Differential sensitivities to hypoxia by two anoxia-tolerant marine molluscs: A biochemical analysis

The metabolic responses to a series of low oxygen tensions were compared for two species of Mediterranaean bivalves,Mytilus galloprovincialis andScapharca inaequivalvis. Whereas both species have well-developed and similar tolerances of anoxia, the metabolic responses ofS. inaequivalvis to low oxygen tensions indicate a substantially greater tolerance of hypoxia. Compared withM. galloprovincialis, the responses ofS. inaequivalvis included the ability to maintain a constant oxygen consumption down to a much lower pO₂ value (ca. 1.7 vs 3.4 ppm), and a lower critical pO₂ for the recruitment of fermentative pathways of ATP production (ca. 1 vs 3 ppm). Furthermore, a graded increase in the output of anaerobic products (succinate, alanine) occured at oxygen tensions below 3 ppm inM. galloprovincialis and reached a maximum at 1.6 ppm whereas inS. inaequivalvis the net accumulation of anaerobic products at the lowest oxygen tension tested (0.5 ppm) was still substantially less than the level of production output in complete anoxia. This suggests that fermentative pathways are maximally activated at all oxygen tensions below 1.6 ppm inM. galloprovincialis whereas rates of anaerobic pathways are still less than maximum at 0.5 ppm inS. inaequivalvis. These results indicate that in situations of declining oxygen tensions, such as occur due to eutrophication,M. galloprovincialis would not only begin to experience metabolic stress at higher oxygen tensions thanS. inaequivalvis but would experience greater stress at any given pO₂. Such differences in hypoxia tolerances may explain the success of the recently introducedS. inaequivalvis in out-competing the nativeM. galloprovincialis in the Adriatic Sea.This research was conducted at the Consorzio di Studi, Richerche ed Interventi sulle Risorse Marine, Viale Amerigo Vespucci 2, I-47042 Cesenatico (FO), Italy. Delta Institute for Hydrobiological Research, Yerseke, The Netherlands. Publication no. 554     

An investigation on the anaerobic metabolism of Nephtys hombergii (Annelida: Polychaeta)

In Nephtys hombergii Savigny (Annelida: Polychaeta: Errantia), succinate and less pronounced L-alanine are accumulated as endproducts during an initial phase of anaerobiosis. In this phase aspartate is utilized as substrate for anaerobic energy production in addition to glycogen. Prolonged anaerobiosis results in the formation of propionate and acetate which, to a large extent, are excreted into the water. The concentrations of aspartate and succinate were found to remain unchanged down to a Pw_{O 2}(oxygen partial pressure in incubation water) of approximately 20 torr, indicating a fully aerobic metabolism. At a Pw_{O 2}of 15 torr characteristic changes can be observed: energy production becomes partially anaerobic. At a Pw_{O 2}of 7 torr the level of aspartate is largely reduced, but the accumulation of succinate is still significantly less than at anoxia. [In the habitat of N. hombergii, the intertidal mud flats, a Pw_{O 2}in the range from 0 torr (Pamatmat, 1968) to 12 torr (Wells and Warren, 1975) was measured.] The activities of some important enzymes involved in anaerobic energy production (among them pyruvate kinase, phosphoenolpyruvate carboxykinase and lactate dehydrogenase) were measured. Although N. hombergii has a high glycolytic capacity, no lactate dehydrogenase was detected. Instead strombine and alanopine dehydrogenases were present in comparatively high activities.     

Effects of oxygen and inorganic carbon concentrations on the photosynthetic quotients of marine algae

The photosynthetic quotients of the marine prymnesiophyte Pavlova lutheri and the marine dinoflagellate Glenodinum sp. were measured at different concentrations of dissolved oxygen and inorganic carbon. Dissolved oxygen concentration appeared to be the most important factor controlling the photosynthetic quotient. Photosynthetic quotients generally were between 1.0 and 1.8 at oxygen concentrations less than saturation, were approximately 1.0 at oxygen saturation, and generally were from 0.1 to 1.0 at oxygen concentrations greater than saturation. The photosynthetic quotients greater than 1.0 were not caused by lipid synthesis. They may have been partially caused by the presence of KNO₃ rather than an ammonium salt in the growth media. The lowered photosynthetic quotients at higher oxygen concentrations were probably caused by algal photorespiration.Contribution No. 202 from the Department of Biology, The Pennsylvania State University; 202 Buckhout Laboratory, University Park, Pennsylvania 16802, USA     

Effects of arsenic on survival and metabolism of Crangon crangon

Metabolic changes and toxic effects in Crangon crangon (collected during summer 1989 near Helsingør, Denmark and at Kulhuse in the Isefjord) were observed during exposure to different arsenate concentrations. Survival, respiration, and concentrations of hemolymph glucose, muscle glycogen andadenine nucleotides were measured. C. crangon was very tolerant of arsenate; survival was only affected at arsenate concentrations >25 ppm. Small individuals (0.2 to 0.29 g wet weight) were less tolerant than medium-sized and large individuals (0.5 to 0.59 g and 0.8 to 0.89 g wet weight), e.g. at 50 ppm the LT₅₀ for small individuals was 180 h compared to 343 h and 360 h in middle-sized and large individuals. The relation between size and toxicity is probably due to the change in surface/volume ratio. Under clean conditions the respiration of C. crangon showed a clear relation between routine MO₂ (O₂ in mlg^-1 h^-1) and wet weight (r²=0.89; P<0.001). During shortterm exposure to arsenate (10 and 50 ppm), the respiratory rates (MO₂) seemed to decrease in a size-dependent manner. The concentrations of blood glucose and muscle glycogen did not change in response to arsenate exposure. The concentrations of adenylate phosphates were high and constant throughout the experiments. C. crangon had maximal adenylate energy charge (AEC) values of ca. 0.8. No decrease in AEC due to the exposure to arsenate was observed.     

The toxicity of heavy metals to embryos of the American oyster Crassostrea virginica

The acute toxicity of 11 heavy metals to embryos of the American oyster Crassostrea virginica was studied and the concentrations at which 50% of the embryos did not develop were determined. The most toxic metals and their LC₅₀ values were mercury (0.0056 ppm), silver (0.0058 ppm), copper (0.103 ppm) and zinc (0.31 ppm). Those metals that were not as toxic and their LC₅₀ values were nickel (1.18 ppm), lead (2.45 ppm) and cadmium (3.80 ppm). Those metals that were relatively non-toxic and their LC₅₀ values were arsenic (7.5 ppm), chromium (10.3 ppm) and manganese (16.0 ppm). Aluminum was non-toxic at 7.5 ppm, the highest concentration tested.     

Nephtys hombergii, a free-living predator in marine sediments: energy production under environmental stress

Nephtys hombergii is a free-living, burrowing predator in marine sediments. The worm is, therefore, exposed to various environmental conditions which tube-dwelling polychaetes of the same habitat most likely do not encounter. The worms have to survive periods of severe hypoxia and sulphide exposure, while at the same time, they have to maintain agility in order to feed on other invertebrates. N. hombergii is adapted to these conditions by utilising several strategies. The species has a remarkably high content of phosphagen (phosphoglycocyamine), which is the primary energy source during periods of environmental stress. With increasing hypoxia, energy is also provided via anaerobic glycolysis (pO₂<7 kPa), with strombine as the main end-product. Energy production via the succinate pathway becomes important only under severe hypoxia (<2 kPa), suggesting a biphasic response to low oxygen conditions which probably is related to the worm's mode of life. The presence of sulphide resulted in a higher anaerobic energy flux and a more pronounced energy production via glycolysis than in anoxia alone. Nevertheless, after sulphide exposure under anaerobic conditions of <24 h, N. hombergii is able to recover completely. Although N. hombergii appears to be well adapted to a habitat with short-term fluctuations in oxygen and appearance of hydrogen sulphide, its high energy demand as a predator renders it likely to limit its survival in an environment with longer lasting anoxia and concomitant sulphide exposure. Received: 28 May 1997 / Accepted: 21 June 1997     

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.     

Studies on factors affecting survival of nile fish in the Sudan. III. The effect of oxygen

Tilapia nilotica L. were exposed to different levels of ambient oxygen concentrations for 24 h periods. Nitrogen was bubbled through the water to reduced the oxygen concentration to the respective test level. When testing concentrations of oxygen between 2.5 and 0.4 ppm over 24 h periods of exposure, the median tolerance limits (TLM) over 24 h were attained at 1.41 ppm of oxygen.     

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.     

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).     

Depression of feeding and growth rates of the seastar Evasterias troschelii during long-term exposure to the water-soluble fraction of crude oil

To test the effect of petroleum hydrocarbons on predation by the seastar Evasterias troschelii (Stimpson, 1862) on the mussel Mytilus edulis (L.), we exposed the predator with the prey to six concentrations of the water-soluble fraction (WSF) of Cook Inlet crude oil. Seastars and mussels were collected at Auke Bay, Alaska, in November 1980. During a 28 d exposure in a flow-through system, seastars were more sensitive to the WSF than mussels: the LC₅₀ for the seastars was 0.82 ppm at Day 19 and, although no mussels were exposed to WSF for more than 12 d, none died. Daily feeding rates (whether in terms of number of mussels seastar^-1 d^-1 or dry weight of mussels seastar^-1 d^-1) were significantly reduced at all concentrations above 0.12 ppm. At 0.20, 0.28 and 0.72 ppm WSF, daily feeding rates (in terms of dry weight of mussels) were, respectively, 53, 37, and 5% of the control rate; at the two highest concentrations (0.97 and 1.31 ppm WSF), the seastars did not feed. Seastars at concentrations greater than 0.12 ppm WSF grew slower than individuals from the control group and the 0.12 ppm-treatment group combined. These laboratory results show that E. troschelii is more sensitive to chronic low levels of the WSF of crude oil. The possibility that such oil pollution could reduce predation and permit M. edulis to monopolize the low intertidal zone of southern Alaska remains to be studied.     

Respiratory quotient and calcification of Nautilus macromphalus (Cephalopoda: Nautiloidea)

Respiration and calcification were investigated in the ectocochleate cephalopod Nautilus macromphalus Sowerby. Specimens were collected off New Caledonia, in October 1991, and kept at the Nouméa Aquarium until December 1991. The respiratory quotient and calcification rate of 5 individuals were measured during 14 short term incubations (63 to 363 min). Oxygen uptake was recorded with a polarographic oxygen sensor. CO₂ flux and calcification were calculated from changes in pH and alkalinity (alkalinity-anomaly technique). Several methods were used to compute the respiratory quotient (RQ); a functional regression indicated an RQ of 0.74. CaCO₃ exchange rates were linearly related to respiratory quotient, calcification occurring in individuals with a low RQ. CaCO₃ uptake from the surrounding water was noncontinuous. From the highest CaCO₃ uptake, maximum growth rate was estimated as 7.1 mg shell wt h^- (=61 g yr^-1).     

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.     

New clues for freshwater eels (<Emphasis Type="Italic">Anguilla</Emphasis> spp.) migration routes to eastern Madagascar and surrounding islands

A total of 4,172 freshwater eels have been collected by electrofishing in upper estuaries from Madagascar (East coast), Mascarene (Réunion and Mauritius Is.), Comoros (Mayotte Is.) and Seychelles (Mahé and Praslin Is.) Archipelagos, between October 2003 and February 2006. Eel species composition in the sampling stations was contrasted between eastern Madagascar (Anguilla mossambica 96.0%, A. marmorata 3.9% and A. bicolor bicolor 0.2%), the Comoros (A. marmorata 56.1% and A. bicolor bicolor 43.9%), the Mascarene (A. marmorata 91.4%, A. bicolor bicolor 5.4% and A. mossambica 3.2%) and the Seychelles Archipelagos (A. bicolor bicolor 100.0%). This gradient in species composition, even concerning the short time-range of our sampling, argued for separate migration routes between species. A total of 168 eels were aged by reading their otolith microstructure, and otolith growth rates were calculated from pre-leptocephalus stage (post-hatching) to metamorphosis, until freshwater check. For all species, mean otolith growth rate (OGR) was related to specific migration routes: A. bicolor bicolor is distributed in the lowest latitudes and showed the highest OGR during leptocephalus stage, whereas A. mossambica, endemic of the Malagasy area, has the most southern distribution and showed the lowest OGR. OGR during leptocephalus stage was negatively correlated to the leptocephalus stage duration, showing a decrease of global metabolism with time, classical in leptocephali. This relationship was found significant for A. marmorata and A. mossambica, probably because all these larvae crossed successively the same environments, but not for A. bicolor bicolor, probably because their larvae crossed different pelagic environments, opening the hypothesis of larvae from different origins.     

Calcification in the articulated coralline alga Corallina pilulifera,with special reference to the effect of elevated CO2 concentration

Calcification in Corallina pilulifera Postels et Ruprecht displayed diurnal variations in aerated (350 ppm CO₂) culture media, with faster rates during the light than during the dark period. Addition of CO₂ (air+1250 ppm) inhibited calcification. This was attributable to the decreased pH resulting from CO₂ addition. Both photosynthesis and calcification were enhanced in seawater, with elevated dissolved inorganic carbon concentrations at a constant pH of 8.2.     

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.     

Monitoring toxicity of an azo dye methyl red and a heavy metal Cu,using plant and animal bioassays

Toxicities of an azo dye methyl red and a heavy metal copper (Cu) were quantified, using growth and mortality as end points, in four plant species and three animal species by subjecting them to short-term (4 days for animals, 10 days for plants) static bioassays. Lemna aequinoctialis Welwitch (EC₅₀: 7–16 ppm) was found to be the most sensitive species for methyl red, Ceratophyllum demersum L. (EC₅₀: 25 ppm) and Lactuca sativa L. (EC₅₀: 56 ppm) were intermediate, while Oryza sativa L. shows reduction in seedling vigor (9–27%) of <50%, being the least sensitive amongst the tested plant species. Methyl red toxicity is almost 3–5-fold higher in growing medium (pH = 5.8–6.0), even at high nutrient levels, while Cu toxicity is higher in nutrient-poor alkaline medium at alkaline pH (8.3–8.7; EC₅₀: Ceratophyllum = 104–200 ppb; Lemna = 100–170 ppb) compared to nutrient-rich acidic medium (pH = 5.4–5.7; EC₅₀: Ceratophyllum = 2600–3175 ppb; Lemna = 4350–4715 ppb). Rice tolerance (EC₅₀: 6500 ppb) was found to be higher than hydrophytes while lettuce was most tolerant to Cu. Fish sensitivity toward the test chemicals was almost parallel to Ceratophyllum and Lemna [Gambusia affinis Baird and Gerard (LC₅₀: 250 ppb for Cu) and Poecilia reticulata Peters (LC₅₀: 24 ppm for methyl red)]. Similar to the plants, dye toxicity increased markedly (LC₅₀: 7 ppm) in the acidic medium (pH = 6.0). Amongst the tested organisms, Daphnia was found to be most sensitive to methyl red (EC₅₀: 6 ppm) while its sensitivity to Cu (EC₅₀: 230 ppb) was similar to hydrophytes and fish. Initially, the combination of dye and Cu (at their sublethal concentrations) had additive effects in duckweed, while dye concentration ruled afterward. These results indicate that hydrophytes and animals are equally sensitive toward the test chemicals. Dye toxicity in hydrophytes and fish was pH dependent, while in the case of Cu, it is related to the nutrient status of the growth medium of plants.     

Effects of feeding rate on conversion efficiency and chemical composition of the fish Tilapia mossambica

Average daily rations of 14, 35 or 58 mg Tubifex tubifex worm per gram of the fish Tilapia mossambica Peters represent maintenance, optimum or maximum feeding levels. At these levels, conversion efficiency (K ₁) is 5,9 or 24%. An amount of 65 mg worm/g fish/day, when fed under experimental conditions, is converted with the poorest conversion efficiency (4%). Test individuals fed at 11 to 49 mg worm/g fish/day show a decreasing trend in water content (78.1 to 74.8%), and an increasing trend in fat content (32.1 to 44.2%). Below or above this feeding rate range, water content increases, while fat content decreases. The range of individual variations in fat content is nearly 3 times greater than that of ash and 15 times greater than that of water. Test individuals starved for 60 days lose 2.1 mg dry body weight/g/day. This loss is contributed by calorifically equivalent amounts of fat and protein. The endogenous loss of nitrogen by these individuals averaged 0.18 mg N/g body weight/day.     

Behavioural and physiological responses of the Norway lobster,<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> (Crustacea: Decapoda), to sulphide exposure

The Norway lobster, Nephrops norvegicus, is moderately tolerant of sulphide [the lethal time to 50% mortality (LT₅₀) was 22.5 h when exposed to 500 M sulphide] but, whenever possible, it attempts to avoid the presence of sulphide in its immediate vicinity. Any sulphide entering the animal is oxidized to thiosulphate, which accumulates in the haemolymph and in the tissues. During exposure to low concentrations of sulphide, the rate of oxygen consumption is maintained or even enhanced even though the lobsters become quiescent. The apparent increase in oxygen consumption is probably due to the oxidation of sulphide to thiosulphate. At higher concentrations, oxygen consumption decreases, perhaps because of the inhibitory effect of sulphide on electron transport, and N. norvegicus resorts to anaerobic metabolism as indicated by the accumulation of lactate in the haemolymph and in the tissues.Communicated by L. Hagerman, Helsingør     

Adaptation of the polychaete worm Scoloplos armiger to hypoxic conditions

The anaerobic metabolism of the intertidal polychaete Scoloplos armiger, its recovery from anaerobiosis and the importance of anaerobic energy production during low tide in the field were investigated. Under anaerobic conditions S. armiger produces energy in the same manner as Arenicola marina, a prototype of an euryoxic invertebrate from the intertidal. Energy is produced from the phosphagen stores and from the breakdown of glycogen to volatile fatty acids, mainly propionate and to a lesser extend acetate. However, S. armiger cannot reduce its energy demand to the same degree as A. marina. This and the relatively small pool of glycogen may be the reason for its only moderate resistance to anoxia. The recovery from anaerobiosis proceeds in S. armiger significantly slower than in A. marina. S. armiger is able to maintain a fully aerobic metabolism down to a Pw_{O 2}of ca. 20 torr and even at a Pw_{O 2}of 10 torr a partly aerobic metabolism was retained. In the field during low tide S. armiger ascends into the oxidative layer, where it is able to maintain an aerobic metabolism even at parts without remaining puddels on the surface.