enzymes, LDH and PK. LDH activity was most variable, and differed by 3 orders of magnitude between the most active shallow-living species and certain deep-sea fishes likely to have only minimal capacities for active locomotion. Superimposed on the depth-related patterns was a high degree of interspecific variation (up to 20-fold) in enzymic activity among species from any given range of depth of occurrence. Variation of both LDH and PK activities, noted for shallow- and deep-living fishes, seems to be largely accounted for by differences in feeding habits and locomotory performance. Active pelagic swimmers have much higher activities of LDH and PK than, for example, deep-living sit-and-wait predators. Benthopelagic fishes like rattails and the sablefish have the highest activities found among deep-living fishes, suggesting that these species engage in relatively active food-searching behavior compared to most other deep-sea fishes. The activity of CS, an enzyme of the citric acid cycle and an indicator of aerobic metabolism, varied little among species. Thus, the large interspecific variation in glycolytic potential (LDH and PK) among species is not associated with a similar variation in aerobic metabolism of white muscle. The much higher and more variable activity of MDH relative to CS suggests that, in addition to its function in the citric acid cycle, MDH may play an important role in redox balance in fish white muscle. In a comparison of white muscle composition between the shallow- and deep-living species, water content did not differ significantly, but protein content was significantly higher in shallow- than in deep-living fishes (211 and 130 mg g^-1 wet wt of muscle, respectively). The differences in muscle protein content are small relative to the differences between shallow- and deep-living species in LDH, PK and MDH activities. Thus, depthrelated differences in muscle enzymic activity are caused by factors other than enzyme dilution. Enzyme activities (LDH, PK and CS) in brain tissue were relatively constant among species regardless of depth of occurrence or feeding and locomotory habits. Habitat and lifestyle do not seem to influence the demands for overall metabolic function in brain. The utility of muscle enzymic activity data for making predictions about the ecological characteristics of difficult-to-observe, deep-living, fishes is discussed.     

Depth-related enzymic activities in muscle,brain and heart of deep-living pelagic marine teleosts

The activities [units^-1 wet weight tissue] of lactate dehydrogenase (LDH), pyruvate kinase (PK), malate dehydrogenase (MDH) and isocitrate dehydrogenase (IDH) in white skeletal muscle, brain and heart of 24 pelagic teleost fishes were determined. In addition, for several of the foregoing species, citrate synthase (CS) activities were examined in white skeletal muscle. In muscle, the activities of all these enzymes decrease exponentially with increasing minimum depth of occurrence of the species; this decrease closely parallels the decrease in respiratory rate found previously for these same species. The decline in enzymic activity with increasing minimum depth of occurrence is much greater than the decline in body protein content of the whole fishes, suggesting a disproportionately rapid fall in enzyme concentration in comparison to contractile and structural protein concentrations. The similar reductions in activities of both glycolytic (LDH and PK) and citric acid cycle (CS, MDH and IDH) enzymes with depth indicate that both standard and active metabolisms of deeper-dwelling species are reduced relative to shallower-dwelling forms. There is no suggestion of increased anaerobic capacity with depth or in relation to species, occurrence in the oxygen minimum layer. In brain and heart, there is no significant decrease in enzymic activity with increasing minimum depth of occurrence. These two tissues have similar capacities for metabolism in most fishes, when comparisons are based on enzymic activity per gram wet weight of tissue.     

Biochemical indices of aerobic and anaerobic capacity in muscle tissues of California elasmobranch fishes differing in typical activity level

Biochemical indices of aerobic and anaerobic metabolic capacity were measured in white myotomal muscle of eight marine elasmobranch fish species representing a broad range of locomotor performance, and in red myotomal muscle and heart of three of those species. The objectives were to determine if metabolic capacities vary with typical fish activity level, to compare the endothermic mako shark with ectothermic pelagic sharks, and to compare elasmobranchs with teleosts in order to test the hypothesis that elasmobranchs have lower aerobic capacities, metabolic rates, and swimming speeds. In white myotomal muscle, activities of the enzymes citrate synthase (an index of aerobic capacity), pyruvate kinase, and lactate dehydrogenase (LDH, an index of anaerobic capacity) covaried with typical activity level, and the ability to tolerate intracellular acidification (nonbicarbonate buffering capacity) corresponded with LDH activity. Enzyme activities in red muscle and heart did not show a consistent pattern with respect to fish activity. In comparison with ectothermic sharks, the mako shark had greater aerobic and anaerobic capacities in white muscle, but no significant differences were found in red muscle or heart. This pattern has also been found in teleosts. Thus, endothermic fishes elevate the temperature of red muscle, a tissue specialized for high aerobic performance, whereas white muscle biochemical characteristics are adjusted to support high rates of contraction both aerobically and anaerobically. Muscle enzymic activities of elasmobranchs and teleosts with comparable locomotor habits are similar, thus refuting the hypothesis that elasmobranchs are sluggish, with lower metabolic capacities than teleosts.     

Metabolic and molecular stress responses of the gilthead seabream Sparus aurata during long-term exposure to increasing temperatures

Tolerance to a changing climate regime and persistence in the natural environment depends on the limited capacity to acclimate to changing temperatures. The present study aimed to identify and characterize thermal limits of the Mediterranean fish Sparus aurata as well as the processes providing heat protection during exposure to high temperatures. Processes studied included heat shock protein expression, protein kinase activity and metabolic adjustments. Molecular responses were addressed through the expression of Hsp70 and Hsp90 and the phosphorylation of stress-activated protein kinases, p38 mitogen-activated protein kinase (p38 MAPK) and cJun-N-terminal kinases (JNKs). Thermal impacts on metabolic capacities were assessed by studying the maximum activities of citrate synthase (CS), malate dehydrogenase (MDH) and 3-hydroxyacyl CoA dehydrogenase (HOAD) as well as pyruvate kinase (PK) and lactate dehydrogenase (L-LDH). The expression of Hsp70 and hsp90 was activated when the fish were exposed to temperatures beyond 20°C. Increased phosphorylation of p38 MAPK and JNKs indicated the parallel activation of MAPK signaling cascades and the potential involvement of MAPKs in the induction of Hsp genes. Exposure to extreme temperatures beyond 24°C caused an increase in the enzymatic activity of PK and LDH indicating an enhanced glycolytic potential.     

Effects of nutrition and temperature on metabolic enzyme activities in larval and juvenile red drum,Sciaenops ocellatus,and lane snapper,Lutjanus synagris

The metabolic enzyme activities were determined in larvae of red drum, Sciaenops ocellatus, and lane snapper, Lutjanus synagris, to determine the effect of temperature and nutrition on metabolic enzyme activities and to evaluate if metabolic enzyme activities are useful in assessing the feeding condition of larval fish. During experiments conducted during the spring of 1990, lactate dehydrogenase (LDH) activities in both red drum and lane snapper were approximately an order of magnitude lower than values typical for adult fish; LDH and citrate synthase (CS) activities increased during early developmental stages, but nutritional effects were apparent. Clear differences (up to 4-fold) between well-fed and starving fish were evident in both LDH and CS activity in red drum. Differences between well-fed and poorly fed larvae were evident until 9 d after hatching. Lane snapper larvae reared at a 25°C had significantly lower LDH activities than larvae reared at 28°C.     

The effect of pressure on muscle lactate dehydrogenase activity of some deep-sea and shallow-water fishes

The activity of crude muscle lactate dehydrogenase (LDH) of several species of bathypelagic and shallow-water fishes has been measured at pressures between 1 and 578 atm and at temperatures of 15° and 25°C. No relationship has been found between the effect of pressure on enzyme activity and the hydrostatic pressure of the organism's environment. Applied hydrostatic pressure reduced activity at both temperatures. The decrease at 25°C was double the decrease at 15°C in LDH from shallow-water fishes. However, enzymes from 2 bathypelagic fishes showed approximately the same reduction at both temperatures. Thus, the interaction of temperature and pressure was less in deep-sea than in shallow-water fish LDH. Decreasing temperature and increasing pressure would both reduce the activity of LDH. That is, deep-sea conditions are noncompensatory in this instance. It is possible that the dissociation of the effects of temperature and pressure could be an adaptive feature of deep-sea life.This paper is a portion of a thesis submitted to the Graduate School, University of Georgia, in partial fulfillment for the degree of Master of Science.     

Benthic community respiration in the N.W. Atlantic Ocean: in situ measurements from 40 to 5200 m

Benthic community respiration was measured in situ at 9 stations along the Gay Head-Bermuda transect from depths of 40 to 5200 m. Three methods were used; bell jar respirometers, grab respirometers, and free vehicle respirometers. Benthic community respiration rates spanned three orders of magnitude, decreasing from 21.5 ml O₂ m^-2 h^-1 at 40 m in November to 0.02 ml O₂ m^-2 h^-1 at 5200 m. Rates decreased two orders of magnitude between 40 and 1800 m and then significantly declined again between the continental rise (3650 m) and the abyssal plain stations. Predictive equations for benthic community respiration along the transect reflect a strong correlation with depth of water. Of lesser significance are the correlations with water temperature, dissolved oxygen, benthic animal biomass, surface primary productivity and sediment organic matter. Calculations show that annual benthic respiration can utilize 1 to 2% of the surface primary productivity. Of the 2 to 7% organic carbon fixed at the surface which supposedly reaches the bottom, only 15 to 29% is utilized by the benthic community at 2200, 3000, and 3650 m. The energy requirements of other biological components of deep-sea benthic communities, such as benthopelagic and macro-epibenthic animals, not included in these measurements, must also be considered in calculating a balance of carbon.Contribution from Scripps Institution of Oceanography.     

Depth zonation and metabolic adaptation in Dover sole,Microstomus pacificus,and other deep-living flatfishes: factors that affect the sole

Flatfishes of Monterey Bay, central California, undergo species replacements with increasing depth along a transect from 100 m on the continental shelf down to a depth of 1400 m on the continental slope. The Dover sole, Microstomus pacificus, differs from the other local flatfish species by undergoing an extensive ontogenetic vertical migration, occupying all depth zones at different life stages, and having its maximum spawning biomass in the oxygen minimum zone between 600 and 1000 m. Size-activity relationships and depth-activity relationships for the glycolytic enzyme lactate dehydrogenase (LDH) and for two enzymes associated with aerobic metabolism, malate dehydrogenase and citrate synthase (CS), were examined in white-muscle tissue of shallow-living, deep-living and ontogenetically-migrating species. Scaling coefficients (b) for weight-specific enzyme activity (log activity)=a+b (log wet weight), varied in sign as well as magnitude for fishes living at different depths. In the shallow-living California halibut Paralichthys californicus, LDH scaled positively (0.39) and CS scaled negatively (-0.15) with size, a pattern observed previously for most shallow-water fish species. The permanently deep-living species, the deepsea sole Embassichthys bathybius, differed in that both LDH and CS scaled strongly negative (-2.0 and-1.5, respectively). For the ontogenetically migrating Dover sole Microstomus pacificus, there was a shelf-slope transition. For the shelf specimens (200 m), LDH scaled positive (0.11) and CS negative (-0.29) and for the slope specimens (400 m), LDH scaled negative (-0.65) and CS strongly negative (-0.63). Rex sole, Glyptocephalus zachirus, showed a similar shelf-slope transition. Intraspecific depth-enzyme activity differences were not incremental, but changed abruptly between the continental shelf stations (100 to 200 m) and the continental slope (400 to 1400 m). Based on comparisons with laboratory-maintained individuals, the decline in the metabolic capacity of the white muscle of Dover sole is a phenotypic response to the low food and oxygen conditions of the continental slope. Contrary to expectation, anaerobic capacity (LDH activity) decreased in response to low oxygen conditions, suggesting that in a permanently hypoxic environment such as the oxygen minimum zone the metabolic strategy may be to not incur an oxygen debt that would be difficult to pay back.     

Lipid composition and partitioning of deepwater chondrichthyans: inferences of feeding ecology and distribution

The composition of lipids and fatty acids was determined for the livers, muscle, pancreas, kidney and stomach fluids of deepwater chondrichthyan species (including 11 squaliformes, 3 chimaeriformes, 1 hexanchiforme and 3 carcharhiniformes) caught as bycatch from continental waters off south-eastern Australia. The lipid class, fatty acid and fatty alcohol composition differed markedly in each tissue and in each species. The lipid and fatty acid composition of large, lipid-rich (38–70% wet weight, ww) livers demonstrated the multifunctional role of this organ in: lipid distribution, storage and biosynthesis, and buoyancy regulation. In the liver, the importance of certain lipids (including squalene, diacylglyceryl ethers, triacylglycerols and to a lesser extent wax esters) as mediators of buoyancy varied according to lifestyle and habitat. Less variability was observed in the muscle profiles, characterized by low lipid content (<1.0% ww) and high relative levels of polar lipids (>70%). The lipid and fatty acid profiles of the kidney and pancreas showed the highest intraspecific variability, suggesting these organs also have complex roles in lipid storage and metabolism. Overall intra- and interspecific differences in the tissue fatty acid profiles could be related to differences in a number of factors including phylogeny, habitat (depth), buoyancy regulation and diet and presumably also reflect different ecological roles. The lipid and fatty acid profiles are the first published for Rhinochimaera pacifica, Chimaera lignaria and Figaro boardmani and the first to demonstrate interspecific variation in lipid profiles of various tissues of deepwater chondrichthyans. The application of multivariate analysis to lipid class and fatty acid tissue profiles in chondrichthyans inferred dietary differences and metabolic preferences between species and habitats. These results have important implications for the future use of fatty acids as dietary tracers in chondrichthyan research.     

Temporal patterns in diversity and species composition of deep-sea benthic copepods in bathyal Sagami Bay,central Japan

Little is known about temporal changes in the diversity and species composition of deep-sea metazoan meiofauna and their relationships with changes in the food supply. Those changes were studied for benthic copepod assemblages based on 2-year time-series data at a bathyal site in Sagami Bay (1430 m depth), central Japan, where annual fluctuation in the abundance of benthic foraminiferans was previously observed. Species diversity of benthic copepods at the site was as high as, or slightly higher than, that observed at other deep-sea sites, but did not fluctuate temporally through the study period. Multivariate analyses did not reveal any clear seasonal or directional change occurring over the longer term in their species composition, although there was some consistent pattern. These results indicate a lack of, or only weak, seasonality in the diversity and species structure of the deep-sea benthic copepod assemblages, even though the fresh organic food supply fluctuates seasonally. They also suggest that there are differences between copepods and foraminiferans in the response to changes in environmental factors, and that spatial differences in the composition of copepod communities are greater than temporal ones at this deep-sea site.Communicated by T. Ikeda, Hakodate     

Influence of depth and season on the diet of the deep-water aristeid Aristeus antennatus along the continental slope (400 to 2300 m) in the Catalan Sea (western Mediterranean)

The composition of the diet of the deep-sea aristeid shrimp Aristeus antennatus (Risso, 1816) was determined based on the analysis of 1578 stomach contents. Samples were collected using bottom trawls during 1984 to 1989 along the deep continental slope (380 to 2266 m) in the Catalan Sea (Western Mediterranea). A. antennatus displayed a high feeding activity on a large variety of endobenthic and epibenthic invertebrates, whereas benthopelagic prey were supplementary to its diet. The shrimp's diet varied mainly as a function of depth. Various preferred prey items on the middle slope (Calocaris macandreae, Cirolana borealis, Abra longicallus) disappeared from its diet below 1300 m, coinciding with the transtion boundary between different taxocenoses in the Catalan Sea. Less mobile prey and inert remains (mainly pteropods) contributed a progressively larger share of the diet of Aristeus antennatus with increasing depth. The importance of seasonality in the changes in diet decreased with increasing depth, and was relatively high only in the upper middle slope (down to 1000 m depth), where two seasonal dietary groups were detected. The most distinct seasonal changes in diet were among benthopelagic prey. The diet of A. antennatus in the submarine canyons mainly consisted of endobenthic prey (large polychaetes, ophiuroids). The higher stomach-fullness values and less diversified diet in this area were probably related to high productivity in the submarine canyons.     

Metabolic cold adaptation in Antarctic fishes: evidence from enzymatic activities of brain

To evaluate the concept of metabolic cold adaptation (MCA) in fishes, we compared - in brain, red muscle, and white muscle of Antarctic notothenioid fishes and tropical/subtropical fishes - the activities of two enzymes of ATP-generating pathways, citrate synthase (CS), an indicator of citric acid cycle activity (aerobic metabolism), and lactate dehydrogenase (LDH), an indicator of potential for ATP production through anaerobic glycolysis. Brain was chosen because, unlike locomotory muscle, its metabolic activity is not likely to be influenced by a species' level of activity or nutritional status, so MCA should be readily observed if present. CS and LDH activities in brain exhibited a high level of MCA, but compensation to temperature was not complete (48% for CS; 46% for LDH). CS and LDH activities in red and white muscle varied widely among species, according to the general level of locomotory activity. The 'mode of life'-related enzymatic activities in locomotory muscle show that study of MCA at the level of whole organism metabolism is fraught with difficulties and experimental ambiguities. In contrast, the low variation among species within each group in enzymatic activities in brain, and the large differences between groups in CS and LDH activity, show that brain is an excellent study system for evaluating metabolic compensation to temperature.     

硫化物胁迫对日本沼虾呼吸代谢和能量代谢酶的影响

研究了硫化物暴露后日本沼虾细胞色素氧化酶(CCO)、琥珀酸脱氢酶(SDH)、延胡索酸还原酶(FRD)和乳酸脱氢酶(LDH)4种呼吸代谢酶及能量代谢酶-精氨酸激酶(AK)活性的变化规律.将日本沼虾暴露于0.6 mg·L~(-1)、2mg·L~(-1)的2个硫化物质量浓度组和不含硫化物的对照组水体中,暴露后0、2、12、24、48 h和解除暴露后48 h取肝胰腺和肌肉组织进行酶活性分析.结果显示:肝胰腺和肌肉组织SDH和CCO活性随硫化物质量浓度升高或暴露时间延长而显著降低(P<0.05).FRD、LDH和AK活性随硫化物质量浓度升高或暴露时间延长而显著升高(P<0.05).解除硫化物暴露后48 h,各质量浓度组酶活性与对照组无显著差异.上述酶活性还存在组织差异,肝胰腺呼吸代谢酶活性高于肌肉中相应酶活性,而AK活性与此相反.结果表明,硫化物胁迫导致日本沼虾有氧呼吸代谢减弱,无氧呼吸代谢增强,并动用其能量贮存物质磷酸精氨酸,产生更多ATP以适应外界不良环境.     

Metabolism,enzymic activities and cold adaptation in Antarctic mesopelagic fishes

Several species of Antarctic mesopelagic fishes that have different minimal depths of occurrence but the same environmental temperature were collected in November–December 1983 and in March 1986 between 0 and 1 000 m in the open water near the marginal ice zone in the vicinity of 60°S 40°W (1983) and 65°S 46°W (1986), and oxygen consumption rate (V _{O 2}) and the activity of two metabolic enzymes, lactate dehydrogenase (LDH, an indicator of the anaerobic potential of locomotory muscle) and citrate synthase (CS, an indicator of citric acid cycle activity or aerobic potential), were determined. In four dominant species, whole-individual oxygen-consumption rate (y, ml O₂ individual^–1 h^–1) varied with weight (X, g) according to the equation y=aX ^b, with b values falling between 0.889 and 1.029. The relation of weight-specific LDH activity (y, U g^–1 wet wt) with weight (x, g) was also described by the equation y=aX ^b, with b values varying between 0.229 and 1.025. Weight-specific CS activity declined with weight, with b values from-0.031 to-0.369. V _{2 O}, LDH activity and CS activity all declined markedly with increased species' minimum depth of occurrence (the depth below which 90% of a species' population lives). Comparisons with previous studies on ecologically equivalent species of the California Borderland indicate that depth-related decreases in metabolism are the result of adapted traits of deeper-living species, not declining temperature within the water column. The metabolic rate of Antarctic mesopelagic fishes is approximately twice that of California species at equivalent temperatures; similar rates were found at the normal habitat temperatures of the two groups. Thus, a well-developed compensation for temperature is present in the Antarctic fishes: cold adaptation. Differences in enzymic activity among species, and among different sized individuals of a species are related to differences in metabolic rate and locomotory capacity. Enzymic indices can be used to estimate metabolic rates and evaluate ecological parameters such as predatory strategies and niche separation.     

Bathymetric patterns of genetic variation in a deep-sea protobranch bivalve, Deminucula atacellana

The origin of the deep-sea benthic fauna is poorly understood and represents an enormous gap in our understanding of basic evolutionary phenomena. One obstacle to studying evolutionary patterns in the deep sea has been the technical difficulty of measuring genetic variation in species that are typically minute, rare, and must be recovered from extreme depths. We used molecular genetic techniques to quantify variation in the 16S rRNA mitochondrial gene within and among populations of the common protobranch bivalve Deminucula atacellana (Schenck, 1939). We analyzed 89 individuals from nine samples collected in the 1960s along a depth gradient from 1100 to 3800 m in the western North Atlantic. Genetic variability within populations is much lower than between populations, and peak haplotype numbers occur near the center of its depth distribution. Continental slope (<2500 m) and rise (>2500 m) populations were genetically distinct despite the lack of any obvious topographic or oceanographic features that would impede gene flow. These findings indicate that the deep-sea macrofauna can have strong population structure over small (134 km) spatial scales, similar to that observed in shallow-water and terrestrial organisms. This surprisingly high biodiversity at the genetic level affords the potential for adaptation and evolutionary diversification, the ultimate historical causes of high species diversity in the deep-sea benthos. Received: 24 July 1997 / Accepted: 26 January 1998     

Spatial changes in the distributions of deep-sea “Cerviniidae” (Harpacticoida, Copepoda) and their associations with environmental factors in the bathyal zone around Sagami Bay, Japan

To estimate species turnover rates on scales of several tens of km in deep-sea benthic animals, we analyzed spatial and inter-annual changes in species diversity and composition of cerviniids, a typical group of deep-sea harpacticoids, at stations in and around Sagami Bay, central Japan. Associations with environmental factors were also investigated. Generally, bathymetrical patterns in diversity of benthos are unimodal and peak at depths of 2,000–3,000 m. In Sagami Bay, cerviniid diversity did not follow this trend; both species richness and evenness were negatively correlated with water depth. Multivariate analyses [detrended correspondence analysis (DCA) and non-metric multi-dimensional scaling] suggested that temporal changes in species composition of cerviniids are smaller than spatial changes that occur on horizontal scales of several tens of km. Community structure does not change completely on these scales in the bathyal zone around Sagami Bay. DCA also showed that bathymetrical changes in species composition can be regulated by certain factors associated with water depth.     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.     

In situ filtering and ingestion rates of deep-sea benthic boundary-layer zooplankton in the Santa Catalina Basin

In situ rates of filtration, particulate ingestion, and carbon ingestion of deep-sea benthic boundary-layer zooplankton were determined in December 1984 in the Santa Catalina Basin, at 1 300 m depth in the California Borderland, by a short-term radioisotope-incorporation technique. Zooplankton were collected at 1 or 50 m above the bottom with an opening-closing net system on a submersible, and incubated at depth with labelled amino acids in special cod-end chambers. Concentrations of particulate material and particulate organic carbon in the ambient water were also measured. The zooplankton had a median weight-specific filtration rate of 12.4 ml (mg dr. wt)^-1 h^-1 and a median carbon ingestion rate of 5.4 g C (mg dr. wt)^-1 h^-1. Filtration rates were not significantly different from those in similar experiments in the north Atlantic at 2 175 m depth or Narragansett Bay in the winter, although the medians of the deep-sea experiments were lower than for the Bay. In the Santa Catalina Basin, rates from experiments at 1 m above the bottom in more turbid water were not significantly different from those at 50 m above the bottom in clearer water. These deep-sea benthic boundary-layer zooplankton may have the potential to respond to food pulses, and their relatively high ingestion rates suggest that they could have significant effects on particulate, chemical, and bacterial processes in the near-bottom water column.     

Benthic oxygen uptake and carbon cycling under aphotic and resource-limiting conditions in a submarine cave

To establish relationships between organic input to the benthos and decreases in benthic population biomass and density, benthic oxygen uptake was measured in an oligotrophic submarine cave in the northwestern Mediterranean Sea (Marseille, France), on seven separate occasions in 1987, using an in situ bell-jar respirometer. Oxygen uptake was measured in both the outer twilight section and the dark inner section of the cave during an annual survey (seven recording periods from February 1987 to November 1988). The mean annual benthic oxygen uptake was 80.9 litres O₂ m^–2 yr^–1 for the twilight outer section and 15.5 litres O₂ m^–2 yr^–1 for the dark inner section. The results are discussed and the biogeochemical budget for particulate organic carbon at the sediment-water interface calculated. Respiration rates (expressed as carbon equivalents), together with previously published data on vertical fluxes and burial of organic carbon, revealed that anaerobic pathways accounted for 14% and aerobic pathways for 86% of the total benthic metabolism in the outer part of the cave. In the inner section of the cave, degradation of organic carbon occurred only through aerobic degradation, indicating a strongly carbon-limited ecosystem. The low respiration rates recorded in the dark section were similar to values recorded for some oligotrophic deep-sea environments (1 000 to 2 000 m). Such budgets are essential preliminary steps in order to accurately model benthic metabolic pathways. The determination of annual fluxes linked to the acquisition of long-term data will yield better knowledge of the recycling processes at the water-sediment interface.     

Effects of long-term acclimation to environmental hypercapnia on extracellular acid–base status and metabolic capacity in Mediterranean fish <Emphasis Type="Italic">Sparus aurata</Emphasis>

In the context of future scenarios of anthropogenic CO₂ accumulation in marine surface waters, the present study addresses the effects of long-term hypercapnia on a Mediterranean fish, Sparus aurata. By equilibration with elevated CO₂ levels seawater pH was lowered to a value of 7.3, close to the maximum pH drop expected in marine surface waters from atmospheric CO₂ accumulation. Intra- and extracellular acid–base parameters as well as changes in enzyme profiles were studied in red and white muscles and the heart under both normocapnia and hypercapnia. The activities of pyruvate kinase (PK), lactate dehydrogenase (L-LDH), citrate synthase (CS), malate dehydrogenase and and 3-hydroxyacyl CoA dehydrogenase (HOAD) reflect the pathways and capacity of oxidative processes in metabolism. Long-term hypercapnia caused a transient reduction in blood plasma pH (pH_e) as well as in intracellular pH (pH_i). Compensation of the acidosis occurred through increased plasma and cellular bicarbonate levels. Changes in enzymatic activities, especially the increase in the activity of L-LDH, paralleled by a drop in CS activity in white and red muscles reflect a shift from aerobic to anaerobic pathways of substrate oxidation during long-term acclimation under hypercapnia. The present results suggest that moderate environmental hypercapnia changes the metabolic profile in tissues of S. aurata. Consequences for slow processes like growth and reproduction potential as well as potential harm at population, species and ecosystem levels require further investigation.