Acid-base status in the sea urchins Psammechinus miliaris and Echinus esculentus (Echinodermata: Echinoidea) during emersion

The acid-base status of two sea urchins, Psammechinus miliaris (Gmelin) and Echinus esculentus (L.) during experimental emersion has been investigated. Sea urchins were collected from the Firth of Clyde between August and September 1987. The carbon dioxide capacity of the coelomic fluid of P. miliaris was greater than that of E. esculentus, although both were low and only marginally greater than that of sea water. The pH of the coelonic fluid was also low (7.05 to 7.17) and was influenced mainly by the internal partial pressure of CO₂ (P_{CO 2}). Acid-base disturbance in the coelomic fluid of both species during emersion, although minimal, was more pronounced in E. esculentus than in P. miliaris and was due primarily to an increase in the internal P_{CO 2}, although there was an increase in the concentration of L-lactate in the coelomic fluid of E. esculentus. The coelomic fluid of both species was in a state of perfectly compensated respiratory acidosis. An increase in the concentration of divalent ions (Ca²⁺ and Mg²⁺) may be related to the dissolution of the test as a source of carbonate buffer.     

Oxygen dependent sulfide detoxification in the lugwormArenicola marina

The lugwormArenicola marina L. oxidizes entering sulfide to thiosulfate. After 8 h of normoxic incubations with sulfide concentrations of 0.2 to 1.0 mmoll^-1 thiosulfate in the coelomic fluid amounted up to about 4 mmoll^-1 whereas sulfite concentrations were 100-fold lower and no accumulation of sulfate in the coelomic fluid was found. The sulfide oxidation was highly oxygen dependent. An increase of oxygen partial pressure ( ) in the medium was followed by enhanced thiosulfate production and by a decrease of sulfide concentration in the coelomic fluid. Under normoxia, the sulfide oxidation rate was sufficient to compensate the influx of sulfide into the coelomic fluid when the sulfide concentration in the medium was below 0.33 mmoll^-1. When external sulfide was raised beyond this level, sulfide up to 5 moll^-1 in the coelomic fluid appeared. Succinate in the body wall tissue was low as long as no sulfide appeared in the coelomic fluid, indicating the maintenance of an aerobic metabolism. The oxidation of sulfide to thiosulfate was localized in the mitochondria of the body wall tissue. The oxygen consumption of mitochondria was stimulated by the addition of sulfide. The mitochondrial sulfide oxidation rate depended on the amount of mitochondrial protein and followed a Michaelis-Menten kinetic. An apparentK _m of 0.68±0.29 moll^-1 and aV _max of 41.9±22.3 nmol min^-1 mg^-1 protein was calculated. Sulfide was stoichiometrically oxidized to thiosulfate with 1 mol sulfide consuming 1 mol oxygen. Sulfide oxidation was not inhibited by sulfide concentrations as high as 100 moll^-1. At low concentrations of cyanide or azide, when respiration without sulfide was already inhibited, sulfide oxidation could still be stimulated, tentatively indicating the existence of an alternative terminal oxidase. Specimens examined in the present study were collected near St. Pol de Leon, France, from 1989 to 1992.     

Anaerobic and aerobic energy metabolism in ovaries of the sea urchin Strongylocentrotus droebachiensis

The gonads of sea urchins undergo large changes in mass during their gametogenic cycle. In addition, they have relatively low aerobic capacities and are poorly perfused by the circulatory system and thus are continually hypoxic or anoxic. The present study of Strongylocentrotus droebachiensis investigates seasonal changes in the relationships among mass of the ovaries, pH and P_O2 of the perivisceral coelomic fluid which bathes the ovaries, and partitioning of ovary energy metabolism into its anaerobic and aerobic components. S. droebachiensis were collected at Blue Hill Falls, Maine, USA, from August 1982 to March 1984. We found that from 76 to 92% of the heat dissipated by isolated ovaries of the sea urchin S. droebachiensis derives from anaerobic energy metabolism at partial pressures of oxygen prevailing in vivo. Ovaries from S. droebachiensis have the capacity to produce large amounts of lactate under imposed anoxia, but lactate accounts for only 37% of the total anoxic heat dissipation, which suggests that other end products of anaerobiosis are present. Seasonal changes in pH and P_O2 of the perivisceral coelomic fluid can be explained by a complex relationship among changes in temperature, reproductive condition, and anaerobic and aerobic metabolism in the ovaries, gut and body wall. Seasonal changes in the buffering capacity of the perivisceral coelomic fluid must be determined before the effects of respiratory and metabolic acid production on the acid-base status of the coelomic fluid can be fully understood.     

Opsonin-like molecule found in coelomic fluid of a sea cucumber, Holothuria leucospilota

Opsonins, a group of serum proteins present in vertebrates, are known to bind to the surface of many kinds of pathogens and tag them as targets for phagocytosis. Opsonin-like molecules have been suggested to exist in several invertebrates. In this paper, we report the discovery of a new opsonin-like molecule in the coelomic fluid of the sea cucumber Holothuria leucospilota. Opsonic activity in cell-free coelomic fluid was characterized by sodium dodecyl sulfate–polyacrylamide gel electrophoresis, two-dimensional electrophoresis and gel filtration. Phagocytic uptake of yeast cells and latex beads by amoebocytes was quantified by microscopic analysis. It was found that a 20 kdalton protein bound to and enhanced phagocytosis of heat-killed yeast cells. The results of Western blot analysis suggest that both amoebocytes and spherule cells produce this opsonin-like molecule. Received: 22 October 1998 / Accepted: 4 March 2000     

Cysteine-glutathione disulfide, the sperm-release pheromone of the marine polychaete Nereis succinea (Annelida: Polychaeta)

Summary. In the marine polychaete Nereis succinea (Frey & Leuckart 1847) a sex pheromone was isolated from the coelomic fluid of sexually mature females and identified by NMR studies and independent synthesis. This pheromone is released by the females during reproduction together with eggs and coelomic fluid into the free water column and induces sperm release of surrounding males. Its structure was ascertained as L-cysteine-glutathione disulfide. It exhibited a response threshold of 0.6 · 10⁻⁷ M. Received 15 July 1997; accepted 25 November 1997.     

Metabolic and blood characteristics of the hydrothermal vent tube-worm Riftia pachyptila

Specimens of the hydrothermal vent pogonophoran Riftia pachyptila Jones were collected by submersible at a depth of 2 600 m at the 21°N hydrothermal vent site on the East Pacific Rise (20°50N, 109°06W) in April and May of 1982. The worms were maintained in pressurized aquaria for up to 45 d for metabolic studies. Consumption of O₂ was regulated down to low P_{O 2} (oxygen partial pressure) values; O₂ consumption rates were 0.63 and 1.12 mol g^-1 wet wt h^-1 at 2.5° and 8°C, respectively; such rates were comparable to those previously measured for other pogonophorans. Intact specimens of R. pachyptila (including bacterial symbionts) did not consume significant amounts of CH₄ from the environment. The respiratory quotients, in the absence of added sulfide, indicated that metabolism was mainly heterotrophic. High rates of uptake of dissolved amino acids were recorded for one specimen. The total [CO₂] in the vascular blood and the Hb-containing coelomic fluid were high. Under anaerobic conditions, there were equilibrium distributions of pH, total [CO₂] and sulfide concentrations between the vascular blood and the coelomic fluid, apparently because these metabolites were readily exchanged between the two compartments. The vascular blood bound neither CH₄ nor H₂. However, sulfide was reversibly bound by both the vascular blood and coelomic fluid; because this binding depended strongly on pH (with a maximum at about 7.5), HS^- was probably the molecular species bound. Under anaerobic, but not aerobic conditions, the trophosome bound substantial amount of sulfide; thus, the high concentrations of sulfide in the trophosome may have resulted mainly from sulfide bound to sulfide oxidases under anaerobic conditions. The coelomic fluid had a relatively low buffering capacity (2.2 mmol CO₂pH^-1).     

Burrow environment and coelomic fluid characteristics of the echiuran worm Urechis caupo from populations at three sites in northern California

The burrow microhabitat and physiology of the echiuran worm Urechis caupo at a high-density site (Elkhorn Slough, California, USA), were investigated from 1987 to 1990 to determine physical and chemical conditions, worm density and distribution, and coelomic fluid characteristics such as heme composition, pO₂, pH, and coelomocyte volume. During tidal exposure, worm burrows at this site exhibited, on average, 52% air-saturated water, 11M burrow-water sulfide, 85 g/mg wet weight sediment sulfide, and salinity and osmolalities similar to those of seawater. These conditions are compared to those of another California site, Bodega Bay, which had slightly lower oxygen concentrations, but higher water and sediment-sulfide levels. A more limited comparison to a third site, Princeton Harbor, California, is included. Worms from Bodega Bay, the higher sulfide site, had greater concentrations of hematin, a non-globin heme compound contained in the coelomocytes, and exhibited a greater tolerance to sulfide in the laboratory. These data are consistent with the hypothesis that hematin is a sulfide-detoxifying agent that may enhance survival of U. caupo in the sulfide-rich mudflat environment.     

Uptake,binding and clearance of divalent cadmium in Glycera dibranchiata (Annelida: Polychaeta)

The bloodworm Glycera dibranchiata Ehlers, 1968 accumulates cadmium through the general body surface and the intestine. Absorption through the gut accounts for cadmium which rapidly binds to coelomic proteins. Intracoelomic injection of ¹⁰⁹Cd demonstrates that cadmium binds readily to hemoglobin and other proteins. The degree of cadmium binding is pH-dependent. The apparent pK of binding sites in body wall and musculature homogenates is 5.39. Cadmium ions injected into the coelom at 7 g g^-1 tissue increase proline incorporation rates into the positively charged hemoglobin (cathode fraction) by 15-fold in 3 days. A 3-fold increase of proline incorporation was observed in the anode hemoglobin fraction over the same time period. Radioactivity in various protein fractions decreases at different rates after injection of ¹⁰⁹Cd. Comparisons between the function of mammalian metallothionein and the coelomic fluid proteins of G. dibranchiata as a detoxification mechanism are discussed.     

Role of ovarian follicle cells in vitellogenesis and oocyte resorption inCapitella sp. I (Polychaeta)

Capitella sp. I has a short generation time, is polytelic and produces broods of relatively large, yolky eggs at intervals of as short as 5–7 d at 20°C. The follicle cells surrounding vitellogenic oocytes in the medial portion of the ovary contain large nuclei, extensive arrays of rough endoplasmic reticulum and Golgi complexes. The coelomic surfaces of follicle cells in the outer layer are covered by blade-like microvilli, which project into the coelom. In the dorsal and lateral portions of the ovary, where mature oocytes are found, the follicle cells are covered by flat, stellate-shaped, peritoneal cells that resemble the podocytes of many invertebrate excretory organs and of the mammalian glomerulus. Changes occurring in the ovary during resorption of oocytes in the middle to advanced stages of vitellogenesis are described. Oogonia and previtellogenic oocytes are not resorbed and perhaps subsequently undergo vitellogenesis. The ovary in this highly opportunistic polychaete species may play a significant regulatory role in the control of reproduction by producing yolk precursors, regulating their uptake from the coelomic fluid via the activity of the follicle cells and by resorbing mature oocytes in response to stress.     

Carbon dioxide use by chemoautotrophic endosymbionts of hydrothermal vent vestimentiferans: affinities for carbon dioxide, absence of carboxysomes, and δ13C values

The hydrothermal vent vestimentiferans Riftia pachyptila Jones, 1981 and Ridgeia piscesae Jones, 1985 live in habitats with different abundances of external CO₂. R. pachyptila is found in areas with a high input of hydrothermal fluid, and therefore with a high [CO₂]. R. piscesae is found in a range of habitats with low to high levels of hydrothermal fluid input, with a correspondingly broad range of CO₂ concentrations. We examined the strategies for dissolved inorganic carbon (DIC) use by the symbionts from these two species. R. pachyptila were collected from the East Pacific Rise (9°50′N; 104°20′W) in March 1996, and R. piscesae were collected from the Juan de Fuca Ridge (47°57′N; 129°07′W) during September of 1996 and 1997. The differences in the hosts' habitats were reflected by the internal pools of DIC in these organisms. The concentrations of DIC in coelomic fluid from R. piscesae were 3.1 to 10.5 mM, lower than those previously reported for R. pachyptila, which often exceed 30 mM. When symbionts from both hosts were incubated at in situ pressures, their carbon fixation rates increased with the extracellular concentration of CO₂, and not HCO₃ ⁻, and symbionts from R. piscesae had a higher affinity for CO₂ than those from R. pachyptila (K _1/2 of 7.6 μM versus 49 μM). Transmission electron micrographs showed that symbionts from R. piscesae lack carboxysomes, irrespective of the coelomic fluid [DIC] of their host. This suggests that the higher affinity for CO₂ of R. piscesae symbionts may be their sole means of compensating for lower DIC concentrations. The δ¹³C values of tissues from R. piscesae with higher [DIC] in the coelomic fluid were more positive, opposite to the trend previously described for other autotrophs. Factors which may contribute to this trend are discussed. Received: 24 September 1998 / Accepted: 12 May 1999     

Venoms from six species of marine fish: lethal and hemolytic activities and their neutralization by commercial stonefish antivenom

A specimen of the stonefish Synanceja verrucosa was captured in Okinawa in March 1988. Live specimens of the scorpionfish Inimicus japonicus were purchased from the Tokyo Central Wholesale Market in November 1988 and those of four species of zebrafish Pterois lumulata, P. volitans, P. antennata and Dendrochirus zebra from an aquarium in December 1988. Crude venoms were extracted from dorsal spines of the six species. All venoms exhibited lethal activity against mice and hemolytic activity specific for rabbit erythrocytes. The lethal activity (or hemolytic activity) of each venom was very unstable to freezing, lyophilization and heating. Both lethal and hemolytic activities of S. verrucosa venom were remarkably neutralized by the commerical stonefish (Synanceja trachynis) antivenom. This antivenom was also effective, to some extent, in counteracting the other venom activities. The neutralizing capacities of the antivenom were calculated to be 7310 LD₅₀ ml^-1 for S. verrucosa venom and 1 220 to 2 990 LD₅₀ ml^-1 for the other venoms. Results of neutralization tests suggest that venoms from the six species were comparable in terms of antigenecity.     

Effects of pressure,temperature and oxygen on the oxygen consumption rate of the Midwater copepod Gaussia princeps

In this study we assessed the amount of lead accumulated in the body of a grazing mollusc by transfer from its algal food in laboratory experiments, and compared these results with the amounts found in naturally occurring molluscs and seaweed. Near La Jolla, California (USA), where the concentration of lead in seawater is probably less than 0.08 g l^-1, most of the naturally occurring Egregia laevigata contains less than 0.4 g Pb g^-1 wet weight. The total body masses, without shells, of juvenile Haliotis rufescens fed on this seaweed for 3 to 6 months showed similar concentrations. When, however, E. laevigata is placed for 1 to 6 days in seawater to which lead has been added (0.1 or 1.0 mg l^-1) both the seaweed and the abalone subsequently fed with it accumulate proportionally larger amounts of lead. After 6 months, young abalone fed on E. laevigata pretreated with 1.0 mg Pb l^-1 accumulated up to 21 g Pb g^-1 wet weight. This amount of lead had no apparent consequences on the growth or activity of the molluscs. Analyses of 6 different organs from adult abalone showed that the lead was selectively concentrated in the digestive gland. In the foot (muscle tissue), which is the part normally consumed by humans, only negligible amounts were found.     

Timing of septal formation in two species of Nautilus based on radiometric and aquarium data

Growth rates of four Nautilus scrobiculatus Lightfoot and three N. pompilius Linnaeus captured in Papua New Guinea in 1985 were measured using a chronology based on the naturally occurring radionuclides ²¹⁰Pb and ²¹⁰Po. The age of the las septum in mature specimens of both species was more than one year, whereas the time of septal formation in a submature specimen of N. pompilius was ca 180 d. The ²¹⁰Po/²¹⁰Pb method was also used to determine the growth rates of four immature N. pompilius captured in the Philippines in 1983 before and after introduction into an aquarium. Introduction into the aquarium was marked by a dramatic decrease in the activity of ²¹⁰Pb of the septa, the formation of a black line on the exterior of the shell, and a reduction in the spacing of adjacent septa in two specimens. Based on the number of septa formed in the aquarium, the average time of septal formation ranged from 50 to 80 d. These estimates were consistent with those calculated from the ²¹⁰Po/²¹⁰Pb activity ratios of the septa that formed in the aquarium, thus providing a check on the radiometric method. The average time of septal formation just prior to introduction into the aquarium appears, within the uncertainty of the radioactive measurements, to have been similar to that in the aquarium, indicating no dramatic alteration in the growth rate of these specimens.     

Alkaline phosphatase and carbonic anhydrase activity associated with arm regeneration in the seastarAsterias forbesi

A sharp increase in alkaline phosphatase (AP) activity occurred 7 d after arm autotomy inAsterias forbesi. This activity remained elevated at 20 d post-autotomy, during which time active ossicle-calcification was occurring. By 60 d post-autotomy, AP activity had returned to the level of non-regenerating individuals. Conversely, carbonic anhydrase (CA) activity exhibited no initial change in response to arm autotomy. By Day 60 a 50% increase in CA activity was observed. AP is localized in the outer membranes and associated microvilli of the coelomic epithelium and is not associated directly with the tissues involved in calcification. CA activity was highest in the tissues containing the ossicles, but was also present in the non-calcifying tube feet. These data suggest that inA. forbesi AP is involved in cell division and differentiation associated with wound-healing and the initiation of the regeneration process, and that CA may facilitate the ongoing maturation process, of newly formed ossicles.     

重金属Cr(VI)、Pb及Cu胁迫对双齿围沙蚕体腔细胞的DNA损伤

为探讨重金属Cr(VI)、Pb以及Cu对沙蚕体腔细胞DNA的毒性效应,以双齿围沙蚕为受试动物,重金属按不同剂量水平,Cr(VI):10、100和200 mg·L~(-1),Pb:5、50和100 mg·L~(-1),Cu:1、10和20 mg·L~(-1),分别胁迫沙蚕24 h,以不加任何重金属离子的海水为对照,采用单细胞凝胶电泳技术,检测其体腔细胞DNA损伤程度。结果表明,与空白对照组相比,3种重金属离子的各浓度组都能引起沙蚕体腔细胞DNA损伤,且3种重金属胁迫浓度与细胞DNA损伤程度之间存在显著的剂量-效应关系。双齿围沙蚕可以作为单细胞凝胶电泳的实验材料用于重金属所致环境污染的生物监测指示生物。     

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.     

Interactive accumulation of mercury and selenium in the sea starAsterias rubens

Interactions between mercury and selenium accumulation and subcellular binding inAsterias rubens (L.), collected in 1987 from Lille Bælt at Middelfart, Funen, Denmark, were investigated in laboratory experiments. Sea stars exposed to 10µg Hg l^–1 for 30 d accumulated mercury in body wall, tube feet and stomach linearly with time at 1.2, 1.2 and 0.5µg Hg g^–1 dry wt d^–1, respectively. Mercury was accumulated in pyloric caeca and coelomic fluid initially at 1.4µg Hg g^–1 dry wt d^–1 and 9.4 ng Hg ml^–1 d^–1, respectively; after 10 d uptake rates decreased. Sea stars exposed to 75µg Se-SeO ₃ ^{- -} l^–1 accumulated selenium linearly with time over 30 d in the stomach, pyloric caeca, tube feet and body wall at 2.0, 1.2, 1.2 and 0.6µg Se g^–1 dry wt d^–1. Sea stars exposed to 75µg Se-SeO ₄ ^{- -} l^–1 maintained selenium levels in the coelomic fluid at 75µg Se l^–1 over 30 d. Exposure to selenate did not alter the selenium concentrations in the tissues. Sea stars exposed concurrently to 75µg Se-SeO ₃ ^{- -} and 10µg Hg l^–1 accumulated more mercury and selenium in tube feet and body wall than did sea stars exposed to the two elements alone. In pyloric caeca and stomach concurrent exposure reduced accumulation of both elements. Mercury was bound predominantly in the insoluble fraction of the tissues, and soluble mercury was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Ca. half of the selenium recovered was bound in the insoluble fraction, and soluble selenium was bound in proteins of high (> 70 kilodaltons) or very low (< 6000 daltons) molecular weight. Interaction between the two elements was exerted predominantly in the insoluble fraction of the tissues.     

A population study of the polychaete Capitella capitata at plymouth

A population of Capitella capitata (Fabricius) was sampled from January, 1973 to February, 1974. Size of the worms was measured and sexual maturity determined. There is an extensive breeding season, spawning occurring asynchronously throughout the year. Females containing ripe oocytes were almost always present, but most females at any one time contained immature oocytes. A high percentage of males contained active sperm. Oocytes are not released into the coelomic fluid until almost fully developed, and all are released at a single spawning. A large number of eggs are produced at each spawning and there is some evidence to suggest that larval development may be completely benthonic. Gametes first develop in 7 to 8 month old worms and take about 4 months to complete development. Analyses of the female population structure suggests that there are 3 breeding classes. The growth rate is estimated at 30 mm per year, and few worms are believed to survive longer than 2 years.     

Characterization of alkaline phosphatase and organic phosphorous utilization in the oceanic dinoflagellate Pyrocystis noctiluca

Phosphate depleted Pyrocystis noctiluca (Murray) Schuett 1895 has at least one phosphomonoesterase (EC 3:1:3:1) which is triphasic between 0.1 and 222 mol P. The enzyme has a broad temperature range with maximum activity at 50 °C and a Q₁₀ of 1.4 to 1.5. A break in the Arrhenius plot at 35 °C implies the enzyme is membrane-bound. Cytological staining of whole cells and cell fractionation studies (showing 26 times higher specific activity in the particulate compared with the cytoplasmic fraction) suggest the enzyme is plasmalemma-bound. The enzyme has an absolute metal requirement which would be satisfied by Mg⁺⁺ but not Mn⁺⁺, Zn⁺⁺, Fe⁺⁺, or Co⁺⁺ at seawater concentrations. Alkaline phosphatase is a stable enzyme whose activity is not altered by inhibitors of protein synthesis. Orthophosphate inhibition of enzyme activity was largely eliminated in the presence of these inhibitors. Apparently, a protein induced by PO₄ ^3-, rather than PO₄ ^3- itself, inhibits alkaline phosphatase. Cell-free alkaline phosphatase can hydrolyze a variety of phosphate esters and linear polymers of inorganic phosphorus as well as disolved organic phosphorus from tropical oceanic waters. These same hydrolysable organic and inorganic phosphorus compounds support the axenic culture growth of P. noctiluca, suggesting that naturally occurring hydrolysable organic phosphorus compounds may also support the growth of this alga.     

Circulation of perivisceral fluid in the sea urchin Lytechinus variegatus

The principal driving mechanism of perivisceral-fluid circulation was investigated in the echinoid Lytechinus variegatus by metal-clad hot-wire anemometry. Analysis of 23 flow profiles from 13 individuals collected from 1 to 3 m water depth at Tarpon Springs and Sarasota, Florida (USA) in 1984 indicates that for this species, the ciliated epithelium is an ineffective driving mechanism for mixing of the perivisceral fluid. Instead, the Aristotle's lantern is the major driving force of circulation of the perivisceral fluid, with velocity fluctuations of 1.5 mm s^-1 amplitude in the cavity center during intermittent periods of activity. These fluctuations rapidly decay as the ciliated surfaces are approached. Spectral analysis of velocity time-series reveals predominant energy-containing velocity fluctuations between 0.007 and 0.05 Hz. The circulation, which can be described as non-turbulent forced mixing, has revealed four different flow patterns which appear to be correlated with specific animal activities. A modified eddydiffusion model predicts that oxygen transport through the cavity over a 2 cm path length requires more than 20 h. Calculation of the oxygen flux to a ripe ovary suggests that supply does not meet ovarian oxygen demand in this case. Alternative pathways of oxygen transport invoking advection in perivisceral-fluid eddies and along the ciliated coelomic surfaces are compatible with the observed flow patterns and lead to much shorter oxygen-transport time scales of the order of 1 to 10 min.