The cardiac responses of the scallop Pecten maximus (L.) to respiratory stress

Heart activity of Pecten maximus (L.) has been recorded during various forms of experimentally induced respiratory stress. There was considerable variation in the responses of individual scallops but bradycardia generally occurred in response to all forms of respiratory stress, with the rate of fall in heart rate dependent upon the severity of hypoxia.When oxygen tension declined slowly in a closed respirometer there was regulation of both heart rate and oxygen consumption. The critical tension, P_c, for oxygen consumption lay between 70 and 80 mm Hg, and corresponded with a slight regulatory upswing of the heart rate, whereas the P_c for heart rate was much lower at 20–30 mm Hg. Sudden transfer to deoxygenated water for 3 h resulted in very rapid bradycardia and there was a rapid recovery and initial overshoot of the normal rate on return to well-oxygenated sea water. Aerial exposure for 3 h produced more gradual bradycardia followed by gradual recovery on return to sea water.The results of this work are compared in some detail with previous work on other species of bivalve from different geographical areas and habitats, and the mechanisms controlling cardiac and respiratory regulation are discussed. It is concluded that there are few clear-cut general differences between littoral and sublittoral species in their behavioural and physiological adaptations to hypoxia; the main distinguishing feature of littoral-adapted species is their ability to control air-gaping. Changes in heart activity generally indicate variations in metabolic rate, the speed at which the metabolic rate may be altered reflecting the degree of adaptation to the littoral environment.     

Oxygen consumption and the respiratory responses to declining oxygen tension in the crab Ebalia tuberosa (Pennant) (Crustacea: Decapoda: Leucosiidae)

Oxygen consumption and its relationship to body weight and activity have been examined in Ebalia tuberosa (Pennant). Ebalia showed only very limited ability to regulate their oxygen consumption under conditions of declining P_O2. Respiratory independence was markedly affected by the level of activity of the animals. When inactive, respiratory independence was maintained down to a critical P_O2 of only about 130 mm Hg while when active, the crabs behaved as perfect conformers. Inactive Ebalia responded to a decrease in PO₂ by increasing the pumping activity of the scaphognathites and heart. This increased activity continued even at oxygen tensions lower than the P_c. The rate of reversals of the ventilatory currents did not change with decreasing P_O2. In the active crabs, the pumping activity of the scaphognathites and heart, and the rate of current reversals were higher than in the inactive animals, and a decrease in PO₂, did not bring about any further change in any of these responses. When exposed to hypoxic conditions for long periods initially inactive crabs survived longer than did active animals.     

Anaerobic capacity of a crustacean sensitive to low environmental oxygen tensions,the freshwater amphipod Gammarus pulex (L.)

We investigated the accumulation of haemolymph L-lactate in response to anoxia, progressive hypoxia and activity, in a species that rarely encounters low PO₂ in its natural environment, the freshwater amphipod Gammarus pulex (L.). Individuals survived <1.5 h anoxia and accumulated L-lactate in the haemolymph at a rate of 10.26 mmol l^–1 h^–1. No lactate was excreted into the medium during this time. Despite the fact that most individuals of G. pulex were oxyconformers, exposure to acutely declining PO₂s did not result in a switch to anaerobic pathways until PO₂ < 4.4 kPa. Even then the concentrations accumulated remained very low until PO₂ < 0.46 kPa. There was no accumulation of L-lactate in the haemolymph as a result of exhaustive activity. Consequently, it was suggested that (a) it cannot be automatically assumed that anaerobic metabolism is switched on only at the point at which oxyregulation breaks down, (b) that the poor anaerobic capacity observed plays little, if any, role in the metabolic response to hypoxia by Gammarus pulex.     

Priming responses of leek (Allium porrum L.) seeds to different dissolved oxygen levels in the osmoticum

Leek seeds (Allium porrum L.) cv. Winterreuzen were treated for 5 days in stirred bioreactors containing a -1.0 MPa polyethylene glycol (PEG 6000) solution. The level of dissolved oxygen was controlled from zero to approximately two and a half times that of saturation with respect to atmospheric air by bubbling through 0₂/N₂ gas blends from 0% O₂ to 50% O₂. The treatment with 0% O₂ (i.e. bubbling with N₂) gave no reduction in germination time compared to untreated seeds after allowing for the time for water imbibition. As the proportion of oxygen was increased, a rapid reduction was observed until a concentration equivalent to air was used. Further increases to the maximum, produced only a marginal further improvement. The reduction in germination times for seeds that had been cabinet dried was longer for all treatments due to the time required for re-imbibition of water but the general response to priming was the same as with undried seeds. Only dried seeds treated with air or higher oxygen concentrations showed enhanced synchronicity of germination times. The percentage germination of seeds from all treatments was the same, including the treatment with nitrogen gas. These tests help to explain the recently reported advantages of using enriched air for seed priming and are one of the necessary protocols for ensuring satisfactory bulk priming.     

Physiological responses to emersion in the intertidal green crab,Carcinus maenas (L.)

The green shore crab, Carcinus maenas, undergoes on average 6?h periods of emersion during each low-tide cycle during the summer months. Under those conditions, the crab is cut off from its normal water environment and is exposed to potential stress from a suite of environmental and physiological changes: dehydration, compromised gas exchange and resultant internal hypoxia and hypercapnia, thermal stress, and ammonia toxicity. This study examined the comprehensive responses of the green crab in water and to a 6?h emersion period laboratory simulation of a tidal cycle followed by a 1?h re-immersion period, measuring indicators of dehydration, hemolymph osmolality, oxygen uptake, hemolymph acid–base status, heart and ventilatory rate, and hemolymph ammonia and ammonia excretion. Green crabs showed physiological responses of varying magnitude to 6?h of emersion. Individuals were found in the field exclusively under rocks and large clumps of seaweed where temperatures were approximately half those of exposed surfaces and relative humidity was about twice as high as ambient air. During emersion, crabs lost less than 5% of their wet weight, and hemolymph osmolality did not increase significantly. Oxygen uptake continued in air at about 50% of the control, aquatic values; and the gills continued to be ventilated by the scaphognathite, albeit at lower rates. Hemolymph lactate concentrations increased, indicating a shift to a greater reliance on anaerobic metabolism to support energetic needs. A slight acidosis developed in the hemolymph after 1?h of emersion, but it did not increase thereafter. Ammonia concentrations in the hemolymph were unchanged, as ammonia was volatilized by the gills and excreted into the air as NH₃ gas. These results show that the green crab copes with emersion by seeking refuge in microhabitats that mitigate the changes in the physical parameters of intertidal emersion. Physiologically, desiccation is avoided, cardio-respiratory processes are maintained at reduced levels, and hemolymph acid–base balance is minimally affected. Ammonia toxicity appears to be avoided by a shift to excreting NH₃ gas directly or indirectly to air.     

The combined effects of temperature,salinity, and declining oxygen tension on oxygen consumption in the marine pulmonate Amphibola crenata (Gmelin, 1791)

Oxygen consumption of Amphibola crenata (Gmelin) was measured in various salinity-temperature combinations (< 0.1‰ to 41‰ salinity and 5 to 30°C) in air, and following exposure to declining oxygen tensions. In all experimental conditions, respiration varied with the 0.44 power of the body weight (sd = 0.14). The aquatic rate was consistently higher than the aerial rate of oxygen consumption, although at 30 °C the two rates were similar. Oxygen consumption increased with temperature up to 25 °C in all salinities; the lowest values were recorded at temperatures below 10 °C and at 30 °C in the most dilute medium. At all exposure temperatures, the oxygen consumption of Amphibola decreased regularly with salinity down to 0.1 ‰, and following exposure to concentrated sea water (41‰). Salinity had the least effect at 15 °C which was the acclimation temperature. In general, all of the temperature coefficients (Q₁₀ values) were low, < 1.65. However, Q₁₀ values above 2.8 were recorded at a salinity of 17.8‰ between 10 and 15 °C. Oxygen consumption of all size classes of Amphibola was more temperature dependent in air than in water and small individuals show a greater difference between their aerial and aquatic rates than larger snails. The rates of oxygen consumption in declining oxygen tensions were expressed as fractions of the rates in air saturated sea water at each experimental salinity-temperature combination. The quadratic coefficient B₂ becomes increasingly more negative with both decreasing salinity and temperatures up to 20 °C. At higher temperatures (25 and 30 °C) the response is reversed such that O₂ uptake in snails becomes increasingly independent of declining oxygen tensions at higher salinities. On exposure to a salinity of 4‰, Amphibola showed no systematic response to declining oxygen tension with respect to temperature. The ability of Amphibola to maintain its rate of oxygen consumption in a wide range of environmental conditions is discussed in relation to its potential for invading terrestrial habitats and its widespread distribution on New Zealand's intertidal mudflats.     

Rates of oxygen consumption by Thais (Nucella) Lapillus (L.)

Oxygen consumption by Thais varied seasonally with higher values in summer than in winter. This seasonal difference was due in part to the effects of temperature and in part to those of feeding. During feeding, rates of oxygen consumption were high, but declined in the period between meals. There was little evidence of acclimation of oxygen consumption to changes in temperature; low (winter) rates of consumption were more sensitive to increases in temperature than were high (summer) rates. A polynomial expression, including terms for temperature and ‘time since last meal’, was derived for the constant a′ in the allometric equation relating oxygen consumption (_o2) to dry body weight: _o2 = a′.W^0.511.     

Xylem sap flow as a major pathway for oxygen supply to the sapwood of birch (Betula pubescens Ehr.)

The role of xylem sap flow as an aqueous pathway for oxygen supply to the wood parenchyma of Betula pubescens saplings was investigated. Using micro‐optode sensors the oxygen status of the sapwood was quantified in relation to mass flow of xylem sap. Sap flow was gradually reduced by an increasing oxygen depletion in the root space. The effect of sap flow on radial O₂ transport between stem and atmosphere was assessed by a stoichiometrical approach between respiratory CO₂ production and O₂ consumption. Restriction of sap flow set in 36.5 h after the onset of O₂ depletion, and was complete after 71 h. Interruption of sap flow drastically increased the O₂ deficit in the sapwood to 70%. Sap flow contributed about 60% to the total oxygen supply to the sapwood. Diurnal O₂ flow rates varied between 3 and 6.3 nmol O₂ m^?2 leaf area (LA) s^?1 during night‐ and daytime, respectively. Maximum O₂ flow rates of 20 nmol O₂ m^?2 LA s^?1 were reached at highest sap flow rates of 5.7 mmol H₂O m^?2 LA s^?1. Sap flow not only affected the oxygen status of the sapwood but also had an effect on radial O₂ transport between stem and atmosphere.     

Respiratory responses of Oreochromis niloticus (Pisces, Cichlidae) to environmental hypoxia under different thermal conditions

The oxygen uptake ( V O₂), breathing frequency ( f _R), breath volume ( V _S.R), gill ventilation ( V G) and oxygen extraction (%) from the ventilatory current of four groups of Oreochromis niloticus during graded hypoxia were measured under the following acclimation temperatures: 20. 25. 30 and 35°C. The critical oxygen tensions ( P O₂), determined from V O₂ v. P O₂ of inspired water at each experimental temperature were, respectively. 19±1±3±1. 18±0±4±9, 29±7± 4±1 and 30±2± 0.6 mmHg. The f _R remained nearly constant during the reductions of O₂ at all the temperatures studied. V G increased discretely from normoxic levels until the P O₂ was reached, below which it assumed extremely high values (17-fold higher or more). The increases observed in V G resulted, at all the acclimation temperatures, in an elevation in V _S.R rather than in f _R. The extraction of O₂ decreased gradually from normoxia until the P O₂ was reached, below which an abrupt reduction of extraction was recorded, except at 35°C when fish showed a gradual reduction in extraction just below the tension of 80 mmHg.     

Effects of digenean trematodes and heterotrophic bacteria on mortality and burying capability of the common cockle Cerastoderma edule (L.)

Population dynamics of marine invertebrates is controlled by a variety of abiotic and biotic factors. Among these, some have received lesser attention from marine ecologists because of their ‘discrete’ nature. This is the case of parasitism and bacterial load. In the present study, we focused on the role that both digenean trematodes and heterotrophic aerobic bacteria might play in the mortality and burying behaviour of cockles, Cerastoderma edule.The bivalves were sampled monthly during 1 year from two sites in Arcachon Bay (French Atlantic coast). Mortality rates were assessed after transferring in the laboratory normally buried and unburied (i.e. found lying at the sediment surface at low tide) cockles. Their digenean and bacterial loads were determined for both positions (normally buried and unburied). Mortality rate was significantly higher for cockles found out of the sediment at low tide, suggesting that this abnormal position was a prelude to cockles' death. Comparison of digenean load of cockles showed no significant difference between buried and unburied bivalves. In contrast, bacterial load was significantly higher in unburied cockles than in normally buried animals. The effect of high concentration of a marine bacterial strain (Pseudomonas fluorescens) on cockles' burying behaviour and mortality was tested in the laboratory. Results showed that these bacteria could trigger the emergence of animals from the sediment but did not cause cockles' death. These field observations and laboratory experiments suggest that bacteria, rather than digenean trematodes, could play a role in the emergence of cockles and, hence, affect their survival in the wild.     

Wound-stimulated moulting and calcification responses in the adult cirripede Balanus balanoides (L.)

Earlier studies of hormonal action in adult barnacles have involved the injection of crustecdysone into the basal mantle space through a hole drilled in the shell. The drilling and injection result in increased moulting activity and in calcification within the mantle tissue around the site of injection. A preliminary investigation of the nature and causes of these responses has been carried out. The increase in moulting activity is distinct from that induced by injected crustecdysone and can be sustained over several months, the level of stimulation depending upon whether the adults are subjected to single or multiple injections. The associated calcification extends from the shell wall and develops as a layer around an area of wound tissue at the injection site. It appears that these moulting and calcification activities are healing responses to wounding and that the processes involved may be similar to those of other arthropods. Moulting in barnacles is apparently controlled by an ecdysone hormonal system and these results suggest that such a system may be implicated in the control of calcification.     

The control of respiration in wheat (Triticum aestivum L.) leaves

The aim of this work was to discover whether the respiration of wheat (Triticum aestivum L. cv. Huntsman) leaves, transferred to darkness after 7 h photosynthesis, showed an initial period of wasteful respiration. For young and old leaves, CO₂ production and O₂ uptake after 7 h photosynthesis were up to 56% higher than at the end of an 8-h night. The maximum catalytic activities of citrate synthase (EC 4.1.3.7), aconitase (EC 4.2.1.3), fumarase (EC 4.2.1.2) and cytochrome-c oxidase (EC 1.9.3.1) at the end of the day did not differ from those at the end of the night. Changes in the contents of glucose 6-phosphate, fructose-1,6-bisphosphate, dihydroxyacetone phosphate, and -ketoglutarate did not as a group parallel the changes in the rate of respiration. The detailed distribution of label from [U-¹⁴C] sucrose supplied to leaves in the dark was similar at the end of the day and the end of the night. No correlation was observed between the rates of leaf respiration and extension growth. It is argued that the higher rate of respiration at the beginning of the night cannot be attributed to wasteful respiration.Abbreviation RQ respiratory quotient We thank Dr H. Thomas and Professor C.J. Pollock, Institute for Grassland and Environmental Research, Plas Gogerddan, Aberystwyth, UK for their generous help in measuring leaf extension. R.H.A. thanks the Science and Engineering Research Council for a studentship.     

The reproductive response of the sea urchins Paracentrotus lividus (G.) and Psammechinus miliaris (L.) to a hyperproteinated macrophytic diet

The sea urchins Paracentrotus lividus and Psammechinus miliaris are submitted to the same environmental conditions in the Bay of Brest. The relationship between seasonal changes in food source quality and their gonad production was investigated in reproducing experimentally these conditions. In a first stage two macroalgae (Palmaria palmata and Laminaria digitata) were tested. P. miliaris showed a stronger preference for P. palmata and over a year-long experiment both urchins progressively preferred P. palmata. Seasonal variations in the chemical composition of P. palmaria were observed in the Bay of Brest: total carbohydrates were important and the relative maximum (about 50%) was reached between February and August; the lipid level was low and had a relative maximum of about 1% in June and August. Total protein in P. palmaria was high compared to other seaweeds: the maximum value (25%) was observed in June, which was probably due to the maintenance of nitrogen nutrient in the bay.In the second stage of the study, seasonal changes in biochemical components of ingestion and absorption of the two sea urchins were followed in the laboratory using a monospecific diet of P. palmaria. The patterns of total carbohydrates and lipid absorption were very similar for both sea urchin species. Carbohydrates were absorbed strongly and uniformly, year round. Lipid absorption mimicked the lipid nutrient pattern in the food source. Only changes in protein absorption varied slightly between the two urchin species. Protein absorption was maximal for both species in February and June, but the quantity of absorbed protein was significantly higher in P. miliaris than in P. lividus during February. This increase was concomitant with protein storage in the sea urchin gonads, which peaked in February for P. miliaris and in June for P. lividus. P. lividus had a higher gonad production efficiency, based on gonad yield. The comparison between in situ data and the experimental results suggests that an algal diet more nitrogenous than the in situ algal food source would benefit the herbivorous P. lividus, rather than the more omnivorous species P. miliaris. Although P. milaris has been described as a species with large gonad production potential, P. lividus appears to be a more suitable species for echiniculture conditions.     

Brown shrimp (Crangon crangon, L.) functional response to density of different sized juvenile bivalves Macoma balthica (L.)

Variability in infaunal bivalve abundance in the Wadden Sea is largely determined by recruitment variability. Post-settlement, but pre-recruitment bivalve mortality is high and related to the occurrence of their most abundant predator, the brown shrimp Crangon crangon. To investigate if the mortality patterns of newly settled bivalves can be explained by the foraging behavior of brown shrimp, we carried out experiments on shrimp functional response to three size classes of juveniles of the Baltic Tellin Macoma balthica. The functional response curves for all three prey sizes (0.62 mm, 0.73 mm, and 0.85 mm) were the hyperbolic Holling's type II. The attack rate was highest for the smallest prey size (a = 0.31, medium and large prey a = 0.22); the handling time was longest for the largest prey size (T_h = 29 s, small and medium prey T_h = 15 s). Thus, a large body size is advantageous for the bivalves over the whole density range. Knowledge of individual foraging behavior is needed to model predation mortality of bivalves. The consumption rates in the experiment were theoretically high enough to account for M. balthica mortality in the field.     

Photosynthetic and morphological responses of eelgrass (Zostera marina L.) to a gradient of light conditions

Zostera marina L. (eelgrass) from Great Bay Estuary, New Hampshire and Maine (USA), was transplanted in outdoor mesocosms and subjected to four light treatments (100, 58, 34 and 11% surface irradiance, SI) between May and September 2003 to investigate the relationship between light availability and the growth and survival of eelgrass. Evaluating eelgrass seedlings and adult mature plants demonstrated no differences in photosynthetic response after 22 days of acclimation. During at least the first 19 days of shading, maximum electron transport rate (ETR_max) rate of eelgrass did not differ significantly between light treatments. After 40 days, a significant reduction in ETR_max and minimum saturating light was observed in plants growing at 34% SI and below. Morphological responses exhibited a linear increasing trend with greater light. 34% SI exhibited drastic reductions (to less than 25% of control) in rhizome growth, shoot density, shoot production, number of nodes per plant and plant weight at the end of the study (81 days). Shoot to root ratio at 34% SI increased by > 50%. Plants shaded to 58% SI showed no significant difference from the control in plant parameters except an increased rate of rhizome elongation. Our results link the lower shoot densities with shading to the slow growth rate of horizontal rhizomes and a total lack of lateral expansion at 11% SI. ETR_max declined over time in plants at 11% SI resulting in 81% mortality, no lateral branching and no morphological development, indicating that the minimum light required for long-term eelgrass growth and survival is greater than the previously suggested 11% SI. We demonstrate that eelgrass plants at these latitudes can persist at light levels of 58% SI and above, and are light-limited at 34% SI and below.     

The incidence of autotomy in Carcinus maenas (L.)

A total of 1023 individuals of the common shore crab, Carcinus maenas (L.), were obtained from the environs of Whitby Harbour and Robin Hood's Bay. The bulk of the sample was taken from the sublittoral zone, but crabs from a sheltered shore were included. Each crab was measured, sexed and examined for the incidence of autotomized and regenerated limbs.There was a positive correlation between the incidence of autotomy and carapace width for crabs in both the sublittoral zone and the exposed intertidal zone. In sublittoral crabs the males suffered a higher incidence of autotomy than did the females and the incidence of cheliped autotomy was higher than for the walking limbs. The discrepancy between the loss of the chelipeds and walking limbs was reflected in a similar disparity between the occurrence of the regenerated chelipeds and walking limbs.Crabs on an exposed shore suffered a higher incidence of autotomy than did those from a sheltered shore. There were more crabs with cumulative limb losses than would be expected which suggests that once a crab enters into a state of autotomy it becomes increasingly vulnerable to the loss of another limb. Only 1 % of the population of sublittoral crabs would be expected to carry the loss of five limbs at once. No crabs were found with six or more limbs missing.     

The behavioural responses and tolerance of freshwater benthic cyclopoid copepods to hypoxia and anoxia

The response of four benthic cyclopoid copepods,Acanthocyclops viridis (Megacyclops viridis) (Jurine, 1820),Macrocyclops albidus (Jurine),Eucyclops agilis (Koch, Sars) (Eucyclops serrulatus) (Fischer, 1851) andParacyclops fimbriatus (Fischer), to hypoxia and anoxia was investigated. All of these species died within six hours when confronted by totally anoxic conditions, but all survived four days at oxygen saturation levels as low as 25%. Males succumbed to the effects of anoxia more rapidly than the larger females of each species, and larger species survived for shorter periods than smaller species. In artificially stratified columns, where the lower layer was anoxic, all four species displayed an upward migratory response towards oxygenated conditions. Where the artificial hypolimnion was hypoxic, however, the migratory response was not observed. The results suggest that some benthic copepods cope with seasonal anoxia in eutrophic stratified lakes by migration rather than the various physiological adaptations shown by planktonic and semi-planktonic species.     

Comparison of oxygen consumption in freshwater mussels (Unionidae) from different habitats during declining dissolved oxygen concentration

The rate of oxygen consumption (OC) of 9 species of freshwater mussels was measured under declining dissolved oxygen (DO) concentrations. The effects of temperature for some species also was investigated. The pattern of the OC vs. DO curve for each species was used in a hyperbolic model to compare abilities to regulate OC under low oxygen conditions. At 24.5 °C, Pyganodon grandis (from lakes), Amblema plicata and Quadrula pustulosa (from mud or sand in large rivers), Elliptio complanata (from pool areas in rivers), and Elliptio fisheriana and Elliptio lanceolata (from bank margins of rivers) were better able to maintain OC under low DO than were Villosa iris and Villosa constricta (which inhabit riffles) and Pleurobema cordatum (found in rivers with moderate flow). Villosa iris was especially sensitive to low oxygen conditions. The ability to maintain normal OC at low DO was improved considerably at 16.5 °C for V. iris, P. grandis and E. complanata. It is concluded that oxygen regulation ability appears to be related to the degree of hypoxia a species normally experiences in its habitat type, and it is enhanced at low temperature. The measurement of OC vs. DO may be a useful technique for estimating DO water quality criteria for endangered species because it is noninvasive.     

Seasonal respiratory responses to temperature and hypoxia in relation to burrowing depth in three intertidal bivalves

Respiratory responses to temperature and hypoxia in relation to burrowing depth were determined for winter- (W-C) and summer-conditioned (S-C) individuals of , , and . These bivalves occur sympatrically on sand-flats but display different sediment burrowing depths (. , <2 cm; . , 3–8 cm; . , 5–20 cm). A range depths over which daily temperature variation and O₂ concentration decline rapidly from surface values. Species thermal tolerance limits were found to decrease and to be more greatly temperature compensated with burrowing depth. Oxygen consumption rate (VO₂ increased with temperature to 25°C in . , but was, thereafter, regulated (Q10 1.0) up to 40°C while VO₂ increased with temperature in . and . until thermally streased at 25° to 30°C. The deposit feeding . does not acclimate VO₂ to temperature while . and . , both suspension feeders, show “reverse” acclimation [VO₂ (W-C) < VO₂ (S-C)] that conserves overwintering energy stores. The shallow burrowing . and deposit feeding . rarely experience hypoxia and are poor to non-regulators of VO₂ in reduced O₂ concentrations. In contrast, when winter-conditioned, . is a moderate regulator of VO₂, the degree of regulation increasing in S-C individuals which are exposed to higher levels of hypoxia.