Oxygen and acid-base status of the sea urchin Psammechinus miliaris during environmental hypoxia

The respiratory and acid-base responses to hypoxia of the sea urchin Psammechinus miliaris Gmelin have been studied both in the laboratory and in the field. Sea urchins were collected from the Clyde Sea area, Scotland, between March and July 1993. Individual urchins were unable to regulate their oxygen uptake during hypoxia. The partial pressure of oxygen (P_{O 2}) in the coelomic fluid was lower than ambient P_{O 2} during normoxia but still decreased with a decrease in environmental P_{O 2}. There was a significant increase in pH and in the concentrations of bicarbonate, magnesium and calcium in the coelomic fluid, but a significant decrease in the partial pressure of carbon dioxide (P_{CO 2}), in response to declining P_{O 2}s. The disturbance of respiratory parameters in urchins accompanying hypoxic exposure in rock pools was similar to that observed in the laboratory, although not as severe as might have been expected on the basis of the laboratory experiments.     

The hypoxia avoidance behaviour of juvenile Atlantic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis> L.) depends on the provision and pressure level of an O<Subscript>2</Subscript> refuge

The frequency of low O₂ (hypoxia) has increased in coastal marine areas but how fish avoid deleterious water masses is not yet clear. To assess whether the presence and oxygen pressure (PO₂) level of an O₂ refuge affects the hypoxia avoidance behaviour of fish, individual Atlantic cod (Gadus morhua L.) were exposed to a range of O₂ choices in a 2-way choice chamber at 11.4°C over two different experiments. Cod in the first experiment were allowed access to a fixed O₂ refuge (fully air-saturated seawater) whilst oxygen pressure (PO₂) on the other side was reduced in steps to a critically low level, i.e. 4.3 kPa—a point where cod can no longer regulate O₂ consumption. Under these conditions, cod did not avoid any level of hypoxia and fish swimming speed also remained unchanged. In contrast, strong avoidance reactions were exhibited in a second experiment when fish were again exposed to 4.3 kPa but the safety, i.e. PO₂, of the refuge was reduced. Fish not only spent less time at 4.3 kPa as a result of fewer sampling visits but they also swam at considerably slower speeds. The presence of an avoidance response was thus strongly related to refuge PO₂ and it is unlikely that cod, and possibly other fish species, would enter low O₂ to feed in the wild if a sufficiently safe O₂ refuge was not available. It is therefore hypothesized that the feeding range of fish may be heavily compressed if hypoxia expands and intensifies in future years.     

Respiration of Sepia officinalis during embryonic and early juvenile life

Adult Sepia officinalis L. were caught in June 1984, in the coastal waters of Wimereux (France). Deposition of the eggs took place in the seawater aquaria of the Station Marine. The oxygen consumption of S. officinalis was measured during embryonic and juvenile development. Aerobic metabolism occurs as soon as the early embryonic Stage 21. Oxygen diffuses through the initially thick egg shell; the oxygen level in the perivitelline liquid reaches a maximal value just before hatching (116.7±6.9 mm Hg). Hatchings display only a slight increase in oxygen consumption compared to embryos in the last stage of development. Respiration experiments with 40 d old juveniles showed that oxygen consumption increases with temperature, but is not affected by photoperiod. Experiments under increasing hypoxia revealed that S. officinalis juveniles are good regulators and maintain a constant oxygen consumption in the range of 4 to 7 mg O₂l^-1. Juveniles successfully recover from an hypoxic stress of 2 mg O₂l^-1 maintained for 1 h. This suggests that the respiratory pigments (pre-hemocyanins) of 40 d-old juveniles have a high oxygen affinity and/or that these juveniles have the ability to adapt to anaerobic conditions.     

Effects of hypoxia on the respiratory and circulatory regulation of Nephrops norvegicus

The burrowing decapod Nephrops norvegicus (L.) was kept under various degrees of hypoxia in order to measure respiration, heart rate, scaphognathite rate, haemolymph oxygen content and pH. An emergence reaction to hypoxia occurred only in dim light (<10^-2 m-c) or darkness, but after 10 d of moderate hypoxia the decapods showed no emergence response at all. The weight specific respiration of quiescent individuals was relatively low and increased only slightly in hypoxia (P_wO₂=40 torr). Heart rate, about 50 beats min^-1, changed little during hypoxia, down to P_wO₂=40 torr, whereas scaphognathite rates rose from about 60 beats min^-1 at normoxia to peak at 120 beats min^-1 at P_wO₂=40 torr. The oxygen extraction efficiency (E) remained at 20 to 30% during the first hour of hypoxia then rose gradually to maximum values of 30 to 40%. A small respiratory alkalosis of the blood became evident only after 4h of hypoxia (P_wO₂=50 torr). Normoxic postbranchial O₂ tensions (P_aO₂) were low (25–30 torr) and showed only a small decline during hypoxia. Over 10 to 13 d in moderate hypoxia an effective biosynthesis of 0.024 mM haemocyanin individual^-1 d^-1 occurred in fed decapods, whereas controls (normoxic) showed no significant change in pigment levels. A linear relationship between oxygen carrying capacity and haemocyanin concentration was found. It is contended that N. norvegicus is better able to cope with periodic exposure to hypoxia when food of sufficient quantity and quality is available.     

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.     

Effect of salinity on hypoxia tolerance of resting green crabs, Carcinus maenas, after feeding

Mechanisms that can influence the tolerance of hypoxia in brackish waters were studied in resting and fed crabs, Carcinus maenas, at 15?°C. Mortality, blood oxygenation, acid-base status and lactate concentration were analysed in fed crabs held in full-strength normoxic seawater (32.5‰?S) and then transferred for 24?h to a partial pressure of oxygen (P_o2) of 3?kPa (1.4?mg?l^?1) and various salinities (17, 12.5, 10, 8‰?S). At salinity levels >10‰, fed crabs tolerated P_o2 values as low as 3?kPa in the ambient water and 0.5?kPa in their arterial blood for 24?h without switching to anaerobic metabolism. Only below 10‰?S did their blood-lactate content rise, leading to their death despite the fact that their blood O₂-content was twice the control value measured in full-strength normoxic seawater and their blood P_o2 did not decrease below values recorded at higher salinity levels. Addition of CO₂ to 8‰?S water (CO₂ partial pressure increasing from 0.1 to 0.3?kPa) decreased blood-lactate production and mortality, suggesting that at 10‰?S impairment of the O₂ supply is limited by an excessive blood O₂-affinity. The results are discussed in terms of the distribution (?10‰?S) of C. maenas along salinity gradients in estuaries and bays.     

Comparative studies on the metabolism of shallow-water and deep-sea marine fishes. IV. Patterns of aerobic metabolism in the mesopelagic deep-sea fangtooth fish Anoplogaster cornuta

Measurements have been made of oxygen consumption rates O₂ of 10 specimens of the mesopelagic deep-sea fangtooth fish Anoplogaster cornuta. Determinations were made at 1 atm pressure, at temperatures of 3°, 7°, and 10°C, at dissolved oxygen concentrations ranging from near saturation to zero, with the fish swimming at low, controlled speeds. Weight-specific O₂ were uniformly low. They showed Q₁₀'s of 2.5 and 1.3, respectively, in the temperature ranges 3° to 7°C, and 7° to 10°C, at dissolved oxygen concentrations above 2 ml (standard temperature and presusure, STP)/1. Measurable O₂ continued in these fish at dissolved oxygen concentrations down to the lowest levels detectable with our instruments. At 7°C the average critical oxygen tension (P _c ) for the entire group was near 35 mm Hg. However, there is a statistically significant positive slope to the regression line relating O₂ to P _c for individual fish. The physiological and ecological significance of these results is discussed, particularly with reference to thermal effects and to the basis for survival by A. cornuta in the oxygen minimum layers of the eastern Pacific Ocean.     

Comparative studies on the metabolism of shallow-water and deep-sea marine fishes. V. Effects of temperature and hydrostatic pressure on oxygen consumption in the mesopelagic zoarcid Melanostigma pammelas

Measurements have been made of routine oxygen consumption rates ( ) of the mesopelagic deep-sea zoarcid fish Melanostigma pammelas. Determinations were made over ecologically relevant ranges of 3 variables; temperature (3° to 10°C), hydrostatic pressure (1 to 170 atm), and oxygen partial pressure (1 to 160 mm Hg). Weight-specific s were uniformly low. Of the 3 test variables, only temperature had significant metabolic effects within the ranges studied. Q₁₀'s were 6.75 between 3° and 5°C. 1.47 between 5° and 7°C, and 17.4 between 7° and 10°C. These Q₁₀'s were constant over the hydrostatic pressure range studied. Between 3° and 7°C the fish regulated their rates of oxygen consumption down to PO₂'s comparable to those occurring in their natural environments (6 to 12 mm Hg). The showed no capacity to tolerate anoxic conditions. The physiological and ecological significance of these results is discussed, particularly with reference to thermal effects and to the basis of survival of this fish in the oxygen minimum layers of the eastern Pacific Ocean. Since it is possible to maintain M. pammelas in the laboratory for extended periods of time (over 12 months) it could serve as the basis for many interesting studies of deep-sea fish biology.     

Does oxygen deficiency modify the functional response of Saduria entomon (Isopoda) to Bathyporeia pilosa (Amphipoda)?

The functional response of the predatory isopod Saduria entomon to the prey amphipod Bathyporeia pilosa was measured in normoxia (95% O₂ saturation), moderate hypoxia (45% O₂ saturation) and hypoxia (35% O₂ saturation) in aquarium experiments. The prey densities tested ranged from 400 to 8000 ind m⁻². Prey density influenced consumption rates of S. entomon in normoxia and 45% O₂ saturation, but there was no difference between consumption rates at these two oxygen levels. Nevertheless the form of functional response differed. In normoxia S. entomon showed a positively density-dependent functional response to B. pilosa, indicating a potentially stabilizing effect on the prey population. In moderate hypoxia the variance in consumption increased, decreasing the statistical power to distinguish between response models. The functional response of S. entomon in moderate hypoxia was best described with a density-independent response, characterized as destabilizing for the prey population. In hypoxia (35% O₂) predation by S. entomon did not respond to increasing prey density, as almost no amphipods were eaten at this oxygen level. The results are discussed in terms of the usability of theoretical models to examine predator–prey relationships in stressful environments. Received: 26 April 1997 / Accepted: 20 May 1997     

The respiratory metabolism of Tilapia mossambica (Teleostei)

Tilapia mossambica (Teleostei) weighing 5 to 80 g were acclimated at 30°C to salinities of 0.4 (tap water), 12.5 (50% sea water) and 30.5 (100% sea water). Their respiration was measured at routine activity and the partial pressure of ambient oxygen gradually reduced from 250 to 50 mm Hg. Respiration is salinity-dependent; the proportionate ability to use oxygen in any one salinity is — above the critical pO₂ —the same in all experimental groups. This ability is a function of temperature and increases from 15° to 30°C, becoming temperature independent from 30° to 40°C as long as the pO₂ remains above 150 mm Hg. At 50 mm Hg pO₂, the limiting effect of oxygen causes a decrease in metabolic rate. This limiting effect is minimal in 80 g fish kept in an isotonic medium (12.5 S), allowing greater scope for activity and a higher rate of oxygen uptake.     

The burrows and physiological adaptations to a burrowing lifestyle of Natatolana borealis (Isopoda: Cirolanidae)

The isopod Natatolana borealis Lilljeborg constructs U-shaped burrows in soft mud, the bore of which closely approximates the width of the occupant. Within artificial burrows, the isopods are largely quiescent and often adopt a position close to one of the burrow openings. Conditions within burrows constructed in the laboratory are moderately hypoxic [11.7 to 14.9 kPa (88 to 112 torr)], with isopods showing discontinuous irrigation behaviour (pleopod beating). Rates of oxygen consumption (measured at 10°C) are maintained approximately constant over a wide range of oxygen partial pressure (P_{O 2}) due, in part, to a pronounced increase in pleopod beat rate. Values for the critical partial pressure of oxygen (P_c), the P_{O 2} at which can no longer be maintained independent of P_{O 2}, were 2.0 to 3.3 kPa (15 to 25 torr). N. borealis can survive lengthy periods (65 h at 5°C) of anoxia, during which there is a significant reduction in the carbohydrate concentration and an increase in the l-lactate concentration of the tissues. The oxygencarrying capacity of the haemolymph of N. borealis was low. The haemocyanin showed a relatively high oxygen affinity [P₅₀=0.39 kPa (2.99 torr) at 10°C at the in vivo pH of 7.80] and a pronounced Bohr effect (-1.22). These characteristics may be advantageous to a burrowing mode of life and also for the conditions likely to be encountered in fish carcasses into which they burrow en masse to feed.     

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.     

Blood oxygen-binding characteristics of bigeye tuna (Thunnus obesus), a high-energy-demand teleost that is tolerant of low ambient oxygen

 We found blood from bigeye tuna (Thunnus obesus) to have a significantly higher O₂ affinity than blood from other tunas. Its P₅₀ (partial pressure of oxygen, P_O2 required to reach 50% saturation) was 1.6 to 2.0 kPa (12 to 15 mmHg) when equilibrated with 0.5% CO₂. Previous studies employing similar methodologies found blood from yellowfin tuna (T. albacares), skipjack tuna (Katsuwonus pelamis), and kawakawa (Euthynnus affinis) to have a P₅₀ of 2.8 to 3.1 kPa (21 to 23 mmHg). These observations suggest that bigeye tuna are more tolerant of low ambient oxygen than other tuna species, and support similar conclusions derived from laboratory whole-animal studies, depth-of-capture data, and directly-recorded vertical movements of fish in the open ocean. We also found the O₂ affinity of bigeye tuna blood to be essentially unaffected by a 10 C° open-system temperature change (as is the blood of all tuna species studied to date). The O₂ affinity of bigeye tuna blood was, however, more affected by a 10 C° closed-system temperature change than the blood of any tuna species yet examined. In other words, bigeye tuna blood displayed a significantly enhanced Bohr effect (change in log P₅₀ per unit change in plasma pH at P₅₀) when subjected to the inevitable changes in partial pressure of carbon dioxide (P_CO2) and plasma pH that accompany closed-system temperature shifts, than when subjected to changes in plasma pH accomplished by changing P_CO2 alone. In vivo, the resultant large decrease in O₂ affinity (i.e. the increase in P₅₀) that occurs as the blood of bigeye tuna is warmed during its passage through the vascular counter-current heat exchangers ensures adequate rates of O₂ off-loading in the swimming muscles of this high-energy-demand teleost. Received: 12 March 1999 / Accepted: 18 December 1999     

Effects of environmental parameters on the oxygen consumption of four marine invertebrates: a comparative factorial study

The oxygen consumption curves of two decapod crustaceans (Palaemon serratus, Penaeus monodon) and two prosobranch molluscs (Trunculariopsis trunculus, Nassarius mutabilis) have been detected in the entire pO₂ interval from 0 to 160 mmHg, under different conditions of temperature and salinity. From the experimental curves, physiological parameters such as the initial oxygen consumption velocity, the Q ₁₀ values and the oxygen independence indices have been measured. The latter parameters have been obtained using normalised plots which allow their better evaluation. The effects of temperature, salinity and oxygen partial pressure on the oxygen-consumption features have been studied using a factorial experimental plan which allows measurement of the effects of each experimental variable as well as the effects of synergistic interactions between different variables. Received: 27 March 2000 / Accepted: 13 November 2000     

Some effects of hypoxia and medium ammonia enrichment on efflux rates and circulating levels of ammonia inNephrops norvegicus

Eutrophication has been reported for autumn months in regions of the Kattegat/Skagerrak, causing stress to bottom-living organisms. The present studies, undertaken in April (1989), investigated the effects of hypoxia and high ammonia levels in the burrowing decapodNephrops norvegicus (L.). The net ammonia efflux rates and circulating ammonia levels at 6 and 12°C, at normoxia [partial pressure of O₂ in the water (torr),P _wO₂ = 155 torr)] and hypoxiaP _wO₂ = 24 torr) in normal seawater and ammonia-enriched (300µmol ammonia l^–1) seawater were examined. The hourly weight-specific efflux rates were very variable and in all groups included some individuals which showed periods of no net efflux, or even a net uptake of ammonia. At each temperature, net efflux-rate differences due to treatments were not significant (P>0.05; ANOVA, in all cases) and only the differences between the net efflux rates of the normoxic groups were significantly affected by temperature (P<0.05; ANOVA). Circulating ammonia levels were also variable, and at 6°C the ammonia-enriched groups had significantly higher weight-specific blood ammonia content values than the normoxic group (P<0.05 in both cases). A net uptake of ammonia occurred in ammoniaenriched conditions — probably along a reversed NH ₄ ⁺ gradient, as downhill pNH₃ gradients were maintained in all groups — and may represent the only means by which some branchial efflux of ammonia could proceed.     

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.     

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.     

Essential fatty acids influence metabolic rate and tolerance of hypoxia in Dover sole (Solea solea) larvae and juveniles

Dover sole (Solea solea, Linneaus 1758) were raised from first feeding on brine shrimp (Artemia sp.) with different contents and compositions of the essential fatty acids (EFA) arachidonic acid (ARA, 20:4n − 6); eicosapentaenoic acid (EPA, 20:5n − 3), and docosahexaenoic acid (DHA, 22:6n − 3), and their metabolic rate and tolerance to hypoxia measured prior to and following metamorphosis and settlement. Four dietary Artemia preparations were compared: (1) un-enriched; (2) enriched with a commercial EFA mixture (Easy DHA SELCO Emulsion); (3) enriched with a marine fish oil combination (VEVODAR and Incromega DHA) to provide a high ratio of ARA to DHA, and (4) enriched with these fish oils to provide a low ratio of ARA to DHA. Sole fed un-enriched Artemia were significantly less tolerant to hypoxia than the other dietary groups. Larvae from this group had significantly higher routine metabolic rate (RMR) in normoxia, and significantly higher O₂ partial pressure (PO₂) thresholds in progressive hypoxia for their regulation of RMR (P _crit) and for the onset of agitation, respiratory distress and loss of equilibrium. Metamorphosis was associated with an overall decline in RMR and increase in P _crit, but juveniles fed on un-enriched Artemia still exhibited higher P _crit and agitation thresholds than the other groups. Sole fed un-enriched Artemia had significantly lower contents of EFA in their tissues, both before and after settlement. Thus, enriching live feeds with EFA has significant effects on the respiratory physiology of sole early life stages and improves their in vivo tolerance to hypoxia. We found no evidence, however, for any effect of the ratio of ARA to DHA.     

Anaerobic metabolism in the shrimp Crangon crangon exposed to hypoxia,anoxia and hydrogen sulfide

The brown shrimp Crangon crangon was collected in the Penzé estuary, Bretagne, in April 1994 and exposed to hypoxia, anoxia and combinations of hypoxia and sulfide. Exposure to sulfide induced total anaerobic metabolism even at an oxygen saturation which would otherwise permit totally aerobic metabolism. In addition to preventing aerobic metabolism there was a direct toxic effect of sulfide. Haemocyanin oxygen affinity (p₅₀) values from non-stressed C. crangon were relatively low. The p₅₀ values all exceed those where environmentally induced lactate accumulation occurs. It seems unlikely that lactate is an affinity modulating factor under environmental hypoxia.     

Influence of environmental factors on burrow irrigation and oxygen consumption in the mudflat invertebrate Urechis caupo

We examined burrow irrigation activity by the mudflat worm Urechis caupo in response to suspended food, ambient hypoxia (down to 3.3 kPa PO₂), hydrogen sulfide exposure (up to 100 µmol l^-1), and short-term temperature change (range 10-22°C). In normoxic, nutrient-free water at 14°C, O₂ consumption ( [(M)\dot]O₂ ) \left( {\dot M{\rm O}_2 } \right) was 45 nmol min^-1 g^-1, water flow rate ( [(V)\dot]_W ) \left( {\dot V_{\rm W} } \right) was 27 ml min^-1 (0.66 ml min^-1 g^-1), frequency of peristaltic waves (F_P) was 2.6 contractions min^-1, stroke volume (SV) was 11 ml, and O₂ extraction coefficient (EO₂) was 0.27. Adding suspended food to the burrow water occasionally elicited stereotypical feeding behavior but had no effect on any measured variables during nonfeeding periods. Hypoxia greatly decreased [(M)\dot]O₂ \dot M{\rm O}_2 (75% reduction at 3.3 kPa PO₂) but did not affect [(V)\dot]_W \dot V_{\rm W} , F_P, SV, or EO₂. Sulfide at 50 µmol l^-1 or less had no effect on burrow irrigation activity, whereas 100 µmol l^-1 sulfide decreased [(V)\dot]_W \dot V_{\rm W} by 58% and F_P by 50% but had no effect on SV. Temperature strongly affected [(V)\dot]_W \dot V_{\rm W} (Q₁₀ of 1.9 from 10°C to 22°C). We propose that U. caupo's ability to live in the hypoxic, sulfidic mud of productive mudflat environments, combined with its very efficient mucous net, allows it to process much less water for feeding than other suspension-feeding invertebrates. This, in turn, necessitates an efficient O₂ extraction mechanism, which is provided by the water lung activity of U. caupo's unique hindgut.