Activity and ventilation-induced variations of postbranchial oxygen tensions in the isopod Saduria entomon

Variations in postbranchial oxygen tensions were studied in vivo in the brackish water isopod Saduria entomon collected from the Baltic Sea in 1992. Haemolymph oxygen tensions were highly influenced by activity. Resting isopods showed small and regular variations in haemolymph oxygen tensions, while a sudden burst of activity caused an immediate reduction in oxygen tension. Periods of no ventilation caused anoxic haemolymph in less than 3 to 4 min. Haemolymph oxygen tension responds rapidly, within seconds to minutes, to changes in isopod activity. Oxygen uptake rates calculated from oxygen tension difference, gill area and membrane thickness were compared with measured values. Permeability studies showed that only 30% of the gill area was effective in oxygen transfer. The present study has confirmed earlier measured oxygen uptake rates and reevaluated the role of the gills as respiratory organs. Postbranchial oxygen tension was established as a function of external steady state oxygen tensions and the intrinsic diffusive conductance was estimated.     

Effects of environmental factors and body size on rates of oxygen consumption in Archaeomysis grebnitzkii and Neomysis awatschensis (Crustacea: Mysidae)

The rates of oxygen consumption in relation to oxygen tension, temperature, salinity and body size were determined for Archaeomysis grebnitzkii Czerniavsky and Neomysis awatschensis (Brandt). Oxygen uptake was regulated by both species down to an oxygen tension of about 50 mm Hg (30% of saturation value); below this level, it was related to the oxygen tension in the medium. Oxygen tension below 20 mm Hg was lethal. A statistically significant interaction was noted for the effects of species, temperature, and salinity. In general, oxygen-uptake rate decreased with decreasing salinity in both species. The effect of size is expressed as a power function of body weight. The regression coefficients of oxygen consumption on body weight, 0.70 for A. grebnitzkii and 0.62 for N. awatschensis, were found to be significantly different from each other.Contribution No. 726 from the Department of Oceanography, University of Washington, Seattle, USA.     

Effect of temperature and food on the heart beat,ventilation rate and oxygen uptake of Mytilus edulis

The heart beat, ventilation rate and oxygen uptake of Mytilus edulis L. were measured simultaneously, in response to changes in temperature and food level. There was no thermal acclimation of heart-beat frequency or amplitude to temperatures from 5° to 25°C. Oxygen consumption and ventilation rate acclimated to 10°, 15° and 20°C, but not to 25°C. Starvation reduced the rate of oxygen uptake and heart-beat frequency to a standard level and, in response to food, the ventilation rate and oxygen consumption immediately increased to an active level. Feeding was maintained after the initiation of active metabolism, and during the following 10 days the heart-beat frequency gradually increased to the level characteristic of fed individuals. There was no direct correlation between the rate of oxygen consumption and heart rate, and an apparent absence of a close nervous coupling between ventilation rate and heart rate in M. edulis.     

Comparative study of the effects of temperature,salinity and oxygen tension on the rates of oxygen consumption of nauplii of different strains of Artemia

The effects of temperature, salinity and oxygen tension on the rates of oxygen consumption of three different strains of Artemia nauplii have been studied. When acclimated to a salinity of 30, nauplii from each of the three strains were able to maintain approximately constant rates of oxygen consumption over a wide range of oxygen tension. The ability to maintain respiratory independence during hypoxia was reduced, however, with an increase in either temperature or salinity. Nauplii of two of the strains (parthenogenetic diploid and tetraploid) showed a progressive increase in the rate of oxygen consumption with increasing temperature up to 35°C. Nauplii of the bisexual strain appeared to be less tolerant of exposure to temperatures >30°C, since at higher temperatures their oxygen consumption declined slightly. The differences between the nauplii of the different strains in their physiological responses to changing environmental conditions appear to correlate well with their seasonal occurrence in the field.     

Calorimetric studies on the energy metabolism of an infaunal bivalve,Abra tenuis,under normoxia,hypoxia and anoxia

Direct calorimetry was employed to measure the energy metabolism of infaunal bivalves, Abra tenuis, collected from a tidal lagoon in the Fleet, southern England, in June 1989, at various oxygen partial pressures. A significant anaerobic component (i.e., 20% of total metabolic rate) was detected under normoxia, presumably brought about by the intermittent ventilatory activity of this bivalve under these conditions. Under hypoxia (2.3 to 10 kPa, or 11 to 48% of full air saturation), however, the energy metabolism was maintained fully aerobic; the measured heat equivalent of oxygen uptake was not significantly different from the theoretical ranges for fully aerobic catabolism. Under anoxia, the rate of heat dissipation was reduced to 5–6% of the normoxic rate of heat dissipation. This conserves energy expenditure and would thus increase resistance of A. tenuis to anoxia or emersion. Physiological compensation by A. tenuis under conditions of declining oxygen tension involved a marked increase in ventilation rate. Comparison between fed and starved individuals indicated that costly physiological processes, such as digestion, absorption and growth declined at 10 and 5 kPa and were arrested at P_{O 2} (oxygen partial pressure) levels below 2.3 kPa. The present study provides evidence that there are no major differences between the metabolic responses of epifaunal suspension-feeding (eg. Mytilus edulis) and infaunal deposit-feeding (eg. A. tenuis) bivalves when exposed to environmental hypoxic stress.     

Detection and avoidance of hypoxic water by juvenile Callinectes sapidus and C. similis

Detection and avoidance of hypoxic water by juvenile blue crabs, Callinectes sapidus Rathbun and C. similis Williams were observed under laboratory conditions. Hypoxia avoidance was quantified within an avoidance chamber using response time (time in minutes before a crab's initial entry into the normoxic side of the avoidance chamber after the introduction of hypoxic water into the chamber), total time (amount of time spent by each crab at the hypoxic end of the chamber during the final 50 min of each 60 min treatment), activity (total distance travelled by a crab during a treatment), percent avoidance and preferred oxygen tension as indices of measurement. The mean preferred oxygen tension for C. sapidus was 112 torr oxygen (range = 98 to 125 torr) and for C. similis was 108 torr oxygen (range = 82 to 121 torr). Both species were able to detect hypoxic water and remain at an optimum oxygen tension. When percent avoidance, response time and total time were used as measures of avoidance, C. similis was found to detect and avoid hypoxic water at 0, 25 and 50 torr oxygen tension but not at any higher levels of hypoxia. C. sapidus, on the other hand, did not initiate any significant avoidance behavior even when the water was completely anoxic. Both species of crabs were more active at the higher oxygen tensions than in the more hypoxic levels. Behavioral responses associated with short-term hypoxic exposure included increased activity with the introduction of hypoxic water into the chamber, frequent movement of the eye-stalk, avoidance of hypoxic water by slowly crawling out of the hypoxic zone, restless and erratic movements, and rapid movements of the crabs' antennae. We conclude that C. similis can better detect and avoid hypoxia than C. sapidus.     

Respiratory impairment by water-borne copper and zinc in the edible crab cancer pagurus (L.) (Crustacea: Decapoda) during hypoxic exposure

Specimens of the edible crab Cancer pagurus (L.) collected in the Skagerrak, Denmark, between June and August 1990 were examined in the laboratory. Impairment of respiratory function after pre-exposure (7 d) to sub-lethal concentrations of Cu and Zn (0.4 mg l^-1) was only detectable during hypoxic exposure [PO₂ (partial pressure of oxygen) =60 torr]. This was indicated by a decrease in the transfer factor (TO₂), due principally to an increase in the PO₂ differential across the gills. Cu and Zn exposure did not cause significant changes in ventilation or perfusion although there was some indication that cardiac output may increase in respiratory-impaired individuals. After 28 d exposure no difference was noted in the respiratory responses to hypoxia of treated and untreated crabs. It is concluded that respiratory impairment was due to an increase in the diffusion barrier thickness at the gills and that this was reversible even during continued exposure to trace metal contamination.     

Adjustment of the components of energy balance in the gastropod Crepidula fornicata in response to thermal acclimation

Synchronous measurements were made of the routine rate of oxygen consumption and the clearance rate of Phaeodactylum tricornutum at different exposure temperatures by specimens of the suspension-feeding gastropod Crepidula fornicata which had been acclimated to temperatures between 10° and 25°C. The results show that the cost of activity (l O₂ consumed h^-1/ml seawater cleared h^-1) increases dramatically in individuals exposed to short-term increases of temperature up to 30°C, especially in limpets acclimated to 10°C. The process of thermal acclimation, however, results in two compensatory adjustments in energy expenditure and uptake which profoundly affect the energetics of water transport. Firstly, the routine oxygen consumption shows lateral translation of the rate-temperature curve which results in the maintenance of a relatively uniform energy expenditure despite an increase in acclimation temperature from 10° to 25°C. Secondly, because of the form of the rate-temperature curve for filtration by C. fornicata, lateral translation in response to warm acclimation results in an increase in the maximal clearance rate. Lateral translation of the rate-temperature curves for feeding rates and for oxygen consumption in response to thermal acclimation may thus be linked to maintain a balance between energy gain and expenditure. In this way, the greatly increased cost of activity which would occur with increase of temperature in the absence of acclimation is evaded. The minimal maintenance energy requirement, and hence the greatest scope for growth and reproduction, is then adjusted to coincide with temperatures prevailing in the environment.     

Oxygen consumption of the West African blood clamAnadara senilis

The West African blood clamAnadara senilis (L.) is one of a small number of bivalve species which have haemoglobin in their haemolymph. In the investigation reported here, in which the musselMytilus edulis (L.) was also studied for comparative purposes, it was shown thatA. senilis was an oxygen regulator between oxygen tensions corresponding to 50 and 100% air saturation. Oxygen uptake increased by about 128% after 24 h at zero pO₂; although the haemoglobins ofA. senilis were found to be responsible for about 34% of oxygen uptake at 25°C (a proportion unaffected by pO₂), there was clearly insufficient haemoglobin present in the haemolymph to act as an effective oxygen store during prolonged exposure to anoxic conditions.     

Effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immune condition of southern king crab Lithodes santolla (Molina, 1782) (Decapoda, Lithodidae)

Episodes of hypoxia are common in the marine environment, and their ecological effects depend, in part, on their severity and duration. Many species of decapod crustaceans reside in areas with fluctuating oxygen regimens. Physiological mechanisms enhance the ability of these crustaceans to cope with acute episodes of hypoxia. Southern king crab, Lithodes santolla, fishery is important in the south of South America, and some data describe fishing zones with low dissolved oxygen (DO) levels (3.5 mgO₂ l⁻¹, i.e., 8.3 kPa). Our main objective was to evaluate the effect of dissolved oxygen level on respiratory metabolism, nutritional physiology, and immunological condition of L. santolla juveniles. Individual animals were exposed for 10 days to different oxygen tensions (2.1, 4.2, 8.5, 12.7, and 21.1 kPa) to quantify the oxygen consumption rate; thereafter, blood oxyhemocyanin (Hc), protein concentration, as well as hemocytes, were sampled. Freeze-dried animals were dissected, and digestive gland metabolites (glycogen, protein, glucose, cholesterol, acylglycerol, and lactate) and digestive enzyme activity (general protease, trypsin, and chymotrypsin), as well as gill lactate dehydrogenase (LDH) activity, were quantified. In the present study, Lithodes santolla showed a critical oxygen tension between 4 and 9 kPa, indicating that this crab species is more sensitive to DO than other crustacean species. Protein and Hc concentrations followed a similar pattern to that of oxygen consumption. Digestive gland glycogen and protein concentration did not change after 10 days at different oxygen exposures, but glucose, cholesterol, and acylglycerol concentrations decreased linearly and proportionally to the available oxygen in the water. As in other decapods, chymotrypsin showed over 90% of the total quantified proteases activity. Chymotrypsin activity together with total proteases and trypsin was not affected by the environmental oxygen tension. Gill LDH and digestive gland lactate followed a similar increase at lower environmental oxygen tension but dropped sharply at the lowest tension (2.1 kPa). Dissolved oxygen affected also the immune system through reduction of hemocytes. This could provide a critical window for opportunistic pathogens to become established when crabs are exposed to hypoxic conditions. L. santolla juveniles show a moderate tolerance to low oxygen availability by modifying the concentration of hemolymph proteins, mainly OxyHc, some digestive gland metabolites, and by activating the anaerobic metabolism. This allows L. santolla juveniles to inhabit temporarily low oxygen zones in the deep ocean and suggests an advantage for culture conditions.     

Factors influencing egg size in the gammarid amphipod Gammarus insensibilis

Developmental and seasonal changes in egg volume were examined in a population of the amphipod Gammarus insensibilis Stock occurring on the south coast of England, towards its northern limit of distribution. Results showed a marked increase in egg volume during development (2.9 times by Egg Stage V), resulting from water uptake and from the conversion of yolk reserves into structural elements. The maximum rate of increase coincides with the period of organ and limb development. At hatching, after initial rupture of the egg membrane by urosome spines, egg volume increases rapidly over a short period (15 to 20 min) by a further 30% (uptake rate 3.6×10^–5 mm³s^–1), followed by a post-hatching decrease in juvenile volume. Increase in size at hatching is the result of drinking by embryos, although changes in body-wall permeability may contribute. Females carrying eggs in an advanced stage of development exhibit egg-collecting behaviour. This is seen as an adaptation to an increased likelihood of egg loss with increase in volume of the brood as hatching approaches. Seasonal changes in Stage I (early) egg size are marked in this species, with winter eggs as much as 60% greater in volume than summer eggs. Egg size is inversely related to the temperature during oocyte development. A simple model has been derived to account for the observed seasonal pattern in egg size. The consequences of seasonal variation in egg and juvenile size are considered.     

Oxygen uptake and heat production in a metabolic conformer (Littorina irrorata) and a metabolic regulator (Uca pugnax)

The heat production of Littorina irrorata and Uca pugnax in air was measured with a twin calorimeter while oxygen tension was measured with a pO₂ electrode. Both L. irrorata, an oxyconformer, and U. pugnax, a metabolic regulator, showed a rapid decrease in oxygen uptake (below 1.3 mm Hg in L. irrorata and 13.4 mm Hg in U. pugnax) while heat production decreased more slowly. Consequently, during the period of minimum oxygen uptake, the oxycalorific coefficient increased from about 4.8 for both species to an average value of 8.3 in L. irrorata and 19.9 cal ml^-1 O₂ in U. pugnax, indicating the onset of anaerobic metabolism and accumulation of metabolic and products. Above their respective critical pO₂, the oxycalorific coefficient was the same as the commonly used conversion factor of 4.8 cal ml^-1 O₂. From one time interval to the next, however, the coefficient varied from 3.8 to 5.4 in L. irrorata and from 2.9 to 6.0 in U. pugnax, indicating, that the processes of oxygen consumption and heat production are more or less independent of each other and usually not in phase.     

Effect of temperature on specific dynamic action in the common octopus,<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> (Cephalopoda)

Feeding causes an increase of metabolic rate, which initially escalates rapidly, reaches a peak value and then gradually declines to the pre-feeding rate. This phenomenon, termed specific dynamic action (SDA), reflects the energy requirements of the behavioral, physiological and biochemical processes that constitute feeding. The effect of temperature on SDA of the common octopus, Octopus vulgaris, was evaluated, by measuring the temporal pattern of the oxygen consumption rates of octopuses, after feeding, at two constant temperatures, 20°C and 28°C. At 20°C, the relative increase in the oxygen consumption rate after feeding (relative SDA) was significantly greater than at 28°C. The peak of the relative SDA occurred 1 h after feeding, and it was 64% at 20°C and 42% at 28°C. However, the SDA absolute peak, SDA duration (9.5 h) and SDA magnitude (the integrated postprandial increase in oxygen uptake) did not differ significantly between the two temperatures, indicating that the energetic cost of feeding was the same at both temperatures. The SDA response in O. vulgaris was much faster than it was in polar species, which have extended SDA responses due to low temperatures, and was also relatively fast in relation to the response in other temperate species, which is probably connected to the remarkably high growth rates of the species. A possible explanation of the observed summer migration of large octopuses from shallow to deeper areas is given, based on the effect of temperature on the energetic requirements of octopuses.     

Acute temperature and low dissolved oxygen tolerances of brachyuran crab (Cancer irroratus) larvae

Tolerance limits to acute temperature, and temperature and low dissolved oxygen stresses were determined for 5 zoeal stages and megalops of Cancer irroratus Say. The acute temperature limits for a 120 min exposure were all approximately 29.0°C, with little interstage, variation, while those for 240 min exposure ranged from 27.3° to 28.5°C. More interstage variation was shown when temperature and low dissolved oxygen stress were combined, with low oxygen tolerance decreasing as temperature increased. The first, second, and fourth zoeal stages display similar patterns of response. The third and fifth stages show a similar response, but are different from the former stages. The megalops is relatively insensitive to changes in oxygen concentration with temperature. The larval stages did not show a progressive increase in tolerance to temperature or low dissolved oxygen with development. Tolerance to these factors may be related to size, stage of development and activity level. The larval stages have capacities to tolerate a wider range of these factors than they require in the natural environment.     

Critical thermal maxima of two species of estuarine fish

Critical thermal maxima (CTM) were determined for winter flounder (Pseudopleuronectes americanus, Walbaum) and scup (Stenotomus chrysops, L.), acclimated to ambient water temperature or constant increments above ambient. The acclimation regimes simulated a natural environment by tracking the temperature fluctuations in Point Judith Pond, a small Rhode Island estuary. An elevation in CTM from 30.2° to 35.6°C was observed for scup as the mean acclimation temperature increased from 14.8° to 22.2°C. CTM for flounder increased from 26.4° to 32.4°C over an acclimation range of 4.2° to 23.0°C. The use of CTM studies for predicting the effects of heated effluents is discussed.     

Effect of low dissolved oxygen concentrations on the ventilation rhythm of three tubicolous crustaceans,with special reference to the phenomenon of intermittent ventilation

The pleopod ventilatory activity of the tube-dwelling amphipods Corophium volutator (Pallas) and C. arenarium Crawford is intermittent. Unlike the free-living amphipod Marinogammarus obtusatus (Dahl), which increases its rate of pleopod beating, the pleopod beat rate of both Corophium species is unaffected by low oxygen concentrations. However, the nature of the intermittent ventilation rhythm is altered so that, at low oxygen concentrations, the pleopods of Corophium beat almost continuously. It is suggested that the intermittent ventilation of Corophium can be interpreted as an adaptation to a tubicolous existence. The peracaridean Tanais chevreuxi Dollfus has 2 independent ventilatory systems, the maxilliped epipodites and the pleopods. The former, which respond to low oxygen concentrations, are thought to be the principal respiratory organs of T. chevreuxi, while the pleopods, which show no response to low oxygen tensions, are believed to be primarily concerned in filter feeding.     

Uptake of dissolved organics by marine bacteria as a function of fluid motion

A mass transfer analysis predicts that fluid motion can increase the assimilation of dissolved organics by attached compared to free-living microorganisms under certain conditions. To test this we examined the effect of advective flow and fluid shear on the uptake of model compounds (leucine and glucose) by natural assemblages of heterotrophic bacteria, collected from Roosevelt Inlet, Delaware Bay (USA), in 1989. We found that [³H]leucine uptake by cells held in fluid moving at 20 to 70 m d^–1 was eight times larger than uptake by cells at a velocity of 3 m d^–1. This effect was only observed at low leucine concentrations (ca. 1 nM), when uptake was likely not saturated. When we added leucine at concentrations expected to saturate leucine uptake (ca. 11 nM), fluid motion past cells did not affect uptake. Fluid flow past bacteria did not increase [³H]glucose uptake, and laminar shear rates of 0.5 to 2.1 s^–1 did not increase either glucose or leucine uptake by suspended bacteria. These results indicate that fluid motion increases bacterial uptake of certain lowmolecular-weight dissolved organics only when the microorganism exists in an advective flow field. As predicted from a mass transfer model, fluid shear rates in natural systems are too low to affect bacterial uptake of such compounds.     

Oxygen transfer in a high compacting multiphasic reactor (H.C.M.R.)

H.C.M.R. is a fluidised multiphasic reactor. In this reactor the rate of oxygen transfer was studied in order to characterise its efficiency as a system of aerobic treatment.

The studied parameters are: The size of the reactor, the volume of the particles, the oxygen transfer rate.

Several experiments were carried out with a variation of the above parameters and the results of the experiments are summarised as follows: The overall mass transfer rate coefficient k ₁ a increases with increasing of upflow gas rate (U_g ); The overall mass transfer rate coefficient k ₁ a decreases with increasing of the inert support volume (V_s); The overall mass transfer rate coefficient values from cylinder shaped particles were higher than those for disc shaped particles.     

Electrocardiogram of a marine fish,Pagrus major,exposed to red tide plankton,Chattonella marina

Examinations of the electrocardiogram of Pagrus major exposed to Chattonella marina, a planktonic organism causing red tide, were made to determine the physiological effects on fish. The heart rate decreased as a result of the extention in the interval between T and P waves. The decrease in heart rate with the extended intervals between T and P waves was also recognized in the condition of decreased dissolved oxygen. Also, the decrease in heart rate of the fish exposed to the red tide occurred while the fish was struggling. This reduction seems to be caused by the strong tension of the vagal nerve. Upon exposure to C. marina at high cell concentration, the heart beat at a very low frequency after 30 min. The very low heart rate is expected to limit seriously the oxygen uptake by the gill, because the cardiac output is probably very low in this situation.     

Histological study of the effects of starvation on reared and wild-caught larval stone flounder,Kareius bicoloratus

Starvation tolerance of laboratory-reared larval stone flounder, Kareius bicoloratus, was examined at different temperatures and salinities during the winters of 1984, 1985 and 1986. Starvation tolerance decreased with increased temperature and exhibited low values with high salinities. The highest starvation tolerance observed at low salinity was just before the metamorphosis stage. Starvation tolerance showed little change until larvae were 11 d. It increased with age thereafter. Epithelial cell heights of the digestive tract and cell diameters of pancreas and liver were measured histologically in reared stone flounder during growth and starvation. These values decreased markedly in the starved condition. Aldehyde fuchsin positive granules in the rectal epithelium also disappeared during the short starved period. The nutitrional condition of wild-caught stone flounder larvae, collected in January and February 1986 from the Matsukawa-ura inlet, Fukushima, Japan, was also examined. Eighty percent of larvae were estimated to be in fed condition just before sampling. The changes in cell heights of digestive organs agreed with this estimate. These histological methods seem to be useful in assessing the nutritional condition of marine fish larvae.