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.     

Chemical composition of midwater fishes as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

The variation with depth in water, lipid, protein, carbon and nitrogen contents (% wet weight) of 42 species of midwater fishes, collected in November 1976 off the west coast of Oahu in the Hawaiian Archipelago, was measured. The Hawaiian fishes show significant relationships between these components and depth of occurrence. The slopes of these relationships are not significantly different from those reported for midwater fishes from off California, USA. However, the fishes from Hawaii have significantly lower lipid levels and higher protein levels than do the species from off California. The deep-living Hawaiian species (500 m and deeper) have significantly lower lipid (% wet weight), but there is no significant difference in protein (% wet weight). The difference in lipid contents at all depths appears to be an evolved characteristic, with the greater lipid levels off California being selected for by greater spatial and temporal variation in the food supply for these fishes off the California coast than off Hawaii. The higher protein contents in the shallow-living Hawaiian fishes appear to reflect greater muscle power selected for in these fishes by the greater water clarity, and therefore greater reactive distances, in the surface layers off Hawaii. These conclusions support the general hypothesis that the lower protein contents of bathypelagic fishes are not directly selected by food limitation at depth, but rather result from the relaxation of selection for rapid-swimming abilities at greater depths due to the great reduction at greater depths in the distance over which visual predator-prey interactions can take place. The lower lipid levels in the deeper-living species are apparently made possible by the reduced metabolic rates of these species which reduces their need for energy stores.     

Triacylglycerols and wax esters in the lipids of deep midwater teleost fishes of the Southern California Bight

Total lipid contents and the proportions of triacylglycerols and wax esters were examined in 23 species of deep-sea fishes collected between 1977 and 1980 in the Santa Catalina, San Clemente and San Pedro Basins off Southern California, USA. Mean total lipid content ranged from 1 to 22% of wet weight. Triacylglycerols accounted for 1 to 91% of total lipids present and were more abundant than wax esters in the lipids of all but a single species. Wax ester contents ranged from 0 to 24% of total lipids but in 20 species were 10% of the lipids. These results do not substantiate the previously suggested idea that wax esters are important constituents of the lipids of deep-sea fishes. Moreover, the hypothesis that wax ester content increases with increasing depth of occurrence is not supported. To assess the importance of the lipids present in bouyancy regulation, the water content and swimbladder inflation of these species were also examined. Neither total lipids nor wax esters appear to play an important role in buoyancy regulation in the majority of these fishes. Eight species have gas-filled swimbladders. Of the species lacking inflated swimbladders, 7 have watery bodies (87 to 95% H₂O) and 6 with low lipid (8.3%) and water (85%) contents may need to swim continuously to maintain their position in the water column. Lipids, largely in the form of triacylglycerols, may be important in the buoyancy of three species with lipid contents>10% of wet weight. Two of these species also possess inflated swimbladders. The reasons for the production of large quantitites of was esters by certain deep-sea teleosts, especially gonostomatids and myctophids, remain unknown.     

Crossing the frontier: vertical transit rates of deep diving cormorants reveal depth zone of neutral buoyancy

Knowing the depth zone of neutral buoyancy of divers is important because buoyancy can determine how animals manage their energy budget. In this study, we estimate the depth zone of neutral buoyancy of free-ranging cormorants for the first time, using time-depth recorders. We discovered that vertical ascent rates of 12 Crozet and 15 Kerguelen diving blue-eyed shags (respectively Phalacrocorax melanogenis and P. verrucosus) slowed down considerably at the 50–60 m depth zone. We suggest this was due to birds trying to reach the surface from that point upwards using reduced locomotor activity because the force of buoyancy becomes greater than the force of gravity at that depth. The results show a shift of this depth zone in relation to maximum targeted dive depth, suggesting cormorants may control buoyancy through respiratory air volume adjustment. Interestingly, 60 m is close to the maximum depth zone reached by these two species during dives lasting 4 min, their estimated behavioural aerobic dive limit. This suggests that the decision to swim deeper has a direct consequence on the energy budget, with time spent recovering at the surface (time thus lost to foraging) strongly increasing relative to the preceding time of submergence. Resources found in deeper waters must be of sufficient quantity or quality to justify crossing the frontier of physical neutral buoyancy.     

Proximate composition and energy content of mesopelagic fishes from the eastern Gulf of Mexico

The proximate composition of 33 species of mesopelagic fishes collected from the eastern Gulf of Mexico during May–June 1984, July–November 1985 and January 1986 was determined. Water level increased and ash-free dry weight (% dry weight) and protein level (% wet weight) decreased with increasing species' minimum depth of occurrence (that depth below which 90% of a species' population lives). Lipid level (% wet weight or % ash-free dry weight), did not change with species' minimum depth of occurrence. Skeletal ash level (% wet weight) generally decreased with minimum depth of occurrence, whereas carbohydrate level did not change with depth. The variable water level, low lipid level, and high protein level in eastern Gulf of Mexico fishes resulted in a low energy content. These results are generally characteristic of fishes from warm, stable regions of low productivity, such as the eastern North Pacific Gyre. The constant food supply provided by a stable environment may obviate the need for large lipid reserves, in contrast to colder waters at higher latitudes where food availability is seasonal. In addition, the large energy requirements for diel excursions into high-temperature surface waters by the many vertically migrating fishes of this region may influence lipid deposition. The relatively high protein level found in migrators compared to that in non-migrators or weak migrators indicates that locomotory demands influence the percentage of protein found in Gulf fishes. The lack of a positive correlation between protein level and the food availability of a zoogeographic region, suggested in previous studies, is supported here.     

Proximate and elemental composition and energy content of mesopelagic crustaceans from the Eastern Gulf of Mexico

The proximate and elemental chemical compositions of 25 species of pelagic decapod and mysid crustaceans collected from the eastern Gulf of Mexico (27°N; 86°W, 1984 to 1989) was examined. Water level ranged from 63 to 95% and increased slightly with species' increased depth of occurrence. Protein levels were generally high (1.5 to 18.3% wet wt, WW; 27.6 to 62.4% ash-free dry wt, AFDW) and comprised the primary organic component in the majority of species. Protein, both as % WW and % AFDW, decreased with increased depth of occurrence. In contrast to protein, lipid levels were low (0.5 to 8.9% WW; 5.7 to 60.9% AFDW), and increased with increased depth of occurrence. Carbon and nitrogen best mirrored measured lipid and protein levels when considered as non-protein carbon and non-chitin nitrogen, respectively. C:N ratios increased with increased depth, consistent with changes in protein and lipid with depth. Because of their compositional attributes, resident Gulf of Mexico species have a low total wet weight energy content relative to species from more productive regions. Energy content showed no significant trend with depth. Vertical migration patterns were distinctly different between shallow-and deep-living gulf species and these differences were largely responsible for the relationships observed between composition and depth. In migrating species, the protein and nitrogen content were higher, the lipid and carbon contents and C:N ratio lower, than in non-migrating species. Three deep-living species of the genus Acanthephyra were found to be compositionally atypical, resembling shallow, migrating types rather than other deep-living, non-migratory species.     

Calorific values of benthic marine algae and their postulated relation to invertebrate food preference

Data are given on the kcal/g dry weight, percent ash, and kcal/ash-free g of dry weight for 1 marine diatom species, 70 macroscopic benthic algae, and 1 marine tracheophyte species. For 41 of these, the data have also been converted to kcal/g wet weight. Calorific values, though relatively homogeneous within genera, appear to be influenced by phyletic affinity, water purity, or depth of immersion, and such ecological properties as growth form, generation time, and relative susceptibility to herbivory. Seasonal factors, portion of the plant combusted and growth rate appear to have little effect. The food preferences of various invertebrate herbivores seem to have evolved more in response to an availability factor than to absolute food value.     

Standing crop and leaf area index of the sublittoral Laminaria species near Helgoland

Standing crop values of Laminaria species sampled between 0 to 2 m depth in the sublittoral of Helgoland, a small rocky island in the southern North Sea, are similar to those reported by other workers from Norwegian or Scottish coasts. Below 2 m water depth, however, standing crop values are found to be reduced to a far greater extent with increasing depth than in the Norwegian or Scottish areas studied. Maximum standing crop values are not found in Laminaria digitata and L. saccharina growing just below the low water mark of spring tides, but in the upper L. hyperborea forest at 2 m depth. Population density and leaf area index (LAI) decrease with increasing depth (but not the average weight of individuals constituting the L. hyperborea forest). It is suggested that the depth at which LAI becomes 1 represents an important ecological threshold, since growth rates of L. hyperborea specimens appear to decrease at greater depths but not to increase above the threshold. Self-shading might equalize, to a certain extent, the quantity of underwater irradiance available to an individual phylloid at all depths at which a close vegetation of Laminaria species occurs.     

Biomass of the invertebrate megabenthos from 500 to 4100 m in the northeast Atlantic Ocean

Data are presented on the biomass of the invertebrate megafauna at 22 stations on the continental slope in the Porcupine Seabight (PSB) (northeast Atlantic Ocean). Samples were collected between 1980 and 1982. Several units of biomass are used, all of which illustrate a decrease by a factor of about 30 from 500 to 4100 m. Lognormal curves were fitted to the data, the gradients of which were very similar for all biomass units and similar to the value for a transect down the continental slope in the western Atlantic. Biomass levels in the PSB are compared with those from other deep-sea environments. Some published values are more than ten times higher than the values reported here, while others are less than a tenth. The reasons for these differences and trends are discussed in terms of food supply. Sampling variability was examined at two stations, but by chance one (at 1300 m) appeared to encompass a sharp faunal discontinuity of the dominant fauna and the other (at 4000 m) contained very small numbers of large animals. For these reasons, sample variability was high at the repeat stations. Suspension-feeders and crustaceans dominated the biomass at upper-slope depths, while echinoderms were dominant on the middle and lower slope. As a result of this phyletic change, there was a small but insignificant decrease in mean body weight with increasing depth. Within phyla there was also a small but insignificant decrease with depth. If large species are excluded from the biomass/depth regression, the gradient changes considerably, demonstrating the increasing importance of small species at greater depths. The size distribution of megafaunal biomass was examined at several stations. This indicated that the megafauna form a functional group distinct from the macrofauna, just as the macrofauna are distinct from the meiofauna.     

Relationship of life-history patterns to depth in deep-water caridean shrimps (Crustacea: Natantia)

Deep-water shrimps, distributed on the steep outer reef slopes of tropical Pacific Islands, were obtained by setting baited traps in depths down to 850 m in the vicinity of Laucala Bay, Fiji, over the period 1979 to 1983. Life-history variables were estimated and interspecific comparisons made between Parapandalus serratifrons, Plesionika longirostris, Heterocarpus ensifer, H. gibbosus, H. sibogae, H. laevigatus (Pandalidae) and Saron marmoratus (Hippolytidae) which have different depth distributions. Results suggested that, interspecifically, reproductive lifespan tended to increase with increasing depth of distribution and that mean egg sizes of each species increased with increasing depth. Brood sizes tended to increase inconsistently with depth, although relative brood size (number of eggs per “standar” 15 g individual) appeared unrelated to depth. Although annual reproductive effort varied inconsistently with depth, reproductive effort totalled over the lifespan of the Heterocarpus species tended to increase with increasing depth. Some of the findings are counter to competition-based ecological theory, and it is proposed that adult predation decreases with increasing depth, allowing deeper-water species to have an extended lifespan, an increased degree of iteroparity, and a corresponding increase in lifetime reproductive effort. Further, we suggest that the probabilities of larval survival, which appear to decrease with increasing depth, are offset by the production of larger eggs by deeper-water species.     

Food availability as a selective factor on the chemical compositions of midwater fishes in the eastern North Pacific

In September and October 1980 we examined the relationships between food availability, depth, and chemical composition among 12 midwater fish species, from three adjacent areas of the eastern North Pacific: the eastern gyre, the California Current, and the transitional region between them. By comparing trends in chemical composition across a geographical productivity gradient, the influence of food availability could be examined both with and without depth as a dependent variable. In general, caloric density, lipids, and water content showed consistent trends along both vertical and horizontal gradients of food availability. Lipids and caloric contents were lowest among bathypelagic species and among fishes from the gyre. Water content was highest in the gyre and among deeper-living species. While protein content as a percentage of the total wet weight declined with depth, there was no clear trend among mesopelagic species between stations. Protein as percent of the total organic content showed no decrease with depth, but was significantly higher in the gyre among mesopelagic species. These findings suggest that compositional trends in water, lipids, and caloric content are correlated with food availability, regardless of any depth-related factors. In contrast, protein did not vary consistently with food availability and thus may be linked to other factors.     

Oxygen consumption of midwater fishes and crustaceans from the eastern Gulf of Mexico

Oxygen consumption was measured as a function of temperature, oxygen partial-pressure (P_O2)and species depth of occurrence for twenty-three species of midwater fishes and crustaceans collected from the eastern Gulf of Mexico from June 1981 to July 1985. Q₁₀s (7° to 20°C) of 3.90 and 3.24 were recorded for myctophid and non-myctophid fish groups, respectively, while values of 2.22, 2.19, 2.19 and 2.54 were calculated for sergestid, penaeid, carid and euphausiid crustacean groups, respectively. Q₁₀s were consistent for species within each group. All of the species tested regulated their oxygen consumption to P_O2levels normally encountered within the eastern Gulf. Values of critical partial pressure (P_c) ranged from 20 to 40mm Hg and increased slightly with increasing temperature and respiration rate. Declining respiration with increasing minimum depth of occurrence was primarily a function of temperature alone. Changes in size, dry weight and water content contributed only a small fraction to the observed decrease. This finding contrasts with studies from the eastern Pacific Ocean, where temperature is a minor contributor to changes in respiration rate with depth.     

Distribution,abundance and survival of juvenile hermatypic corals (Scleractinia) and the importance of life history strategies in the parent coral community

The distribution and abundance of juvenile corals were examined at depths from 3 to 37 m on the reefs of Curaçao and Bonaire (Netherlands Antilles). Juveniles of Agaricia agaricites were most abundant (60.6%), followed by Helioseris cucullata (8.3%). The large massive corals such as Montastrea annularis, M. cavernosa and branched species such as Madracis mirabilis and Acropora palmata had few juveniles. This, combined with species characteristics, shows that these species employ very different life history strategies. In some species the abundance of juveniles over the reef paralleled that of larger colonies, but not for example in Agaricia agaricites. The composition of the coral community was apparently no direct function of juvenile abundance. A change in angle of settlement of A. agaricites juveniles with increasing depth, from vertical to horizontal surfaces, seems to reflect the preferred light intensity. We studied the survival of juvenile corals during a half-year period. One-third remained unharmed, one-third died or disappeared, and one-third was limited in growth by factors such as spatial competition. This was the same for all depths, but factors influencing survival varied with depth.     

Observation and collection of deep-reef fishes off the coasts of Jamaica and British Honduras (Belize)

Over 60 species of reef fishes were observed from submersibles between 50 and 305 m depth, and 25 species were collected from the same depth range using fish traps and submersibles. Short transects gave relative abundance of some species with depth. Many shallow-water coral-reef fishes reach depths far below the lower growth limits of reef-building corals. A true “deep-reef” fauna exists, but the juveniles of some of these species are found at depths of less than 50 m.     

The occurrence of wax esters in crustaceans from the North-east Atlantic Ocean

The lipids of 2 species and 1 group of copepods, 4 species of euphausiids, 2 species of mysids, and 6 species of decapods, taken from different depths in the North-eastern Atlantic Ocean and Western Mediterranean Sea, have been analysed specifically for their wax ester content. The mid-water animals living below 500 m contained large amounts of wax ester (30 to 80% total lipid), while those animals living in the upper 600 to 700 m or on the sea floor had only small wax ester fractions (5 to 10% total lipid). It is suggested that the wax esters are laid down by those animals living in areas of low food density, where energy conservation is important, partly to achieve a more neutral buoyancy. In addition, it would appear that wax esters can function as energy reserves.     

Feeding ecology of the loggerhead sea turtle Caretta caretta in the Northwestern Gulf of Mexico

Digestive tract contents collected from carcasses of 82 loggerhead sea turles (Caretta caretta) found on the south Texas coast from 1986 through 1988 were examined. Benthic invertebrates were the predominant prey. Sea pens (Virgularia presbytes), crabs, and mollusks accounted for 94% of the dry weight of the digestive tract samples. Temporal changes in the percent occurrence and percent dry weight of sea pens, crabs, and mollusks in the digestive tract samples were significant. Loggerheads fed primarily on sea pens during spring, then primarily on crabs during summer and fall. The increase of crabs in the loggerhead diet paralleled the annual increase in the abundance of crabs in the Gulf of Mexico. Sea pens were located nearshore at depths of 6 to 12 m and had a disjunct distribution. Present address: Texas A&M University, Department of Biology, College Station, Texas 77843-3258, USA     

Metabolic rates of midwater crustaceans as a function of depth of occurrence off the Hawaiian Islands: Food availability as a selective factor?

During July of 1983, 1986, and 1987, we measured rates of oxygen consumption of 234 individuals of 17 species of midwater crustaceans (orders Decapoda, Mysidacea, and Euphausiacea) off the Hawaiian islands at depths from the surface to greater than 1200 m. The routine metabolic rates declined with increasing depths of the species' occurrence to an extent greater than could be accounted for by depth-related changes in body size or water temperature. Most species appeared able to regulate their oxygen consumption down to the lowest oxygen partial pressures found in their depth range (20 mm Hg O₂), but did not regulate to such low oxygen partial pressures as did similar midwater crustaceans off California, where oxygen levels reach as low as 6 mm Hg. Metabolic rates of the shallower-living, but not the deepest-living Hawaiian crustaceans were significantly higher than those of Californian crustaceans. This is interpreted as indicating that the metabolic rates of midwater crustaceans are not adapted specifically to differing levels of primary production and that the decline with depth of metabolic rates in these species is not the result of food limitation at depth. The data are, however, consistent with the hypothesis that lower metabolic rates at depth are due to the relaxation of selection pressures relating to visual predation near the surface.     

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.     

Recherches sur la situation trophique d'un groupe d'organismes pélagiques (Euphausiacea). III. Potentiel alimentaire du groupe

The position of a group of pelagic organisms among the food webs depends upon (1) its abundance in the biomass; (2) the factors allowing its utilization by predators; among these, the size distribution of animals and their bathymetric repartition appear to prevail. In the equatorial and southtropical Pacific Ocean, euphausiids represent 8% of the total macroplanktonic and micronektonic biomass, i.e., approximately 13 g/1000 m³ wet weight (somewhat less in oligotrophic tropical zones, a little more in the richer equatorial belt); Euphausia diomedae accounts for more than 50% of this biomass. Individuals measuring 9 to 18 mm in total length (4 to 37 mg wet weight) constitute 84% of the whole biomass of the group; the importance of each species in terms of size groups is discussed. Depth distribution is examined at the specific level. By night, 75% of the biomass concentrates in the 0 to 160 m water layer (ca. 10 g/1000 m³ wet weight; main species Euphausia diomedae), 22% between 160 and 300 m (3 g/1000m³ main species Nematoscelis spp.), and 3% (0.3g/1000 m³) in deeper layers. During the day, the only species abundant at depths less than 400 m belong to the genera Stylocheiron and Nematoscelis; specimens smaller than 15 mm and 20 mg remain at depths shallower than 200 m, individuals between 15 and 22 mm (20 to 65 mg) appear around 200 to 400 m, and larger animals are restricted to depths below 400 m.     

Growth parameters of deep-water decapod crustaceans in the Northwestern Mediterranean Sea: a comparative approach

 Relative and absolute growth were studied in 17 species of deep-water decapod crustaceans, spanning nine families of six different infra-orders, in the Northwestern Mediterranean Sea. The overall maximum abundance of these species lay between 200 m and 750 m (i.e. upper- and mid-slope species). Relative and absolute growth rates were compared by contrasting the slopes of the size–weight relationships for the different species and calculating the von Bertalanffy growth-equation parameters asymptotic length (L _∞ ) and growth rate (k). The size–weight relationships differed significantly as function of the species' life habits. The results revealed a significant decrease in weight relative to size in mesopelagic species (which carry out diel vertical migrations), an almost isometric relationship between size and weight in the less mobile nektobenthic species, and a significant increase in weight relative to size in strictly benthic species. The mean allometric coefficient for each group increased significantly from mesopelagic to benthic species. However, no general trend was observed in the growth-performance index, Φ (an index used to compare absolute growth rates between species, as a function of habit and depth of maximum abundance for all species combined), suggesting that the deep-water decapod crustaceans studied have similar absolute growth rates. Nevertheless, comparison of growth-parameter and growth-performance index values within families did reveal differences. Mesopelagic species of the families Sergestoidae and Pasiphaeidae showed slightly increased growth rates with increasing depth of distribution. Nektobenthic species of the genus Plesionika followed a trend opposite to that shown by mesopelagic species, with a higher growth rate for the shallowest-dwelling species (P. heterocarpus) than the deepest-dwelling species (P. acanthonotus). Taking growth as one of the major components of an organism's energy budget, the growth rates for the decapod crustacean species in this study were significantly lower than those reported in the literature for shallow-water penaeid crustacean species (which are distributed in higher-temperature habitats than deep-water Mediterranean crustaceans) and higher than those reported for mesopelagic myctophid fish species. Hence, the well-defined growth trends shown by deep-water decapod crustacean species in the Northwestern Mediterranean Sea, compared to the less well-defined trends in the other taxa, is discussed in the framework of the overall dynamics of their ecosystem. Received: 25 May 1998 / Accepted: 27 September 1999