On-line determination of respiration rates of aquatic organisms in a mono-phase oxystat at steady-state dissolved oxygen tensions

On-line determination of respiration rates at steady-state dissolved oxygen tensions was performed in a thermostatted measuring chamber equipped with a galvanic oxygen electrode. The measuring chamber is designed to completely exclude gas bubbles from the liquid phase and is named a mono-phase oxystat. Additions of oxygenated water to maintain steady-state dissolved oxygen tensions are controlled by a computer, and the respiration rate is determined from the rate of these additions. An external loop with a flow cell in a spectrophotometer makes it possible to determine and control the concentration of algal cells in the mono-phase oxystat and allowed simultaneous determination of respiration and filtration rates of filter-feeding invertebrates. Because the system maintains steady-state conditions the experiments do not suffer from time limitations. The response time of the system is negligible compared to the response time of other open systems. Received: 7 November 1996 / Accepted: 6 December 1996     

Far-field vortex structures in mountain wakes

Laboratory experiments on scaled mountain wakes have provided the first demonstration that indicates the existence of two distinct mechanisms for the formation of the long-lived vortex structures that have been frequently observed in satellite imagery. In a series of laboratory experiments the well-known von Kármán vortex shedding mechanism has been observed at low values of Froude number (corresponding to low speed flow and strong stratification). At high Froude numbers the initial highly turbulent wake evolves into a vortex pattern similar to the far-wake dipole eddies previously observed in laboratory studies of submerged stratified wakes. In satellite imagery vortex structures have been observed to persist for hundreds of kilometers. In the present laboratory experiments far-wake dipole eddies were observed to persist for at least 200 diameters downstream, which corresponds to ~4,000?km when scaled to actual mountains. When generated at multiple sites along a continental mountain chain, these eddy structures could play a significant role in the development and variability of multi-scale weather and climate patterns.     

Metabolism of shallow and deep-sea benthic crustaceans and echinoderms in Hawaii

Little is known about the metabolism of deep-living, benthic invertebrates, despite its importance in estimating energy flow through individuals and populations. To evaluate the effects of depth and broad taxonomic group/locomotory mode, we measured the respiration rates of 25 species of benthic decapod crustaceans and 18 species of echinoderms from the littoral zone to the deep slope of Hawaii. Specimens were collected by hand, trap, or submersible and maintained in the laboratory at temperatures close to ambient temperatures recorded at the time of collection. After acclimatization to laboratory conditions, oxygen consumption was measured for each individual in closed chambers. Overall, crustaceans had higher metabolic rates than echinoderms, and within the crustaceans, caridean shrimps had higher rates than crabs and lobsters. These differences are probably related to locomotory mode and general levels of activity. At in situ environmental temperatures, metabolic rates of deeper-living invertebrates are much lower than those of shallower living species, but this decline is explained by changes in temperature. When the data were compared with similar data sets collected off California and in the Mediterranean, Hawaiian crabs, lobsters, and echinoderms had lower metabolic rates than similar species in the other regions after adjustments for temperature were made. Some of these differences could be methodological. Regional food web models should use broad taxonomic groupings and region-specific data when possible.     

Aquatic metabolism and ecosystem health assessment using dissolved O2 stable isotope diel curves.

Dissolved O2 concentration and delta18O-O2 diel curves can be combined to assess aquatic photosynthesis, respiration, and metabolic balance, and to disentangle some of the confounding factors associated with interpretation of traditional O2 concentration curves. A dynamic model is used to illustrate how six key environmental and biological parameters interact to affect diel O2 saturation and delta18O-O2 curves, thereby providing a fundamental framework for the use of delta18O-O2 in ecosystem productivity studies. delta18O-O2 provides information unavailable from concentration alone because delta18O-O2 and saturation curves are not symmetrical and can be used to constrain gas exchange and isotopic fractionation by eliminating many common assumptions. Changes in key parameters affect diel O2 saturation and delta18O-O2 curves as follows: (1) an increase in primary production and respiration rates increases the diel range of O2 saturation and delta18O-O2 and decreases the mean delta18O-O2 value; (2) a decrease in the primary production to respiration ratio (P:R) decreases the level of O2 saturation and increases the delta18O-O2 values; (3) an increase in the gas exchange rate decreases the diel range of O2 saturation and delta18O-O2 values and moves the mean O2 saturation and delta18O-O2 values toward atmospheric equilibrium; (4) a decrease in strength of the respiratory isotopic fractionation (alphaR closer to 1) has no effect on O2 saturation and decreases the delta18O-O2 values; (5) an increase in the delta18O of water has no effect on O2 saturation and increases the minimum (daytime) delta18O-O2 value; and (6) an increase in temperature reduces O2 solubility and thus increases the diel range of O2 saturation and delta18O-O2 values. Understanding the interplay between these key parameters makes it easier to decipher the controls on O2 and delta18O-O2, compare aquatic ecosystems, and make quantitative estimates of ecosystem metabolism. The photosynthesis to respiration to gas exchange ratio (P:R:G) is better than the P:R ratio at describing and assessing the vulnerability of aquatic ecosystems under various environmental stressors by providing better constrained estimates of ecosystem metabolism and gas exchange.     

Diel periodicity in cellular chlorophyll content in marine diatoms

Intracellular chlorophyll a content is one of the many measurable parameters which displays a diel rhythm in marine phytoplankton. In asynchronous laboratory cultures of the diatom Skeletonema costatum, cellular chlorophylls a and c exhibit periodicities which closely follow the light-dark cycle and are not the result of cell division. The laboratory cultures also exhibit diel rhythms in cellular flourescence properties and carbon: chlorophyll a ratios. The occurrence of similar patterns of cellular flourescence, carbon: chlorophyll a ratios, and in situ flourescence in diatom-dominated natural phytoplankton communities suggests the possibility of diel rhythms in cellular chlorophyl a content in diatoms in the sea. The data also suggest that the observed periodicity in cellular chlorophyll content is regulated by the diel light cycle and that the co-occurrence of synchronous or phased cell division would only modify the observed periodicity.This research was performed under the auspices of the United States Department of Energy under Contract No. EY-76-C-02-0016     

Physiological basis of extreme growth rate differences in the spat of oyster (Crassostrea gigas)

Juvenile oysters (Crassostrea gigas) (produced in November 2009) reared under uniform hatchery conditions for 4 months were selected for extreme growth rate differences by repeatedly taking larger and smaller individuals to achieve weight differences >30× between fast (F) and slow (S) growers. The physiological basis of differential growth was analyzed in experiments in June 2010, where components of energy gain (clearance and ingestion rates and absorption efficiency), energy loss (metabolic rates) and resulting scope for growth (J h^?1) were compared for groups of F and S oysters fed three different ration levels (≈0.5, 1.5 and 3.0 mg of total particulate matter L^?1). In both F and S oysters, a higher food ration promoted asymptotic increases in energy gain rates through regulatory adjustments to clearance rates, which maintained similar absorption efficiencies across the food concentrations. No significant differences were found between growth groups in mass-specific physiological rates (i.e., per unit of body mass). However, the scaling of these rates to a common size in both groups using allometric coefficients derived for C. gigas revealed higher energy gain rates coupled with lower metabolic costs of growth in fast growers. Thus, appropriate size-standardization is essential in accounting for observed differences in growth rate. Present results are in accordance with previous reports on other bivalve species on the physiological processes underlying endogenous growth differences, suggesting that the same interpretation can be applied to the extremes of these differences.     

Dynamic fate modeling of γ-hexachlorocyclohexane in the lower reaches of the Liao River

A QWASI model dependent on temperature is parameterized to describe the long-term fate of persistent organic pollutants (POPs) in the Liao River. The model parameters, namely fugacity capacity, degradation rate, and transfer coefficient, are profoundly affected by temperature. This model is used to simulate the fate of γ-hexachlorocyclohexane (γ-HCH) in the lower reaches of the Liao River from 1998 to 2008. Modeling results show that γ-HCH fugacity capacities in air, water, and sediment increase as temperature decreases, and the transfer and transformation rate coefficients increase as temperature increases. The variations of transfer and transformation parameter D values depend on fugacity capacities, and transfer and transformation coefficients simultaneously. The performance of the model is evaluated by comparing the predicted and observed concentrations in the water and sediment of the Liao River. The predicted values agree well with the observed value in the order of magnitude, in most cases within the factor of 3. It is believed that the model is appropriate for simulating the long term fate of POPs in the Liao River.     

Daily photosynthesis,respiration, and carbon budgets in a tropical marine jellyfish (Mastigias sp.)

From measured diel photosynthesis and respiration rates, using oxygen electrodes, estimates of carbon flux between symbiotic algae (zooxanthellae) and host animal are presented for the marine scyphomedusan Mastigias sp. from a marine lake in Palau, Western Caroline Islands, during February and March 1982. The carbon budgets calculated for these lake medusae indicate that carbon fixed photosynthetically by zooxanthellae and made available to the host may satisfy up to 100% of the host's daily metabolic carbon demand (CZAR). The stable carbon isotope (¹³C) signature of the mesogleal carbon of lake Mastigias sp. was close to that of the zooxanthellae, supporting the interpretation that while these medusae may feed holozoically, some of their carbon comes from their symbionts. The diel photosynthesis, respiration, and preliminary estimates of carbon budgets of three individuals of another ecotype of Mastigias sp. collected from nearby oceanic lagoons are also given. Photosynthesis of lagoon medusae was generally greater than that for lake medusae of similar size, and lagoon medusae were phototrophic with respect to carbon, with commensurately greater CZAR values. Carbon translocated from the symbiotic algae also may contribute to the growth requirements of both lake and lagoon medusae. From carbon flux data, the lake jellyfish were estimated to contribute about 16% to the total primary productivity of their marine lake habitat.     

An energy budget for Porites porites (Scleractinia)

An energy budget for Porites porites (Pallas) was determined for specimens from 10 m depth on the Fore Reef of Discovery Bay, Jamaica, between July 1984 and July 1985. Evidence for habitual zooplankton ingestion was not obtained, and P. porites appears to be largely autotrophic. Out of the daily photosynthetically fixed energy, 26% is used for animal respiration and growth, 22% for zooxanthellae respiration and growth, and <1% for colony reproduction as mature planulae; 45% remains unaccounted for. Colony respiration, net photosynthesis, colony skeleton and tissue growth, zooplankton ingestion, reproductive effort and energy content of tissues were measured. Energy loss as continuous mucus secretion was not detected, but may occur by an alternative route via mucus tunics, which occur periodically in situ and in the laboratory. The energy budget suggests that a considerable excess of photosynthetically fixed energy is produced on an ideal sunny day at 10 m depth. This surplus may be required for periodic rather than continuous energy demands, or may be essential to survive less-than-ideal days, when net photosynthetic input is reduced.Contribution No. 357 of the Discovery Bay Marine Laboratory, University of the West Indies     

The influence of fresh weight and water temperature on metabolic rates and the energy budget of Meretrix meretrix Linnaeus

Meretrix meretrix L. was held in the laboratory under simulated natural conditions to measure specific physiological parameters of its energy budget. O₂ consumption rate, NH₃ excretion rate (NR), ingestion rate, faeces excretion rate and scope for growth (SFG) were negatively related in an exponential manner to the fresh weight of the clams at all water temperatures, while almost all metabolic rates of the clams were positively related in an exponential or e-exponential manner with water temperature. However, the co-relationship between metabolic rates and water temperature was not as close as that between metabolic rates and fresh weight of the clam. The combined effect of fresh weight and water temperature was observed on all metabolic rates except for NR and SFG. At all culture temperatures and for all fresh weights of clams used, respiration took the largest percentage of ingested energy (41.5–51.2%), faeces excretion was second (31.0–42.3%), growth third (12.1–15.5%) and urine production last (2.1–5.6%).     

Physiological ecology of the clonal corallimorpharian <Emphasis Type="Italic">Corynactis californica</Emphasis>

For clonal taxa, the reduced genetic variability associated with clonal proliferation is hypothesized to reduce the ability to respond to variable conditions, unless a general-purpose genotype (GPG) confers success in multiple environments. In this study, Corynactis californica (Carlgren 1936) from the subtidal of California was used as a model system to test the hypothesis that clones dampen fluctuations in fitness through a GPG that facilitates phenotypic plasticity. To achieve this goal, tissue composition, respiration, excretion, and growth were compared among clones of C. californica at one site, and a reciprocal transplant experiment was used to test the response of clones to differing conditions at two sites. All experiments were completed at Santa Catalina Island (N 33°25′, W 118°30′) between April and September 1991. Clones at a single site differed significantly in multiple traits, varying as much as 1.6-fold in protein content, 3.4-fold in respiration, and 3.5-fold in excretion. Interestingly, while tissue growth was the most labile trait (differing up to 35.4-fold among clones), polyp fission rates were not significantly different among clones, in part because fission continued even though tissue growth was unable to restore polyp size in between divisions. Partial energy budgets revealed that the majority (47–90%) of the daily energy expenditure was accounted for by respiration, 13–47% by growth, and 0.3–14% by excretion. In the transplant experiment, reaction norms revealed strong effects of the environment on some traits but not others, notably with growth differing between sites in a pattern that differed among clones, and excretion differing between sites; neither respiration nor fission were affected by transplantation. Partial energy budgets revealed that the energy allocation to respiration varied between sites in a pattern that differed among clones, and a similar trend was evident for tissue growth. Together, these results demonstrate that clones of C. californica have markedly different phenotypes and exploit phenotypic plasticity to maintain relatively constant fission rates, even though tissue growth varies greatly among clones and between environments. While these findings support the GPG hypothesis for clones of C. californica—at least based on relative fitness achieved through asexual proliferation—this conclusion depends on the extent to which polyps are successful when they have low rates of tissue growth.     

In situ thermal dynamics of shallow water corals is affected by tidal patterns and irradiance

We studied the diel variation of in situ coral temperature, irradiance and photosynthetic performance of hemispherical colonies of Porites lobata and branching colonies of Porites cylindrica during different bulk water temperature and tidal scenarios on the shallow reef flat of Heron Island, Great Barrier Reef, Australia. Our study presents in situ evidence that coral tissue surface temperatures can exceed that of the surrounding water under environmental conditions typically occurring during low tide in shallow reef or lagoon environments. Such heating may be a regular occurrence on shallow reef flats, triggered by the combined effects of high irradiance and low water flow characteristic of low Spring tides. At these times, solar heating of corals coincides with times of maximum water temperature and high irradiance, where the slow flow and consequent thick boundary layers impede heat exchange between corals and the surrounding water. Despite similar light-absorbing properties, the heating effect was more pronounced for the hemispherical P. lobata than for the branching P. cylindrica. This is consistent with previous laboratory experiments showing the evidence of interspecific variation in coral thermal environment and may result from morphologically influenced variation in convective heat transfer and/or thermal properties of the skeleton. Maximum coral surface warming did not coincide with maximum irradiance, but with maximum water temperature, well into the low-tide period with extremely low water flow in the partially drained reef flat, just prior to flushing by the rising tide. The timing of low tide thus influences the thermal exposure and photophysiological performance of corals, and the timing of tidally driven coral surface warming could potentially have different physiological impacts in the morning or in the afternoon.     

Effect of elevated temperature on aerobic respiration of coral recruits

Metabolic rates provide a valuable means to assess the condition of early life stages of scleractinians, but their small biomass creates a signal-to-noise problem in a confined respirometer. To avoid this problem, measurements of the oxygen diffusion boundary layer (DBL) and Ficks first law were used to calculate the respiration rate of coenosarc tissue on recruits (i.e., colonies 5–14 mm diameter) of Porites lutea (Edwards and Haime, 1860) exposed to two temperatures at a flow speed of 0.6 cm s^–1. All experiments were completed in Moorea, French Polynesia, between November and December 2003. At 26.8°C, the DBL was 565±55 µm thick, the oxygen saturation adjacent to the tissue was 80±3%, and the mean respiration of the coenosarc was 1.2±0.1 µl O₂ cm^–2 h^–1 (all values mean ± SE, n=10). Exposure to 29.7°C for 24–48 h did not affect the DBL thickness but significantly reduced the oxygen saturation adjacent to the tissue (to 74%) and increased the mean respiration rate by 35%. As the small corals differed slightly in size, in a uniform flow speed they experienced dissimilar flow environments as characterized by the Reynolds number (Re), thereby creating the opportunity to test the flow dependency of respiration. At 26.8°C, respiration and Re were unrelated, but at 29.7°C, the relationship was positive and statistically significant. Thus, respiration of small corals may not be mass transfer limited at low temperature, but relatively small increases in temperature may result in an increased metabolic rate leading to mass transfer limitation and flow-dependent rates of respiration.Communicated by J.P. Grassle, New Brunswick     

Latitudinal patterns of shell thickness and metabolism in the eastern oyster Crassostrea virginica along the east coast of North America

The goal of this study was to quantify growth and metabolic responses of oysters to increased temperatures like those that will occur due to global warming. Impact of temperature on eastern oyster (Crassostrea virginica) shell growth and metabolism was investigated by sampling 24 sites along the eastern North American seaboard ranging from New Brunswick, Canada, to Florida, USA, in March and August 2013. There was a positive correlation between oyster shell thickness and site temperature. At southern sites, shells were up to 65 % thicker than at the northernmost site, likely due to higher precipitation of CaCO₃ in warmer water. This was supported by laboratory experiments showing that thicker shells were produced in response to temperatures 2, 4, and 6 °C above ambient seawater temperatures (8–14 °C) in Connecticut, USA. Field experiments with oyster respiration were conducted during winter and summer at 13 sites to compare responses to thermal stress with latitude. Respiration rates were much higher during summer than winter, but the combination of summer and winter data fell along the same exponential curve with respect to temperature. At all sites, temperature-specific metabolic rates at elevated temperatures were lower than predicted, indicating significant seasonal acclimatization by C. virginica.     

Nutrient transfer in a marine mutualism: patterns of ammonia excretion by anemonefish and uptake by giant sea anemones

Many symbioses involve multiple partners in complex, multi-level associations, yet little is known concerning patterns of nutrient transfer in multi-level marine mutualisms. We used the anemonefish symbiosis as a model system to create a balance sheet for nitrogen production and transfer within a three-way symbiotic system. We quantified diel patterns in excretion of ammonia by anemonefish and subsequent absorption by host sea anemones and zooxanthellae under laboratory conditions. Rates of ammonia excretion by the anemonefish Amphiprion bicinctus varied from a high of 1.84 μmole g⁻¹ h⁻¹ at 2 h after feeding, to a basal rate of 0.50 μmole g⁻¹ h⁻¹ at 24–36 h since the last meal. Conversely, host sea anemones Entacmaea quadricolor absorbed ammonia at a rate of 0.10 μmole g⁻¹ h⁻¹ during the daytime in ammonia-enriched seawater, but during the night reduced their absorption rate to near zero, indicating that ammonia uptake was driven by zooxanthella photosynthesis. When incubated together, net ammonia excretion was virturally zero, indicating that host anemones absorbed most of the ammonia produced by resident fish. Adult anemonefish weighed about 11 g under laboratory conditions, but on the coral reef may reach up to 64 g, resulting in a maximal potential ammonia load of >200 μmole h⁻¹ produced by two adult fish during daylight hours. In contrast, host sea anemones weighed about 47 g in the laboratory, but under field conditions, large individuals may reach 680 g, so their maximal ammonia clearance rates may reach about 70 μmole h⁻¹ during the daytime. As such, the ammonia load produced by adult anemonefish far exceeds the clearance rate of host anemones and zooxanthellae. Ammonia transfer likely occurs mainly during the daytime, when anemonefish consume zooplankton and excrete rapidly, and in turn the zooxanthellae are photosynthetically active and drive rapid ammonia uptake. We conclude that zooplanktivorous fishes that form mutualisms with coral reef cnidarians may serve as an important link between open water and benthic ecosystems, through the transfer of large quantities of nutrients to zooxanthellate hosts, thus enhancing coral reef productivity.     

Respiration and nitrogen excretion of zooplankton. IV. The influence of starvation on the metabolism and the biochemical composition of some species

Changes in the respiration, ammonia excretion and biochemical composition were studied for three species of starving zooplankton (Calanus finmarchicus, Sagitta elegans, and Acartia clausi). Over the period of starvation, the respiration rate of all three species followed the same pattern of an initial decrease followed by a more or less constant level. A similar pattern was observed for the ammonia excretion rate of S. elegans and A. clausi, whereas C. finmarchicus excretion appeared to oscillate between high and low levels of protein catabolism. Study of the biochemical changes showed that C. finmarchicus consumed primarily lipids, and at times proteins, to meet its energy requirement whereas S. elegans and A. clausi primarily used protein. Variations in the elemental composition as well as the O:N ratio confirmed that C. finmarchicus alternated between periods of protein-dominant catabolism and lipid-dominant catabolism during starvation. No similar change in catabolism was observed in the two other species. The results are discussed in terms of physiological mechanisms of resistance to starvation and were used to calculate the energy budget of S. elegans and C. finmarchicus during the period of total starvation. The significance of such budgets is discussed and some of the sources of error examined.Bedford Institute of Oceanography Contribution.     

Turbulence and waves inside flexible-wall systems designed for biological studies

Simultaneous flow measurements were performed by hot-wire anemometry inside and outside (a) a closed bag, and (b) a flow-through system designed for metabolic experiments of Fucus vesiculosus communities. Since water movement is considered an important parameter in such biological studies, the walls of the systems were flexible in order to establish flow conditions within the enclosed water body comparable to those in the natural environment. In situ experiments in the Baltic Sea at a water depth of 2 m showed that energy spectra inside the systems were comparable to those outside for a variety of flow and wave conditions. Thus, biological data from glexible wall systems, carefully designed to meet specific natural flow requirements, can be taken as reliable input data for natural ecosystem modeling.Contribution No. 147 of the Joint Research Programme 95 Interaction Sea-Sea Bottom, Kiel University, Kiel (FRG), and Hawaii Institute of Geophysics Contribution No. 803.     

Algal symbiosis: A mathematical analysis

Host and algal symbion growth can be described by an iterative model which incorporates utilization efficiencies of host and symbiont. This model predicts that, with input of organic matter to the host and at very low host and algal utilization efficiences coupled with efficient recycling of nutrients between the host and symbionts, production of organic matter by the system can be increased by 2–3 orders of magnitude over that of a system comprised of only autotrophs and heterotrophs. Energy available for growth and respiration by the host is 1–2 orders of magnitude over that available to a heterotroph without symbionts. Algal symbiosis is highly advantageous in oligotrophic environments where radiant energy is abundant, growth-limiting nutrients are scarce and only concentrated in organic matter, and much energy must be expended to capture that organic matter.     

Scavenging deep-sea amphipods: Effects of food odor on oxygen consumption and a proposed metabolic strategy

In situ respiration rates as a response to the odor of food were measured for two species of scavenging amphipods, Paralicella caperesca from 3 650 m in the western North Atlantic Ocean and Orchomene sp. B from 1 300 m in the Santa Catalina Basin off southern California (USA). In addition, complementary laboratory starvation/respiration rates for a shallow-water species, Orchomene sp. A, were determined. Initial elevated O₂ consumption rates were found for up to 8 h in all deep-sea amphipods exposed to bait odor, followed by a period of lowered respiration equivalent to rates observed in individuals not exposed to bait. Orchomene sp. A revealed a response similar to that observed in the deep-sea species. A metabolic strategy is proposed whereby scavenging amphipods efficiently utilize large episodic organic falls in the food-limited environment of the deep sea. This strategy involves (1) the ability to withstand long periods of starvation, (2) rapid response to an organic fall, (3) rapid location of the organic fall, (4) maximal rate of food consumption with maximal quantity ingested, and (5) efficient utilization of the consumed food. Each of these attributes are explored with the expected and observed mechanisms employed to achieve them.     

Oxygen microoptodes: a new tool for oxygen measurements in aquatic animal ecology

We describe two applications of a recently introduced system for very precise, continuous measurement of water oxygen saturation. Oxygen microoptodes (based on the dynamic fluorescence quenching principle) with a tip diameter of ~50 µm, an eight-channel optode array, an intermittent flow system, and online data registration were used to perform two types of experiments. The metabolic activity of Antarctic invertebrates (sponges and scallops) was estimated in respiration experiments, and, secondly, oxygen saturation inside living sponge tissue was determined in different flow regimes. Even in long-term experiments (several days) no drift was detectable in between calibrations. Data obtained were in excellent correspondence with control measurements performed with a modified Winkler method. Antarctic invertebrates in our study showed low oxygen consumption rates, ranging from 0.03-0.19 cm³ O₂ h^-1 ind.^-1. Oxygen saturation inside living sponge specimens was affected by flow regime and culturing conditions of sponges. Our results suggest that oxygen optodes are a reliable tool for oxygen measurements beyond the methodological limits of traditional methods.